Some physiological and growth responses of kiwi fruit (Actinidia chinensis) to flooding

The effects of long-term flooding on the growth of six-month-old Actinidia chinensis Planch cv. Abbot plants and some effects on stomatal behaviour and leaf water relations were examined under controlled conditions for 28 days. Flooding caused stomatal closure and decreases in transpiration rate, xylem water potential, osmotic potential and turgor potential. Flooding also caused inhibition of the dry weight increase of leaves plus stems and of roots, chlorosis and necrosis of leaves, production of hypertrophied lenticels and the appearance of a small number of adventitious roots on the submerged portions of the stems. Rapid and partial stomatal closure by flooding may not only be due to the passive mechanical response which follows leaf dehydration, since flooded plants showed an increase in xylem water potential and osmotic potential during the first days of the experiment. The marked intolerance of Actinidia chinensis to flooding has been a serious barrier to its culture in poorly drained soils, hence careful irrigation management is required.     

Are Roots a Source of Abscisic Acid for the Shoots of Flooded Pea Plants?

Flooding the soil for 2–5 d decreased stomatal conductancesof pea plants (Pisum sativum L., cv. Sprite) with six or sevenleaves. This coincided with slower transpiration, increasedleaf water potentials and increased concentrations of abscisicacid (ABA) in the leaves. No increase in ABA was found in theterminal 20 mm of roots of flooded plants over the same timeperiod. Small stomatal conductances associated with increases in foliarABA were also found in plants grown in nutrient solution whenaeration was halted, causing the equilibrium partial pressuresof dissolved oxygen to fall below 05 It Pa. No increase in ABAconcentration in young secondary roots of the non-aerated plantswas detected after 24, 48 or 72 h, even when the shoot, thepresumed site of deposition for any ABA from the roots, wasremoved 5–6 h before analysis. Similarly, ABA concentrations in roots were not increased whenthe nutrient solution was de-oxygenated by continuous purgingwith nitrogen gas. The abscisic acid concentration in leaf epidermis,the tissue most likely to be the recipient of any ABA movingin the transpiration stream from oxygen-deficient roots, waslower than in the remaining parts of the leaf when examinedin the mutant Argenteum which possesses easily removable epidermallayers. It is concluded that the leaves of plants subjectedto flooding of the soil or oxygen shortage in the root environmentare not enriched substantially with ABA from the roots. A moreprobable source of this growth regulator is the leaf itself. Key words: Pisum sativum, flooding, roots, hormones, aeration stress, abscisic acid, Argenteum mutant     

Effect of flooding and temperature regime on growth and stomatal resistance ofBetula platyphylla var.japonica seedlings

Summary Both flooding and low temperature reduced height and stem diameter growth; leaf initiation; growth of leaves, stems, and roots; and lowered root-shoot ratios of 112-dayoldBetula platyphylla var.japonica seedlings. Flooding also induced leaf scorching and abscission. Growth was reduced much more by flooding than by low temperature. Interactive effects of flooding and temperature were shown on height growth, leaf initiation and expansion, and dry weight increment of leaves, stems and roots. The amount of growth reduction by flooding and low temperature was greater when based on analysis of dry weight increment of leaves, stems, and roots, than on their relative growth rates. The greater reduction of growth by flooding than by low temperature was associated with fewer and smaller leaves, more leaf injury, more stomatal closure, and greater decay of roots in flooded plants. Flooding and low temperature appeared to reduce growth by somewhat different physiological mechanisms. Research supported by the College of Agricultural and Life Sciences, University of Wisconsin, Madison, WI, USA and by Yamagata University, Tsuruoka, Japan. McIntyre-Stennis Project 2599.     

Positive and Negative Messages from Roots Induce Foliar Desiccation and Stomatal Closure in Flooded Pea Plants

Flooding the soil for 5–7 d caused partial desiccationin leaves of pea plants (Pisum sativum. L. cv. ‘Sprite’).The injury was associated with anaerobiosis in the soil, a largeincrease in the permeability of leaf tissue to electrolytesand other substances, a low leaf water content and an increasedwater saturation deficit (WSD). Desiccating leaves also lackedthe capacity to rehydrate in humid atmospheres, a disabilityexpressed as a water resaturation deficit (WRSD). This irreversibleinjury was preceded during the first 4–5 d of floodingby closure of stomata within 24 h, decreased transpiration,an unusually large leaf water content and small WSD. Leaf waterpotentials were higher than those in well-drained controls.Also, there was no appreciable WRSD. Leaflets detached fromflooded plants during this early phase retained their watermore effectively than those from controls when left exposedto the atmosphere for 5 min. Stomatal closure and the associated increase in leaf hydrationcould be simulated by excising leaves and incubating them withtheir petioles in open vials of water. Thus, such changes inflooded plants possibly represented a response to a deficiencyin the supply of substances that would usually be transportedfrom roots to leaves in healthy plants (negative message). Ionleakage and the associated loss of leaf hydration that occurswhen flooding is extended for more than 5 d could not be simulatedby isolating the leaves from the roots. Appearance of this symptomdepended on leaves remaining attached to flooded root systems,implying that the damage is caused by injurious substances passingupwards (positive message). Both ethylene and ethanol have beeneliminated as likely causes, but flooding increased phosphorusin the leaves to concentrations that may be toxic. Key words: Pisum sativum, Flooding, Foliar desiccation, Stomata, Ethylene     

Can early wilting of old leaves account for much of the ABA accumulation in flooded pea plants?

When pea plants (Pisum sativum L.) were subjected to flooding,abscisic acid (ABA) content in shoots and roots increased upto 8-fold in the following days and stomatal conductance significantlydecreased. Although young leaves of flooded plants had a slightlyhigher water potential than those of the unflooded plants, oldleaves had lower water potential and lost turgor at the timewhen a substantial ABA increase was detected. In plants wherethe old leaves were clipped off, flooding did not cause anyABA increase during 7 d of the experimental period, except underconditions of higher transpiration demand, when the increasein ABA content was both delayed and small in scale (only I-fold).When intact plants were flooded and ABA was assayed separatelyin both old and young leaves, the ABA increase in old leavespreceded that in young leaves. Evidence here suggests that theflooding-induced ABA increase mainly results from the wiltingof old leaves. This suggests that young leaves may be protectedfrom wilting by ABA originating in old leaves under unfavourableenvironmental conditions. Key words: Waterlogging, soil flooding, ABA, leaf water relations, pea, Pisum sativum     

Influence of flooding with diluted sea water on morphological and physiological characteristics in hexaploid triticale

The influence of flooding with tap water, and with 10 and 20% sea water on the growth, yield and physiological characteristics in hexaploid triticale (X Triticosecale Wittmack) were analysed. The plants were grown in a glasshouse and flooded at the booting stage for 19 d. Flooding seriously affected the growth, yield and physiological activities of the plants. However, the growth and yield were greater in plants flooded with diluted sea water than in plants flooded with tap water. The amount of newly developed roots, photosynthetic rate and accumulation of mineral nutrients in the leaves was greater in sea water-flooded plants as compared to tap water-flooded plants. Oxygen concentrations in the diluted sea water were lower than in the tap water 6 d from the beginning of flooding. Leaf water potentials in the plants flooded with diluted sea water were lower than in the plants flooded with tap water until 7 d from the beginning of flooding when the position reversed. Transpiration was slower and stomatal resistance was greater in flooded plants, and were similar irrespective of water quality.     

Senescence-Associated Changes during Long-day-induced Leaf Senescence and the Nature of the Graft-transmissible Senescence Substance in Kleinia articulata

Schwabe, W. W. and Kulkarni, V. J. 1987. Senescence-associatedchanges during long-day-induced leaf senescence and the natureof the graft-transmissible senescence substance in Kleinia articulata.— J. exp. Bot. 38: 1741–1755. The long-day-induced senescence in Kleinia articulata leaveswas characterized by a loss in fresh and dry weight, in therate of leaf expansion and progressive loss of chlorophyll inthe detached rooted leaves. Ultrastructural examination of mesophyllcells of leaves from plants grown in continuous light showedthat osmiophilic globules accumulating in the chloroplasts werethe first visible sign of senescence in the organdies. Thesefirst signs of senescence could be detected in very young leavesof plants in continuous light, even before the leaves had expanded.Attempts were made to study the cause of this photoperiodicsenescence which, from previous work, appeared to involve agraft-transmissible substance. Leaves in continuous light showed reduced stomatal opening andextracts from them had very much higher activity in the Commelinastomatal closure assay (ABA-like activity ?) compared with non-senescingleaves grown in short days (8 h). However, even if all the activitywere due to ABA, this on its own does not appear to be the senescencesubstance because a much longer exposure to continuous lightwas required to induce irreversible senescence than to reachmaximum stomatal closure promoting activity in the bioassay.Moreover, severe water stress (high ABA?) did not lead to senescenceunless combined with continuous light or ethylene treatment.It is postulated that while ABA may play an important role inKleinia leaf senescence its lethal effect may not be realizedunless ethylene-induced membrane changes may synergisticallyassist. Key words: Leaf senescence, ABA, Daylength, stomatal movement, Kleinia     

Patterns of Stomatal Behaviour, Transpiration, and CO2 Exchange in Pea Following Infection by Powdery Mildew (Erysiphe pisi)

A comparison was made of stomatal behaviour, and related phenomena,between leaves of garden pea (Pisum sativum cv. Feltham First)inoculated with powdery mildew fungus (Erysiphe pisi) and uninfectedleaves on healthy plants. Twenty four hours after inoculation,stomata opened more widely in the light in infected leaves thanin healthy leaves. Thereafter, stomatal opening was progressivelyreduced by infection and stomata failed to close completelyin the dark until, 7 d after inoculation, all movements ceasedand stomata remained partly open. Transpiration in the lightfollowed closely the pattem of stomatal opening and, after anearly increase compared with healthy controls, was progressivelyreduced by infection. Evidence is presented that transpirationfrom the fungus was less than the reduction in transpiraationfrom the leaf which was caused when development of the myceliumincreased the boundary layer resistance of the leaf. Seven daysafter inoculation, transpiration in the dark was greater frominfected leaves than from healthy leaves because of partly openstomata in the dark. Net photosynthesis in infected leaves was reduced within 24h of inoculation to a level below that found in healthy leavesand thereafter it declined progressively. The initial reductionwas due to a transient increase in photorespiration, for whenthe glycolate pathway was inhibited by a 2% O₂ concentrationthere was no difference between the (gross) photosynthetic ratesof healthy and infected leaves. Changes in photorespirationrate were confirmed from the interpretation of the CO₂ burston darkening. Reduced stomatal opening was a contributory causeof the reduction in net photosynthesis in the later stages ofinfection. Since the rate of gross photosynthesis, but not therate of photorespiration, of infected plants fell below thatof healthy plants, and infected plants had a higher rate ofrelease of CO₂ in the dark than healthy plants from the thirdday after inoculation onwards, infected plants consume an increasinglygreater proportion of their photosynthate in respiratory processesthan do healthy plants. The CO₂ compensation point of infectedplants increased at every time of sampling after inoculation.     

A Physiological Approach to the Analysis of Crop Growth Data: 3. The Effects of Repeated Short Term Soil Water Deficits on the Growth of Spaced Plants of the Legume, Macroptilium atropurpureum cv. Siratro

A physiological analysis of crop growth, reported in the previouspapers in this series, is extended to deal with the growth ofspaced plants, and applied to the growth data for M. atropurpureumsubject to several successive cycles of water deficit. Differences in growth between stressed and unstressed plantscan be quantitatively attributed to a reduction in the photosyntheticactivity of the leaves of stressed plants induced by increasingtheir leaf water deficits, and to an increase in the rate ofleaf abscission by the stressed plants. Macroptilium atropurpureum, growth analysis, water stress, photosynthesis, leaf abscission, stomatal conductance     

Photosynthesis affects following night leaf conductance in Vicia faba

Night-time stomatal opening in C₃ plants may result in significant water loss when no carbon gain is possible. The objective of this study was to determine if endogenous patterns of night-time stomatal opening, as reflected in leaf conductance, in Vicia faba are affected by photosynthetic conditions the previous day. Reducing photosynthesis with low light or low CO₂ resulted in reduced night-time stomatal opening the following night, irrespective of the effects on daytime stomatal conductance. Likewise, increasing photosynthesis with enriched CO₂ levels resulted in increased night-time stomatal opening the following night. Reduced night-time stomatal opening was not the result of an inability to regulate stomatal aperture as leaves with reduced night-time stomatal opening were capable of greater night-time opening when exposed to low CO₂. After acclimating plants to long or short days, it was found that night-time leaf conductance was greater in plants acclimated to short days, and associated with greater leaf starch and nitrate accumulation, both of which may affect night-time guard cell osmotic potential. Direct measurement of guard cell contents during endogenous night-time stomatal opening will help identify the mechanism of the effect of daytime photosynthesis on subsequent night-time stomatal regulation.     

Involvement of plant growth substances in the alteration of leaf gas exchange of flooded tomato plants

Ethylene, abscisic acid, and cytokinins were tested for their ability to either induce or prevent the changes which occur in gas exchange characteristics of tomato (Lycopersicon esculentum Mill. cv. Rheinlands Ruhm) leaves during short-term soil flooding. Ethylene, which increases in the shoots of flooded plants, had no effect on stomatal conductance or photosynthetic capacity of drained plants. Abscisic acid, which also accumulates in the shoots of flooded plants, could reproduce the stomatal behavior of flooded plants when sprayed on the leaves of drained plants. However, photosynthetic capacity of drained plants was unaffected by abscisic acid sprays. Cytokinin export from the roots to the shoots declines in flooded plants. Spray applications of benzyladenine increased stomatal conductance in both flooded and drained plants. In addition, the decline in photosynthetic capacity during flooding was largely prevented by supplementary cytokinin applications. The possible involvement of these growth substances in modifying leaf gas exchange during flooding is discussed.     

The Influence of Leaf Water Status and ABA on Leaf Growth and Stomata of Phaseolus Seedlings with Hypoxic Roots

The objective of this study was to assess the relative rolesof leaf water status and root-sourced signals in mediating beanleaf responses to root hypoxia. To do so, the roots of beanplants under varied VPD (0.95 kPa to 0.25 KPa) were made hypoxic.Under all conditions, leaf growth rates and stomatal conductanceswere reduced. There was a transitory decline in leaf water potentialat high VPD which accounted for the initial reduction in leafgrowth rates and stomatal conductance. At low VPD, no waterdeficits were detected. Leaf growth inhibition and reduced stomatalconductance under low VPD treatments were unrelated to leafwater status and must be induced by some other factor. In vitrogrowth of leaf discs was reduced by xylem sap collected fromhypoxic roots. Exogenously applied ABA, at high concentrationsin KCl and sucrose, or at low concentrations diluted in xylemsap from aerated plants, inhibited in vitro growth of leaf discs.Applications of ABA in the transpiration stream reduced stomatalconductance.     

The Role of Endogenous Abscisic Acid in the Response of Plants to Stress

When a continuous stream of warm air (38°C) was directedon to the leaves of dwarf bean seedlings they wilted and thengradually regained turgor. This process of adaptation was accompaniedby an increasing abscisic acid (ABA) level in the leaves andan increase in leaf resistance (R_L). It is suggested that theleaf-water deficit induced by the warm-air treatment causedthe increase in ABA level and that the latter was responsiblefor stimulating stomatal closure, enabling the plants to regainfull turgor. A similar type of adaptation, brought about byan increased level of ABA in the leaves, is believed to occurin tomato, dwarf bean, and wheat plants when they are flooded.Predictably, in rice, a species adapted to a flooded environment,seedlings showed no increase in ABA level as a result of flooding. It is proposed that adaptation may involve the formation ofan equilibrium between ABA and its conjugate form (i. e. theglucose ester). The ABA-conjugate was observed to disperse slowlyfrom leaves recovering from a water deficit and therefore itmay act as a metabolic ‘back-stop’, enabling the‘free’ ABA level to remain high for a period evenwhen the leaves have regained turgor. Abscisic acid appears to be responsible for alleviating theeffects of water stress in plants, making it possible for plantsto pass through periods of stress with little harm.     

Ineffectiveness of cytokinin-induced chlorophyll retention in hypostomatous leaf discs

Kinetin retarded the decrease in chlorophyll content in leafdiscs from 5 species of plants with amphistomatous leaves, wherethe upper surface was exposed to air, but not in Rumex acetosera.When leaf discs were floated so that the lower surface was exposed,the effect of kinetin was less evident. Kinetin also stimulatedtranspiration in leaf discs from Nicotiana tabacum (amphistomatous),but not in leaf discs from Paederia chinensis (hypostomatous).Nor kinetin did retard chlorophyll breakdown in this specieswhen leaf discs were floated so that the stomatal surface wasin contact with the solution. The ineffectiveness of cytokininsin chlorophyll retention in leaf discs from hypostomatous leaveswas not due to reduced uptake of benzylaminopurine-¹⁴C. Chlorophyll retention was severely inhibited by coating theleaf surface with vaseline either in the presence or absenceof kinetin. Leaf discs floated on a solution exposed to CO₂-lessair retained more chlorophyll than those in normal air. Thereis thus a close relationship between stomatal opening (as measuredby stimulation of transpiration) and chlorophyll retention,as influenced by cytokinins. It is suggested that cytokinin-induced chlorophyll retentionand odier effects on leaf tissues could be mediated throughits effects on stomatal opening. (Received January 22, 1976; )     

Characterization of a wilty sunflower (Helianthus annuus L.) mutant II. Water relations, stomatal conductance, abscisic acid content in leaves and xylem sap of plants subjected to water deficiency

The response of w-1, a wilty sunflower (Helianthus annuus L.)mutant, to water stress is described in comparison with thecontrol line (W-1). Detached leaves of w-1 strongly dehydratedduring the first 30 min without significant changes in leafconductance, whereas W-1 responded rapidly to water loss byreducing stomatal aperture. After 2 h stress ABA increased slightlyin w-1, while W-1 leaves showed a 20-fold increase. When waterstress was imposed to potted plants by water withholding, w-1quickly dehydrated, and lost turgor, while W-1 maintained positiveturgor values for a longer period. Wild-type plants respondedto small changes in leaf water potential by accumulating ABAand by closing stomata, whereas in the mutant significant changesin ABA content and in stomatal conductance were found only atvery low water potentials. In another experiment in which waterwas withheld under high relative humidity, when soil water contentstarted to decrease W-1 rapidly closed stomata in the absenceof any change in leaf water status and the reduction in conductancewas paralleled by a rise in xylem sap ABA concentration. Bycontrast the mutant started to accumulate ABA in the xylem sapand to close stomata when soil water content and leaf waterpotential were dramatically reduced. The low endogenous ABAlevels and the inability to synthesize the hormone rapidly eitherin the leaves or in the roots seem to be responsible for thehigh sensitivity of w-1 to water stress. Key words: ABA, Helianthus annuus L, water relations, stomatal conductance, drought, wilty mutant     

Physiology of the parasitic association between maize and witchweed (Striga hermonthica): is ABA involved?

Striga hermonthica (Del.) Benth. is an obligate hemiparasiticangiosperm which can cause severe losses of yield in cerealcrops in the semi-arid tropics. The effects of this parasiteon the growth and stomatal conductance of three varieties ofmaize (Zea mays L.) during the first 6 weeks of the associationhave been studied. From 24 d after planting (DAP), infectedplants were significantly shorter than uninfected controls.When the plants were harvested 45 DAP, infected plants had fewerfully expanded leaves, less leaf biomass and less pseudo-stembiomass than uninfected controls. However, the parasitized plantshad more root biomass and hence a higher root:shoot ratio thanuninfected controls. The stomatal conductance of infected hostswas severely inhibited by comparison with that in uninfectedplants. The possibility that abscisic acid (ABA) may be involved inthe regulation of the parasitic association was investigated.ABA concentrations in leaf tissue of maize (cv. Cargimontana)and S. hermonthica were determined by radioimmunoassay. Whilethere was a difference between cultivars in the extent of theresponse, the concentrations of ABA were significantly higherin infected maize plants than in the uninfected controls. InS. hermonthica, leaf tissue ABA concentration was found to bean order of magnitude higher than in the host leaf tissue. Detachedleaves of S. hermonthica which were dehydrated at room temperatureuntil they had lost 10–20% of their fresh weight containedthree times the ABA concentration of control leaves. This suggeststhat leaves of S. hermonthica can synthesize or re-mobilizeABA in response to water deficit. It is not yet known whetherthis contributes to the higher concentration in infected hosts,but the results suggest that ABA has a role in this parasiticassociation. Key words: Striga hermonthica, abscisic acid, growth, parasitic angiosperm, stomatal conductance     

Stomatal Closure in Flooded Tomato Plants Involves Abscisic Acid and a Chemically Unidentified Anti-Transpirant in Xylem Sap

We address the question of how soil flooding closes stomata of tomato (Lycopersicon esculentum Mill. cv Ailsa Craig) plants within a few hours in the absence of leaf water deficits. Three hypotheses to explain this were tested, namely that (a) flooding increases abscisic acid (ABA) export in xylem sap from roots, (b) flooding increases ABA synthesis and export from older to younger leaves, and (c) flooding promotes accumulation of ABA within foliage because of reduced export. Hypothesis a was rejected because delivery of ABA from flooded roots in xylem sap decreased. Hypothesis b was rejected because older leaves neither supplied younger leaves with ABA nor influenced their stomata. Limited support was obtained for hypothesis c. Heat girdling of petioles inhibited phloem export and mimicked flooding by decreasing export of [14C]sucrose, increasing bulk ABA, and closing stomata without leaf water deficits. However, in flooded plants bulk leaf ABA did not increase until after stomata began to close. Later, ABA declined, even though stomata remained closed. Commelina communis L. epidermal strip bioassays showed that xylem sap from roots of flooded tomato plants contained an unknown factor that promoted stomatal closure, but it was not ABA. This may be a root-sourced positive message that closes stomata in flooded tomato plants.     

Effects of flooding on carbohydrate and ABA levels in roots and shoots of alfalfa

Alfalfa is sensitive to waterlogging, and its yields are significantly reduced under this condition. We investigated the effects of soil flooding on free abscisic acid (ABA) accumulation in shoots and roots of alfalfa in relation to plant growth and stomatal conductance responses. The production of dry matter in alfalfa was significantly affected by flooding mainly as a result of a rapid reduction in root growth. Shoot dry matter accumulation was maintained during the first 10 d of treatment and started to decline thereafter. Foliar concentration of the major mineral elements (N, P, K) was reduced by flooding, whereas only K concentration decreased in roots of flooded plants. Regrowth declined with duration of flooding and was less than 50% of controls after 2 weeks. While no changes in ABA concentration could be detected in flooded roots, an increase was noted within a few days in leaves when compared to unflooded controls. This increase in free ABA coincided with the accumulation of large quantities of starch in leaves and a rapid decline in leaf stomatal conductance. Our results support the suggestion that leaf ABA originates from the leaf itself and may be accumulating along with starch as a result of reduced translocation to the roots. Our observation of large accumulations of sucrose in flooded roots agrees with previous reports that supply of carbohydrates is not a limiting factor to root anaerobic metabolism in flooded alfalfa.     

Flooding responses inZea mays L.

Summary This study was undertaken to determine the specific mechanisms of flooding injury inZea mays L., especially the role of water deficit. Maize plants in soil were artificially flooded in greenhouse and field, both with and without exchanging the soil solution with new water and nutrients. Plants in solution culture were rapidly stressed by replacing aeration gas with nitrogen. In all cases of measurable short term response, root and leaf growth rates decreased within 1 to 12 hours, and stomatal resistance increased 2 to 3 days later. Both growth rates and stomatal resistance recovered spontaneously during the flooding period. Over the long term (more than a week), growth rates were inhibited only when the soil solution was not periodically changed and Kjeldahl measured nitrogen deficiency was evident in the leaves. None of the above effects was associated with high root resistance to water uptake, plant water deficit, or a change in the osmotic balance of the leaves. The only water stress associated with the experiments was found immediately after draining plants that had been flooded for 2 weeks. In this instance, the preflooding root systems appeared damaged.     

Potassium Loss from Stomatal Guard Cells at Low Water Potentials

The potassium content of guard cells and the resistance to viscousflow of air through the leaf were determined in sunflower (Helianthusannuus) subjected to low leaf water potentials under illuminatedconditions. In intact plants desiccated slowly by withholdingwater from the soil, large losses in guard cell K occurred asleaf water potentials decreased. Leaf viscous resistance increased,indicating stomatal closure. Similar results were obtained whendetached leaf segments were desiccated rapidly. Upon rehydrationof leaves, no stomatal opening was observed initially, despiteleaf water potentials at predesiccated levels. After severalhours, however, re-entry of K occurred and stomata became fullyopen. Turgid leaf segments floated on an ABA solution showedlosses of guard cell K and closure of stomata as rapidly andcompletely as those brought about by desiccation. It is concludedthat stomatal closure at low water potentials under illuminatedconditions is not controlled solely by water loss from the tissuebut involves the loss of osmoticum from the guard cells as well.This in turn decreases the turgor difference between the guardcells and the surrounding cells, and closing occurs.