Effects of Water and Nutrient Availability on Water Relations, Gas Exchange and Growth Rate of Mature Plants and Resprouts of Arbutus unedo L.

This study examines the effects of water supply and nutritionon the water status, gas exchange and growth of mature plantsand resprouts of Arbutus unedo, a Mediterranean evergreen shrubadapted to drought and poor nutrition. Mature plants of A. unedorespond to irrigation with increased leaf water potential duringsummer drought, but they show a very conservative use of waterand they do not increase leaf conductance. There is also a verysmall increase in net photosynthesis and growth, which doesnot significantly increase productivity. Resprouts of A. unedo increase water potential, leaf conductance,transpiration rate, net photosynthesis and growth rate in responseto watering, showing a less conservative use of water than matureplants. Increased growth rates, both in mature plants and resprouts,are likely to be due to the higher cell turgor caused by improvedleaf water potential, rather than to increased photosynthesis. The only effect of nutrient addition on mature plants is anincrease in leaf nutrient content, and other aspects of thephysiology and growth of resprouts were unaffected. We thereforeconclude that water is a more limiting factor than nutrientsfor mature plants and resprouts of A. unedo growing in the studyarea. These results support previous data which indicate thathigher growth rates in resprouts than in mature plants of A.unedo are mainly the result of a higher water availability.Copyright1994, 1999 Academic Press Arbutus unedo L., strawberry tree, resprouts, water stress, nutrient availability, water relations, gas exchange, growth rate, regeneration     

Growth and Survival Responses of Rumex Species to Flooded and Submerged Conditions: The Importance of Shoot Elongation, Underwater Photosynthesis and Reserve Carbohydrates

Plants of Rumex thyrsiflorus Fingerh., R. crispus L. and R.maritimus L., which are zoned along a gradient of elevationin a river foreland ecosystem, and differ in their flood-tolerance,were subjected to different flooding levels. Under conditionsof soil flooding, the growth rates of the flood-tolerant R.crispus and R. maritimus were as high as under drained conditions,but that of the flood-intolerant R. thyrsiflorus was halved.Upon submergence, the low elevation species R. maritimus showedrapid shoot elongation; when elongation resulted in a protrusionof leaves above the water surface, the plants survived. Alternatively,underwater photosynthesis also led to a 100% survival of submergedR. maritimus plants, provided that enough inorganic carbon wasmade available in the water. This could be attributed in partto the use of photosynthetically-derived oxygen for root respiration;in a hydroculture experiment, with 5.0 mM CO₂ in the water inthe shoot environment, photosynthetically-derived oxygen contributedmore than 50% to root oxygen consumption at low oxygen concentrationsin the root environment. The intermediately elevated species R. crispus appeared to bemuch more tolerant towards conditions of prolonged total submergence:older plants survived eight weeks submergence in the dark. Thisresponse was explicable in terms of a dormancy-strategy, whichis characterized by a slow consumption of carbohydrates storedin the tap-root. The differential responses of R. maritimusand R. crispus to total submergence reveal the limitations offlood-tolerance and reflect the different life-histories ofthe species. Key words: Photosynthesis, Rumex, submergence, carbohydrates, growth rate, shoot elongation     

The Combined Effects of Desiccation and Irradiance on Mosses from Xeric and Hydric Habitats

The combined effects of desiccation and irradiance on the physiologyof the sand dune moss Tortula ruraliformis (Besch.) Grout andthe minerotrophic flush moss Dicranella palustris (Dicks.) Crundw.ex. E. F. Warb (D. squarrosa (Starke) Schp.) were studied. Damageas a result of desiccation in the dark, measured by loss ofprotein and the relative accumulation of thiobarbituric acid(TBA) reactive products (which gives an estimation of lipidperoxidation), was greater in D. palustris. Desiccation alonehad no effect on the total concentrations of chlorophyll andcarotenoids in either species. Water loss resulted in the cessationof measurable photosynthetic oxygen evolution in both species.Respiration was less sensitive to desiccation than was photosynthesis.A combination of irradiance and water stress prevented any recoveryof photosynthesis during subsequent rehydration in D. palustris,but suppressed recovery only marginally (at the highest irradiance)in T. ruraliformis. The loss of protein, chlorophyll, and carotenoids,and lipid peroxidation were all substantially increased in D.palustris desiccated in the light, but these same conditionsresulted in only minimal damage of T. ruraliformis. Continuousexposure to high irradiance was less deleterious to desiccatedthan hydrated T. ruraliformis. The data are discussed in relationto the habitat preferences of the two species, and also in relationto possible causal factors in the initiation of damage. Key words: Desiccation, mosses, oxidative damage, photo-oxidation     

External and Internal CO2 Transport in Lemanea: Interactions with the Kinetics of Ribulose Bisphosphate Carboxylase

The rate of photosynthesis by the freshwater alga Lemanea mamillosais proportional to CO₂ concentration, virtually to the pointof saturation, and inversely proportional to the radius of thethallus. By contrast, the CO₂ response curve of very thin slicesof the thallus is a rectangular hyperbola with a (lower) halfsaturation concentration of 10 mmol m^–3. For the intactplant, the kinetics of CO₂ fixation are strongly masked by internalCO₂ transport limitations, although the maximum rate of photosynthesisis probably determined by the rate of supply of ribulose bisphosphate(RuBP). The flow of water over the alga becomes turbulent atwater velocities greater than about 90 mm s^–1 and thethallus stretches significantly at higher water velocities.In its natural habitat, therefore, the external unstirred layerwill be thin (< 10 µm) and the thallus will be stretched,leading to rapid external and increased internal rates of CO₂transport from the bulk solution. The estimated maximum rateof CO₂ transport is commensurate with the maximum rate of photosynthesis(i.e. the rate of supply of RuBP). Key words: Transport limitations, Kinetics of CO₂ fixation     

The Influence of Water Stress on the Photosynthesis of a Simulated Sward of Perennial Ryegrass

The rate of canopy photosynthesis, single leaf photosynthesis,leaf resistance to gaseous exchange, and leaf water potentialof simulated swards of perennial ryegrass (Lolium perenne cv.S24) in a controlled environment, were determined during a periodof increasing water stress and recovery from that stress. Canopyphotosynthesis did not decline immediately water was withheldbut continued at an undiminished rate for several days; thereafterit fell rapidly, particularly at first. As water stress increasedsuccessive relationships between canopy photosynthesis and irradiancebecame more curved, indicating that the effect of water stressincreased with increasing irradiance. After the swards werere-watered canopy photosynthesis rose, most rapidly during thefirst 24 h. In general, the pattern of change of leaf waterpotential was similar to that of canopy photosynthesis, althougha more detailed examination of this relationship showed it tobe hysteresial; in particular, the fall in leaf water potentialpreceded that of canopy photosynthesis. Single leaf photosynthesisappeared to be the main agent through which water stress influencedcanopy photosynthesis although in the more severely stressedswards (leaf water potentials of about—15 bars) some leaftissue died and so limited the recovery of canopy photosynthesis.The leaf resistance to gaseous diffusion increased with increasingwater stress, as did the CO₂ compensation point, thereby influencingsingle-leaf photosynthesis and through it canopy photosynthesis.     

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)     

The Effect of Soil Water and Aeration on Seed Germination

The time rate of germination and the final germination percentageof Oryzopsis holciformis decreased with increasing water stress.The optimum matric potential for germination was–0.005bar in coarse sand and –0.5 bar in sandy loam soil. Thisdiscrepancy was explained by changes in the rate of water-supplyto the seed, as determined by the area of contact between seedand germination medium, and by the hydraulic conductivity ofthe medium. At high soil moisture potentials germination also decreased.Such a decrease was not found at equivalent osmotic potentials.It seems that this decrease in germination was brought aboutby the thickening of the water films around the seeds, whichinterfered with oxygen diffusion. This assumption was supportedby determinations with Pt electrodes, and by previous work ongermination at lowered oxygen concentrations.     

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     

Autunmal sedimentation of Microcystis spp. as result of an increase in carbohydrate ballast at reduced temperature

Autumnal sedimentation of the Microcystis population was studiedin Lake Nieuwe Meer (Amsterdam, The Netherlands). In summer,Microcystis formed a high percentage of the total phytoplanktonin the water column, but a low percentage in sedimentation traps.The reverse was found during September and October, with a highpercentage in the sedimentation traps, but a low percentagein the water column. The decrease in the numbers of Microcystiscolonies coincided with a decrease in waler temperature. Inexperiment with a strain of Mirocystis, isolated from Lake NieuweMeer, the percentage of total colonies that were sinking increasedin a few days to 100% after a shift from 20C to 15.3, 13.0or 10.5C. The gas vesicle volume in the cells remained constantduring the incubations. Sinking of the colonies resulted froman increased glycogen content. Calculation of carbon (C) flowsduring the first 2 days of the incubation at reduced temperatureshowed that the glycogen accumulation was the result of a muchlower rate of protein synthesis during the light period at thelower temperatures. Although the photosynthetic rate itselfdecreased at reduced temperature, it resulted in more fixedCO₂ being stored as glucose. Because the respiratory rate alsodecreased (with an almost similar decrease to that of photosynthesis),glycogen accumulated at lower temperatures. It was calculatedthat after an incubation period of-I week at reduced temperature,the rate of photosynthesis had decreased by 10.1% of the valueat 20C per 1C, while the rate of respiration had decreasedonly 1.8%. It is proposed that there is a feedback mechanismin which an increasing concentration of glycogen inhibits photosynthesisand stimulates respiration.     

14C Fixation and Translocation in two Clones of Sugarcane with Contrasting Rates of Sucrose Uptake in vitro

The rates of ¹⁴CO₂ fixation and translocation of ¹⁴C labelledassimilates were measured in field experiments at two timesof the day in two sugar-cane clones known to have differentrates of sucrose uptake in vitro but the same weight of leafper unit weight of cane. The rate of ¹⁴CO₂ fixation and the velocity and rate of translocationwere significantly greater at both times in the clone with thehigher rate of sucrose uptake in vitro. The velocities of translocationwere 2.18 and 2.36 cm/min^–1 for the clone with high sucroseuptake and 1.46 cm min^–1 at both times in the clone withlow uptake. It is suggested that among sugar-cane clones the ability oftheir canes to store sugar may play a part in determining theirrates of photosynthesis and translocation.     

The Effect of Temperature on Photosynthesis of Ryegrass and White Clover Leaves

The rate of photosynthesis of leaves of perennial ryegrass (Loliumperenne L.) and white clover (Trifollum pratense L.) grown atdifferent temperatures was measured at a range of temperatures.There was a small effect of the temperature at which a leafhad grown on its photosynthetic rate, but a large effect ofmeasurement temperature, especially in bright light, where photosyntheticrates at 15°C were about twice those at 5°C. It appearsthat temperature could affect sward photosynthesis in the field.Ryegrass and clover had similar photosynthetic rates which respondedsimilarly to temperature. Lolium perenne L., ryegrass, Trifolium pratense L., white clover, photosynthesis, temperature, irradiance     

Inhibition of photosynthesis in olive trees (Olea europaea L.) during water stress and rewatering

The effect of high levels of natural light on leaf photosynthesisin olive trees (Olea europaea L. var. Coratina), grown in potsoutdoors in the summer and subjected to water, stress, was studied.Net photosynthetic rates reached maximum values early in themorning in both control and stressed plants and subsequentlydeclined gradually. This inactivation of photosynthetic activitywas accompanied by changes in the fluorescence characteristicsof the upper intact leaf surface. The maximum fluorescence yield(Fp) and the ratio Fv/Fp decreased at midday especially in water-stressedplants, but the initial fluorescence (Fo) rose to a maximumvalue at midday and declined again in the afternoon. In controlplants the values of maximum fluorescence Fp and the ratio Fv/Fpincreased again in the afternoon and had recovered almost completelyby 8 p.m. as the leaf water potential recovered. In stressedplants this diurnal recovery was not complete, so that the photosyntheticrates and the ratio Fv/Fp declined gradually during the developmentof water stress. These results indicate that in olive treessubjected to severe water stress the non-stomatal componentof photosynthesis was affected and perhaps a light-dependentinactivation of the primary photochemistry associated with photosystemII (PSII) occurred. Four to five days after rewatering severelystressed plants, the predawn leaf water potential, net photosyntheticrates and chlorophyll fluorescence indices recovered only partially. Key words: Olea europaea, photosynthesis, water stress, chlorophyll a fluorescence, inhibition of photosynthesis     

Glycolate excretion by Anacystis nidulans: effect of HCO3- concentration, oxygen concentration and light intensity

Anacystis nidulans exhibited a light-dependent glycolate excretionwhich occurred at a maximum rate in the absence of added HCO₃^–and in the presence of high oxygen concentration. At high lightintensity excretion was linear from the beginning of the illuminationperiod. Low light intensity simply delayed the onset of excretionbut did not affect the rate of excretion once it began. Glycolateexcretion was sensitive to photosynthesis inhibitors but wasinsensitive to isonicotinyl hydrazide. (Received September 19, 1972; )     

Influence of Temperature and O2 Concentration on Photosynthesis and Light Activation of Ribulosebisphosphate Carboxylase Oxygenase in Intact Leaves of White Clover (Trifolium repens L.)

Detached leaves of white clover (Trifolium repens L.) were keptfor 1 h under various conditions of temperature, oxygen concentrationand light intensity. Rates of photosynthesis were measured whereappropriate and then ribulosebisphosphate carboxylase oxygenase(RuBPCO) was extracted rapidly and its initial activity measuredimmediately. The extracted activity increased with increased intensity ofillumination of the leaves. Where leaves were pretreated atlow light intensity, the lower the temperature of the leavesthe higher the extracted activity of RuBPCO. At high light intensitytemperature did not affect the activity of subsequently extractedRuBPCO but the light intensity which was necessary for maximumactivity increased with temperature. Activity of RuBPCO fromleaves pretreated in the dark was least when CO₂ was low andtemperature high. Leaves, pretreated at low temperatures andhigh light intensity in 20% O₂, yielded higher activity in extractsthan leaves pretreated under similar conditions but in 2% O₂.A relatively weak temperature response of photosynthesis atlow irradiances was associated with a decrease in extractableRuBPCO activity with increasing temperature. A strong temperaturedependence of the oxygen inhibition of photosynthesis was associatedwith lower extractable RuBPCO activity in leaves pretreatedat low oxygen concentration at low temperatures. With leavesfrom plants grown at low temperatures prior to treatment ofleaves, oxygen inhibition of photosynthesis was less temperaturedependent and activity of RuBPCO in extracts was not decreasedby low O₂ at low temperatures. Differences in the activationof RuBPCO appear to influence photosynthesis and account foran absence of oxygen inhibition of photosynthesis at low temperaturesin plants grown in warm conditions. Key words: Ribulosebisphosphate carboxylase oxygenase activation, Photosynthesis, Temperature, O₂ effect, White clover     

Gas Exchange and Water Relations of the Root Hemi-parasiteSantalum albumL. in Association with Legume and Non-legume Hosts

This paper examines foliar nitrogen (N) levels, photosynthesis,transpiration, water use efficiency and tissue water relationsof the xylem-tapping root hemi-parasiteSantalum albumin potculture with various N₂-fixing woody hosts, a non-fixing host(a eucalypt), or in the absence of a host. Foliar N concentrationsofSantalumwere significantly greater than corresponding hostsand higher when on N₂-fixing hosts than on the eucalypt, orwithout a host. Strong positive relationships were evident inSantalumbetweenfoliar N concentration, rates of net photosynthesis and instantaneouswater use efficiencies. Photosynthesis rate and water use efficiencyofSantalumwere generally lower than in corresponding hosts,but transpiration rates were not noticeably different betweenassociations.     

Nitrate Nutrition and Temperature Effects on Wheat: Photosynthesis and Photorespiration of Leaves

Lawlor, D. W., Boyle, F. A., Young, A. T., Keys, A. J. and Kendall,A. C. 1987. Nitrate nutrition and temperature effects on wheat:photosynthesis and photorespiration of leaves.—J. exp.Bot. 38: 393–408. Photosynthetic and photorespiratory carbon dioxide exchangeby the third leaf of spring wheat (Triticum aestivum cv. Kolibri),was analysed for plants grown at 13/10 °C (day/night temperature)and 23/18 °C with two rates of nitrate fertilization (abasal rate, — N, and a 4-fold larger rate, +N) and, insome experiments, with two photon fluxes. Net photosynthesiswas greatest at the time of maximum lamina expansion, and forleaves grown with additional nitrate. Maximum rate of photosynthesis,carboxylation efficiency and photochemical efficiency at maturitywere slightly decreased by nitrate deficiency but photosystemactivity was similar under all conditions. As leaves aged, photosynthesisand photochemical efficiency decreased; carboxylation efficiencydecreased more than photochemical efficiency particularly withbasal nitrate. Low oxygen increased the carboxylation and photochemicalefficiencies, and increased the maximum rate of assimilationby a constant proportion in all treatments. Photorespiration,measured by CO₂ efflux to CO₂-free air, by ¹⁴CO₂ uptake, andfrom compensation concentration, was proportional to assimilationin all treatments. It was greater, and formed a larger proportionof net photosynthesis, when measured in warm than in cold conditionsbut was independent of growth conditions. Assimilation was relatedto RuBPc-o activity in the tissue. Relationships between photosynthesis,photorespiration and enzyme complement are discussed. Key words: Wheat, leaves, nitrate nutrition, temperature effect, photosynthesis, photorespiration     

Effects of mute swan grazing on a keystone macrophyte

1. This study describes the early summer foraging behaviour of mute swans (Cygnus olor) on the River Frome, a highly productive chalk stream in southern England in which Ranunculus penicillatus pseudofluitans is the dominant macrophyte. 2. A daily maximum of 41 ± 2.5 swans were present along the 1.1 km study reach during the study period (late May to the end of June). The river was the primary feeding habitat. Feeding activity on the river at dawn and dusk was much lower than during daylight, but we cannot rule out the possibility that swans fed during the hours of darkness. 3. The effects of herbivory on R. pseudofluitans biomass and morphology were quantified. Biomass was lower in grazed areas and swans grazed selectively on leaves in preference to stems. A lower proportion of stems from grazed areas possessed intact stem apices and flowering of the plant was reduced in grazed areas. 4. A model, based on the swans’ daily consumption, was used to predict the grazing pressure of swans on R. pseudofluitans. The model accurately predicted the number of bird days supported by the study site, only if grazing was assumed to severely reduce R. pseudofluitans growth. The proportion of the initial R. pseudofluitans biomass consumed by a fixed number of swans was predicted to be greater when the habitat area was smaller, initial R. pseudofluitans biomass was lower and R. pseudofluitans was of lower food value. 5. We concluded that the flux of N and P through the study reach was largely unaffected by swan activity. The quality of R. pseudofluitans mesohabitat (the plant as habitat for invertebrates and fish) was significantly reduced by grazing which also indirectly contributed to reduced roughness (Manning's n) and by inference water depth. Wetted habitat area for fish and invertebrates would also be lowered over the summer period as a consequence of the reduction in water depth. It was estimated that, while grazing, an individual swan may eat the same mass of invertebrates per day as a 300‐g trout. 6. There is a need to manage the conflict between mute swans and the keystone macrophyte, R. pseudofluitans, in chalk streams, and the modelling approach used here offers a potentially useful tool for this purpose.     

Effect of Oxygen on Photosynthetic 14CO2 Fixation in Chroomonas sp. (Cryptophyta) III: Effect of Oxygen on Photosynthetic Carbon Metabolism

Photosynthetic carbon metabolism was studied with Chroomonassp. cells in which the rate of photosynthesis was inhibitedunder both an anaerobic condition and high concentrations ofoxygen. The time course of ¹⁴C-incorporation into photosyntheticproducts showed that 3-phosphoglycerate was the initial productof photosynthetic CO₂ fixation in Chroomonas sp. cells. During5-min photosynthesis, a considerable amount of ¹⁴C was incorporatedinto the insoluble fraction (mostly cryptomonad starch), andoxygen predominantly affected ¹⁴C-incorporation into this fraction.Although ¹⁴C-incorporation into intermediates of the photorespiratorypathway increased with increasing O₂ concentration, the amountswere much less than expected from the degree of oxygen inhibition.It is noteworthy that ¹⁴C-dihydroxyacetone phosphate accumulatedduring photosynthesis only under the anaerobic condition, whereasthe levels of the other phosphate esters were scarcely affectedby the oxygen concentration. Ribulose-1,5-bisphosphate carboxylase from Chroomonas sp. wascompetitively inhibited by oxygen, and its K_m(CO₂) value wassimilar to those of terrestrial C₃ plant enzymes. (Received November 19, 1984; Accepted May 20, 1985)     

Effect of elevated CO2 and nutrient status on growth, dry matter partitioning and nutrient content of Poa alpina var. vivipara L.

Poa alpina var. vivipara L. was grown in an atmosphere containingeither 340 or 680 µmol CO₂ mol^–1 within controlledenvironment chambers. The available nutrient regime was variedby altering the supply of nitrogen and phosphorus within a completenutrient solution. At a high, but not low, N and P supply regime,elevated CO₂ markedly increased growth. Differences betweennutrient supply, but not atmospheric CO₂ concentration, alteredthe allometric relations between root and shoot. Net photosynthesisof mature leaf blades and leaf N and P concentration were reducedin plants grown at the elevated CO₂ concentration. The question was asked: is it possible to ascribe all of theseeffects to elevated CO₂ or are some due to nutrient deficiencycaused by dilution with excess carbon? Several criteria, includingthe nutrient content of sink tissue, root:shoot allometry andthe use of divalent cations to estimate integrated water flowsare suggested in order to make this distinction. It is concludedthat only at a low supply of N and P₁ and elevated CO₂ concentration,was low leaf N concentration due to induced nutrient deficiency.The data are consistent with a model where the capacity of sinksto use photosynthetically assimilated carbon sets both the rateof import into those sinks (and thus rate of export from sourceleaves) and the rate of photosynthesis of source leaves themselves. Key words: Poa alpina L., growth, photosynthesis, carbohydrate, export, nitrogen, phosphorus     

The Effect of Nitrogen Fertilization and Irrigation Frequency on Photosynthesis of Potatoes (Solanum tuberosum)

Gas exchange parameters for the potato (Solanum tuberosum L.,cv. ‘Desiree’) crop were measured under field conditions.Nitrogen nutrition was found to have a positive effect on photosynthesisthroughout the growing season. The gas exchange data and thechlorophyll content in the leaves indicated that this effectwas essentially in the mesophyll. Stomatal resistance increasedas photosynthetic rate decreased, but substomatal CO₂ concentration(Ci) was not affected by nitrogen treatments. Long term effects of water deficit on photosynthetic capacitywere also mostly mesophyllic in origin. A reduction in photosyntheticrate was observed after a period of moisture stress and alsoafter relief of drought by irrigation, indicating a limitedcapacity for recovery. A short term afternoon increase in stomatalresistance, associated with lower Ci values, occurred afterlong periods without irrigation. Leaf water potential was affectedby irrigation frequency, but less strongly than the gas exchangeparameters. Mesophyll activity and stomatal behaviour were bothaffected by water stress and by nitrogen deficiency, whereasCi values were not affected in most cases. Solanum tuberosum L, photosynthesis, mesophyll, stomata, irrigation frequency, nitrogen deficiency, chlorophyll, leaf water potential