The Relationship between Transpiration, Root Water Uptake, and Leaf Water Potential

The effect of changing the transpiration rate on leaf waterpotential and water balance has been examined to show if permeabilityof the plant (predominantly the roots) is constant or varieswith the transpiration rate. Measurements of leaf effectivethickness, water potential, transpiration, and uptake of waterby roots were made on sunflower, barley, and maize plants grownin solution culture and subjected to a range of atmosphericconditions and root treatments: cooling, low osmotic potential,and removal of part of the root system. Leaf water potential changed little under a wide range of atmosphericconditions and rates of water flux in the three species, sothat the root permeability to water increases as the rate oftranspiration, and therefore flow across the root surface, increases.Equality between uptake and loss of water and thereby maintenanceof constant leaf water potential is assisted by stomatal changes,which appear to be in response to conditions at or in the rootrather than a direct response to changes in bulk leaf waterpotential.     

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.     

断根处理对银杏树体水力特征及生长的影响

以8年生银杏为试材,采用不同程度的断根处理,测定处理后树体超声（ultrasound acoustic emission,UAE）信号、枝条栓塞程度（percentage loss of hydraulic conductance,PLC）、树干液流通量、气孔导度、蒸腾速率以及叶片水势等生理指标,分析断根对银杏树体水力特征及生长状况的影响.结果表明:断根处理后,在短时间内有大量UAE信号产生,且随断根程度的加重而增大,在6 h出现最大值,随后逐渐降低;枝条的PLC在起初的12 h迅速增加,24 h后增速趋于停滞;银杏树干液流通量、叶片气孔导度、蒸腾速率和水势明显降低,各处理间差异显著.经统计分析,累积UAE（cumulative number of UAE, cUAE）信号量与PLC呈现显著的正相关关系,与液流通量和叶片水势呈负相关关系.次年的单叶面积、新生枝条长度显著下降,下降程度随断根处理的加重而加剧.断根处理在短时间内增加了银杏管胞气穴数量和木质部栓塞的程度,影响到水分输送和叶片蒸腾,最终对叶片和枝条的生长产生负面影响.     

Identifying soybean lines differing in gas exchange sensitivity to humidity

The ratios of root length and root weight to leaf area differed within and between cultivars of soybean. Plants with low ratios of root length or weight to leaf area had leaf conductances and net photosynthetic rates more reduced by a given increase in the leaf to air water vapour pressure difference around a single leaf than plants with high ratios. Plant and root system conductances to water were estimated as transpiration rate per unit leaf area divided by the difference between substrate and leaf water potentials, and by the rate of water flow through pressurised root systems. These conductances were greater in plants with large, as compared with small, root systems per unit leaf area. Cultivar rankings in sensitivity of gas exchange to humidity were consistent in controlled environment chambers and in field tests.     

Hydraulic conductance as a factor limiting leaf expansion of phosphorus-deficient cotton plants

Suboptimal levels of phosphorus (P) strongly inhibited leaf expansion in young cotton (Gossypium hirsutum L.) plants during the daytime, but had little effect at night. The effect of P was primarily on cell expansion. Compared to plants grown on high P, plants grown on low P had lower leaf water potentials and transpiration rates, and greater diurnal fluctuations in leaf water potential. Hydraulic conductances of excised root systems and of intact transpiring plants were determined from curves relating water flow rate per unit root length to the pressure differential across the roots. Both techniques showed that low P significantly decreased root hydraulic conductance. The effects of P nutrition on hydraulic conductance preceded effects on leaf area. Differences in total root length, shoot dry weight, and root dry weight all occurred well after the onset of differences in leaf expansion. The data strongly indicate that low P limits leaf expansion by decreasing the hydraulic conductance of the root system.     

Responses of Net Photosynthesis and Conductance to Independent Changes in the Humidity Environments of the Upper and Lower Surfaces of Leaves of Sunflower and Soybean

A dual-surface leaf chamber was used to investigate the responsesof net photosynthesis and leaf conductance to independent changesin the humidity environments of the upper and lower surfacesof leaves of sunflower and soybean. In sunflower decreasingthe humidity around the upper leaf surface while maintainingthat of the lower surface constant and high reduced both thephotosynthetic rate and the conductance of the lower surface.These reductions could not be attributed to changes in bulkleaf water potential since the transpiration rate of the wholeleaf remained constant. Similarly, the reductions were not relatedto localized water deficits in the lower epidermis or lowermesophyll since the transpiration rate of the lower surfacewas reduced. Possible mechanisms whereby the gas exchange characteristicsof the lower leaf surface of sunflower respond to the humidityenvironment of the upper surface are discussed. In contrastto sunflower, the photosynthetic rate of the lower surface ofsoybean was insensitive to the humidity environment of the uppersurface. In leaves of sunflower grown under a moderate temperature anda medium light level, simultaneous decreases of humidity atboth leaf surfaces reduced the photosynthetic rate of the wholeleaf without affecting the substomatal partial pressure of CO₂.In contrast, with leaves developed under a cool temperatureand a high light level, both the photosynthetic rate and thesubstomatal partial pressure of CO₂ were reduced. Evidently,the occurrence in sunflower of the response pattern suggestinga non-stomatal inhibition of photosynthesis by low humiditydepends upon the environment during growth. The possibilitythat this non-stomatal inhibition may be an artifact due toan error in the assumption of water vapour saturation withinthe leaf airspace is considered. Key words: Vapour pressure deficit, photosynthesis, conductance, non-stomatal inhibition, Helianthus annuus, Glycine max     

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     

Long-term transpiration change with rainfall decline in a Mediterranean Quercus ilex forest

In the Mediterranean basin, precipitation is expected to decline as a consequence of climate change. The response of a Quercus ilex forest in southern France to such a decline in water availability was studied using a 4-year throughfall exclusion experiment. Seasonal courses of sap flow and leaf water potential were obtained from 2004 to 2007 and used to characterize tree water relations in a control and a dry treatment. The experiment reduced the average precipitation input to the soil by 29%, and resulted in a 23% reduction in annual transpiration. Soil water potential was significantly lower in the dry treatment only during summer drought, but transpiration was reduced all year round even during well-watered periods. Despite a tight stomatal control over transpiration, whole-tree hydraulic conductance was found to be lower in the trees growing in the driest conditions. This reduction in water transport capacity was observed jointly with a reduction in leaf transpiring area. Canopy leaf area decreased by 18% in the dry treatment as a consequence of the throughfall exclusion, which was found to validate the ecohydrological equilibrium theory.     

The Transpiration rate of Abies georgei and the Correlation with Enviromental Factors

The transpiration rate of Abies georgei (20 years) was determined at sample plots on subalpine in Northwest of Yunnan Province. The result of annual change shows that the rate is from lower to higher monthly. The transpiration rate reached maximum value of 195.88 mgH20·gdw-1·h-1 in July, then, it decreases gradually in a open-grown sample leaf of the current year. The phenomenon of “noon-rest” shows that the curve for day change of transpiration rate is the form of double peaks. Statistical analysis proves that the variance of transpiration rate is influenced mainly by the environmental factors, particularly by temperature, relative humidity and light intensity. The correlations between transpiration rate and temperature, light intensity are positive and that between transpiration rate and relative humidity is negative. The variances of the transpiration rate on the elevations are different. The transpiration rate decreases with the increase of elevation in the natural forest area. Change is less in whole day at high elevation and light wave too. The rate of transpiration in clear cut over area is higher than in the area under deep canopy of a subalpine natural forest, they are 186.15 and 123.97 respectively Lastly, the correlation between transpiration rate and environmental factors and plant setting are discussed in this paper.     

Influence of low air humidity and low root temperature on water uptake, growth and aquaporin expression in rice plants

The effects of low air humidity and low root temperature (LRT) on water uptake, growth and aquaporin gene expression were investigated in rice plants. The daily transpiration of the plants grown at low humidity was 1.5- to 2-fold higher than that at high humidity. LRT at 13°C reduced transpiration, and the extent was larger at lower humidity. LRT also reduced total dry matter production and leaf area expansion, and the extent was again larger at lower humidity. These observations suggest that the suppression of plant growth by LRT is associated with water stress due to decreased water uptake ability of the root. On the other hand, the net assimilation rate was not affected by low humidity and LRT, and water use efficiency was larger for LRT. We found that low humidity induced coordinated up-regulation of many PIP and TIP aquaporin genes in both the leaves and the roots. Expression levels of two root-specific aquaporin genes, OsPIP2;4 and OsPIP2;5, were increased significantly after 6 and 13 d of LRT exposure. Taken together, we discuss the possibility that aquaporins are part of an integrated response of this crop to low air humidity and LRT.     

Cyclic Variations in Plant Properties under Constant Environmental Conditions

Cyclic fluctuations in stomatal aperture, transpiration rate and leaf water potential under constant environmental conditions have been investigated in intact plants of cotton, pepper, and sunflower. Stomatal aperture and transpiration rate were least when leaf water potential was high and were greatest when leaf water potential was low. Lowest leaf water potential values lagged behind the occurrence of highest transpiration rates, and high overall resistance to water flow occurred in cycling plants. Both of these are considered essential for the occurrence of persistent cyclic behaviour. Hydropassive opening of stomates as the leaves wilted facilitated cycling in cotton and pepper, but not in sunflower, where hydropassive opening did not occur. The roots were identified as the site of the major resistance to water flow in the plant and further experiments directly showed the importance of this root resistance in initiating cycling by causing water stress in the leaves as the stomates opened. Root resistance varied diurnally, becoming increasingly important at night. Root resistance naturally rose to high levels in cotton. High levels were induced in pepper or sunflower by having the roots in deionized water for several days or by anoxia. Quantitative measurements of overall plant resistance were made from leaf water potential and transpiration rate data. The results are discussed and it suggested that plant resistance may indirectly be of importance in the movement of water from the plant to the air.     

Response of the water status of soybean to changes in soil water potentials controlled by the water pressure in microporous tubes

Water transport through a microporous tube-soil-plant system was investigated by measuring the response of soil and plant water status to step change reductions in the water pressure within the tubes. Soybeans were germinated and grown in a porous ceramic 'soil' at a porous tube water pressure of -0.5 kpa for 28 d. During this time, the soil matric potential was nearly in equilibrium with tube water pressure. Water pressure in the porous tubes was then reduced to either -1.0, -1.5 or -2.0 kPa. Sap flow rates, leaf conductance and soil, root and leaf water potentials were measured before and after this change. A reduction in porous tube water pressure from -0.5 to -1.0 or -1.5 kPa did not result in any significant change in soil or plant water status. A reduction in porous tube water pressure to -2.0 kPa resulted in significant reductions in sap flow, leaf conductance, and soil, root and leaf water potentials. Hydraulic conductance, calculated as the transpiration rate/delta psi between two points in the water transport pathway, was used to analyse water transport through the tube-soil-plant continuum. At porous tube water pressures of -0.5 to-1.5 kPa soil moisture was readily available and hydraulic conductance of the plant limited water transport. At -2.0 kPa, hydraulic conductance of the bulk soil was the dominant factor in water movement.     

Unusual stomatal behaviour on partial root excision in wheat seedlings

The excision of four out of five primary roots of wheat (Triticum durum Desf.) seedlings often leads to an enhanced rate of transpiration. Surprisingly this enhancement could be maintained for several hours after root excision and was particularly likely to occur at low irradiances or high atmospheric humidity. This long‐term enhancement could not be explained in terms of conventional hydropassive stomatal effects. Elevated rates of transpiration were associated with and possibly caused by increased cytokinin concentrations in shoots of plants with partially excised roots. The single root remaining after excision was able to maintain an adequate water uptake for the continued enhanced transpiration, after only a short transient reduction in leaf water content. The enhanced capacity for water uptake by the remaining root was confirmed by measuring the water flow from detached roots at negative hydrostatic pressure. Even without additional suction, flow from the reduced root system increased about 1.5 h after the start of treatment, suggesting an increase in membrane permeability for water. Although abscisic acid (ABA) concentrations in the roots increased after the root excision treatment, there was no evidence for any enhanced concentration in the xylem sap. The possible role that this accumulation of ABA in roots may have in the apparent increase in hydraulic conductivity after root excision is discussed.     

Effects of Extended Periods of Osmotic Stress on Water Relationships of Pepper

Pepper plants grown to uniform size in a controlled environment were subjected to an osmotic stress for periods of 1 to 10 days. Polyethylene glycol 400 was used as the osmotic agent. Leaf area of the plants, grown under uniform conditions, was proportional to the weight of the plants. This relationship was not altered by reduction in rate of growth due to a decrease in osmotic potential of the nutrient solution. The rate of transpiration of the pepper plants decreased as the osmotic potential of the nutrient solution was decreased. The reduction in rate of transpiration was most rapid when the osmotic potential was reduced from ?0.5 to ?7.5 bars. There was continued reduction in the rate of transpiration with change in potential to ?12.5 bar but this change was less than that at the higher potentials. The rate of transpiration remained at a reduced rate for as long as the plants were growing in the solution with low osmotic potential. Alternating the osmotic potential of the nutrient solution between ?0.5 and ?5.0 bar did not change the response to the ?5.0 tension. The reduction in rate of transpiration resulting from the lowering of the osmotic potential by addition of NaCl was similar to that produced by addition of polyethylene glycol. Water potential, osmotic potential, relative water content and stomatal movement were all in dynamic equilibrium with the water content of the leaves. The water content of the leaves was regulated by the supply and demand. In these investigations the demand remained constant. The supply was altered by decreasing the difference in water potential between leaf and substrate and by an increase in resistance to flow of water in the roots as a result of the decrease in osmotic potential of the nutrient solution.     

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.     

Plant and soil resistance to water flow in faba bean (Vicia faba L. major Harz.)

An experiment was conducted to determine soil and plant resistance to water flow in faba bean under field conditions during the growing season. During each sampling period transpiration flux and leaf water potential measured hourly were used with daily measurements of root and soil water potential to calculate total resistance using Ohm's law analogy. Plant growth, root density and soil water content distributions with depth were measured. Leaf area and root length per plant reached their maximum value during flowering and pod setting (0.31 m² and 2200 m, respectively), then decreasing until the end of the growing period. Root distribution decreased with depth ranging, on average, between 34.2% (in the 0–0.25 m soil layer) and 18.1% (in the 0.75–1.0 m soil layer). Mean root diameter was 0.6 mm but most of the roots were less than 0.7 mm in diameter. Changes in plant and soil water potentials reflected plant growth characteristics and climatic patterns. The overall relationship between the difference in water potential between soil and leaf and transpiration was linear, with the slope equal to average plant resistance (0.0165 MPa/(cm³ m^-1 h^-1 10^-3). Different regression parameters were obtained for the various measurement days. The water potential difference was inversely related to transpiration at high leaf stomatal resistance and at high values of VPD. Total resistance decreased with transpiration flux in a linear relationship (r=−0.68). Different slope values were obtained for the different measurement days. Estimated soil resistance was much lower than the observed total resistance to water flow. The change from vegetative growth to pod filling was accompanied by an increase in plant resistance. The experimental results support previous findings that resistance to water flow through plants is not constant but is influenced by plant age, growth stage and environmental conditions. A more complex model than Ohm's law analogy may be necessary for describing the dynamic flow system under field conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

Modeling soil water movement with water uptake by roots

Soil water movement with root water uptake is a key process for plant growth and transport of water and chemicals in the soil-plant system. In this study, a root water extraction model was developed to incorporate the effect of soil water deficit and plant root distributions on plant transpiration of annual crops. For several annual crops, normalized root density distribution functions were established to characterize the relative distributions of root density at different growth stages. The ratio of actual to potential cumulative transpiration was used to determine plant leaf area index under water stress from measurements of plant leaf area index at optimal soil water condition. The root water uptake model was implemented in a numerical model. The numerical model was applied to simulate soil water movement with root water uptake and simulation results were compared with field experimental data. The simulated soil matric potential, soil water content and cumulative evapotranspiration had reasonable agreement with the measured data. Potentially the numerical model implemented with the root water extraction model is a useful tool to study various problems related to flow transport with plant water uptake in variably saturated soils. This revised version was published online in June 2006 with corrections to the Cover Date.     

Modelling the impact of heterogeneous rootzone water distribution on the regulation of transpiration by hormone transport and/or hydraulic pressures

Aims

A simulation model to demonstrate that soil water potential can regulate transpiration, by influencing leaf water potential and/or inducing root production of chemical signals that are transported to the leaves.

Methods

Signalling impacts on the relationship between soil water potential and transpiration were simulated by coupling a 3D model for water flow in soil, into and through roots (Javaux et al. 2008) with a model for xylem transport of chemicals (produced as a function of local root water potential). Stomatal conductance was regulated by simulated leaf water potential (H) and/or foliar chemical signal concentrations (C; H?+?C). Split-root experiments were simulated by varying transpiration demands and irrigation placement.

Results

While regulation of stomatal conductance by chemical transport was unstable and oscillatory, simulated transpiration over time and root water uptake from the two soil compartments were similar for both H and H?+?C regulation. Increased stomatal sensitivity more strongly decreased transpiration, and decreased threshold root water potential (below which a chemical signal is produced) delayed transpiration reduction.

Conclusions

Although simulations with H?+?C regulation qualitatively reproduced transpiration of plants exposed to partial rootzone drying (PRD), long-term effects seemed negligible. Moreover, most transpiration responses to PRD could be explained by hydraulic signalling alone.     

Changes in hydraulic conductance of citrus trees following a reduction in wetted soil volume

Abstract The effcct of the transition from fully to partially wetted soil voluine on transpiration rate and hydraulic conductance of mature citrus trees was examined in a 23-year-old, coninicrcial, sprinklerirrigated, Shanio u t i orange orchard. I rriga t i on frequency was determined by the rate of water loss from the soil, a s measured by neutron probes. The hydraulic conductance of tlic tree was coniputed from the rclationship between sap flow i n the trunk and leaf water potential. The diurnal valucs of leaf water potential and sap flow shifted towards lower levels as tlie water stored in the root zone was depleted. In the fully wetted soil volume the tree hydraulic conductance remained constant throughout the irrigation period, from June to Novcniber. However, partial wetting of the soil volume (40%) caused a reduction in the hydraulic conductance of the tree. Tlie decreased hydraulic conductance is attributed to tlie permanent interruption of water transport in part of tlie root system. Tlie rcsults of tlie experiment suggest that despite tlie increase of irrigation frequency, partial wetting intensifies water stress in tlie trees.     

Intervention de l'oxygène atmosphériqoe sur l'absorption d'eau chez Helianthus annuus

The role of atmospheric oxygen on root water absorption in Helianthus annuus . The effect of atmospheric anoxia on root water absorption was studied. The experiments were carried out on intact young sunflowers in controlled temperature, light and gas environment; roots were kept in aerated nutrient solution at constant temperature. The evolution of root water absorption and transpiration rate was measured continuously. Before the experiment, the plant was preconditioned at a high transpiration rate by illumination or by CO₂ free air in darkness. Then the atmospheric oxygen was suppressed for 1 h, after which the normal conditions were restored.
In anoxia and darkness, the root water absorption cannot balance transpiration, so that an important water stress develops in the plant; the light compensates this effect through the photosynthetic oxygen. The supply of oxygen, in darkness or in light, immediately removes inhibition of stomatal closure and of root water absorption. Two mechanisms control water absorption by roots: the fast one occurs in the leaves and the slower one cannot develop without the root system.