Evaluation of the transpiration rate of lotus using the stem heat-balance method

To reveal the mechanism of transpiration by hydrophytes in the field, it is necessary to evaluate the transpiration rate without the effect of the evaporation from the water surface. In order to test the suitability for evaluating the transpiration rate of lotus (Nelumbo nucifera Gaertn.) leaves in the field, stem heat-balance method was applied and the obtained sap-flow rate was compared with the transpiration rate measured by weighing and with the overall canopy evapotranspiration rate by means of the eddy covariance technique. The transpiration rate estimated with the sap-flow measurements showed good agreement with that obtained from the weighing method. Lotus has many air canals in its petiole to carry oxygen-rich air to the rhizome and methane- and carbon dioxide-rich air back to the atmosphere, but there was little effect of the mass flow of air through these canals on the sap-flow rates. In the field observations, the canopy evapotranspiration rate (0.28 mm h⁻¹ at maximum) was nearly equal to the sum of the transpiration rate from all sunlit leaves (0.30 mm h⁻¹), and the contribution of the transpiration from shaded leaves and evaporation from the water surface was considered to be minor in the seasons when the leaves were fully developed. Evaluation of bulk leaf conductance revealed that the conductance in the leaf boundary layer of lotus could be low (ca. 0.23 mol m⁻² s⁻¹) because of its large leaf area. The low conductance in the leaf boundary layer would increase leaf temperature, which, in turn, would generate air circulation within the plant's ventilation system. Because there was a linear relationship between transpiration rate and the leaf-to-air vapor-pressure deficit, with no apparent maximum, high vapor-pressure deficits (3.4 kPa at maximum) did not appear to cause significant stomatal closure in lotus plants. The stomata of lotus leaves play a role as air inlets to carry oxygen-rich air to the rhizome, so their low sensitivity would help to increase air intake.     

Anatomy of the gas canal system of Nelumbo nucifera

Nelumbo nucifera (Gaertn.) grows by extending a creeping rhizome through anaerobic sediments. Nodes form at intervals along the rhizome, each producing a single leaf, and gas canals channel air from the leaves throughout the petioles and rhizomes. The gas flow pathway was mapped by casting the canals in growing shoots with silicone and by blowing air through complexes of rhizomes and petioles. Air from a leaf flows to a rhizome through one of two petiolar canal pairs, joining with the lowermost of three canal pairs in the rhizome through a chamber in the node. The lowermost canal pair links these nodal chambers along the length of a rhizome, allowing air from a node to flow both forward, toward a growing shoot, and backward, toward preceding leaves. These linked chambers also connect with the middle pair of canals on their proximal side, enabling flow to proceed backward along the rhizome to an adjacent node. A chamber in the next node then diverts the flow into the upper canal pair. This pair leads to a third node and chamber from which the air vents to the atmosphere through the second petiolar canal pair. Thus, pressurised air from one leaf must flow backward through two nodes before it returns to the atmosphere. Forward flow also ventilates a shoot's growing tip, with air from the lowermost canal pair entering a chamber in the developing node which, as described above, connects with the middle canal. This allows the air to reverse direction at the tip and enter the vent flow pathway.     

A two-way gas transport system in Nelumbo nucifera

Abstract The aquatic vascular plant Nelumbo nucifera Gaertn. is able to improve its oxygen supply to the submerged and buried organs by a thermo-osmotic gas transport. Investigations with tracer gas and oxygen measurements have shown that thermo-osmotic gas transport exists in N. nucifera when there is a temperature difference between the lacunar air of the leaves and the surrounding atmosphere. The gas transport was increased by up to 935% when a temperature difference of 2.9 ± 1.0 K was detected. Lacunar pressure of up to 166 ± 44 Pa was measured in both young and old leaves. In contrast to the flow-through ventilation system recently described for Nuphar lutea and Nymphoides peltata, a two-way flow in separate air canals in the petioles of both young and old Nelumbo leaves may carry oxygen-rich air down to the rhizome and excess air back to the atmosphere. Anatomical investigations have shown that, in Nelumbo, the two largest air canals of the petiole end directly under the mesh system of the centre plate. These large air canals are proposed to be predominant in the upward flow of air in sunlight. The other air canals of the petiole veer into the leaf blade well below the centre plate. The gas flow system through fresh leaves may carry as much as 10.3 ± 4.5 cm³ air per minute to the buried rhizome.     

Stomatal responses to changes in vapour pressure difference between leaf and air

We observed that stomata of Gossypium hirsutum, Glycine max and Xanthium strumarium respond to a change in vapour pressure difference between leaf and air at ambient partial pressures of CO₂ and below the CO₂ compensation point. Our report is at variance with a recent report (J. A. Bunce 1997, Plant, Cell and Environment 20, pp. 131–135) that stomatal sensitivity of leaves to a change in vapour pressure difference between leaf and air was eliminated when gas exchange measurements were made at near-zero carbon dioxide partial pressures (0–5 Pa).     

不同荷花品种出泥部分与根状茎生长性状和部分生理指标的比较及相关性分析

在盆栽条件下,对不同荷花(Nelumbo nucifera Gaertn.)品种〔藕莲品种'大紫红'('Dazihong')、花莲品种'中国红·上海'('Zhongguohongshanghai')和花藕兼用莲品种'逸仙莲'('Yixianlian')〕出泥部分(叶和花)和泥下部分(根状茎)的生长性状和部分生理指标进行了比较研究,并对3个荷花品种出泥部分与根状茎间的生长性状和部分生理指标进行了相关性分析.结果表明:'大紫红'在整个生长发育期不开花;'中国红·上海'开花时间相对较早(6月上旬),花重瓣且色泽鲜艳;'逸仙莲'开花时间相对较晚(7月上中旬),花单瓣呈白色.3个荷花品种中,'大紫红'立叶数最少,立叶面积最大,根状茎膨大明显,根状茎中可溶性糖和可溶性蛋白质含量最高;'中国红·上海'立叶数最多,立叶中叶绿素含量最高,根状茎节间长度最长,但根状茎膨大不明显;'逸仙莲'立叶数居中,叶柄最长,根状茎膨大明显,根状茎中可溶性蛋白质、可溶性糖、淀粉和维生素C含量介于其他2个荷花品种之间.相关性分析结果表明:荷花立叶面积与根状茎节间长度呈显著负相关,而与根状茎中可溶性糖、可溶性蛋白质及维生素C含量呈极显著或显著正相关;叶柄长度与根状茎直径和根状茎节间质量呈显著正相关;立叶中叶绿素含量与根状茎节间长度和根状茎中淀粉含量呈显著正相关,而与根状茎中可溶性糖含量呈显著负相关.研究结果显示:根据供试3个荷花品种的立叶面积、叶柄长度、立叶中叶绿素含量可间接判断根状茎的生长发育及营养品质状况,并可进一步应用于快速选育花藕兼用莲品种.     

LEAF SURFACE WETNESS AND GAS EXCHANGE IN THE POND LILY NUPHAR POLYSEPALUM (NYMPHAEACEAE)

As part of a continuing study of the effects of leaf surface wetness on gas exchange, the occurrence of leaf surface wetting by dewfall and associated effects on photosynthesis were evaluated for floating and aerial leaves of the pond lily Nuphar polysepalum Engelm. Because of nighttime radiation exchange with a cold sky, high humidity, and the presence of adaxial stomata, we predicted that pond lily leaves would be particularly susceptible to wetting events such as dewfall. A substantial reduction in net photosynthesis (up to 20%) occurred for leaves that were experimentally misted to simulate leaf wetting by dewfall. Aerial leaves remained below dewpoint temperatures for long periods on clear nights. However, floating leaves rarely approached dewpoint temperatures at night because minimum nighttime temperatures of leaves were up to 10 C warmer than air temperature. Thus, floating leaves of N. polysepalum did not experience dew formation primarily because of strong thermal coupling to a substrate (water) that was much warmer than air temperature at night. This coupling to a warmer substrate prevented a potentially strong inhibition of photosynthetic CO₂ exchange the following morning.     

Allelopathic effect of Nymphaea lotus L. on growth and yield of cultivated rice around Lake Manzala (Nile Delta)

Lotus infestation of ricefields is a major cause of crop failure and decrease of grain yield in the newly reclaimed cut-off areas around lake Manzala, Egypt. This study provides insights in the allelopathic effect of Nymphaea lotus L. on rice (Oryza sativa cavr. Giza-177). Lotus rhizome extracts were inhibitory to seed germination and seedling growth of rice. The degree of inhibition was dependent on extract type and concentration. Ethanol and water extracts were more inhibitory than chloroform extracts. The phenolic fraction of ethanol extracts showed the highest inhibitory effects. In a target (rice)-neighbour (lotus) pot experiment, rice dry mass and relative growth rate were dependent on its age and on lotus rhizome density, with decreased growth at increased lotus density. Field data on infested and non-infested ricefields demonstrated a decreased leaf area index and yield in infested fields. Identification of the potential allelochemical compounds by gas chromatography/mass spectrometry revealed the presence of allelopathic phenolics in lotus rhizomes.     

Knudsen-transitional flow and gas pressurization in leaves of nelumbo

Pressures in gas spaces of leaves of the lotus Nelumbo are higher than ambient pressure. The pressurization capacity of leaves was studied as a function of leaf temperature, and the composition of air entering evacuated leaves was used to calibrate the pore sizes which determine flow in these leaves. The adaxial side of the leaf of Nelumbo has two distinct regions in terms of gas exchange characteristics. There is a region of relatively high mean pore diameter in the center of the leaf opposite the point of petiole insertion. Gas exchange between the remainder of the leaf (>99% by area) and the atmosphere is restricted by “pores” with an effective mean diameter less than 0.03 micrometer. As a result, a flowthrough ventilation operates within each leaf. Air enters the leaf across the expanse of the lamina, and escapes back to the atmosphere through the highly porous region at the center of the lamina.     

Effect of leaf water deficit on stomatal and nonstomatal regulation of net carbon dioxide assimilation

The effect of leaf water deficit on net CO₂ assimilation was studied under two conditions: in one, the stomata were allowed to contribute to the regulation of CO₂ assimilation; in the other, air was forced through the leaf at a constant rate to overcome the effects of change in stomatal resistance accompanying changes in leaf water deficit. When the stomata were allowed to regulate the gaseous diffusive resistance of the leaf, CO₂ assimilation decreased with increasing leaf water deficit. However, when air was forced through the leaf, the rate of assimilation was not inhibited by increasing leaf water deficit. The results indicate that the inhibition of net CO₂ assimilation with increasing leaf water deficit is a consequence of an increase in the diffusive resistance to gas exchange and not of a change in apparent mesophyll resistance.     

Stomatal responses to changes in humidity in plants growing in the desert

Summary The stomata of plants growing in the Negev Desert, namely the stomata of the mesomorphic leaves of Prunus armeniaca, the xeromorphic stems of Hammada scoparia, and the succulent leaves of Zygophyllum dumosum, respond to changes in air humidity. Under dry air conditions diffusion resistance increases. Under moist air conditions diffusion resistance decreases. When the stomata close at low air humidity the water content of the apricot leaves increases. The stomata open at high air humidity in spite of a decrease in leaf water content. This excludes a reaction via the water potential in the leaf tissue and proves that the stomatal aperture has a direct response to the evaporative conditions in the atmosphere. In all species the response to air humidity is maintained over a period of many hours also when the soil is considerably dry. The response is higher in plants with poor water supply then in well watered plants. Thus for field conditions and for morphologically different types of photosynthesizing organs the results confirm former experiments carried out with isolated epidermal strips.     

Effects of oil on internal gas transport, radial oxygen loss, gas films and bud growth in Phragmites australis

Background and Aims

Oil pollution of wetlands is a world-wide problem but, to date, research has concentrated on its influences on salt marsh rather than freshwater plant communities. The effects of water-borne light oils (liquid paraffin and diesel) were investigated on the fresh/brackish wetland species Phragmites australis in terms of routes of oil infiltration, internal gas transport, radial O₂ loss (ROL), underwater gas films and bud growth.

Methods

Pressure flow resistances of pith cavities of nodes and aerenchyma of leaf sheaths, with or without previous exposure to oil, were recorded from flow rates under applied pressure. Convective flows were measured from living excised culms with oiled and non-oiled nodes and leaf sheaths. The effect of oil around culm basal nodes on ROL from rhizome and root apices was measured polarographically. Surface gas films on submerged shoots with and without oil treatment were recorded photographically. Growth and emergence of buds through water with and without an oil film were measured.

Key Results

Internodes are virtually impermeable, but nodes of senesced and living culms are permeable to oils which can block pith cavity diaphragms, preventing flows at applied pressures of 1 kPa, natural convective transport to the rhizome, and greatly decreasing ROL to phyllospheres and rhizospheres. Oil infiltrating or covering living leaf sheaths prevents humidity-induced convection. Oil displaces surface gas films from laminae and leaf sheaths. Buds emerge only a few centimetres through oil and die.

Conclusions

Oil infiltrates the gas space system via nodal and leaf sheath stomata, reducing O₂ diffusion and convective flows into the rhizome system and decreasing oxygenation of phyllospheres and rhizospheres; underwater gas exchange via gas films will be impeded. Plants can be weakened by oil-induced failure of emerging buds. Plants will be most at risk during the growing season.Key words: Phragmites australis, oil pollution, convective flow, pressure flow resistance, phyllosphere oxygenation, rhizosphere oxygenation, underwater gas films, bud emergence, stomata, pith cavity diaphragms, leaf sheath aerenchyma, rhizome aeration     

莲的生态解剖学研究

莲叶有三种形式:沉水叶无气孔;浮水叶气孔仅分布于腹面;挺水叶形大、气孔数多。器官中气室和气道发达。排水器位于叶腹面的中央,除常敞开着外,与气孔相似,水分可从此排出。莲具有星散的维管束、无形成层,只在根中有导管,胚根简化,不定根发达。花早期发育在水中,后由柄生长延伸出水面。果实中具有三个功能不同的孔。果皮上的表皮层及栅状细胞层发达。     

A field portable system for the measurement of gas exchange of leaves under natural and controlled conditions: examples with field-grown Eucalyptus pauciflora Sieb. ex Spreng. ssp. pauciflora, E. behriana F. Muell. and Pinus radiata R. Don.

Abstract A field portable system is described which measures the response of gas exchange of one leaf to changes in environmental parameters under controlled conditions, and which simultaneously measures the gas exchange of another leaf as the climatic parameters vary naturally. The system consists of two independently operating cuvettes. It enables detailed studies of photosynthesis and stomata/transpiration of leaves attached to the plant in their natural position. It provides control of temperature, humidity, CO₂ and oxygen concentration (or, alternatively, of other gases) as well as of light. Infrared gas analyzers for CO₂ and H₂O are used which allow similar time constants for the measurement of the two gases. Examples of a diurnal course of gas exchange of a leaf in its natural exposition and of experiments with steady-state responses of gas exchange are presented. In Eucalyptus pauciflora Sieb. ex Spreng. ssp. pauciflora, a set of response curves of CO, assimilation (A) to CO₂, as measured at various leaf temperatures and light levels, shows carboxylation efficiency to be light saturated at the lower photon irradiances the lower the leaf temperature is. Carboxylation efficiency is maximal at 25°C. At ambient CO, partial pressure stomata open in a way that CO₂ assimilation occurs at a rate found within the curvature region of the CO₂ response function of A. The light-independent CO² compensation point as a function of temperature is presented. Applying a combined heat/low humidity pulse (15 or 60 min) on leaves of Eucalyptus behriana F. Muell. or Pinus radiata R. Don, respectively, leads to a lower level of intercellular carbon dioxide partial pressure (C_i) during the decline in A and leaf conductance to water vapour (g). A lower C_i level is maintained during recovery of A and g, A almost reaching the pre-pulse level but not g. The existence of an after-effect indicates that the response to the combined high temperature/low humidity pulse is a multi-step process.     

Changes in stomatal frequency, stomatal conductance and cuticle thickness during leaf expansion in the broad-leaved evergreen species, Eucalyptus regnans

Aspects of leaf anatomical and physiological development were investigated in the broad-leaved evergreen species, Eucalyptus regnans F.Muell. Newly emergent leaves were tagged in the field and measured for stomatal conductance while a subset was collected every 14 days for the measurement of stomata and cuticle over a 113-day period. Cuticle thickness increased during leaf expansion, the increase following a sigmoid curve. Stomatal frequency (no. mm⁻²) decreased from 56 to 113 days after leaf emergence. The frequency of both immature and intermediate developmental stages of stomata also decreased over this time, but the total number of stomata per leaf remained relatively constant. Stomatal conductance (g _s) of young expanding leaves increased during expansion, and was significantly linearly correlated with stomatal frequency (excluding immature stomata), and with cuticle thickness. The progressive increase in g _s in young developing leaves was contrary to the observed changes in structural characteristics (increased cuticle thickness and decreased stomatal frequency). This increase in g _s with development may be related to the progressive increase in number of mature stomata with larger apertures and, therefore, a higher total pore area in fully expanded leaves.     

Juvenile Rhus glabra leaves have higher temperatures and lower gas exchange rates than mature leaves when compared in the field during periods of high irradiance

We sought to test the hypothesis that stomatal development determines the timing of gas exchange competency, which then influences leaf temperature through transpirationally driven leaf cooling. To test this idea, daily patterns of gas exchange and leaflet temperature were obtained from leaves of two distinctively different developmental stages of smooth sumac (Rhus glabra) grown in its native habitat. Juvenile and mature leaves were also sampled for ultrastructural studies of stomatal development. When plants were sampled in May-June, the hypothesis was supported: juvenile leaflets were (for part of the day) from 1.4 to 6.0 degrees C warmer than mature leaflets and as much as 2.0 degrees C above ambient air temperature with lower stomatal conductance and photosynthetic rates than mature leaflets. When measurements were taken from July to October, no significant differences were observed, although mature leaflet gas exchange rates declined to the levels of the juvenile leaves. The gas exchange data were supported by the observations that juvenile leaves had approximately half the number of functional stomata on a leaf surface area basis as did mature leaves. It was concluded that leaf temperature and stage of leaf development in sumac are strongly linked with the higher surface temperatures observed in juvenile leaflets in the early spring possibly being involved in promoting photosynthesis and leaf expansion when air temperatures are cooler.     

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.     

Light-flecks cause non-uniform stomatal opening – studies with special emphasis on Fagus sylvatica L.

The appearance of stomatal patchiness in response to rapid (seconds) changes in light has been studied in European beech, Fagus sylvatica L., and, by comparison, in a further 17 different woody species from the understorey of a European beech forest, using a simple water infiltration method. Water infiltrated areoles indicate open stomata. Since infiltration changes optical characteristics of a leaf section it can be analysed by photography, computer-aided image analysis and by weighing. For F. sylvatica clear differences were found between infiltration of cotyledons (no patchy pattern) and any other leaf type. Despite identical cultivation, leaves of the same type and age from different individual plants responded differently to application of 30 s of light after darkness. In contrast, the patchiness patterns were very similar for leaves of the same type originating from the same plant. Infiltration patterns after a light-fleck, observed on different leaves as a series of momentary clusters, probably indicate waves of opening stomata moving across the leaf blade. During and after a 30 s light-fleck infiltration increased and it continued to increase in the dark up to 10 min, indicating increasing stomatal opening over that period. In general, shade leaves became more infiltrated (by weight) than half-shade or sun leaves, due to larger intercellular air spaces. All species, without exception, showed patchy infiltration and, thus, non-uniform stomatal opening. Measuring leaf gas exchange (as ”quasi-steady states” using a fast responding system) during photosynthetic induction resulted in very similar CO₂ responses of net photosynthesis (A/c _i) as in the true steady state, proving that, in shade and half-shade leaves, the presence of stomatal patchiness does not necessarily affect the calculation of intercellular CO₂ concentrations. Causes and consequences of stomatal patchiness are discussed. Received: 18 November 1998 / Accepted: 1 July 1999     

ABSCISIC ACID INDUCES LAND FORM CHARACTERISTICS IN MARSILEA QUADRIFOLIA L.

In aseptically cultured rhizome segments of the aquatic fern Marsilea quadrifolia, the addition of abscisic acid (ABA) to the liquid medium induced development of morphological characteristics distinctive to the land form of the plant. The land-form characteristics induced by ABA included a change in leaf morphology, an increase in the surface area of leaflets, differentiation of stomata and trichomes, elongation of petioles and roots, development of lateral roots, shortening of rhizome, and reduction in the number of leaves and roots formed on each plant.     

Does lateral gas diffusion in leaves matter?

Photosynthesis depends on the diffusion of gaseous CO(2) inside the leaf spaces from the stomatal entry point to the mesophyll cell walls. Although most research considers only the vertical diffusion from stomata on upper and/or leaf lower surfaces, some of the gas will diffuse in the lateral (paradermal) direction. The importance of lateral CO(2) diffusion is reviewed, and the anatomical characteristics of leaves, including the variation of air space volume between species and conditions are discussed. The contribution of the air space conductance to the limitation of photosynthesis by the overall CO(2) diffusion pathway is usually ignored. However, the need to consider three-dimensional diffusion at the small scale of a few stomata is emphasized because stomata are discrete, and separated by 20-300 microm. At the large scale of 100s of micrometres, there may be barriers to CO(2) caused by the vascular tissue, particularly if there are bundle sheath extensions. The possible extent and controls on CO(2) lateral and vertical diffusion in different species and conditions are illustrated using chlorophyll a fluorescence imaging techniques. It is clear that there is a range of effective lateral permeabilities depending on the particular vascular patterns and cell arrangements, and that species cannot be simply divided into homobaric and heterobaric anatomies. Lateral diffusion in more permeable leaves can be sufficient to affect measurements of leaf gas exchange, particularly when fluxes are low, although its contribution to leaf photosynthesis in natural conditions needs clarification.     

Vein cavitation and stomatal behaviour of sunflower (Helianthus annuus) leaves under water limitation

The impact of leaf vein cavitation and embolism on stomatal response and leaf hydraulic conductance was studied in potted plants of sunflower subjected to water limitation. Plant dehydration was achieved either by cutting well‐watered plants near their base and leaving them dehydrating in air or by depriving intact plants of irrigation. The vein cavitation threshold (Ψ_CAV) was estimated in terms of ultrasound acoustic emissions (UAE) from the leaf blade versus leaf water potential (Ψ_L). This was found to be the same (Ψ_CAV ≈ ?0.6 MPa) for leaves of both cut and intact plants where stomata began to close in coincidence with starting vein cavitation. Vein embolism was detected by infiltrating leaves at different Ψ_L with 0.7 mM fluorescein and measuring the percentage fluorescent area as percentage of total leaf surface area. A distinct loss of vein functionality (up to 50%) was found to occur in leaves at progressively decreasing Ψ_L, starting when leaves reached Ψ_CAV. A linear positive relationship with high statistical significance was found to exist between g_L and percentage leaf fluorescent area, thus indicating that stomata were sensitive to vein embolism. The hydraulic conductance (K_L) of the leaf was affected by leaf dehydration less than expected (K_L decreased by about 20% between near full turgor and Ψ_L = ?1.3 MPa). When the extravascular leaf compartment was excluded either by killing cells by immersing leaves in 70% ethanol or by cutting the main leaf venous system through to allow flow to bypass it, K_L turned out to increase 5.5 times, thus suggesting that the high dominance of the hydraulic resistance of the extravascular leaf compartment over the total leaf resistance might buffer or mask possibly large local changes in K_L inducing stomatal closure.