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.     

Short- and Long-Term Inhibition of Respiratory Carbon Dioxide Efflux by Elevated Carbon Dioxide

Dark carbon dioxide efflux rates of recently fully expandedleaves and whole plants of Amaranthus hypochondriacus L., Glycinemax (L.) Merr., and Lycopersicon esculentum Mill. grown in controlledenvironments at 35 and 70 Pa carbon dioxide pressure were measuredat 35 and 70 Pa carbon dioxide pressure. Harvest data and whole-plant24-h carbon dioxide exchange were used to determine relativegrowth rates, net assimilation rates, leaf area ratios, andthe ratio of respiration to photosynthesis under the growthconditions. Biomass at a given time after planting was greaterat the higher carbon dioxide pressure in G. max and L. esculentum,but not the C₄ species, A. hypochondriacus. Relative growthrates for the same range of masses were not different betweencarbon dioxide treatments in the two C₃ species, because highernet assimilation rates at the higher carbon dioxide pressurewere offset by lower leaf area ratios. Whole plant carbon dioxideefflux rates per unit of mass were lower in plants grown andmeasured at the higher carbon dioxide pressure in both G. maxand L. esculentum, and were also smaller in relation to daytimenet carbon dioxide influx. Short-term responses of respirationrate to carbon dioxide pressure were found in all species, withcarbon dioxide efflux rates of leaves and whole plants lowerwhen measured at higher carbon dioxide pressure in almost allcases. Amaranthus hypochondriacus L., Glycine max L. Merr., Lycopersicon esculentum Mill., soybean, tomato, carbon dioxide, respiration, growth     

Effect of Low Relative Humidity on {delta}13C Value in Two C3 Grasses and in Panicum milioides, a C3-C4 Intermediate Species

When grown under conditions of low relative humidity, the C₃–C₄intermediate Panicum milioides, as well as the C₃ grasses Triticumaestivum and Poa pratense, exhibited ¹³C values which were upto 2–7%o less negative than the ¹³C values of the correspondingplants grown at high relative humidity. At both humidity levels,there was no evidence of a substantial contribution of phosphoenolpyruvatecarboxylase to carbon gain in Panicum milioides     

Silica Deposition in Some C3 and C4 Species of Grasses, Sedges and Composites in the USA

We report new information on silica deposition in 15 plant species,including nine grasses, two sedges and four composites. Thesilica depositional patterns found in seven of the grass speciesindicate that they are C₄ plants. However the festucoid grassCortaderia selloana is a C₃ plant with long leaf trichomes andoval silica structures in the leaves. In contrast the panicoidC₄ grasses Chasmathium latifolium, Chasmathium sessiflorum,Imperata cylindrica, Panicum repens, Panicum commutatum andSetaria magna, all produce dumb-bell-shaped silica structuresin the leaves. The chloridoid grasses Spartina patens and Spartinacynosuroides have saddle-shaped structures and no dumb-bellor oval shaped ones. The sedges Rhynchospora plumosa and Scirpuscyperinus were found to have oval phytoliths and may be C₃ plants.Our examination of these and other grasses strongly suggeststhat C₄ grasses tend to produce the same type of silica cells.Grasses and sedges with C₃ type photosynthesis tend to produceoval silica structures. The composite Grindelia squarrosa andsunflowers Helianthus angustifolia, Helianthus atrorubens andHelianthus tuberosus absorb relatively small amounts of siliconand larger amounts of calcium, where both elements deposit inleaf trichomes. We found no clear indicator for the C₃ sunflowersor C₄ types in the Asteraceae. Helianthus tuberosus leaves havemany trichomes on the adaxial surface. These trichomes havea higher concentration of silica than the surrounding leaf surface.Helianthus tuberosus leaves had much higher ash and silica contentsthan those of Helianthus angustifolia and Helianthus atrorubens.The composite Grindelia squarrosa has a usual deposition ofsilica in the basal cells around the guard cells. Silica depositionoften reflects the surface features of a leaf. An exceptionis Scripus cyperinus where the silica structures are deep inthe tissue and do not reflect the surface configurations. Theinforescence of Setaria magna had a 14.64 silica content. Thetufts of white, silky hairs characteristic of Imperata cylindricainflorescence have no silica. C₃ and C₄ plants, silica and ash content, scanning electron microscopy, energy-dispersive X-ray analysis, silicon distribution, spectra of elements in plants, trichomes, silica fibres, phytoliths     

Photosynthetic efficiency of Panicum hians and Panicum milioides in relation to C3 and C4 plants

Panicum hians and Panicum milioides were found to have characteristicsintermediate to those of C₃ and C₄ species with respect to CO₂compensation point, percentage inhibition of photosynthesisby O₂ at various O₂/CO₂ solubility ratios, and water use efficiency.C₄ species have a higher carboxylation efficiency than eitherthe intermediate or C₃ species. During photosynthesis, evenunder 2.5% O₂, C₄ species have a higher affinity for intercellularCO₂ (Km 1.6 µM) apparently due to the initial carboxylationthrough PEP carboxylase. Under low O₂ the intermediate and C₃species had a similar affinity for intercellular CO₂ duringphotosynthesis (Km 5–7 µM) consistent with carboxylationof atmospheric CO₂ through RuDP carboxylase. There were considerablevariation in photosynthesis/unit leaf area at saturating CO₂levels in the species examined which in part is due to differencesin RuDP carboxylase /unit leaf area. The highest rates of photosynthesis/unitleaf area under CO₂-saturating conditions were with the C₃ specieswhich had a correspondingly high level of RuDP carboxylase/unitleaf area. Possibilities for the greater efficiency of P. hiansand P. milioides in comparison to C₃ species in utilizing lowlevels of CO₂ in the presence of atmospheric O₂ are discussed. ¹ This research was supported by the College of Agriculturaland Life Sciences, University of Wisconsin, Madison; and theUniversity of Wisconsin Research Committee with funds from theWisconsin Alumni Research Foundation. (Received June 25, 1977; )     

Influence of High CO2Partial Pressure on Nitrogen Use Efficiency of the C4Grasses Panicum coloratum and Cenchrus ciliaris

Australia's tropical grasslands are dominated by C₄grasses,characterized by their unique biochemistry and anatomy. Twonaturalized C₄grasses (Panicum coloratum and Cenchrus ciliaris)were used to investigate whether high CO₂partial pressure [p(CO₂)] influences photosynthetic nitrogen use efficiency andplant nitrogen use efficiency (PNUE and NUE respectively). Plantswere grown for 30 d with four levels of N at p(CO₂) of 38 or86 Pa. PNUE was calculated from leaf CO₂assimilation rates (A)and leaf N concentrations, and NUE from total leaf N contentand plant dry mass. At each p(CO₂), PNUE and NUE were greaterfor C. ciliaris than for P. coloratum due to higher A and drymass combined with lower leaf N concentrations. Elevatedp (CO₂)increased PNUE of C. ciliaris only. This effect was due to lowerleaf N concentrations (area basis). At high p(CO₂), NUE of C.ciliaris was also greater. This resulted from a 1.6-fold stimulationof dry mass by high p(CO₂). Although dry mass of P. coloratumwas increased 1.2-fold by elevated p(CO₂), its NUE was unaffected.Leaf transpiration rates were halved at elevated p(CO₂), andwe suggest that this factor plays a major role in the growthresponse of C₄grasses to high p(CO₂). Copyright 2001 Annalsof Botany Company Panicum coloratum, Cenchrus ciliaris, nitrogen use efficiency, elevated CO₂, leaf N concentration, growth, photosynthesis     

Structure and Function of Silica Bodies in the Epidermal System of Grass Shoots

Silica (SiO₂.nH₂O) is deposited in large quantities in the shootsystems of grasses. In the leaf epidermal system, it is incorporatedinto the cell wall matrix, primarily of outer epidermal walls,and within the lumena of some types of epidermal cells. This biogenic silica can be stained specifically with methylred, crystal violet lactone, and silver amine chromate. At theultrastructural level, the silica in lumens of silica cells,bulliform cells and long epidermal cells is made up of rodsabout 2.5 µm in length and 0.4µm in width. Ultimateparticles in the rods range from 1 to 2 nm in diameter. In contrast,silica in the cell wall matrix of trichomes and outer wallsof long epidermal cells is not rod-shaped, but rather, formsroughly spherical masses. Detailed analyses are presented on the frequencies of occurrenceof the different types of epidermal cells that contain silicain the leaves of representative C₃ and C₄ grasses. The C₄ grasseshave higher frequencies of bulliform cell clusters, silica cells,and long epidermal cells, whereas the C₃ grasses have higherfrequencies of trichomes. No correlation was found in the frequencyof occurrence of silica bodies in bulliform cells for C₃ grassesas compared with C₄ grasses. Of all the grasses examined, Coix,Oryza, and Eleusine had the highest densities of such bodies,and some taxa had no silica bodies apparent in their bulliformcells. The idea that silica bodies in bulliform cells and silica cellsmight act as "windows’ and trichomes might function as‘light pipes’ to facilitate light transmission throughthe epidermal system to photosynthetic mesophyll tissue belowwas tested. The experimental data presented do not support eitherof these hypotheses. C₂ and C₄ grasses, biogenic silica, light pipes, window hypothesis, silica staining, silica ultrastructure     

Growth and Partitioning in Pascopyrum smithii (C3) and Bouteloua gracilis(C4) as Influenced by Carbon Dioxide and Temperature

This study investigated how CO₂and temperature affect dry weight(d.wt) accumulation, total nonstructural carbohydrate (TNC)concentration, and partitioning of C and N among organs of twoimportant grasses of the shortgrass steppe,Pascopyrum smithiiRydb. (C₃) andBouteloua gracilis(H.B.K.) Lag. ex Steud. (C₄).Treatment combinations comprised two temperatures (20 and 35°C)at two concentrations of CO₂(380 and 750 µmol mol^-1),and two additional temperatures of 25 and 30°C at 750 µmolmol^-1CO₂. Plants were maintained under favourable nutrient andsoil moisture and harvested following 21, 35, and 49d of treatment.CO₂-induced growth enhancements were greatest at temperaturesconsidered favourable for growth of these grasses. Comparedto growth at 380 µmol mol^-1CO₂, final d.wt of CO₂-enrichedP.smithiiincreased 84% at 20°C, but only 4% at 35°C. Finald.wt ofB. graciliswas unaffected by CO₂at 20°C, but wasenhanced by 28% at 35°C. Root:shoot ratios remained relativelyconstant across CO₂levels, but increased inP. smithiiwith reductionin temperature. These partitioning results were adequately explainedby the theory of balanced root and shoot activity. Favourablegrowth temperatures led to CO₂-induced accumulations of TNCin leaves of both species, and in stems ofP. smithii, whichgenerally reflected responses of above-ground d.wt partitioningto CO₂. However, CO₂-induced decreases in plant tissue N concentrationswere more evident forP. smithii. Roots of CO₂-enrichedP. smithiihadgreater total N content at 20°C, an allocation of N below-groundthat may be an especially important adaptation for C₃plants.Tissue N contents ofB. graciliswere unaffected by CO₂. Resultssuggest CO₂enrichment may lead to reduced N requirements forgrowth in C₃plants and lower shoot N concentration, especiallyat favourable growth temperatures. Acclimation to CO₂; blue grama; Bouteloua gracilis ; carbohydrate; climate change; global change; grass; growth; growth temperature optima; nitrogen; N uptake; Pascopyrum smithii; western wheatgrass     

Quantity and Kinetic Properties of Ribulose 1,5-Bisphosphate Carboxylase in C3, C4, and C3-C4 Intermediate Species of Flaveria (Asteraceae)

The kinetic properties of ribulose 1,5-bisphosphate carboxylase(RuBPC) appear to have been modified during evolution of photosynthesisto adjust to changes in substrate availability. C₄ plants areconsidered to have a higher concentration of CO₂ available toRuBPC than C₃plants. In this study, the K_m(CO₂ and catalyticcapacity (k_cat) of RuBPC and the ratio of RuBPC protein to totalsoluble protein from several Flaveria species, including C₃,C₃-C₄ intermediate, and C₄ species, were determined. The C₃and intermediate species had similar K_m(CO₂) values while theC₄ species on average had higher K_m(CO₂) values. The mean ratioof K_cat/K_m for species of each group was similar, supportingthe hypothesis that changes in K_m and K_cat, are linked. Theallocation of total soluble protein to RuBPC was lowest in theC₄ Flaveria species, intermediate in the C₃-C₄ species, andhighest in the C₃ species. The results suggest that during evolutionof C₄ photosynthesis adjustments may occur in the quantity ofRuBPC prior to changes in its kinetic properties. (Received January 4, 1989; Accepted April 11, 1989)     

Silica and Ash in Native Plants of the Central and Southeastern Regions of the United States

Ash and silica contents and depositional patterns were determinedfor different tissues of 11 plants growing in the southeasternand central parts of the USA. Silica content was high in theleaves, sheaths and inflorescences of the grasses studied, especiallyso in the inflorescence of the C₃ grass, Stipa comata Trise.and Rupr. The ash content was especially high in leaves of Polymniauvedalia L., which are also high in calcium. Calcium depositionwas largely in trichomes and in veins of the leaf. Energy-dispersiveX-ray analysis showed that the distribution of the element siliconis closely related to certain epidermal structures such as ridges,cell walls, rows of irregularly-shaped structures lying lenghthwisealong the leaf, dumb-bell shaped structures and trichomes. Thesestructures also correspond to the phytoliths left behind afterdecay of the plant. The C₃ grasses differed from the C₄ in thatthey showed oval structures and produced correspondingly ovalphytoliths. Silicified trichomes (particularly in the C₃ grasses)and long, narrow, silica fibres were common in the inflorescencesof the grasses studied. These sharp particles could be irritatingto oesophageal and other tissues. Similar fibres in other grasseshave been implicated in certain cancers. High silicificationof the inflorescence structures might afford protection forthe seed, as reported for other grasses. C₃ and C₄ grasses, silica and ash content, scanning electron microscopy, energy-dispersive X-ray analysis, silicon distribution, spectra of elements in plants, trichomes, silica fibres, phytoliths     

Effects of elevated carbon dioxide on three montane grass species: I. Growth and dry matter partitioning

Upland grasslands are a major component of natural vegetationwithin the UK. Such grasslands support slow growing relativelystable plant communities. The response of native montane grassspecies to elevated atmospheric carbon dioxide concentrationshas received little attention to date. Of such studies, mosthave only focused on short-term (days to weeks) responses, oftenunder favourable controlled environment conditions. In thisstudy Agrostis caplllaris L.⁵, Festuca vivipara L. and Poa alpinaL. were grown under semi-natural conditions in outdoor open-topchambers at either ambient (340µmol mol^–1) or elevated(680µmol mol^–1) concentrations of atmospheric carbondioxide (CO₂ for periods from 79 to 189 d, with a nutrient availabilitysimilar to that of montane Agrostis-Fescue grassland in Snowdonia,N. Wales. Whole plant dry weight was increased for A. capillarisand P. alpina, but decreased for F. vivipara, at elevated CO₂.Major components of relative growth rate (RGR) contributingto this change at elevated CO₂ were transient changes in specificleaf area (SLA) and leaf area ratio (LAR). Despite changes ingrowth rate at 680 µmol mol^–1 CO₂, partitioningof dry weight between shoot and root in plants of A. capillarisand P. alpina was unaltered. There was a significant decreasein shoot relative to root growth at elevated CO₂ in F. viviparawhich also showed marked discoloration of the leaves and increasedsenescence of the foliage. Key words: Allometry, growth analysis, elevated CO₂, grasses     

A comparison of the impacts of various nitrogen sources on acid-base balance in C3 Triticum aestivum L. and C4 Zea mays L. plants

This work aimed to study the impacts of acquisition and assimilationof various nitrogen sources, i.e. NO₃^–, NH₄⁺ or NH₄NO₃,in combination with gaseous NH₃ on plant growth and acid-basebalance in higher plants. Plants of C₃ Triticum aestivum L.and C₄ Zea mays L. grown with shoots in ambient air in hydroponicculture solutions with 2 mol m^–3 of nitrogen source asNO₃^–, NH₄⁺ or NH₄NO₃ for 21 d and 18 d, respectively,had their shoots exposed either to 320 µg m^–3 NH₃or to ambient air for 7 d. Variations in plant growth (leaves,stubble and roots), and OH^– and H⁺ extrusions as wellas the relative increases in nitrogen, carbon and carboxylatewere determined. These data were computed as H⁺/N, H⁺/C, (C-A)/N,and (C-A)/C to analyse influences of different nitrogen sourceson acid-base balance in C₃ Triticum aestivum and C₄ Zea maysplants. Root growth in dry weight gain was significantly reduced bytreatment with 320 µg m^–3 NH₃ in Triticum aestivumand Zea mays growing with different N-forms, whereas leaf growthwas not significantly affected by NH₃. In comparison with C₃Triticum aestivum, non-fumigated C₄ Zea mays had low ratiosof OH^–/N in NO₃^–3-grown plants and of H⁺/N in NH₄⁺- and NH ₄NO₃-grown plants. Utilization of NH₃ from the atmospherereduced both the OH^–N ratios in NO₃^– -grown plantsand the H⁺/N ratio in NH₄⁺ - and NH₄NO₃ -grown plants of bothspecies. Furthermore, Zea mays had higher ratios of (C-A)/Nin NH₄⁺ - and NH₄NO₃-grown plants than Triticum aestivum. Thismeans that C₄ Zea mays had synthesized more organic anion perunit increase in organic N than C₃ Triticum aestivum plants.Within both species, different nitrogen sources altered theratios of (C-A)/N in the order: NH₄NO₃>NH₄⁺>NO₃^–.Fumigation with NH₃ increased organic acid synthesis in NO₃^–- and NH₄⁺ - grown plants of Triticum aestivum, whereas it decreasedorganic acid synthesis in Zea mays plants under the same conditions.Furthermore, these differences in acid-base regulation betweenC₃ Triticum aestivum and C₄ Zea mays plants growing with differentnitrogen sources are discussed. Key words: Acid-base balance, ammonia, ammonium, nitrate, ammonium nitrate, C₃ Triticum aestivum L., C₄ Zea mays L.     

Effects of Prolonged Darkness on the Sensitivity of Leaf Respiration to Carbon Dioxide Concentration in C3and C4Species

Predicting responses of plant and global carbon balance to theincreasing concentration of carbon dioxide in the atmosphererequires an understanding of the response of plant respirationto carbon dioxide concentration ([CO₂]). Direct effects of thecarbon dioxide concentration at which rates of respiration ofplant tissue are measured are quite variable and their effectsremain controversial. One possible source of variation in responsivenessis the energy status of the tissue, which could influence thecontrol coefficients of enzymes, such as cytochrome-c oxidase,whose activity is sensitive to [CO₂]. In this study we comparedresponses of respiration rate to [CO₂] over the range of 60to 1000 µmol mol^-1in fully expanded leaves of four C₃andfour C₄herbaceous species. Responses were measured near themiddle of the normal 10 h dark period, and also after another24 h of darkness. On average, rates of respiration were reducedabout 70% by the prolonged dark period, and leaf dry mass perunit area decreased about 30%. In all species studied, the relativedecrease in respiration rate with increasing [CO₂] was largerafter prolonged darkness. In the C₃species, rates measured at1000 µmol mol^-1CO₂averaged 0.89 of those measured at 60µmol mol^-1in the middle of the normal dark period, and0.70-times when measured after prolonged darkness. In the C₄species,rates measured at 1000 µmol mol^-1CO₂averaged 0.79 of thoseat 60 µmol mol^-1CO₂in the middle of the normal dark period,and 0.51-times when measured after prolonged darkness. In threeof the C₃species and one of the C₄species, the decrease in theabsolute respiration rate between 60 and 1000 µmol mol^-1CO₂wasessentially the same in the middle of the normal night periodand after prolonged darkness. In the other species, the decreasein the absolute rate of respiration with increase in [CO₂] wassubstantially less after prolonged darkness than in the middleof the normal night period. These results indicated that increasingthe [CO₂] at the time of measurement decreased respiration inall species examined, and that this effect was relatively largerin tissues in which the respiration rate was substrate-limited.The larger relative effect of [CO₂] on respiration in tissuesafter prolonged darkness is evidence against a controlling roleof cytochrome-c oxidase in the direct effects of [CO₂] on respiration.Copyright 2001 Annals of Botany Company Carbon dioxide, respiration, Abutilon theophrasti(L.), Amaranthus retroflexus(L.),Amaranthus hypochondriacus (L.), Datura stramonium(L.), Helianthus annuus(L.), Solanum melongena(L.), Sorghum bicolor(L. Moench), Zea mays     

Leaf Interveinal Distances Corresponding to Anatomical Types in Grasses

Considerable variation in the interveinal distances in leaveswas observed in both C₃ and C₄ grass species. In C₄ species,interveinal distances of the Mestome Sheath type (89.0 µm)were smaller than those of the Parenchyma Sheath type (140.0µm). In contrast, non-Kranz type (C₃ species) had thelargest interveinal distances of all the anatomical types. Variationin the interveinal distances of leaves of grasses is discussedin relation to photosynthetic ability and water use. (Received August 7, 1984; Accepted January 16, 1985)     

Photosynthesis in Panicum milioides, a Species with Reduced Photorespiration

The capacity for C₄ photosynthesis in Panicum milioides, a specieshaving reduced levels of photorespiration, was investigatedby examining the activity of certain key enzymes of the C₄ pathwayand by pulse-chase experiments with ¹⁴CO₂. The ATP$P₁ dependentactivity of pyruvate,P₁ dikinase in the species was extremelylow (0.14–0.18 µmol mg chlorophyll^–1 min^–1).Low activity of the enzyme was also found in Panicum decipiensand Panicum hians (related species with reduced photorespiration)and in Panicum laxum (a C₃ species). The antibody to pyruvate,P₁dikinase caused about 70% inhibition of the ATP$P₁ dependentactivity of the enzyme in P. milioides. The activity of NAD-malicenzyme and NADP-malic enzyme in ^P. ^milioides was equally low(approximately 0.1–0.2 µmol mg chlorophyll^–1min^–1) and similar to the activity in P. decipiens, P.hians and P. laxum. Photosynthetic pulse-chase experiments underatmospheric conditions showed a typical C₃-like pattern of carbonassimilation including the labelling of glycine and serine asexpected during photorespiration. During the pulse with ¹⁴CO₂only about 1% of the labelled products appeared in malate and2–3% in aspartate. During a chase in atmospheric levelsof CO₂ for up to 6 min there was a slight increase in labellingin the C₄ acids. The amount of label in carbon 4 of aspartatedid not change during the chase, indicating little or no turnoverof the C₄ acid via decarboxylation. The results indicate thatunder atmospheric conditions P. milioides assimilates carbondirectly through the C₃ pathway. Photorespiration as indicatedby the CO₂ compensation point may be repressed in the speciesby a more efficient recycling of photorespired CO₂. (Received June 8, 1982; Accepted July 22, 1982)     

C3 grasses have higher nutritional quality than C4 grasses under ambient and elevated atmospheric CO2

Grasses with the C₃ photosynthetic pathway are commonly considered to be more nutritious host plants than C₄ grasses, but the nutritional quality of C₃ grasses is also more greatly impacted by elevated atmospheric CO₂ than is that of C₄ grasses; C₃ grasses produce greater amounts of nonstructural carbohydrates and have greater declines in their nitrogen content than do C₄ grasses under elevated CO₂. Will C₃ grasses remain nutritionally superior to C₄ grasses under elevated CO₂ levels? We addressed this question by determining whether levels of protein in C₃ grasses decline to similar levels as in C₄ grasses, and whether total carbohydrate : protein ratios become similar in C₃ and C₄ grasses under elevated CO₂. In addition, we tested the hypothesis that, among the nonstructural carbohydrates in C₃ grasses, levels of fructan respond most strongly to elevated CO₂. Five C₃ and five C₄ grass species were grown from seed in outdoor open‐top chambers at ambient (370 ppm) or elevated (740 ppm) CO₂ for 2 months. As expected, a significant increase in sugars, starch and fructan in the C₃ grasses under elevated CO₂ was associated with a significant reduction in their protein levels, while protein levels in most C₄ grasses were little affected by elevated CO₂. However, this differential response of the two types of grasses was insufficient to reduce protein in C₃ grasses to the levels in C₄ grasses. Although levels of fructan in the C₃ grasses tripled under elevated CO₂, the amounts produced remained relatively low, both in absolute terms and as a fraction of the total nonstructural carbohydrates in the C₃ grasses. We conclude that C₃ grasses will generally remain more nutritious than C₄ grasses at elevated CO₂ concentrations, having higher levels of protein, nonstructural carbohydrates, and water, but lower levels of fiber and toughness, and lower total carbohydrate : protein ratios than C₄ grasses.     

Physiology and Growth of Wheat Across a Subambient Carbon Dioxide Gradient

Two cultivars of wheat (Triticum aestivum L.), 'Yaqui 54' and'Seri M82', were grown along a gradient of daytime carbon dioxideconcentrations ([CO₂]) from near 350-200 µmol CO₂ mol^-1air in a 38 m long controlled environment chamber. Carbon dioxidefluxes and evapotranspiration were measured for stands (plantsand soil) in five consecutive 7·6-m lengths of the chamberto determined potential effects of the glacial/interglacialincrease in atmospheric [CO₂] on C₃ plants. Growth rates andleaf areas of individual plants and net assimilation per unitleaf area and daily (24-h) net CO₂ accumulation of wheat standsrose with increasing [CO₂]. Daytime net assimilation (P_D, mmolCO₂ m^-2 soil surface area) and water use efficiency of wheatstands increased and the daily total of photosynthetic photonflux density required by stands for positive CO₂ accumulation(light compensation point) declined at higher [CO₂]. Nighttimerespiration (R_N, mmol CO₂ m^-2 soil surface) of wheat, measuredat 369-397 µmol mol^-1 CO₂, apparently was not alteredby growth at different daytime [CO₂], but R_N /P_D of stands declinedlinearly as daytime [CO₂] and P_D increased. The responses ofwheat to [CO₂], if representative of other C₃ species, suggestthat the 75-100% increase in [CO₂] since glaciation and the30% increase since 1800 reduced the minimum light and waterrequirements for growth and increased the productivity of C₃plants.Copyright 1993, 1999 Academic Press Atmospheric carbon dioxide, carbon accumulation, evapotranspiration, light compensation point, net assimilation, respiration, Triticum aestivum, water use efficiency, wheat     

Partial Reduction in Activities of Photorespiratory Enzymes in C3-C4 Intermediates of Alternanthera and Parthenium

The maximum catalytic activities of several photorespiratoryand photosynthetic enzymes were determined in leaf extractsof three C₃–C₄ intermediates (Alternanthera ficoides,A. tenella and Parthenium hysterophorus) and were compared tothose of C₃ (A. sessiles, Pisum sativum) and C₄ (A. pungens,Zea mays and Amaranthus hypochondriacus) species. The activitylevels of key photorespiratory enzymes, glycolate oxidase, catalase,NADH-hydroxypyruvate reductase and glycerate kinase were less(28 to 35% reduced) in intermediates than those of typical C₃species. Similarly, the activities of photorespiratory aminotransferasesin the C₃–C₄ intermediates were also partially reduced(23 to 37% reduction). The activities of phosphoenolpyruvatecarboxylase (PEPC), pyruvate, orthophosphate dikinase and NAD-malicenzyme were higher (2 to 7 times) in leaf extracts of the intermediatesthan those of C₃ species. But the ratios of PEPC/rubisco inthe C₃–C₄ intermediates were more like C₃ than C₄ species.We draw attention to the partial reduction in enzyme activityof photorespiratory metabolism, which could be an importantfactor for restriction of photorespiration in the C₃–C₄intermediate species, in addition to enzyme compartmentationand/or operation of a ‘C₄-like’ cycle Key words: C₃–C₄ intermediates, C₄ pathway, enzyme profile, glycolate metabolism, photorespiration, photosynthesis     

Features of Photosynthesis in Haloxylon species of Chenopodiaceae that are Dominant Plants in Central Asian Deserts

Haloxylon aphyllum and H. persicum of Chenopodiaceae are dominantplants in the continental deserts of the Asian Irano-Turanianregion. The photosynthetic organs, assimilating shoots and leaf-likecotyledons of these two species were studied to characterizetheir photosynthetic types. ¹³C/¹²C isotope ratios, the cellularanatomy of as similating organs, primary photosynthetic products,and activities of carbon metabolism enzymes, RUBP carboxylase,PEP carboxylase, malic enzymes, and aspartate aminotransferase,indicate different pathways of CO₂ fixation in the photosyntheticorgans. Assimilating shoots had attributes of the C₄ photosynthesisentirely, while cotyledons lack Kranz-anatomy and incorporatedCO₂ via C₃ photosynthesis. Cotyledons and seeds had lower     

Dynamic Model of Leaf Photosynthesis with Acclimation to Light and Nitrogen

A simple model of photosynthesis in a mature C₃leaf is described,based on a non-rectangular hyperbola: the model allows the high-lightasymptote of that equation (P_max) to respond dynamically tolight and nitrogen. This causes the leaf light response equationto acclimate continuously to the current conditions of lightand N nutrition, which can vary greatly within a crop canopy,and through a growing season, with important consequences forgross production. Predictions are presented for the dynamicsof acclimation, acclimated and non-acclimated photosyntheticrates are compared, and the dependence of leaf properties onlight and N availability is explored. There is good correspondenceof predictions with experimental results at the leaf level.The model also provides a mechanism for a down regulation ofphotosynthesis in response to increased carbon dioxide concentrations,whose magnitude will depend on conditions, particularly of nitrogennutrition.Copyright 1998 Annals of Botany Company Leaf, photosynthesis, hyperbola, model, C₃, acclimation, light, nitrogen.