Flag Leaf Photosynthesis of Triticum aestivum and Related Diploid and Tetraploid Species

Rates of net photosynthesis of the flag leaves of 15 genotypesof wheat and related species were measured throughout theirlife, using intact leaves on plants grown in the field. At thestage when rates were maximal, they were in general highestfor the diploid species, intermediate for the tetraploidspeciesand lowest for Triticum aestivum (means of 38, 32 and 28 mgCO₂ dm^–2 h^–1 respectively). Rates were stronglynegatively correlated with leaf area, leaf width and the meanplan area per mesophyll cell and positvely correlated with stomatalfrequency and number of veins per mm of leaf width. The differencesamong species in these attributes were mainly related to ploidylevel. It was not possible to determine the relative importanceof each anatomical feature, though the changes in stomatal frequencyhad only slight effects on stomatal conductance and the observeddifferences in rates of photosynthesis were much greater thanwould be expected from those in stomatal conductance alone. There was genetic variation in rates of light dependent oxygenevolution of isolated protoplasts and intact chloroplasts butno difference attributable to ploidy. The mean rate, 91 µmolO₂ mg^–1 chlorophyll h^–1, equivalent to 3.9 mg CO₂mg^-1chlorophyll h^-1 was considerably less than the rate of photosynthesisin comparable intact leaves, which was 7.2 mg CO₂ mg^–1chlorophyll h^–1. The total above-ground dry matter yields were least for thewild diploids T. urartu and T. thauodar and the wild tetraploidT. dicoccoides, but the other wild diploids produced as muchdry matter as the hexaploids. The prospects of exploiting differences in photosynthetic ratein the breeding of higher yielding varieties are discussed. Triticum aestivum L., wheat, Aegilops spp, photosynthesis, stomatal conductance, stomatal frequency, polyploidy     

In-Phase Cycling of Photosynthesis and Conductance at Saturating Carbon Dioxide Pressure Induced by Increases in Water Vapour Pressure Deficit

Bunce, J. A. 1987. In-phase cycling of photosynthesis and conductanceat saturating carbon dioxide pressure induced by increases inwater vapour pressure deficit.—J. exp. Bot. 38: 1413–1420. The leaf to air water vapour deficit was increased suddenlyfrom about 1·0 to 2·5 IcPa for single leaves ofsoybean (Glycine max L. Merr.) plants held at 30 °C, 2·0mmol m ^–2 s^–1 photosynthetic photon flux density(PPFD) and carbon dioxide pressures saturating to photosynthesis.After a lag of about 10 min, photosynthetic rate and stomatalconductance to water vapour began to decrease, and then cycledin phase with each other. The period of the cydes was about20 min. During these cycles the substomatal carbon dioxide pressurewas constant in the majority of leaves examined, and was alwaysabove saturation for photosynthesis. Epidermal impressions showedthat most stomata changed in aperture during the cycles, andthat very few were ever fully closed. Water potential measuredon excised discs changed by at most 0·1 MPa from theminima to the maxima in transpiration rate. In contrast, forleaves of sunflower (Helianthus animus L.) grown at low PPFD,the increase in VPD led to leaf wilting and decreased photosynthesis,followed by recovery of turgor and photosynthesis as stomatalconductance began to decrease. In these leaves photosynthesisand conductance then cycled approximately 180° out of phase.It is suggested that in soybeans decreased leaf conductanceinduced by high VPD provided a signal which decreased the rateof photosynthesis at carbon dioxide saturation by a mechanismthat was not related to a water deficit in the mesophyll. Key words: Photosynthesis, stomatal conductance, cycling, vapour pressure deficit     

The Effect of Shade During Leaf Expansion on Photosynthesis by White Clover Leaves

In three experiments measurements of photosynthesis were madeon single leaves of white clover (Trifolium repens L.) on threecultivars grown in a controlled environment. Plants which had grown under an irradiance of 30 J m^–2s^–1, or in shade within a simulated mixed sward, producedleaves with photosynthetic capacities some 30 per cent lowerthan did plants grown at 120 J m^–2 s^–1 without shade.There were no differences between treatments either in photosynthesismeasured at 30 J m^–2 s^–1, or in respiration ratesper unit leaf dry weight. Respiration per unit leaf area washigher in the plants grown at 120 J m^–2 s^–1, reflectingthe lower specific leaf area of these leaves. There were nodifferences between the three cultivars examined. Leaves which were removed from the shade of a simulated swardshortly after becoming half expanded achieved photosyntheticcapacities as high as those which were in full light throughouttheir development. It is suggested that it is this characteristicwhich enables clover plants growing in an increasingly densemixed sward to produce a succession of leaves of high photosyntheticcapacity, even though each lamina only reaches the top of thesward at a relatively late stage in its development. Trifolium repens L., white clover, photosynthesis, leaf expansion, shade, specific leaf area, stomatal conductance     

Light Distribution and Photosynthesis in Field Crops

In a new model of light distribution in field crops a parameters is the fraction of light passing through unit leaf layer withoutinterception. Radiation profiles measured with solarimetersand photocells give values of s from 0.7 for grasses to 0.4for species with prostrate leaves. Knowing s, leaf transmissionT and leaf-area index L the light distribution in a field cropmay be described by a binomial expansion of the form {s+(I-s)T)^L.To calculate crop photosynthesis at given light intensity thisexpansion is combined with two parameters describing the shapeof the light-response curve of single leaves. Finally, the assumptionthat solar radiation varies sinusoidally allows daily totalphotosynthesis to be estimated from daylength and insolation. The theory predicts about the same potential photosynthesisin a cloudy temperte climate with long days as in a more sunnyequatorial climate with short days. When L < 3 photosynthesisincreases as s decreases, i.e. as leaves become more prostrate;but when L > 5, photosynthesis increases as s increases,i. e. as leaves become more erect. Assuming that respirationis proportional to leaf area, estimated dry-matter productionagrees well with field measurements on sugar-beet, sugar-cane,kale, and subterranean clover. Estimates of maximum gross photosynthesis(for sugar-cane and maize) range from 60 to 9 g m^–2 day^–1depending on insolation.     

Photosynthesis, Water Relations, Temperature and Canopy Structure as Factors Influencing the Growth of Sainfoin (Onobrychis viciifolia Scop.) and Lucerne (Medicago sativa L.)

Measurements of the growth of sainfoin and lucerne were madein the field after cutting on 31 May 1977. Sainfoin reacheda total above-ground dry weight of 408 g m^–2 over thegrowing period of 48 days compared with 598 g m^–2 in lucerne.Final leaf area indices (LAIs) were 2.8 in sainfoin and 6.1in lucerne. The specific leaf areas (SLAs) for sainfoin wereapproximately half those of lucerne throughout the regrowthperiod. The maximum rates of leaf appearance were 0.12 leavesper day in sainfoin and 0.85 leaves per day in lucerne. Themaximum mean rate of plant extension growth for lucerne of 2.12mm h^–1 occurred during the night, whereas, in sainfointhe maximum rate of 1.72 mm h^–1 occurred during the day. Measurements of extinction coefficients for PAR ranged from0.45 to 0.89 in sainfoin and from 0 42 to 0.57 in lucerne. Asthe lucerne crop increased in size leaf water potentials andsolute potentials became more negative. In sainfoin leaf waterpotentials remained remarkably high throughout the growth period,solute potentials decreased and turgor potentials increased.The stomatal conductances of the two species were similar. The photosynthetic capacities and rates of dark respirationper unit leaf area in both species were similar. The rate ofcanopy ‘gross’ photosynthesis at 295 W m^–2was always greater in lucerne than in sainfoin. This was largelya matter of differences between the species in LAI, althoughat higher LAIs the more erect structure of lucerne leads toa better utilization of photosynthetically active radiation. Onobrychis vicifolia Scop, sainfoin, Medicago sativa L., lucerne, photosynthesis, water relations, temperature, canopy structure     

Photoinhibition under Atmospheric O2, the Activation State of RuBP Carboxylase and the Content of Photosynthetic Intermediates in Soybean and Wheat

Ward, D. A. and Drake, B. G. 1987. Photoinhibition under atmosphericO₂, the activation state of RuBP carboxylase and the contentof photosynthetic intermediates in soybean and wheat.—J.exp. Bot. 38: 1937–1948. Associations between photosynthesis, the activation state ofRuBP carboxylase and the contents of photosynthetic intermediateswere compared in soybean and wheat leaves before and after exposureto photoinhibitory treatments in the presence of atmosphericO₂. Exposing attached leaves to a supra-saturating irradiance(3 800 µmol quanta m^{– 2} s^–1) for 2 h in CO_2-freeair decreased carboxylation efficiency and the light-saturatedphotosynthetic rate in air by approximately 50%. Exposure tothe photoinhibitory treatment for periods in excess of 2 h didnot cause a further decrease of photosynthesis in soybean. Althoughphotosynthesis was reduced, the initial and total (fully-activated)activities of ribulose 1,5-bisphosphate carboxylase (RuBPCase)in leaf extracts were unaltered in each species by the photoinhibitorytreatment. This was true for leaves sampled under both air andat a rate-limiting intercellular CO₂ partial pressure (C_i) of75 µPa Pa^–1. The contents of ribulose l,5-bisphosphate(RuBP) and 3-phosphoglyceric acid (3-PGA) were reduced by thephotoinhibitory treatment in soybean leaves sampled in air andat a rate-limiting C_i, although the RuBP/3-PGA ratio was unaffected.The relative reduction of RuBP content in soybean leaves atrate-limiting C₁ was similar to the corresponding reductionof carboxylation efficiency. For wheat,the relative reductionof RuBP content at rate-limiting C_i (–19%) caused by thephotoinhibitory treatment was considerably less than the correspondingdecrease of carboxylation efficiency (–49%).The RuBP/3-PGAratio of wheat was also increased significantly by the photoinhibitorytreatment The significance of these observations to the regulationof CO₂-limited photosynthesis in leaves experiencing photoinhibitionunder atmospheric oxygen is discussed. Consideration is alsogiven to the previous contention that contemporary measurementsof initial activity in crude extracts may provide a spuriousindication of the amount of the enzyme-CO₂-Mg_{2 +} form of RuBPcarboxylase present in the leaf. Key words: Carboxylation efficiency, RuBP carboxylase, photoinhibition, RuBP, 3-PGA     

Relationship between Leaf Structure and Gas Exchange in Wheat Leaves at Different Insertion Levels

Net photosynthesis rate (P_n), stomatal conductance to CO₂ andresidual conductance to CO₂ were measured in the last six leaves(the sixth or flag leaf and the preceding five leaves) of Triticumaestivum L. cv. Kolibri plants grown in Mediterranean conditions.Recently fully expanded leaves of well-watered plants were alwaysused. Measurements were made at saturating photosynthetic photonflux density, and at ambient CO₂ and O₂ levels. The specificleaf area, total organic nitrogen content, some anatomical characteristics,and other parameters, were measured on the same leaves usedfor gas exchange experiments. A progressive xeromorphic adaptation in the leaf structure wasobserved with increasing leaf insertion levels. Furthermore,mesophyll cell volume per unit leaf area (V_mes/A) decreasedby 52·6% from the first leaf to the flag leaf. Mesophyllcell area per unit leaf area also decreased, but only by 24·5%.However, nitrogen content per unit mesophyll cell volume increasedby 50·6% from the first leaf to the flag leaf. This increasecould be associated to an observed higher number of chloroplastcross-sections per mm² of mesophyll cell cross-sectional areain the flag leaf: values of 23000 in the first leaf and 48000in the flag leaf were obtained. P_n per unit leaf area remainedfairly constant at the different insertion levels: values of33·83±0·93 mg dm^–2 h^–1 and32·32±1·61 mg dm^–2 h^–1 wereobtained for the first leaf and the flag leaf, respectively.Residual conductance, however, decreased by 18·2% fromthe first leaf to the flag leaf. Stomatal conductance increasedby 41·7%. The steadiness in P_n per unit leaf area across the leaf insertionlevels could be mainly accounted for by an opposing effect betweena decrease in V_mes/A and a more closely packed arrangement ofphotosynthetic apparatus. Adaptative significance of structuralchanges with increasing leaf insertion levels and the steadinessin P_n per unit leaf area was studied. Key words: Photosynthesis, structure, wheat     

The Control of the Light Acclimation of Photosynthesis in Glycine max: Dependence on Import as Modified by Intraplant Shading

The dependence of photosynthetic capacity on imported and locally-assimilatedsupplies of carbon during leaf development under different irradianceswas investigated in Glycine max. The potential export of carbonto the developing, mainstem trifoliate leaf (source-potential)was restricted non-destructively by shading all lower, sourceleaves (source-shading), while local photosynthesis was modifiedconcurrently by exposing the young leaf to different light levelsduring development. When source-shading was applied below the2nd mainstem trifoliate leaf at the bud stage of development,photosynthetic capacity was unaffected in leaves which had developedunder moderate and low irradiances (500 and 250 µmol PARm ^–2 s^–1 respectively), but was reduced significantlyin leaves developed under a high irradiance (900 µmolPAR m ^–2 s^–1). If source-shading was applied beneaththe 2nd leaf at unfolding, the reduction of photosynthetic capacityunder the high irradiance was relatively minor. The photosyntheticcapacity attained by the 2nd leaf during development under differentirradiances was influenced by the previous light environmentof the whole plant. In contrast to the 2nd leaf, the photosyntheticcapacities of the 1st and 4th mainstem leaves were relativelyunaffected by source-shading, even under the highest light regime.While photosynthetic capacity showed a widespread insensitivityto the light level of the lower region of the canopy, source-shadingreduced final leaf size irrespective of node position or localirradiance during leaf development. These effects were not relatedto differences in daily photosynthesis by the expanding leaf,and are discussed in terms of the source/sink balance of thedeveloping leaf. Key words: Glycine max, source-shading, photosynthetic capacity     

Microclimate, Photosynthesis and Growth of Lucerne (Medicago sativa L.). I. Microclimate and Photosynthesis

Measurements of microclimate and photosynthesis of lucerne var.Europe were made in the field during the spring of 1976. Themaximum rate of canopy gross photosynthesis (14.3 g CO₂ m^–2h^–1, I = ) was 2.5 times greater than that of S 24 perennialryegrass at the same LAI. This difference was due to differencesin individual leaf photosynthesis. The photosynthetic rate ofthe youngest fully expanded leaf of lucerne remained constantthroughout the experimental period at 3.6 g CO₂ m^–2 h^–1(300 W m^–2). Measurements of soil water potential profiles indicated thatlucerne extracted water from the soil to a depth of at least800 mm, with a region of maximum uptake between 400 and 600mm. This capability, with a moderate mean leaf resistance of460 s m^–1, conferred a high assimilation efficiency onlucerne, with a mean water use efficiency of 34 g H₂O lost pergram of carbohydrate assimilated, compared with 200 g H₂O pergram of carbohydrate for S 24. Medicago sativa L, lucerne, photosynthesis, assimilation efficiency     

Carbon Partitioning and Export in Mature Leaves of Pepper (Capsicum annuum)

The partitioning of recently fixed carbon by mature pepper leaveshas been examined over a 10 h photoperiod using a constant specificradioactivity ¹⁴CO₂ labelling technique. Changes in the ratesof carbon partitioning into export, starch, sucrose and hexoseswere examined following changes in irradiance during the photoperiod.Leaves grown under 80 W m^–2 PAR were exposed to this irradiancefor the first 4 h of the photoperiod then the iiradiance wasdecreased. Leaves accumulated sufficient reserves in the first4 h to maintain export at the initial rate (approximately 20µg carbon cm^–2 leaf h^–1) over the following6 h of the photoperiod when the net photosynthesis rate (Pn)was decreased to 10% of the initial rate by the decreased irradiance.Export was initially maintained by the depletion of sucroseand hexose and then by carbon from the degradation of starchin the light. If leaves were exposed to low irradiance at the beginning ofthe photoperiod, then the export rate was linearly related tothe Pn during that period. When Pn exceeded that required tomaintain an export rate of approximately 20 µg carboncm^–2 h^–1, then more carbon was partitioned intostarch. At low initial irradiance, a greater proportion of photosynthatewas partitioned into export rather than starch and at high initialirradiancc the reverse occurred. There was a linear relationship between starch accumulationrate and Pn for all leaves but the relationship between Pn andexport rate was only significant for leaves with low levelsof reserve carbon. The results show that mature pepper leaves subjected to differentirradiances maintain constant export rates through alterationsof carbon partitioning. Export at low Pn is maintained at theexpense of sugar and starch reserves, with partitioning in highirradiance being predominantly to starch. Key words: Carbon partitioning, Starch, Export, Pepper (Capsicum annuum L.)     

Effect of Elevated CO2 on the Photosynthesis, Respiration and Growth of Perennial Ryegrass

Single, seed-grown plants of ryegrass (Lolium perenne L. cv.Melle) were grown for 49 d from the early seedling stage ingrowth cabinets at a day/night temperature of 20/15 C, witha 12 h photoperiod, and a CO₂ concentration of either 340 or680µI 1^–1 CO₂. Following complete acclimation tothe environmental regimes, leaf and whole plant CO₂ effluxesand influxes were measured using infra-red gas analysis techniques.Elevated CO₂ increased rates of photosynthesis of young, fullyexpanded leaves by 35–46% and of whole plants by morethan 50%. For both leaves and whole plants acclimation to 680µI^–1 CO₂ reduced rates of photosynthesis in bothCO₂ regimes, compared with plants acclimated to 340µll^–1. There was no significant effect of CO₂ regime onrespiration rates of either leaves or whole plants, althoughleaves developed in elevated CO₂ exhibited generally lower ratesthan those developed in 340µI I^–1 CO₂. Initially the seedling plants in elevated CO₂ grew faster thantheir counterparts in 340µI I^–1 CO₂, but this effectquickly petered out and final plant weights differed by onlyc. 10%. Since the total area of expanded and unexpanded laminaewas unaffected by CO₂ regime, specific leaf area was persistently13–40% lower in elevated CO₂ while, similarly, root/shootratio was also reduced throughout the experiment. Elevated CO₂reduced tissue nitrogen contents of expanded leaves, but hadno effect on the nitrogen contents of unexpanded leaves, sheathsor roots. The lack of a pronounced effect of elevated CO₂ on plant growthwas primarily due to the fact that CO₂ concentration did notinfluence tiller (branch) numbers. In the absence of an effecton tiller numbers, any possible weight increment was restrictedto the c. 2.5 leaves of each tiller. The reason for the lackof an effect on tillering is not known. Key words: Lolium perenne, ryegrass, elevated CO₂, photosynthesis, respiration, growth, development     

Drought effects on cellular and spatial parameters of leaf growth in tall fescue

The effect of drought and recovery on cellular and spatial parametersof the growth process in tall fescue leaves was studied in twoexperiments. In both experiments plants grown on vermiculiteand maintained in a controlled environment were submitted toa 7 d drought period generated by withholding water. Droughtwas followed by a 3 d recovery period in experiment II. As leafelongation rate (LER) decreased during developing drought boththe growth zone length (initially 40 mm) and the maximum relativeelemental growth rate (initially 0.09 mm mm^–1 h^–1during the dark period of diurnal cycles) within the growthzone declined. But the growth zone still exhibited a lengthof approximately 15 mm when LER approached 0 under severe drought(–2.0 MPa predawn leaf water potential). The growth potentialof the basal 15-mm-long portion of the leaf was conserved duringthe period when drought effected the complete arrest of leafelongation. A (retrospective) analysis of the position-timerelationships of epidermal cells identified on leaf replicas(experiment II) indicated that the cell flux out of the growthzone responded very sensitively to drought. Before drought theflux was maximum at approximately 3.2 cells (cell file h)^–1during the dark period. Flux decreased to 0 when leaf elongationstopped. Flux also varied diurnally both under well-wateredand droughted conditions. In well-watered conditions it wasabout 30% less during the light than the dark period. Cell elongationwas also sensitive to drought. Under well-watered conditionsepidermal cell elongation stopped when cells attained a lengthof approximately 480 µm. During developing drought cellsstopped elongating at progressively shorter lengths. When LERhad decreased to almost nil, cells stopped elongating at a lengthof approximately 250 µn. When drought was relieved followinga 2 d complete arrest of leaf elongation then cells shorterthan 250 µm were able to resume expansion. Following rewateringcell flux out of the growth zone increased rapidly to and abovethe pre-drought level, but there was only a slow increase overtime in the length at which cell elongation stopped. About 2d elapsed until the leaf growth zone produced cells of similarlength as before drought (i.e. approximately 480 µm). Key words: Epidermal cell length, cell flux, (leaf) growth zone, leaf elongation rate, relative elemental growth rate, position-time relationships (path line, growth trajectory), drought, water deficit     

Effects of elevated CO2 concentrations on three montane grass species: III. Source leaf metabolism and whole plant carbon partitioning

Agrostis capillaris L.⁵, Festuca vivipara L. and Poaalpina L.were grown in outdoor open-top chambers at either ambient (340 3µmol mol^–1) or elevated (6804µmol mol^–1)concentrations of atmospheric carbon dioxide (CO₂) for periodsfrom 79–189 d. Photosynthetic capacity of source leaves of plants grown atboth ambient and elevated CO₂ concentrations was measured atsaturating light and 5% CO₂. Dark respiration of leaves wasmeasured using a liquid phase oxygen electrode with the buffersolution in equilibrium with air (21% O₂, 0.034% CO₂). Photo-syntheticcapacity of P. alpina was reduced by growth at 680 µmolmol^–1 CO₂ by 105 d, and that of F. vivipara was reducedat 65 d and 189 d after CO₂ enrichment began, suggesting down-regulationor acclimation. Dark respiration of successive leaf blades ofall three species was unaltered by growth at 680 relative to340 µmol mol^–1 CO₂. In F. vivipara, leaf respirationrate was markedly lower at 189 d than at either 0 d or 65 d,irrespective of growth CO₂ concentration. There was a significantlylower total non-structural carbohydrate (TNC) concentrationin the leaf blades and leaf sheaths of A. capillaris grown at680µmol mol^–1 CO₂. TNC of roots of A. capillariswas unaltered by CO₂ treatment. TNC concentration was increasedin both leaves and sheaths of P. alpina and F. vivipara after105 d and 65 d growth, respectively. A 4-fold increase in thewater-soluble fraction (fructan) in P. alpina and in all carbohydratefractions in F. vivipara accounted for the increased TNC content. In F. vivipara the relationship between leaf photosyn-theticcapacity and leaf carbohydrate concentration was such that therewas a strong positive correlation between photosynthetic capacityand total leaf N concentration (expressed on a per unit structuraldry weight basis), and total nitrogen concentration of successivemature leaves reduced with time. Multiple regression of leafphotosynthetic capacity upon leaf nitrogen and carbohydrateconcentrations further confirmed that leaf photosynthetic capacitywas mainly determined by leaf N concentration. In P. alpina,leaf photosynthetic capacity was mainly determined by leaf CHOconcentration. Thus there is evidence for down-regulation ofphotosynthetic capacity in P. alpina resulting from increasedcarbohydrate accumulation in source leaves. Leaf dark respiration and total N concentration were positivelycorrelated in P. alpina and F. vivipara. Leaf dark respirationand soluble carbohydrate concentration of source leaves werepositively correlated in A. capillaris. Changes in source leafphotosynthetic capacity and carbohydrate concentration of plantsgrown at ambient or elevated CO₂ are discussed in relation toplant growth, nutrient relations and availability of sinks forcarbon. Key words: Elevated CO₂, Climate change, grasses, carbohydrate partitioning, photosynthesis, respiration     

Growth and Nitrogen Fixation of Phaseolus vulgaris L. at Two Irradiances 1. Growth

Plants of Phaseolus vulgaris grown at 7 and 28 W m^–2 showedno differences in rate of development of leaves or flowers.At 7 W m-Z plants had longer internodes, more succulent stemsand leaves, higher ratios of shoot:root and greater leaf areasthat those at 28 W m^–2. These differences were establishedprior to detectable differences in photosynthesis and couldpartly be attributed to an increased proportion of far-red light. Although the final d. wt, carbon content, and fruit yield werehigher at 28 W m^–2, plants at 7 W m^–2 apparentlyhad similar relative growth rates and greater photosyntheticefficiency. Dry weight differences are most easily interpretedas resulting from the establishment of an earlier net carbongain at 28 W m^–2 than at 7 W m^–2.     

Measurement of the Apoplastic Activity of K+ and Cl- in the Leaf Epidermis of Commelina communis in Relation to Stomatal Activity

Bowling, D. J. F. 1987. Measurement of the apoplastic activityof K⁺ and Cl^– in the leaf epidermis of Commelina communisin relation to stomatal activity.–J. exp. Bot. 38: 1351–1355. Ionic activity of K⁺ and Cl^– in the apoplast of the lowerepidermis of the leaf of Commelina communis was measured usingion selective micro-electrodes. Large gradients across the stomatalcomplex were observed which were related to stomatal aperture.On stomatal closure the activity of K⁺ and Cl^– in theapoplast of the guard cell rose from 3·0 mol m^–3to 100 mol m^–3 and 33 mol m^–3 respectively. It wasconcluded that the apoplast is an important pathway for iontransport between the cells. Key words: Stomata, ionic activity, leaves, apoplast     

Control of the Acclimation of Photosynthesis to Light and Temperature in Relation to Partitioning of Photosynthate in Developing Soybean Leaves

Photosynthetic acclimation was examined by exposing third trifoliolateleaves of soybeans to air temperatures of 20 to 30°C andphotosynthetic photon flux densities (PPFD) of 150 to 950µmolphotons m^–2 s^–1 for the last 3 d before they reachedmaximum area. In some cases the environment of the third leafwas controlled separately from that of the rest of the plant.Photosynthesis, respiration and dry mass accumulation were determinedunder the treatment conditions, and photosynthetic capacity,and dry mass and protein content were determined at full expansion.Photosynthetic capacity, the light-saturated rate of net carbondioxide exchange at 25°C and 34 Pa external partial pressureof carbon dioxide, could be modified between 21 and 35 µmolCO₂ m^–2 s^–1 by environmental changes after leaveshad become exporters of photosynthate. Protein per unit leafmass did not differ between treatments, and photosynthetic capacityincreased with leaf mass per unit area. Photosynthetic capacityof third leaves was affected by the PPFD incident on those leaves,but not by the PPFD on other leaves on the plant. Photosyntheticcapacity of third leaves was affected by the temperature ofthe rest of the plant, but not by the temperature of the thirdleaves. Photosynthetic capacity was linearly related to carbondioxide exchange rate in the growth regimes, but not to daytimePPFD. At high PPFD, and at 25 and 30°C, mass accumulationwas about 28% of the mass of photosynthate produced. At lowerPPFD, and at 20°C, larger percentages of the photosynthateproduced accumulated as dry mass. The results suggest that photosynthatesupply is an important factor controlling leaf structural growthand, consequently, photosynthetic acclimation to light and temperature. Key words: Glycine max (L.) Merr., photosynthesis, temperature acclimation, light acclimation, photosynthate partitioning     

Osmotic Stress Temporarily Reverses the Inhibitions of Photosynthesis and Stomatal Conductance by Abscisic Acid--Evidence that Abscisic Acid Induces a Localized Closure of Stomata in Intact, Detached Leaves

Ward, D. A. and Drake, B. G. 1988. Osmotic stress temporarilyreverses the inhibitions of photosynthesis and stomatal conductanceby abscisic acid—evidence that abscisic acid induces alocalized closure of stomata in intact, detached leaves.—J.exp. Bot 39: 147–155. The influence of osmotic stress on whole leaf gas exchange wasmonitored in detached leaves of Glycine max supplied with anexogenous concentration (10^–5 mol dm^–3) of ±abscisicacid (ABA) sufficient to inhibit net photosynthesis and stomatalconductance by 60% and 70%, respectively, under a saturatingirradiance and normal air. Raising the osmotic (sorbitol) concentrationof the ABA solutions feeding leaves elicited rapid and synchronousreversals of the ABA-dependent inhibitions of net photosynthesisand conductance. These reversals reached a peak simultaneously,after which photosynthesis and conductance declined. The magnitudeof the transient stimulations at peak height was dependent uponthe sorbitol concentration of the ABA feeding solution, althoughthe time-course of the transients (half time, 4–6 min)was similar for the different osmotic concentrations applied.Irrespective of transient size the relative changes of photosynthesisand conductance were comparable; consequently the calculatedpartial pressure of CO₂ in the substomatal space (Ci) remainedrelatively constant during the transient phase. In contrastto the ABA-treated leaves, elevating the osmotic concentrationof the distilled water supply feeding control leaves stimulatedconductance to a much greater relative extent than photosynthesis.The co-stimulations of photosynthesis and conductance inducedin ABA-treated leaves by osmotic shock were not due to a restrictionin the transpirational uptake of ABA and occurred irrespectiveof the source osmoticum applied. These data are consistent with the hypothesis that the ABA-dependentinhibition of photosynthesis at constant Ci is an artifact causedby the spatially heterogeneous closure of stomata in responseto ABA. Alternative explanations for the responses are, however,considered. Key words: Abscisic acid, photosynthesis, osmotic stress, Glycine max, stomatal conductance     

Sink Activity Estimation by Sink Size and Dry Matter Increase During the Ripening Stage of Barley (Hordeum vu/gare) and Rice (Oryza sativa)

During the ripening stage of barley and rice, the sink activitywas defined as the dry matter increase per units sink size,leaf area and time, as follows: NAR = A.SinkW+NAR', where NAR is the net assimilation rate (g d.wt dm^–2d^–1);A is the sink activity [g d.wt g^–1d.wt (ear) dm^–2d^–1]; Sink W is ear wt per plant at heading (g d.wt);and NAR' is net assimilation rate excluding the assimilate ofsink organ (g d.wt dm^–2 d^–1). Plant material with 16 combinations of mutually different sink(ear) and source (leaf) size were produced at heading for eachcrop: parts of each leaf and ear were removed to produce fourgrades in barley, and also a part of each leaf was removed producingfour grades for four rice varieties showing different ear size.NAR and NAR' were determined during 26 and 21 d in barley andrice after heading, respectively. Sink activity (A), representedas the assimilation rate induced by the sink organ, was estimatedfrom the relationship between SinkW and NAR using the previousequation. The sink activity was significantly higher in ricewith a value of 0–0.028 g d.wt g^–1 d.wt (ear) dm^–2d^–1 vs. 0–0.0017 in barley, suggesting that therelative role of leaves for grain filling is considerably higherin rice than in barley. The sink activity obtained in the studymight be introduced into a model to predict the yields of barleyand rice. Hordeum vulgare L, barley, Oryza saliva L, rice, dry matter, NAR, sink, source, sink activity, model     

Nitrate Effects on Growth of the First Four Main Stem Leaves of a Range of Temperate Cereals and Pasture Grasses

The effects of different applied NO₃^– concentrations onextension growth and final length and area of leaves 1–4of five cereals and six pasture grasses of temperate originwere examined. Increased applied NO₃^– in the range 0.1–0.5.0mol m^–3 caused decreased duration of growth but increasedgrowth rate and final length of leaves 2–4 of the cerealsAvena saliva, Hordeum vulgare, Secale cereale, x Triticosecaleand Triticum aestivum. For all cereals, increased NO₃^–resulted in increased area of leaves 1–4. Pasture grasseswere supplied either 0.5 or 50 mol m^–3 NO₃^–. Increasedapplied NO₃^– (0.5–5.0 mol m^–3) resulted indecreased duration of growth and increased growth rate and finalarea of leaves 1–4 of Bromus wiltdenowii, leaves 2–4ofFestuca arundinaceae and leaves 3 and 4 of Lolium muitiflorum.In addition, length of leaves 3 and 4 of B. witidenowii increasedwith increased NO₃^–. Increased NO₃^– resulted inincreased area of leaves 2–4 of Dactylis gtomerata andLolium perenne and leaves 3 and 4 of Phalaris aquaiica but hadno effect on extension growth of all three species. Avena sativa L, oat, Hordeum vulgare L, barley, Secale cereale L, rye, x Triticosecale Wittm, triticale, Triticum aestivum L, wheat, Bromus willdenowii Kunth, prairie grass, Dactylis gtomerata L, cocksfoot, Festuca arundinaceae Shreb, tall fescue, Lolium multijlorum Lam, Italian ryegrass, Lolium perenne L, perennial ryegrass, Phalaris aquatica L, nitrate, leaf extension, leaf expansion     

Influence of Elevated CO2 and Temperature on the Photosynthesis and Respiration of White Clover Dependent on N2 Fixation

Single clonal plants of white clover (Trifolium repens L) grownfrom explants in a Perlite rooting medium, and dependent fornitrogen on N₂ fixation in root nodules, were grown for severalweeks in controlled environments which provided two regimesof CO₂, and temperature 23/18 °C day/night temperaturesat 680 µmol mol^–1 CO₂, (C680), and 20/15 °Cday/night temperatures at 340 µmol mol^–1 CO₂ (C340)After 3–4 weeks of growth, when the plants were acclimatedto the environmental regimes, leaf and whole-plant photosynthesisand respiration were measured using conventional infra-red gasanalysis techniques Elevated CO₂ and temperature increased ratesof photosynthesis of young, fully expanded leaves at the growthirradiance by 17–29%, despite decreased stomatal conductancesand transpiration rates Water use efficiency (mol CO₂ mol H₂O^–1)was also significantly increased Plants acclimated to elevatedCO₂, and temperature exhibited rates of leaf photosynthesisvery similar to those of C340 leaves ‘instantaneously’exposed to the C680 regime However, leaves developed in theC680 regime photosynthesised less rapidly than C340 leaves whenboth were exposed to a normal CO₂, and temperature environmentIn measurements where irradiance was varied, the enhancementof photosynthesis in elevated CO₂ at 23 °C increased graduallyfrom approx 10 % at 100 µmol m^–1 s^–1 to >27 % at 1170 µmol m^–2 s^–1 In parallel, wateruse efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 In parallel,water use efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 to approx100 % at the highest irradiance Elevated CO₂, and temperatureincreased whole-plant photosynthesis by > 40 %, when expressedin terms of shoot surface area or shoot weight No effects ofelevated CO₂ and temperature on rate of tissue respiration,either during growth or measurement, were established for singleleaves or for whole plants Dependence on N₂, fixation in rootnodules appeared to have no detrimental effect on photosyntheticperformance in elevated CO₂, and temperature Trifolium repens, white clover, photosynthesis, respiration, elevated CO₂, elevated temperature, water use efficiency, N₂ fixation