Sink-Regulation of Photosynthesis in Relation to Temperature in Sunflower and Rape

Stimulation of the rate of photosynthesis at 2·0 kPaO₂ in comparison with 21 kPa O₂ and carbohydrate accumulationover 4h were measured during exposure of sunflower (Helianthusannuus L.) and rape (Brassica napus L.), grown at 30 °Cand 13 °C, to temperatures between 7 °C and 35 °C.The effect of reducing source: sink ratio by shading on theresponse of photosynthetic rate to temperature was also determined.Stimulation of photosynthesis by 2·0 kPa O₂ in comparisonwith 21 kPa O₂ decreased over 4 h at cool temperatures in sunflowerplants grown at 30 °C but not in rape grown at 30 °C.Stimulation did not decrease over 4 h in plants grown at 13CC. Sucrose was the main carbohydrate accumulated over 4 h;its accumulation increased with decreasing temperature. Starchaccumulation either decreased or remained the same with decreasingtemperature. In plants grown at 30 °C more carbohydrateaccumulated between 8 °C and 21 °C in sunflower thanin rape, but more carbohydrate accumulated at 30 °C in rapethan in sunflower. In plants grown at 13 °C much less carbohydrateaccumulated between 13 °C and 23 °C than in plants grownat 30 °C. Photosynthetic rate in plants grown at 30 °Cexposed to between 20 °C and 35 °C over 32 h (14 h light-10h dark-8 h light), declined over 32 h at 20 °C and 25 °Cin sunflower and at 20 °C in rape. This fall over 32 h,especially at 20 °C in sunflower, was significantly reducedby shading the rest of the plant. Shading had little effecton photosynthetic rate above 25 °C. The work confirms thatlow temperature imposes a sink-limitation on photosynthesiswhich occurs at higher temperatures in sunflower than in rape.This limitation may be relieved by decreasing the source:sinkratio. Key words: Sunflower, rape, photosynthesis, carbohydrates, sink demand, temperature     

Regeneration of Ribulose 1,5-bisphosphate and Ribulose 1,5-bisphosphate carboxylase/oxygenase Activity Associated with Lack of Oxygen Inhibition of Photosynthesis at Low Temperature

The nature of the lack of oxygen inhibition of C₃-photosynthesisat low temperature was investigated in white clover (Trifoliumrepens L.). Detached leaves were brought to steady-state photosynthesisin air (34 Pa p(CO²), 21 kPa p(O₂), balance N₂) at temperaturesof 20°C and 8°C, respectively. Net photosynthesis, ribulose1,5-bisphosphate (RuBP) and ATP contents, and ribulose 1,5-bisphosphatecarboxylase/oxygenase (RuBPCO) activities were followed beforeand after changing to 2·0 kPa p(O₂). At 20°C, lowering p(O₂) increased net photosynthesis by37%. This increase corresponded closely with the increase expectedfrom the effect on the kinetic properties of RuBPCO. Conversely,at 8°C net photosynthesis rapidly decreased following adecrease in p(O₂) and then increased again reaching a steady-statelevel which was only 7% higher than at 21 kPa p(O₂). The steady-staterates of RuBP and associated ATP consumption were both estimatedto have decreased. ATP and RuBP contents decreased by 18% and33% respectively, immediately after the change in p(O₂) suggestingthat RuBP regeneration was reduced at low p(O₂) due to reducedphotophosphorylation. Subsequently, RuBP content increased again.Steady-state RuBP content at 2·0 kPa p(O₂) was 24% higherthan at 21 kPa p(O₂). RuBPCO activity decreased by 22%, indicatingcontrol of steady-state RuBP consumption by RuBPCO activity. It is suggested that lack of oxygen inhibition of photosynthesisat low temperature is due to decreased photophosphorylationat low temperature and low p(O₂). This may be due to assimilateaccumulation within the chloroplasts. Decreased photophosphorylationseems to decrease RuBP synthesis and RuBPCO activity, possiblydue to an acidification of the chloroplast stroma. Key words: Oxygen inhibition, photosynthesis, ribulose bisphosphate carboxylase/oxygenase     

Effect of Temperature on Photosynthesis and Photorespiration of Wheat Leaves

Wheat plants were grown in a controlled environment with daytemperatures of 18 ?C and with 500 µ Einsteins m^–28^–1 of photosynthetically active radiation for 16 h. Beforeanthesis and 2 to 3 weeks after, rates of net photosynthesiswere measured for leaves in 2 or 21% O₂ containing 350 vpm CO₂at 13, 18, 23, and 28 ?C and with 500 µEinsteins m^–2s^–1 of photosynthetically active radiation. Also, underthe same conditions of light intensity and temperature, therates of efflux of CO₂ into CO₂-free air were measured and,for mature flag leaves 3 to 4 weeks after anthesis, gross andnet photosynthesis from air containing 320 vpm ¹⁴CO₂ of specificactivity 39?7 nCi µmol^–1. When the O₂ concentration was decreased from 21 to 2% (v/v)the rate of net photosynthesis increased by 32 per cent at thelowest temperature and 54 per cent at the highest temperature.Efflux of CO₂ into CO₂-free air ranged from 38 per cent of netphotosynthesis at 13 ?C to 86 per cent at 28 ?C. Gross photosynthesis,measured by the ¹⁴C assimilated during 40 s, was greater thannet photosynthesis by some 10 per cent at 13 ?C and 17 per centat 28 ?C. These data indicate that photorespiration was relativelygreater at higher temperatures.     

Effects of NO2 and O3 Alone or in Combination on Kidney Bean Plants (Phaseolus vulgaris L.): Products of 13CO2 Assimilation Detected by 13C Nuclear Magnetic Resonance

The effect of exposure of kidney bean primary leaves to NO₂and O₃, alone or in combination, on the fate of ¹³CO₂ assimilatedby photosynthesis was examined by ¹³C-NMR. There were more than70 peaks appearing in the ¹³C-NMR spectra for substances extractedfrom leaves with 80% ethanol. The 16 relatively well resolvedpeaks corresponded to signals from three sugars, two organicacids and four amino acids. These signals were used to estimatepool sizes and ^l3C incorporation. Exposures to NO₂ and O₃ increased the amounts of sucrose andfructose, but not the incorporation of the ¹³C label during10 min photosynthesis from ¹³CO₂. This suggests the presenceof photo-synthetically inactive pools of sucrose and fructose.Amounts of glycine and serine, and ¹³C incorporation into them,were increased by the exposure to the pollutants. The incorporationof ¹³C into alanine was stimulated by exposure to NO₂, but notby exposure to O₃ alone. The present study shows that with only simple procedures ofsample preparations ¹³C-NMR provides information on the productsof photosynthesis in leaves stressed by the two air pollutants. Key words: NO₂, O₃, Phaseolus vulgaris, CO₂ assimilation, ¹³C-NMR     

Extreme Phosphate Deficiency Decreases the In Vivo CO2/O2 Specificity Factor of Ribulose 1,5-Bisphosphate Carboxylase-Oxygenase in Intact Leaves of Sunflower

Sunflower plants were grown under controlled environmental conditionswith either 0 or 10 mol m^–3 phosphate (Pi). From steady-statemeasurements of gas exchange and chlorophyll fluorescence madeon intact leaves, the in vivo CO₂/O₂ specificity factor (invivo K_sp) of ribulose 1,5-Aisphosphate carboxylase-oxygenase(Rubisco) was determined following two methods based on modelsof C₃ photosynthesis by Brooks and Farquhar (1985) and Peterson(1989). The two methods gave in vivo K_sp values for controlsunflower leaves which were similar to published values forhigher plants. Extreme Pi deficiency decreased in vivo K_sp,in sunflower leaves compared to adequate Pi. This suggests thatPi deficiency affected photorespiration less than photosynthesis.The decrease in in vivo K_sp may be due to a real change in theenzyme kinetics favouring oxygenation more than carboxylationor due to an increase in the number of CO₂ molecules releasedper oxygenation; in which case the observed decrease in thein vivo K_sp determined on intact leaves will not agree numericallywith the true K_sp of Rubisco determined in vitro using purifiedenzyme from the same leaf. We discuss the implications of therelatively large photorespiration in Pi-deficient sunflowerleaves with respect to the increased dissipation of photosyntheticelectrons and photorespiratory recycling of Pi in thechloroplaststroma. Although our results on in vivo K_sp suggested a relativelylarger photorespiratory potential in Pi-deficient than controlsunflower leaves, photosynthesis was insensitive to O₂ in Pi-deficientleaves; the possible reasons for this phenomenon are discussed.Under extreme Pi deficiency, O₂ sensitivity of photosynthesisis not a reflection of the in vivo photorespiratory rates. Determinationof in vivo K_sp of Rubisco is a useful approach to study thephotorespiratory potential of intact leaves. Key words: Chlorophyll fluorescence, phosphate deficiency, photorespiration, photosynthesis, PSII quantum yield, Rubisco specificity factor     

The Effect of Cooling on Photosynthesis, Amounts of Carbohydrate and Assimilate Export in Sunflower

Sunflower plants (Helianthus annuus L.) grown at 30°C werecooled to 13°C in the light in atmospheric CO² or low CO²,or in darkness. Photosynthetic rate at 30°C after coolingwhole plants in atmospheric CO² for 12 h during a photoperiodwas significantly lower than at the start of the photoperiodcompared to plants cooled at low CO², those cooled in the darkand those maintained at 30°C. Amounts of sucrose, hexosesand starch in leaves at 13°C increased throughout a 14 hphotoperiod to levels higher than in leaves at 30°C, whereamounts of sucrose and hexoses were stable or falling after4 h. Carbohydrate accumulation at 13°C during this photoperiodwas more than twice that at 30°C. After three photoperiodsand two dark periods at 13°C carbohydrate levels in leaveswere still as high as at the end of the first photoperiod, butless carbohydrate accumulated during the photoperiods than duringthe first photoperiod, and more was partitioned as starch. Amountsof soluble carbohydrate in roots were greater after 14 h at13°C than in roots of plants at 30°C. Loss of ¹⁴C fromleaves at 30°C as a proportion of ¹⁴CO2 fixed by them at30°C, decreased after exposure of plants to 13°C inthe light for 30 min prior to ¹⁴CO²feeding. Results indicatean effect of cold on the transport process that was light-dependent.It is inferred that the reduction in the proportion of ¹⁴C lostfrom leaves after 10 h cooling was due to reduced sink demand,whereas the rise in the proportion of ¹⁴C lost from leaves after24 h reflects reduced photosynthetic rate. The coincidence ofreduced photosynthetic rate with raised carbohydrate levelsin leaves maintained at 30°C throughout, whilst the restof the plant was cooled to 13°C in the light implies feedbackinhibition of photosynthesis. This may reduce the imbalancebetween source and sink in sunflower during the first days oflong-term cooling. Key words: Temperature, carbon export, carbohydrates, photosynthesis, sunflower     

The Effect of Root Temperature on Growth and Uptake of Ammonium and Nitrate by Brassica napus L. cv. Bien venu in Flowing Solution Culture: II. UPTAKE FROM SOLUTIONS CONTAINING NH4NO3

Macduff, J. H., Hopper, M. J. and Wild, A. 1987. The effectof root temperature on growth and uptake of ammonium and nitrateby Brassica napus L. CV. Bien venu in flowing solution culture.II. Uptake from solutions containing NH₄NO₃.—J. exp. Bot.38: 53–66 The effects of root temperature on uptake and assimilation ofNH₄⁺ and NO₃^– by oilseed rape (Brassica napus L. CV. Bienvenu) were examined. Plants were grown for 49 d in flowing nutrientsolution at pH 6?0 with root temperature decrementally reducedfrom 20?C to 5?C; and then exposed to different root temperatures(3, 5, 7, 9, 11, 13, 17 or 25?C) held constant for 14 d. Theair temperature was 20/15?C day/night and nitrogen was suppliedautomatically to maintain 10 mmol m^–3 NH₄NO₃ in solution.Total uptake of nitrogen over 14 d increased threefold between3–13?C but was constant above 13?C. Net uptake of NH₄⁺exceeded that of NO₃^– at all temperatures except 17?C,and represented 47–65% of the total uptake of nitrogen.Unit absorption rates of NH₄⁺ and of 1?5–2?7 for NO₃^–suggested that NO₃^– absorption was more sensitive thanNH₄⁺ absorption to temperature. Rates of absorption were relativelystable at 3?C and 5?C compared with those at 17?C and 25?C whichincreased sharply after 10 d. Tissue concentration of N in theshoot, expressed on a fresh weight basis, was independent ofroot temperature throughout, but doubled between 3–25?Cwhen expressed on a dry weight basis. The apparent proportionof net uptake of NO₃^– that was assimilated was inverselyrelated to root temperature. The results are used to examinethe relation between unit absorption rate adn shoot:root ratioin the context of short and long term responses to change ofroot temperature Key words: Brassica napus, oilseed rape, root temperature, nitrogen uptake     

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     

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     

The Responses of Net Photosysthesis to Light and Temperature in Spartina townsendii(sensu lato), a C4 Species from a Cool Temperate Climate

Carbon dioxide and water vapour exchanges for single attachedleaves of the temperate C₄ grass Spartina townsendii were measuredunder controlled environment conditions in an open gas-exchangesystem. The responses of net photosynthesis, stomatal resistance,and residual resistance to leaf temperature and photon fluxdensity are described. The light and temperature responses ofnet photosynthesis in S. townsendii are compared to informationon these responses in both temperate C₃ grasses and sub-tropicalC₄ grasses. Adaptation of photosynthesis in this C₄ speciesto a cool temperate climate is indicated both by the light andtemperature responses of net photo-synthesis. Unlike the C₄grasses examined previously, significant rates of net photosynthesiscan be detected at leaf temperatures below 10?C. Rates of netphotosynthesis equal or exceed those reported for temperateC₃ grasses at all of the temperature (5–40?C) and photonflax density (13–2500µmol m^–2 s^–1) conditionsexamined. Maximum rates of net photosynthesis in S. townsendiiare almost double those reported for C₃ herbage grasses. Unliketemperate C₃ grasses, the major limitation to net photosynthesisat low leaf temperatures (10?C and below) is the stomatal resistance,showing that the low residual resistance characteristic of C₄species is maintained in S. townsendii even at low leaf temperatures.     

Photosynthetic carbon metabolism in wild, primitive and cultivated forms of wheat at three levels of ploidy: Role of the glycolate pathway

¹⁴CO₂ assimilation was studied with diploid, tetraploid, hexaploidspecies of the genera Triticum and their wild relatives Aegilops.Attached mature leaves of 3–4 weekold plants were allowedto undergo photosynthesis under air at ambient temperature.The pattern of distribution of ¹⁴C was notably similar in Triticumand Aegilops species whatever the level of ploidy. Sucrose wasthe sink for photosynthetic carbon. ¹⁴C for sucrose synthesis was supplied either through the glycolatepathway by glycolate, the product of the photorespiration orby the Calvin cycle intermediates exported into the cytoplasm.Depending on the species, the glycolate pathway provided 40to 75%of the sucrose ¹⁴C. The higher labeling of sucrose was associated with the greaterparticipation of the glycolate pathway in the wild diploid (DD)A. squarrosa and in the cultivated hexaploid (AABBDD) T. aestivum.The results suggest that the expression of the male D genomeis dominant over the female AB genome in T. aestivum. In T. aestivum under ambient conditions lowering (low temperature)or hindering (1% O₂ ) photorespiration, sucrose labeling decreased,but serine and glycine labeling was favoured. We propose thatin wheat leaves, the role of photorespiration is to drain artof the carbon exported from the chloroplast as glycolate, towardssucrose synthesis. (Received March 16, 1979; )     

The Influence of Temperature on Photosynthesis and Transpiration in ten Temperate Grass Varieties Grown in four Different Environments

The influence of temperature on photosynthesis and transpirationwas studied in ten varieties of Lolium perenne, L. multiflorum,Dactylis glomerata, and Festuca arundinacea from three climaticorigins grown in three different controlled environments (15?C, 72 W m^-2 visible irradiation, 16-h photoperiod; 25 ?C, 72W m^-2 visible irradiation, 16-h photoperiod; and 25 ?C, 180W m^-2 visible irradiation, 16-h photoperiod) and in the glasshousein July/August. The optimum temperature for photosynthesis was influenced primarilyby growth environment; growth at low temperature (15 ?C) resultedin a low optimum temperature, which differed little from varietyto variety. The maximum CO₂-exchange rate was influenced bygrowth environment and by variety. Within a variety, plantsgrown at higher light intensity or lower temperature had a greaterCO₂-exchange rate. Seven varieties showed a negative correlationbetween the optimum leaf temperature and the maximum CO₂-exchangerate. Activation energies for photosynthesis were influencedby growth environment only. There were marked varietal differences in the values of leafresistances (r_a + r_t) obtained from transpiration data at theoptimum leaf temperature for CO₂ exchange. In Lolium, and Dactylisthe Mediterranean varieties had higher leaf resistances thanthe Northern varieties with the maritime varieties intermediate.In general the Dactylis varieties had higher resistances thanthe corresponding Lolium and Festuca varieties. Only at highgrowth temperatures was (r_a+r_l) insensitive to temperature;otherwise an activation energy of about 10 kcal/mole was observed.A negative correlation was found between mean varietal diffusionresistances (r_a+r_l), and corresponding maximum CO₂-exchangerates.     

Photosynthetic Assimilation of 14CO2 by Waterstressed Sunflower Leaves at Two O2 Concentrations and the Specific Activity of Products

Attached leaves of sunflower (Helianthus annuus L. var. Mennonite)with water potentials of –5 to –18 ? 10⁵ Pa, wereexposed for different times to 300 vpm CO₂ containing ¹⁴CO₂and 21 or 1.5% O₂. ¹⁴C accumulated linearly with time in bothO₂ concentrations and at all stresses. 3-Phosphoglyceric acidwas saturated with ¹⁴C after 10 min in unstressed plants atboth O₂ concentrations but with increasing stress the rate ofaccumulation and the specific activity decreased. With decreasingleaf water potential there was accumulation of radioactivityin the glycolate pathway intermediates glycine and serine. Otheramino acids contained a slightly larger proportion of assimilatedcarbon as water potential decreased. The specific activitiesof all compounds were smaller with stress. In contrast to theamino acids less radioactivity accumulated in sugars, organicacids, and sugar phosphates and their specific activities decreasedwith stress. The radioactive labelling patterns and specificactivity measurements are interpreted as showing increased carbonflux in the glycolate pathway and inhibition of the metabolismof serine to sucrose. These changes are related to previousresults showing that with stress photo respiration increasesas a proportion of photosynthesis. Lowering the O₂ concentrationto 1.5% decreased the accumulation of radioactivity in glycineand stopped photorespiration. It increased the amount of radioactivityin serine and sucrose but did not greatly change specific activities.Oxygen effects were independent of water stress. Glycolate pathwaymetabolism is discussed in relation to photorespiration andthe effects of water stress.     

Effects of Nitrogen Nutrition on Photosynthesis in Cd-treated Sunflower Plants

Increased nitrogen supply stimulates plant growth and photosynthesis.Since it was shown that heavy metals may cause deficienciesof essential nutrients in plants the potential reversal of cadmiumtoxicity by increased N nutrition was investigated. The effectson photosynthesis of low Cd (0, 0.5, 2 or 5 mmol m^-3) combinedwith three N treatments (2, 7.5 or 10 mol m^-3) were examinedin young sunflower plants. Chlorophyll fluorescence quenchingparameters were determined at ambient CO₂and at 100 or 800 µmolquanta m^-2 s^-1. The vitality index (R_fd) decreased approx. three-timesin response to 5 mmol m^-3Cd, at 2 and 10 mol m^-3N. The maximumphotochemical efficiency of PSII reaction centres (F_v/ F_m) wasnot influenced by Cd or N treatment. The highest Cd concentrationdecreased quantum efficiency of PSII electron transport (_II)by 30%, at 2 and 10 mol m^-3N, mostly due to increased closureof PSII reaction centres (q_P). Photosynthetic oxygen evolutionrates at saturating CO₂were decreased in plants treated with5 mmol m^-3Cd, at all N concentrations. The results indicatethat Cd treatment affected the ribulose-1,5-bisphosphate (RuBP)regeneration capacity of the Calvin cycle more than other processes.At the same time, the amounts of soluble and ribulose-1,5-bisphosphatecarboxylase/oxygenase (Rubisco) protein increased with Cd treatment.Decreased photosynthesis, but substantially increased Rubiscocontent, in sunflower leaves under Cd stress indicate that asignificant amount of Rubisco protein is not active in photosynthesisand could have another function. It is shown that optimal nitrogennutrition decreases the inhibitory effects of Cd in young sunflowerplants. Copyright 2000 Annals of Botany Company Helianthus annuus L., cadmium, nitrogen, photosynthesis, Rubisco, sunflower     

Effect of CO2 Concentration During Growth on a CO2 Concentrating Mechanism in White Clover as Predicted from Differential 14CO2/12CO2Uptake

Lehnherr, B. M?chler, F. and N?sberger, J. 1985. Effect of CO₂concentration during growth on a CO₂ concentrating mechanismin white clover as predicted from differential ¹⁴CO₂/¹²CO² uptake.-J. exp. Bot. 36: 1835-1841. White clover was grown at 20 and100 Pa p(CO₂). The CO₂ response of net photosynthesis and differentialuptake of ¹⁴CO₂ and ¹²CO₂ by leaves were measured at varioustemperatures and at various O₂ and CO₂ partial pressures andcompared with predictions from ribulose bisphosphate carboxylase/oxygenasekinetics. Discrepancies between the observed gas exchange characteristicsfor the leaves and those predicted from the enzyme kineticswere interpreted as being due to a CO₂ concentrating mechanism.Plants grown at 20 Pa p(CO₂) showed a higher affinity for CO₂than plants grown at 100 Pa p(CO₂) when measured at 10 ?C. Nodifference in affinity was found at 30 ?C. The postulated CO₂concentrating effect was greater in plants grown at low CO₂than in plants grown at high CO₂ concentration and occurredonly at low temperature and low CO₂ partial pressure. It issuggested that plants grown at the lower CO₂ partial pressurehave a higher affinity for CO₂ due to a more efficient CO₂ concentratingsystem than plants grown at the higher CO₂ partial pressure. Key words: Photosynthesis, CO₂, concentration, RuBP carboxylase/oxygenase     

Dynamics of Carbon Photosynthetically Assimilated in Nodulated Soya Bean Plants under Steady-state Conditions 1. Development and Application of 13CO2 Assimilation System at a Constant 13C Abundance

A long-term, steady-state ¹³CO₂ assimilation system at a constantCO₂ concentration with a constant ¹³C abundance was designedand applied to quantitative investigations on the allocationof photoassimilated carbon in nodulated soya bean (Glycine maxL.) plants. The CO₂ concentration in the assimilation chamberand its ¹³C abundance were maintained constant with relativevariances of less than ±0.5 per cent during an 8-h assimilationperiod. At the termination of 8-h ¹³CO₂ assimilation by plantsat early flowering stage, the currently assimilated carbon relativeto total tissue carbon (measured by the degree of isotopic saturation)were for young leaves (including flower buds), 13.9 per cent;mature leaves, 15.7 per cent; stems+petioles, 5.9 per cent;roots, 5.4 per cent and nodules, 6.9 per cent, 48 h after theend of the ¹³CO₂ assimilation period, they were 12.3, 7.5, 7.4,6.8 and 6.1 per cent, respectively. The treatment with a highconcentration of nitrate in the nutrient media significantlydecreased the allocation of ¹³C into nodules. Experiments on¹³CO₂ assimilation by plants at the pod-filling stage were alsoconducted. Labelling by ¹³C was weaker than at the early floweringstage, but an intense accumulation of ¹³C into reproductiveorgans was observed. Glycine max L., nodulated soya bean plants, ¹³CO₂ assimilation, carbon dynamics     

Dynamics of Carbon Photosynthetically Assimilated in Nodulated Soya Bean Plants under Steady-state Conditions 2. The Incorporation of 13C into Carbohydrates, Organic Acids, Amino Acids and some Storage Compounds

Nodulated soya bean (Glycine max L.) plants at the early floweringstage were allowed to assimilate ¹³CO₂ under steady-state conditions,with a constant ¹³C abundance, for 8 h in the light. The plantswere either harvested immediately or 2 d after the end of the¹³CO₂ feeding, divided into young leaves (including flower buds),mature leaves, stems+petioles, roots and nodules; the ¹³C abundancein soluble carbohydrates, organic acids, amino acids, starchand poly-ß-hydroxybutyric acid was determined witha gas chromatography-mass spectrometry. The rapid turnover of ¹³C in the sucrose pools observed in allorgans of the plants showed that sucrose was the principal materialin the translocation stream of primary products of photosynthesis.At the end of the ¹³CO₂ exposure, sucrose in the mature leavesas the major source organs and in the stems+petioles was labelledwith currently assimilated carbon to about 75 per cent, whereasa much higher labelling of sucrose was found in the roots andin the nodules. This suggests the existence of two or more compartmentedpools of sucrose in mature leaves and also in stems+petioles. The relative labelling patterns of individual organic acidsand amino acids were similar in various plant organs. However,the rapid turnover of succinate and glycine was characteristicof nodules. Treatment with a high concentration of nitrate inthe nutrient media increased the turnover rate of amino acidcarbon in shoot organs and roots, while it markedly decreasedthe labelling of amino acids in nodules. The cyclitols, exceptfor D-pinitol, were significantly labelled with assimilated¹³C in mature leaves, but in nodules, the labelling was verymuch less. In the nodules, which were actively fixing atmospheric nitrogen,a large proportion (80–90 per cent) of currently assimilatedcarbon was found as sucrose and starch at the end of the ¹³CO₂feeding. This was also true of the roots. On the other hand,in young growing leaves, the distribution of currently assimilatedcarbon into sucrose, starch and other soluble compounds wasmuch less. This suggests that a large amount of carbon assimilatedby and translocated to young leaves was used to make up structuralmaterials, mainly protein and cell wall polymers synthesis,during the light period. Glycine max L., soya bean, ¹³CO₂ assimilation, carbon metabolism in nodules     

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     

Carbon Loss from the Roots of Forage Rape (Brassica napus L.) Seedlings Following Pulse-labelling with 14CO2

The loss of organic material from the roots of forage rape (Brassicanapus L.,) was studied by pulse-labelling 25-d-old non-sterilesand-grown plants with ¹⁴CO₂. The distribution of ¹⁴C withinthe plant was measured at 0, 6 and 13 d after labelling whilst¹⁴ C accumulating in the root-zone was measured at more frequentintervals. The rates of ¹⁴C release into the rhizosphere, andloss of ¹⁴CO₂ from the rhizosphere were also determined. Thesedata were used to estimate the accumulative loss of ¹⁴C fromroots and loss respiratory ¹⁴CO₂ from both roots and associatedmicro-organisms. Approximately 17-19% of fixed ¹⁴CO₂ was translocatedto the roots over 2 weeks, of which 30-34% was released intothe rhizosphere, and 23-24% was respired by the roots as ¹⁴CO₂. Of the ¹⁴C released into the rhizosphere, between 35-51%was assimilated and respired by rhizosphere micro-organisms.Copyright1993, 1999 Academic Press Brassica napus L., carbon loss, carbon partitioning, microbial nutrition, microbial respiration, forage rape, pulse-labelling, rhizodeposition, root respiration, sand culture     

Photosynthesis of White Clover Leaves as Influenced by Canopy Position, Leaf Age, and Temperature

Microswards of white clover (Trifolium repens L.) were grownin controlled environments at 10/7, 18/13 and 26/21 °C day/nighttemperatures. The vertical distribution of leaves of differentages and their rates of ¹⁴CO₂-uptake in situ were studied. Extending petioles carried the laminae of young leaves throughthe existing foliage. A final position was reached within 1/4to 1/3 of the time between unfolding and death. Newly unfoldedleaves had higher rates of ¹⁴CO₂-uptake per leaf area than olderones at the same height in the canopy. At higher temperatures,the decrease with age was faster. However, the light-photosynthesisresponse of leaves which were removed from different heightsin the canopy varied much less with leaf age than did the ratesof ¹⁴CO₂-uptake in situ. The comparison of the rates of ¹⁴CO₂-uptake in situ with thelight-photosynthesis response curves suggests that young leavesreceive more light than older ones at the same height in thecanopy. This would imply that young white clover leaves havethe ability to reach canopy positions having a favourable lightenvironment. This ability may improve the chances of survivalof white clover in competition with other species. Trifolium repens L., white clover, photosynthesis, canopy, leaf age, ¹⁴CO₂-uptake, ecotypes, temperature