Responses of Soybean Leaf Angle, Photosynthesis and Stomatal Conductance to Leaf and Soil Water Potential

The hypothesis that soil water potential (_s) is better correlatedto heliotropic leaf orientation, photosaturated photosyntheticCO₂ assimilation and stomatal conductance during periods oflimited water availability than is bulk leaf water potential(₁) was examined in greenhouse-grown soybean (Glycine max) plants,submitted to a progressive drought. Paired plants were exposedto either 1000 or 100 µmol m^–2 s^–1 photonflux densities (PFD) for 45–60 mins. The higher irradianceinduced short-term decreases in ₁, due to increased transpiration,while _l in the plant exposed to low PFD did not decrease. Thesechanges in ₁ occurred independently of changes in soil waterstatus. Concurrent to the light treatments, a single attachedleaf from each of the two plants was isolated from the restof the plant by shading, and the pulvinus of its terminal leafletwas exposed to a perpendicular PFD of 500 µmol m–²S^–1. Leaf movement of this leaflet was recorded in responseto this light, until a stable leaflet angle was achieved. Valuesof _s and _l (before and after light treatment), and photosaturatedrates of photosynthesis and stomatal conductance, were thenmeasured on these leaves. Leaflet angle and gas exchange werebetter correlated with _s (r² = 0.50, 0.50 and 0.57 for angle,photosynthesis and conductance, respectively) than with _l especiallywhen _l was the result of short-term, high-light induced changesin leaf water status (r² = 0.36, 0.32 and 0.49, for the sameparameters). Leaflet angle was also correlated with stomatalconductance (r² = 0.61) and photosynthetic rate (r² = 0.60),suggesting a close association between leaf orientation, leafmetabolism and soil water availability. Glycine max (L.) Merr. cv. Essex, soybean, heliotropism, water potential, photosynthesis, stomatal conductance, solar tracking     

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     

Characterization of the Type of Photosynthetic Carbon Dioxide Fixation in Jute (Corchorus olitorius L.)

Activities of photosynthetic and photorespiratory enzymes viz.,ribulose bisphosphate carboxylase, phosphoenol pyruvate carboxylaseand glycolate oxidase from jute (Corchorus olitorius L.; cv.JRO 632) leaves were compared with those from maize (C₄) andsunflower (C₃) leaves. The photosynthetic CO₂ fixation products,the release of ¹⁴CO₂ in light and dark following photosynthesisin ¹⁴CO₂, chlorophyll a: b ratio, gross leaf photosyntheticrate and dry matter production rate were also studied. The resultsshow that jute is a C₃ plant. Key words: Jute, Corchorus olitorius, C₃ photosynthesis     

Acclimation in Seedlings of a Tropical Tree, Bischofia javanica, Following a Stepwise Reduction in Light

Acclimation of fully developed leaves of Bischofia javanicaBlume to shadelight was examined. Seedlings were grown undersimulated daylight (1000 µmol m^–2 s^–1), thentransferred to a simulated shadelight (40 µmol m^–2s^–1). When a high-light leaf was transferred to low light, large negativenet photosynthetic rates (P_m) were recorded. This decrease wasrapid, but within 7 d the rate increased and became equal tothe low-light control leaf. These changes in photosynthesisdid not follow the pattern of changes in stomatal conductance(g_s). Transfer to the low light resulted in a dramatic decreasein leaf weight per unit area (L_w), and most of the decreasesin L_w occurred within 3 d of transfer when the P_m of the transferredleaf was well below that of the low-light control leaf. There was a significant decrease in chlorophyll a in the transferredleaf without an appreciable change in chlorophyll b resultingin a large decrease in the chlorophyll a to chlorophyll b ratio.Leaf chlorophylls per unit area were higher in the transferredleaf than the low-light control leaf. Maximum photosyntheticrate in the transferred leaf was decreased by 40% compared tothat for the high-control leaf, but was almost at the same extenthigher than the low-light control leaf The results are discussedin the context of carbon gain capacity of its seedlings underlight-limiting forest understorey habitats. Bischofia, chlorophylls, light, photosynthesis, shade acclimation, tree seedlings, tropical tree     

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     

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     

Temperature Response of Early Foliar Expansion of Potato and Wheat

We studied the course of early leaf area expansion and specificleaf area (S_LA) in potato (Solanum tuberosum L.) and wheat (Triticumaestivum L.) genotypes and tested whether air temperature explainsdifferences in these courses within different environments.Such knowledge can be used to improve crop growth modelling.The relative rate of leaf area expansion (R_L) of potato andwheat decreased with thermal time, but was nearly linear upto a leaf area index (L) of 1.0. TheR_L (L < 1; mean: 17.9x 10^-3°C^-1 d^-1) of potato showed an interaction betweengenotype and environment, and varied with year. TheR_L (L <1; mean: 7.1 x 10^-3°C^-1 d^-1) of winter wheat was lower thanthat of spring wheat (mean: 10.9 x 10^-3°C^-1 d^-1), and bothvaried with year. S_LAof potato increased nearly linearly withthermal time from 5 to 15 m² kg^-1at 50% emergence, to 20 to25 m² kg^-1at 155°Cd, and then decreased slightly. The S_LAofboth winter and spring wheat began at 16 to 23 m² kg^-1and inmost cases increased slightly with thermal time. In potato,regression parameters of S_LAwith thermal time were affectedby environment (management conditions and year) and genotype;in wheat they were affected by environment (year and site).Treatment effects on R_Lof potato were not correlated with thoseon S_LA , and were only partly correlated for wheat. Thereforewe conclude that the early foliar expansion of potato is associatedwith a strong increase in S_LA , and not so for wheat. For bothcrops the course of early leaf area expansion and ofS_LA withair temperature is not robust over environments and genotypes.The consequences of these results for modelling are discussed.Copyright 2000 Annals of Botany Company Triticum aestivum, spring wheat, winter wheat, Solanum tuberosum, leaf area expansion, specific leaf area, early growth, genotype, environment, modelling     

Effect of Nitrogen Fertilizer on Photosynthesis of Several Varieties of Winter Wheat

The rates of gross photosynthesis of the flag leaf and the nextleaf below (second leaf) in crops of winter wheat were estimatedfrom the ¹⁴C uptake of the leaves after exposure to short pulsesof ¹⁴CO₂. The photosynthetic rates of both leaves during thegrain-filling period decreased with increase in nitrogen fertilizerbecause the intensity of photosynthetically active radiationwas less at the surface of the leaves in the dense crops withadditional nitrogen. In addition, the rate of photosynthesisat saturating light intensity was slightly decreased by nitrogen.The effects of nitrogen, in decreasing the rate of photosynthesisper unit area of leaf and in increasing the leaf-area indexof the top two leaves, were such that the photosynthetic productivityper unit area of land of the flag leaf was increased by nitrogenbut the productivity of the second leaf was unaffected. Applying180 kg N ha^–1 increased the productivity of the top twoleaves by a factor of 2.3 but increased grain yield by only1.8. The photosynthetic productivity of the second leaf duringthe grain-filling period was about half that of the flag leaf. There was no difference in photosynthetic rate per unit areaof leaves of Cappelle-Desprez and Maris Huntsman which couldaccount for the larger yield of the latter cultivar. There wasa slight indication that the leaves of the semi-dwarf cultivarsMaris Fundin and Hobbit photosynthesized faster than those ofMaris Huntsman. Triticum aestivum L., winter wheat, photosynthesis, nitrogen fertilizer     

The Greenhouse Effect: Acclimation of Tomato Plants Growing in High CO2, Photosynthesis and Ribulose-1, 5-Bisphosphate Carboxylase Protein

Tomato plants were grown in solution culture in a controlledenvironment at 20 ?C with a 12 h photoperiod of 400 µmolquanta m^–2 s^–1 PAR with either normal ambient CO₂,approximately 340 vpm, or with 1000 vpm CO₂. The short- andlong-term effects of CO₂ enrichment on photosynthesis were determinedtogether with the levels of ribulose-1, 5-bisphosphate carboxylase(RuBPco) E.C. 4.1.1.39 [EC] protein and activity throughout leafdevelopment of the unshaded 5th leaf above the cotyledons. Thehigh CO₂ concentration during growth did not appreciably affectthe rate of leaf expansion or final leaf area but did increasethe fresh weight per unit area of leaf. With short-term CO₂enrichment, i.e. only during the photosynthesis measurements,the light-saturated photosynthetic rate (P_max) of young leavesdid not increase while those reaching full expansion more thandoubled their net rate of CO₂ fixation. However, with longerterm CO₂ enrichment, i.e. growing the crop in high CO₂, theplants did not maintain this photosynthetic gain. While theCO₂ concentration during growth did not affect the peak in P_maxmeasured in 300 vpm CO₂ or P_max in 1000 vpm CO₂, RuBPco proteinor its activity, the subsequent ontogenetic decline in theseparameters was greatly accelerated by the high CO₂ treatment.Compared with plants grown in normal ambient CO₂ the high CO₂grown leaves, when almost fully expanded, contained only approximatelyhalf as much RuBPco protein and P_max in 300 vpm CO₂ and P_maxin1000 vpm CO₂ were similarly reduced. The loss of RuBPco proteinmay be a major factor associated with the accelerated fall inP_max since it was close to that predicted from the amount andkinetics of RuBPco assuming RuBP saturation. In the oldest leavesexamined grown in high CO₂ additional factors may be limitingphotosynthesis since RuBPco kinetics marginally overestimatedP_max in 300 vpm CO₂ and the initial slope of photosynthesisin response to intercellular CO₂ was also less than expectedfrom the extractable RuBPco. Key words: Lycopersicon esculentum (Mill.) cv. Findon Cross, CO₂ enrichment, acclimation to high CO₂, photosynthesis, RuBPco protein and activity     

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

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

Characterisation of Non-Uniform Photosynthesis Induced by Abscisic Acid in Leaves Having Different Mesophyll Anatomies

The effects of abscisic acid (ABA) on photosynthesis in leavesof Helianthus annuus L. were compared with those in leaves ofVicia faba L. After the ABA treatment, the response of photosyntheticCO₂ assimilation rate, A, to calculated intercellular partialpressure of CO₂, P_i, (A(p_i) relationship) was markedly depressedin H. annuus. A less marked depression was also observed inV.faba. However, when the abaxial epidermes were removed fromthese leaves, neither the maximum rate nor the CO₂ responseof photosynthetic oxygen evolution was affected by the applicationof ABA. Starch-iodine tests revealed that photosynthesis was not uniformover the leaves of H. annuus treated with ABA. The starch contentwas diffferent in each bundle sheath extension compartment (thesmallest subdivision of mesophyll by veins with bundle sheathextensions, having an area of ca. 0.25 mm² and ca. 50 stomata).In some compartments, no starch was detected. The distributionof open stomata, examined using the silicone rubber impressiontechniques, was similar to the pattern of starch accumulation.In V.faba leaves, which lack bundle sheath extensions, distributionof starch was more homogeneous. These results indicate that the apparent non-stomatal inhibitionof photosynthesis by ABA deduced from the depression of A(pⁱ)relationship is an artifact which can be attributed to the non-uniformdistribution of transpiration and photosynthesis over the leaf.Intercellular gaseous environment in the ABA-treated leavesis discussed in relation to mesophyll anatomy. ¹ Present address: Department of Botany, Duke University, Durham,NC 27706, U.S.A. (Received September 30, 1987; Accepted January 13, 1988)     

Afternoon Depression In Photosynthesis in Grapevine Leaves--Evidence for a High Light Stress Effect

A midday depression in net photosynthesis and in stomatal conductancewas observed when leaves of well-watered Vitis vinifera plantswere subjected to a diurnal pattern of variation in leaf temperatureand leaf-to-air water vapour pressure difference similar toa summer day, while photon flux density was kept constant at1450 µmolm^–2 s^–1,. When leaves were kept atconstant leaf temperature (22.5°C) and leaf-to-air watervapour presure difference (8.5 Pa kPa^–1) at the same lightintensity, stomata opened with the onset of illumination andmaximal conductance and photosynthesis values were observedabout 1 h later. Subsequently, conductance and photosynthesisdecreased gradually. Leaf water potential never dropped below{macron}0.3 MPa. Leaves kept under constant environmental conditionsshowed an afternoon decline in photosynthesis at high internalCO₂, in carboxylation efficiency and in maximum conductanceas well as an increase in stomatal sensitivity to CO₂. Whenthe photon flux density during the day was reduced to 750 µmolm^–2 s^–1, the afternoon depression in gas exchangerates was attenuated. To evaluate the possible effects of highlight stress on changes in chloroplastic behaviour we comparedlight response curves of photosynthesis determined with an oxygenelectrode, in the morning and in the afternoon after the plantswere exposed to either high or moderate photon flux densities.A significant depression in photosynthetic capacity was foundby this method in high light treated leaves, but not in leavespreviously exposed to moderate photon flux density. Apparentquantum yield decreased in the afternoon, particularly afterexposure to high light. Maximum chlorophyll a fluorescence at22°C was reduced and the quenching of fluorescence afterreaching the peak was slower in the afternoon than in the morning,especially in high light-treated leaves. Changes in the patternsof chlorophyll fluorescence kinetics were observed after lighttreatment, i.e. in the afternoon, with oscillations either absent(after high light) or significantly reduced (after moderatelight) in comparison to the morning. The significance of theseresults is discussed and it is suggested that a direct inhibitoryeffect of high light at the chloroplast level provides the bestinterpretation for the observed afternoon decline in photosyntheticrate. Key words: Carboxylation efficiency, chlorophyll fluorescence, photosynthetic capacity, quantum yield, stomatal conductance, Vitis vinifera L.     

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     

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     

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     

Photosynthesis and Light Activation of Ribulose 1,5-bisphosphate Carboxylase in the Presence of Starch

Limitation of photosynthesis and light activation of ribulose,1,5-bisphosphate carboxylase (RuBPCO) were examined in the 5thleaf of seedlings of red clover (Trifolium pratense L. cv. Renova)for 5 d following an increase in photosynthetic photon fluxdensity (PPFD) from 200 to 550µmol quanta m^–2 s^–1.Net photosynthesis and its stimulation at 2.0 kPa O₂ initialactivity of rapidly extracted RuBPCO, standard activity of RuBPCOafter incubation of the extracts in the presence of CO₂, Mg²⁺,and inorganic phosphate and contents of soluble protein, starch,soluble sugars, and various photosynthetic metabolites weredetermined. Photosynthesis decreased and starch content increased.No decrease in photosynthesis was found if, when PPFD was increased,all leaves except the investigated 5th leaf were removed, suggestingthat the decrease in photosynthesis was due to accumulated carbohydrates.The stimulation of photosynthesis at 2.0 kPa O₂ did not decreaseand the ratio of the total foliar steady-state contents of triosephosphate to 3-phosphoglycerate increased suggesting that thedecrease in photosynthesis was not due to limiting inorganicphosphate in chloroplasts. Intercellular CO₂ partial pressureand RuBP content were not decreased. Nevertheless, the ratioof photosynthesis to initial RuBPCO activity decreased, suggestingthat the catalysis per active RuBPCO site was decreased. Theincrease in PPFD in the growth cabinet and the PPFD at whichleaves were preconditioned for 1 h, affected not only initialactivity but also the standard activity of RuBPCO. The resultssuggest that a varying proportion of RuBPCO was bound to membranesand was contained in the insoluble fraction of the extracts.A comparison of photosynthesis with extracted RuBPCO activitysuggested that membrane bound RuBPCO did not contribute to photosyntheticCO₂ fixation and that the binding and release to and from membranesmodulated actual RuBPCO activity in vivo. Key words: Photosynthesis, ribulose 1,5-bisphosphate carboxylase, starch     

Further Characteristics of Salt-Dependent Bicarbonate Use by the Seagrass Zostera muelleri

Millhouse, J. and Strother, S. 1987. Further characteristicsof salt-dependent bicarbonate use by the seagrass Zostera muelleri.—J.exp. Bot. 38: 1055–1068. The contribution of HCO^–₃to photosynthetic O₂ evolutionin the seagrass Zostera muelleri Irmisch ex Aschers. increasedwith increasing salinity of the bathing seawater when the inorganiccarbon concentration was kept constant. K_1/2 (seawater salts)for HCO^–₃ -dependent photosynthesis was 66% of seawatersalinity. Both short- and long-term pretreatment at low salinitiesstimulated photosynthesis in full strength seawater. Twentyfour hours pre-incubation of seagrass plants in 3·0 molm^–3 NaHCO₃ resulted in increased photosynthesis at allsalinities, apparently due to stimulation of HCO^–₃ use(K_1/2 (seawater salts) = 26%). V_max (HCO^–₃) was not affectedby low salinity pretreatment. The kinetics of HCO^–₃ stimulationby the major seawater cations was investigated. Ca²⁺ was themost effective cation with the highest V_max (HCO^–₃) andwith K_1/2(Ca²⁺) = 14 mol m^–3. Mg²⁺ was also very effectiveat less than 50 mol m^–3 but higher concentrations wereinhibitory. This inhibition cannot be accounted for solely byprecipitation of MgCO₃. Na⁺ and K⁺ were both capable of stimulatingHCO^–₃ use. Stimulation was in two distinct parts. Up to500 mol m^–3, both citrate and chloride salts gave similarresults (K_1/2(Na⁺) 81 mol m^–3, V_max(HCO^–₃) 0·26µmol O₂ mg^–1 chl min^–1), but use of citratesalts above 500 mol m^–2 caused a second stimulation ofHCO^–₃ use (K_1/2(Na⁺) 830 mol m^–3, V_max(HCO^–₃)0·68 µmol O₂ mg^–1 chl min^–1). V_max(HCO^–₃)for the second-phase Na⁺ or K⁺ stimulation was of the same orderas for Ca²⁺-stimulated HCO^–₃ use. To further characterizesalt-dependent HCO^–₃ use, the sensitivity of photosynthesisto Tris and TES buffers was investigated. The effects of Trisappear to be due to the action of Tris⁺ causing stimulationof HCO^–₃ -dependent photosynthesis in the absence of salt,but inhibition of HCO^–₃ use in saline media. TES has noeffect on photosynthesis. External carbonic anhydrase, althoughimplicated in salt-dependent HCO^–₃ use in Z. muelleri,could not be detected in whole leaves. Key words: Zostera muelleri, HCO^–₃ use, salinity     

准噶尔荒漠早春短命植物的光合特性及生物量分配特点

准噶尔荒漠分布的早春短命植物不仅具有十分独特的生物学特点,而且在荒漠植物群落演替、物种多样性维持及土壤改良与防治水土流失等方面具有重要的生态学价值。该文运用Li-6400开放式气体交换光合作用测定系统,对分布于准噶尔荒漠的16种早春短命植物生长盛期的净光合速率(P_n)、蒸腾速率(T_r)、水分利用效率(WUE)等特征进行了测定,并对其中7种植物与生长相关的生物量分配特征进行了分析。结果表明:1)16种植物的最大P_n、 最大T_r及WUE分别为8.07~35.96 μmol CO₂·m^-2·s^-1、3.16~29.64 mmol H₂O·m^-2·s^-1、0.54~4.26 μmol CO₂·mmol^-1H₂O;种间最大P_n与最大气孔导度(Stomatal conductance, G_s)之间存在正相关关系,其相关系数为0.77(p<0.05),线性回归斜率为26.36 μmol·mmol^-1;从光合速率对胞间CO₂浓度及光量子通量密度的响应曲线来看,这类植物的表观CO₂补偿点均在4~5 Pa之间(28~30 ℃),表观羧化效率为0.64~1.86 μmol CO₂·m^-2·s^-1·Pa^-1,表观量子效率为0.05~0.06。2)从生物量分配来看,所测植物的个体生物量为0.05~0.39 g;单株总叶面积为 3.24~51.40 cm²;单位叶面积干重为0.40~0.77 g·m^-2,根在总生物量中所占比例为5.72%~19.43%,单株叶面积比在2.92~9.00 m²·kg^-1之间。种间根所占生物量的比与对应的WUE之间的比较分析结果表明,二者之间存在显著的正相关关系,其相关系数r为0.93(p<0.01)。这些结果表明,所观测的早春短命植物具有典型的C₃植物特征,相比其它类型的荒漠植物具有较高的单位叶面积P_n、高T_r及低WUE,并且在生长发育过程中表现出很低的根/地上生物量比、较高的叶面积比和单位叶面积干重,说明它们具有相对高的生长速率,这与其生长发育节律相一致,反映了它们与准噶尔荒漠环境相适应的特点。     

Stomatal Responses of Variegated Leaves to CO2 Enrichment

The responses of stomatal density and stomatal index of fivespecies of ornamental plants with variegated leaves grown attwo mole fractions of atmospheric CO₂ (350 and 700 µmolmol^-1) were measured. The use of variegated leaves allowed anypotential effects of mesophyll photosynthetic capacity to beuncoupled from the responses of stomatal density to changesin atmospheric CO₂ concentration. There was a decrease in stomataldensity and stomatal index with CO₂ enrichment on both white(unpigmented) and green (pigmented) leaf areas. A similar responseof stomatal density and index was also observed on areas ofleaves with pigmentation other than green indicating that anydifferences in metabolic processes associated with colouredleaves are not influencing the responses of stomatal densityto CO₂ concentrations. Therefore the carboxylation capacityof mesophyll tissue has no direct influence on stomatal densityand index responses as suggested previously (Friend and Woodward1990 Advances in Ecological Research 20: 59-124), instead theresponses were related to leaf structure. The stomatal characteristics(density and index) of homobaric variegated leaves showed agreater sensitivity to CO₂ on green portions, whereas heterobaricleaves showed a greater sensitivity on white areas. These resultsprovide evidence that leaf structure may play an important rolein determining the magnitude of stomatal density and index responsesto CO₂ concentrations.Copyright 1995, 1999 Academic Press Leaf structure, photosynthesis, stomatal conductance, CO₂, stomatal density, stomatal index     

Gas Exchange and Carbohydrate and Nitrogen Concentrations in Leaves of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) at Different Carbon Dioxide Concentrations and Temperatures

Pascopyrum smithii (C₃) andBouteloua gracilis (C₄) are importantforage grasses native to the Colorado shortgrass steppe. Thisstudy investigated photosynthetic responses of these grassesto long-term CO₂enrichment and temperature in relation to leafnonstructural carbohydrate (TNC) and [N]. Glasshouse-grown seedlingswere transferred to growth chambers and grown for 49 d at twoCO₂concentrations (380 and 750 µmol mol^-1) at 20 and 35°C, and two additional temperatures (25 and 30 °C) at750 µmol mol^-1CO₂. Leaf CO₂exchange rate (CER) was measuredat a plant's respective growth temperature and at two CO₂concentrationsof approx. 380 and 700 µmol mol^-1. Long-term CO₂enrichmentstimulated CER in both species, although the response was greaterin the C₃,P. smithii . Doubling the [CO₂] from 380 to 750 µmolmol^-1stimulated CER ofP. smithii slightly more in plants grownand measured at 30 °C compared to plants grown at 20, 25or 35 °C. CO₂-enriched plants sometimes exhibited lowerCER when compared to ambient-grown controls measured at thesame [CO₂], indicating photosynthetic acclimation to CO₂growthregime. InP. smithii , such reductions in CER were associatedwith increases in TNC and specific leaf mass, reductions inleaf [N] and, in one instance, a reduction in leaf conductancecompared to controls. InB. gracilis , photosynthetic acclimationwas observed more often, but significant changes in leaf metabolitelevels from growth at different [CO₂] were generally less evident.Temperatures considered optimal for growth (C₃: 20 °C; C₄:35 °C) sometimes led to CO₂-induced accumulations of TNCin both species, with starch accumulating in the leaves of bothspecies, and fructans accumulating only inP. smithii. Photosynthesisof both species is likely to be enhanced in future CO₂-enrichedand warmer environments, although responses will sometimes beattenuated by acclimation. Acclimation; blue grama (Bouteloua gracilis (H.B.K.) Lag ex Steud.); leaf nitrogen concentration; nonstructural carbohydrates; photosynthesis; western wheatgrass (Pascopyrum smithii (Rydb.) Love)