Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

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     

Is Partitioning of Dry Weight and Leaf Area Within Dactylis glomerata Affected by N and CO2Enrichment?

We examined changes in dry weight and leaf area within Dactylisglomerata L. plants using allometric analysis to determine whetherobserved patterns were truly affected by [CO₂] and N supplyor merely reflect ontogenetic drift. Plants were grown hydroponicallyat four concentrations of in controlled environment cabinets at ambient (360 µll^–1) or elevated (680 µl l^–1) atmospheric[CO₂]. Both CO₂and N enrichment stimulated net dry matter production.Allometric analyses revealed that [CO₂] did not affect partitioningof dry matter between shoot and root at high N supply. However,at low N supply there was a transient increase in dry matterpartitioning into the shoot at elevated compared to ambient[CO₂] during early stages of growth, which is inconsistent withpredictions based on optimal partitioning theory. In contrast,dry matter partitioning was affected by N supply throughoutontogeny, such that at low N supply dry matter was preferentiallyallocated to roots, which is in agreement with optimal partitioningtheory. Independent of N supply, atmospheric CO₂enrichment resultedin a reduction in leaf area ratio (LAR), solely due to a decreasein specific leaf area (SLA), when plants of the same age werecompared. However, [CO₂] did not affect allometric coefficientsrelating dry weight and leaf area, and effects of elevated [CO₂]on LAR and SLA were the result of an early, transient stimulationof whole plant and leaf dry weight, compared to leaf area production.We conclude that elevated [CO₂], in contrast to N supply, changesallocation patterns only transiently during early stages ofgrowth, if at all. Copyright 2000 Annals of Botany Company Allometric growth, carbon dioxide enrichment, Cocksfoot, Dactylis glomerata L., dry weight partitioning, leaf area ratio, nitrogen supply, shoot:root ratio, specific leaf area     

Compensatory Roles of Nitrogen Uptake and Photosynthetic N-use Efficiency in Determining Plant Growth Response to Elevated CO2: Evaluation Using a Functional Balance Model

We used a modified functional balance (FB) model to predictgrowth response of Helianthus annuus L. to elevated CO₂. Modelpredictions were evaluated against measurements obtained twiceduring the experiment. There was a good agreement between modelpredictions of relative growth rate (RGR) responses to elevatedCO₂and observations, particularly at the second harvest. Themodel was then used to compare the relative effects of biomassallocation to roots, nitrogen (N) uptake and photosyntheticN-use efficiency (PNUE) in determining plant growth responseto elevated CO₂. The model predicted that a rather substantialincrease in biomass allocation to root growth had little effecton whole plant growth response to elevated CO₂, suggesting thatplasticity in root allocation is relatively unimportant in determininggrowth response. Average N uptake rate at elevated comparedto ambient CO₂was decreased by 21–29%. In contrast, elevatedCO₂increased PNUE by approx. 50% due to a corresponding risein the CO₂-saturation factor for carboxylation at elevated CO₂.The model predicted that the decreased N uptake rate at elevatedCO₂lowered RGR modestly, but this effect was counterbalancedby an increase in PNUE resulting in a positive CO₂effect ongrowth. Increased PNUE may also explain why in many experimentselevated CO₂enhances biomass accumulation despite a significantdrop in tissue nitrogen concentration. The formulation of theFB model as presented here successfully predicted plant growthresponses to elevated CO₂. It also proved effective in resolvingwhich plant properties had the greatest leverage on such responses.Copyright 2000 Annals of Botany Company Elevated CO₂, functional balance model, Helianthus annuus L., N uptake, photosynthetic nitrogen use efficiency, root:shoot ratio     

Growth and partitioning of C and fixed N in the shrub legume Acacia littorea in the presence or absence of the root hemiparasite Olax phyllanthi

Nodulated plants of Acacia littorea were pot cultured singlyin minus nitrogen sand culture in the presence or absence ofa transplanted seedling of the root hemiparasite Olax phyllanthiand harvests of cultures made 4 and 8 months after introducingthe parasite. Parasitism decreased host shoot growth while increasingroot growth to a similar extent. Final shoot:root dry weightratio was 2.2 for parasitized versus 4.3 for unparasitized Acacia.Partitioning of fixed N showed 4-fold larger N increments inshoots than roots of unparasitized plants, whereas parasitizedplants lost a small amount of shoot N, made a root gain of Ndouble that of unparasitized plants and lost over half of theirN to Olax. The increment of fixed N in the host:parasite associationwas similar to that of unparasitized Acacia. Data on dry mattergain per unit foliage area and mean CO₂ assimilation rates pershoot of Olax and Acacia (parasitized or unparasitized) werediscussed in relation to an estimated heterotrophic gain ofxylem C from the host equivalent to 40% of the increment ofdry matter C made by the parasite. Growth of Olax was accompaniedby large increases in numbers of haustoria, 9% of which wereattached to root nodules as opposed to roots. Structural andnutritional features of direct parasitism of nodules are described.Models of flow and utilization of C and N in the Acacia:Olaxassociation and unparasitized Acacia are discussed in relationto published data for other host:parasite associations. Key words: Olax phyllanthi, host-parasite relationships, C and N partitioning, Acacia, N₂ fixation     

Effects of P deficiency on the uptake, flows and utilization of C, N and H2O within intact plants of Ricinus communis L.

The influence of P deficiency on the uptake, flow and utilizationof C, N and H₂0 by intact NO₃-fed castor bean plants {Ricinuscommunis L.) was studied over a 9 d period in the middle oftheir vegetative growth. The modelling techniques incorporateddata on net increments or losses of C, N and H₂O in plant parts,photosynthetic gains in and respiratory losses of C, molar C:Nratios of solutes in phloem and xylem sap and transpirationallosses of H₂0. Plant growth was inhibited within 3 d of withholdingP supply and dry matter production was less than one-third ofthe controls. Leaf growth was particularly depressed, whileroot growth was much less affected than that of the shoot. Shoot:rootratio of low-P plants was 1.5 compared with 2.6 under P supply.Over the 9 d study period total plant C and N increased by 560and 47 mmol, respectively, in the controls, but by only 113and 6.9 mmol in the low-P treatment. The particularly low incrementof N in P-deficient plants was due principally to decreasedN0₃^- uptake. Flows of C and N during the study period were markedlydifferent between control and P-deficient plants. The partitioningprofile for C in P-deficient plants showed a dramatic inhibitionof net photosynthesis and attendant photoassimilate flow. Proportionaldownward to upward allocation of carbon increased with increasein sink size of the root relative to shoot. This was reflectedin greater relative allocation of C to root dry matter and rootrespiration than in P-sufficient plants, and suppressed cyclingof C from root to shoot via xylem. Nitrogen intake and xylemtransport to the shoot of P-deficient plants were only 15% ofthe control and, as in the case of C, downward allocation ofN predominated over upward phloem translocation. Apart fromthese severe changes, however, the basic patterns of N flowsincluding xylem-to-phloem and xylem-to-xylem transfer of N werenot changed, a feature highlighting the vital nature of thesetransfer processes even under deficiency conditions. The alterationsin flows and partitioning of C, N and H₂O in response to low-Pconditions are discussed in relation to the corresponding effectsof moderate salt stress in Ricinus and the conclusion is reachedthat changes in nutrient flows under P deficiency were morehighly co-ordinated than when plants experience salt stress.Flow profiles under P deficiency which favour root growth andactivity are viewed as a means for increasing the potentialcapability of the plant to acquire P from the nutrient medium. Key words: Ricinus communis L., P deficiency, carbon, nitrogen, water, partitioning, xylem transport, phloem transport     

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     

Assimilate Partitioning in Red and White Clover Either Dependent on N2 Fixation in Root Nodules or Utilizing Nitrate Nitrogen

During vegetative growth in controlled environments, the patternof distribution of ¹⁴C-labelled assimilates to shoot and root,and to the meristems of the shoot, was measured in red and whiteclover plants either wholly dependent on N₂ fixation in rootnodules or receiving abundant nitrate nitrogen but lacking nodules. In experiments where single leaves on the primary shoot wereexposed to ¹⁴CO₂, nodulated plants of both clovers generallyexported more of their labelled assimilates to root (+nodules),than equivalent plants utilizing nitrate nitrogen, and thiswas offset by reduced export to branches (red clover) or stolons(white clover). The intensity of these effects varied with experiment.The export of labelled assimilate to growing leaves at the terminalmeristem of the donor shoot was not influenced by source ofnitrogen. Internode elongation in the donor shoot utilized nolabelled assimilate. Whole plants of white clover exposed to ¹⁴CO₂ on seven occasionsover 32 days exhibited the same effect on export to root (+nodules),which increased slightly in intensity with increasing plantage. Nodulated plants had larger root: shoot ratios than theirequivalents utilizing nitrate nitrogen. Trifolium repens, Trifolium pratense, red clover, white clover, nitrogen fixation, nitrate utilization, assimilate partitioning     

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     

Growth Analysis of Solution Culture-grown Winter Rye, Wheat and Triticale at Different Relative Rates of Nitrogen Supply

At low nitrogen (N) supply, it is well known that rye has ahigher biomass production than wheat. This study investigateswhether these species differences can be explained by differencesin dry matter and nitrogen partitioning, specific leaf area,specific root length and net assimilation rate, which determineboth N acquisition and carbon assimilation during vegetativegrowth. Winter rye (Secale cereale L.), wheat (Triticum aestivumL.) and triticale (X Triticosecale) were grown in solution cultureat relative addition rates (R_N) of nitrate-N supply rangingfrom 0.03–0.18 d^-1and at non-limiting N supply under controlledconditions. The relative growth rate (R_W) was closely equalto R_Nin the range 0.03–0.15 d^-1. The maximalR_W at non-limitingnitrate nutrition was approx. 0.18 d^-1. The biomass allocationto the roots showed a considerable plasticity but did not differbetween species. There were no interspecific differences ineither net assimilation rate or specific leaf area. Higher accumulationof N in the plant, despite the same relative growth rate atnon-limiting N supplies, suggests that rye has a greater abilityto accumulate reserves of nitrogen. Rye had a higher specificroot length over a wide range of sub-optimal N rates than wheat,especially at extreme N deficiency (R_N=0.03–0.06 d^-1).Triticale had a similar specific root length as that of wheatbut had the ability to accumulate N to the same amount as ryeunder conditions of free N access. It is concluded that thebetter adaptation of rye to low N availability compared to wheatis related to higher specific root length in rye. Additionally,the greater ability to accumulate nitrogen under conditionsof free N access for rye and triticale compared to wheat maybe useful for subsequent N utilization during plant growth.In general, species differences are explained by growth componentsresponsible for nitrogen acquisition rather than carbon assimilation.Copyright 1999 Annals of Botany Company Growth analysis, nitrogen, nitrogen productivity, partitioning, specific root length, Secale cereale L.,Triticum aestivum L., X Triticosecale, winter rye, winter wheat, winter triticale.     

The Influence of Plant Water Deficit on Distribution of 14C-labelled Assimilates in Cacao Seedlings

The relationship between plant water status and distributionof ¹⁴C-labelled assimilates in cacao (Theobroma cacao L.) wasevaluated after ¹⁴CO₂ pulse labelling leaves of seedlings subjectedto varying levels of water deficiency. The proportion of ¹⁴Cexported by source leaves was strongly affected by seedlingwater status. An increasing proportion of labelled assimilatesremained in source leaves at both 24-h and 72-h harvests aswater stress intensity increased. Water stress reduced the distributionof exported label to leaves and to the expanding flush in particularbut increased the proportion of label in stems and roots. Theresults suggest that current photoassimilates may be temporarilystored in source leaves and stems of cacao seedlings duringperiods of plant water deficit. The stress-induced changes inpartitioning of labelled carbon were in concordance with changesin shoot to root biomass ratios, which was likely due to greaterreduction in growth of above-ground organs to that of roots. Theobroma cacao L, assimilate partitioning, cacao, ¹⁴C-photoassimilate, water stress, water potential     

Influence of nitrogen on growth and photosynthesis of a C3 cereal, Oryza sativa, infected with the root hemiparasite Striga hermonthica

Striga hermonthica is a root hemiparasitic angiosperm nativeto the African semi-arid tropics. It is a major weed of C₄ cerealsbut locally it is also an important weed of the C₃ plant, rice[Oryza sativa). Infected rice plants produced 17% and 42% ofthe total biomass of uninfected plants when grown at two differentammonium nitrate concentrations, 1 and 3 mol m^–3, respectively.S. hermonthica prevented grain production at both concentrationsof nitrogen. At the lower concentration no heads were produced.At the higher concentration head weight was only 6% of uninfectedcontrols. S. hermonthica also altered the partitioning of drymatter between plant parts, such that shoot growth was reducedto a greater extent than root growth. As a consequence the root-to-shootratio of infected plants was approximately five times greaterthan that of uninfected control plants. Light saturated ratesof photosynthesis In infected plants were 56% and 70% of thoseof uninfected controls, at low and high nitrogen, respectively.Infection also led to lower values of stomatal conductance althoughthe substom-atal CO₂ concentration was unaffected. Analysisof the response of photosynthesis to substomatal CO₂ concentration(A/C_I curves) demonstrated that lower rates of photosynthesiscould not be solely attributed to lower stomatal conductances.Lower initial slopes and asymptotic rates suggest that bothcarboxylation and processes controlling regeneration of ribulose-1,5-bisphosphate are reduced by infection. The data are discussedwith respect to the influence of S. hermonthica on the growthand photosynthesis of C₄ hosts, where in contrast to the situationwith rice, nitrogen feeding results in a marked alleviationof the effects of the parasite on the host. Key words: Rice, Striga, growth, photosynthesis, nitrogen     

Growth of White Clover, Dependent on N2 Fixation, in Elevated CO2 and Temperature

Clonal plants of white clover (Trifolium repens L ), whollydependent on N₂ fixation, were grown for 6 weeks in controlledenvironments providing either (C680 regime) 23/18 °C day/nighttemperatures and a CO₂, concentration of 680 µmol mol^–1,or (C340 regime) 20/15 °C day/night temperatures and a CO₂,concentration of 340 µmol mol^–1 During the firsthalf of the experimental period the C680 plants grew fasterthan their C340 counterparts so that by week 3 they were twicethe weight this 2 1 superiority in weight persisted until theend of the experiment The faster initial growth of the C680plants was based on an approx 70 % increase in leaf numbersand an approx 30 % increase in their individual area Initially,specific leaf area (cm2 g^–1 leaf) was lower in C680 thanin C340 leaves but became similar in the latter half of theexperiment Shoot organ weights, including petioles and stolons,reflected the C680 plant's better growth in terms of photosyntheticsurface Throughout, C680 plants invested less of their weightin root than C340 plants and this disparity increased with timeAcetylene reduction assays showed that nitrogenase activityper unit nodule weight was the same in both C680 and C340 plantsBoth groups of plants invested about the same fraction of totalweight in nodules Nitrogen contents of plant tissues were similarirrespective of growth regime, but C680 expanded leaves containedslightly less nitrogen and their stolons slightly more nitrogenthan their C340 counterparts However, C680 leaves containedmore non-structural carbohydrate Young, unshaded C680 leavespossessed larger palisade cells, packed more tightly withinthe leaf, than equivalent C340 leaves The reason for the C680regime's loss of superiority in relative growth rate duringthe second half of the experiment was not clear, but more accumulationof non-structural carbohydrate, constriction of root growthand increased self-shading appear to be the most likely causes Trifolium repens, white clover, elevated CO₂, elevated temperature, growth, N₂ fixation, leaf structure     

The Response of Oxygen and Carbon Dioxide Exchanges and Root Activity to Short Term Water Stress in Soybean

The gross and net O₂ evolution together with O₂ uptake, CO₂assimilation, transpiration, shoot dark respiration, root respirationand ion uptake of a soybean plant were studied during 19 d whichincluded two periods of water stress. O₂ uptake was measuredusing ¹⁸O₂ as a tracer. Short term water stress induced immediateand lasting effects: (1) reduction of light interception bywilting, (2) limitation of the total reducing equivalent producedby the electron transport chain, (3) decrease of stomatal conductancereducing both losses of water and the entry of CO₂ for assimilation,(4) relative stimulation of O₂ uptake. The ratio of O₂ uptaketo CO₂ assimilation changed from 1.0 before stress to 1.4 forseveral days after. Root respiration was less affected by thestress than ion uptake and shoot gas exchanges. Key words: Photosynthesis, Photorespiration, Transpiration, Shoot and root respiration, Ion uptake, Water stress, Glycine max. L.     

Wheat Cultivars Respond Differently to a Drying Top Soil and a Possible Non-Hydraulic Root Signal

Research has shown that when plant roots are exposed to a dryingsoil a non-hydraulic (chemical) signal is produced in the rootand transported to the shoot, causing stomatal closure and growthretardation. This study was designed to reveal genetic diversityin wheat response to soil conditions which elicit a root signal,as the first step in the investigation of the genetic controlof the production of and the response to the root signal. Five spring wheat (Triticum aestivum L.) cultivars were establishedin the growth chamber in soil-filled polyvinyl chloride tubes,120 cm long and of an internal diameter of 10·2 cm. Soilwas well fertilized and wet to field capacity at emergence whentwo treatments were imposed: (1) tubes were watered from thetop as needed to eliminate stress (control); and (2) tubes hada constant water table at a soil depth of 100 to 120 cm, withno applied water. Measurements were performed on five dateson leaf water status and stomatal diffusive resistance. Above-groundbiomass and grain yield per plant were determined at maturity. The water table treatment resulted in dry and hard top soilconditions which were previously indicated to elicit a possibleroot signal. Under these experimental conditions, cultivarsdiffered in their leaf water status, stomatal diffusive resistance(R_s) and plant production. In the control treatment, R_s of cultivarsincreased with reductions in their relative water content (RWC)and leaf water potential (LWP), indicating the expected controlof R_s by leaf water status. Under conditions of a drying topsoil, relative water content (RWC) and leaf water potential(LWP) increased in cultivars that had a higher R_s, indicatingthat stomatal activity was controlling leaf water status. Itwas therefore suggested that the drying top soil elicited aroot signal which caused stomatal closure and reduced plantproduction. Under such conditions, two cultivars (Bethlehemand V748) consistently maintained relatively low R_s and highplant production, despite their relatively lower RWC and LWP,as compared with cvs C97, V747 and V652. Limited observationssuggest that in these two cultivars relatively fewer roots mayhave been exposed to the drying top soil, as compared with theother three cultivars. Key words: Triticum aestivum, cultivars, soil moistrue, drought stress, root, root signal, stomata, relative water content, leaf water potential, biomass, yield     

Direct and Maternal Effects of Elevated CO2on Early Root Growth of GerminatingArabidopsis thalianaSeedlings

Individuals ofArabidopsis thaliana, collected in different naturalpopulations, were grown in controlled and elevated CO₂in a glasshouse.Following germination, root growth of progeny of different linesof these populations was studied in control and elevated atmosphericCO₂. No significant direct effect of atmospheric CO₂concentrationcould be demonstrated on root growth. An important parentaleffect was apparent, namely that root length and branching weredecreased in seeds collected from a mother plant which had beengrown in elevated CO₂. This was correlated with smaller seeds,containing less nitrogen. These parental effects were geneticallyvariable. We conclude that CO₂may affect plant fitness via parentaleffects on seed size and early root growth and that the geneticvariability shown in our study demonstrates thatArabidopsispopulationswill evolve in the face of this new selective pressure.Copyright1998 Annals of Botany Company Root growth, root branching, seed, elevated CO₂, natural population,Arabidopsis thaliana, parental effect.     

Temporal and Nutritional Influences on the Response to Elevated CO2 in Selected British Grasses

To investigate the duration of the CO₂ response and its interactionwith mineral nutrition, CO₂-enrichment experiments were performedon four British grasses of differing ecology and functionaltype: Arrhenatherum elatius (L.) Beauv., Festuca ovina L., Festucarubra L. and Poa annua L. Naturally-lit, glasshouse cabinetswere used, with a non-limiting water supply and a daytime meantemperature of 18 °C. Two CO₂ treatments were maintainedat nominal concentrations of 350 and 700 vpm and were combinedfactorially with two levels of balanced mineral nutrition atconductivities of 0·1 and 1 mS cm^-1. Harvests took placeat planting-out, and at 16, 37 and 58 d thereafter. Fitted curves were used to derive instantaneous values of totaldry weight, relative growth rate (RGR), shoot weight fraction(SWF) and unit shoot rate (USR) for all combinations of species,CO₂ level, nutrient level and time of harvesting. At the higher nutrient level there was a reasonably close agreementwith previous estimates of the CO₂ response in the four species.The response, if any, most often arose from an increase in USRbeing accompanied by a less than proportionate decline in SWF.Responses were sustained throughout the period studied. At thelower nutrient level, all species showed a CO₂ response initially,but this declined at a rate which was inversely related to theCO₂-responsiveness of the species at the higher nutrient level. The underlying ontogenetic drift appeared to be markedly towardsadjustment in SWF and away from that of USR. However, this driftwas retarded, suspended or even reversed by low-nutrient conditionsand/or by high CO₂ responsiveness in the species itself.Copyright1995, 1999 Academic Press Climate change, CO₂ enrichment, plant strategies, mineral nutrition, growth analysis, relative growth rate, shoot weight fraction, unit shoot rate, functional equilibria     

Effect of CO2 Concentration on Growth, Carbon Distribution and Loss of Carbon from the Roots of Maize

The effects of three ranges of CO₂ concentration on growth,carbon distribution and loss of carbon from the roots of maizegrown for 14 d and 28 d with shoots in constant specific activity¹⁴CO₂ are described. Increasing concentrations of CO₂ led toenhancement of plant growth with the relative growth rate (RGR)of the roots affected more than the RGR of the shoots. Between16% and 21% of total net fixed carbon (defined as ¹⁴C retainedin the plant plus ¹⁴C lost from the root) was lost from theroots at all CO₂ concentrations at all times but the amountsof carbon lost per unit weight of plant decreased with time.Possible mechanisms to account for these observations are discussed. Key words: Growth, Roots, Carbon loss, [CO₂]     

The Growth and Yield Responses ofFragaria ananassato Elevated CO2and N Supply

Strawberry plants (Fragaria ananassaDuchesne var. Elsanta) weregrown in pots at two concentrations of carbon dioxide (partialpressures of 39 and 56 Pa) and with three rates of nitrogensupply (0.04, 0.4 and 4 mMas nutrient solution) to study theirindividual and interactive effects on plant growth and fruityield. Nitrogen deficiency reduced total dry biomass and relativegrowth rate (RGR), mainly through reductions in leaf area ratio(LAR) and plant N concentration (PNC), although both the netassimilation rate (NAR) and root weight ratio (RWR) increased.Elevated CO₂increased the N productivity (NP) but reduced theLAR. High CO₂increased the fruit yield by 42% at high N supplyand by 17% at low N supply. The CO₂yield enhancement occurredthrough an increase in the flower and fruit number of individualplants. This resulted in an increase in the fruit weight ratio(FWR) of plants at high CO₂. Nitrogen deficiency reduced thefruit yield by about 50% through decreases in fruit size, fruitset and the number of fruits. However, N deficiency increasedthe proportion of total plant dry biomass allocated to fruits.There were no significant interactions between CO₂and N supplyon yield.Copyright 1998 Annals of Botany Company Nitrogen; carbon dioxide; strawberry (Fragaria ananassaDuchesne); fruit yield.     

The Carbon Balance of a Legume and the Functional Economy of its Root Nodules

Budgets for carbon and nitrogen in shoot, root, and nodulesof garden pea (Pisum sativum L.) are drawn up for a 9-d intervalin the life cycle, from data on nitrogen fixation, carbon accumulationin dry matter, respiratory output of plant organs, and organicsolute exchange between shoot and nodulated root. Of the carbon gained photosynthetically by the shoot from theatmosphere 26 per cent is incorporated directly into its drymatter, 32 per cent translocated to the nodules, and 42 percent to the supporting root. Of the nodules’ share, 5per cent is consumed in growth, 12 per cent in respiration,and 15 per cent returned to the shoot via the xylem, as aminocompounds generated in nitrogen fixation. Growth and respirationof the root utilize, respectively, 7 and 35 per cent. The respiratory efficiency of a nodulated root in terms of nitrogenfixation (5.9mg C per mg N₂-N fixed) is found to be very similarto that of an uninoculated root assimilating nitrate (6.2 mgC per mg NO₃-N reduced). The nodules require in growth, respiration,and export 4.1 mg C ( 10.3 mg carbohydrate) for each mg N whichthey fix. The nodules consume 3 ml O₂ for every 1 ml N₂ utilized in fixation. In exporting a milligram of fixed nitrogen the nodules requireat least 0.35 ml of water. Almost half of this requirement mightbe met by mass flow into the nodules via the phloem.