Interaction of soil moisture and elevated CO2 on the above-ground growth rate, root length density and gas exchange of turves from temperate pasture

Interactions between water availability and elevated atmosphericCO₂ concentrations have the potential to be important factorsin determining future forage supply from temperate pastures.Using large turves from an established pasture, the responseof these communities at 350 or 700 l l^–1 CO₂ to a soilmoisture deficit and to recovery from the deficit in comparisonto turves that were well-watered throughout was measured. Priorto this experiment the turves had been exposed to the CO₂ treatmentsfor 324 d. Net CO₂ exchange continued at elevated CO₂ even when the volumetricsoil moisture content was less than 0.10 m³ m^–3 soil;at the same moisture deficit gas exchange at ambient CO₂ waszero. The additional carbon fixed by the elevated CO₂ turveswas primarily allocated below-ground as shown by the maintenanceof root length density at the same level as in well-wateredturves. When the dry turves were rewatered there was compensatorygrowth at ambient CO₂ so that the above-ground growth rate exceededthat of turves that had not experienced a moisture deficit.At the start of this experiment, the turves that were growingat 700 l I^–1 CO₂ had a greater proportion of legume (principallywhite clover, Trifolium repens L.) in the harvested herbage.There was a trend for the legume content at elevated CO₂ tobe reduced under a soil moisture deficit. The results indicate different strategies in response to soilmoisture deficits depending on the CO₂ concentration. At ambientCO₂, growth stopped, but plants were able to respond stronglyon rewatering; while at elevated CO₂ growth continued (particularlybelow-ground), but no additional growth was evident on rewatering.Ecosystem gas exchange measurements taken at the end of theexperiment (after 429 d of exposure to CO₂) showed 33% moreCO₂ was fixed at elevated CO₂ with only a small (12%) and nonsignificantdownward regulation. Key words: Carbon dioxide, climate change, grassland, gas exchange, soil moisture deficit     

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     

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     

Effect of Temperature and Nitrogen Supply on the Growth of Perennial Ryegrass and White Clover. 1. Carbon and Nitrogen Economies of Mixed Swards at Low Temperature

Simulated mixed swards of perennial ryegrass (Lolium perenneL. cv. S23) and white clover (Trifolium repens L. cv. S100)were grown from seed under a constant 10°C day/8°C nighttemperature regime and their growth, and carbon and nitrogeneconomies examined. The swards received a nutrient solution,every second day, which contained either high (220 µgg^–1) or low (40 µg g^–1) nitrate N. The High-N swards had rates of canopy photosynthesis and drymatter production (over the linear phase of growth) similarto those previously shown by mixed swards at high temperature.The Low-N swards grew more slowly; canopy photosynthesis, ata given LAI, was similar to that at High-N but lower LAI's weresustained. Clover increased its contribution to total carbonuptake and total dry weight throughout the period in the Low-Ntreatment and, despite the fact that grass took up most of theavailable nitrate, clover maintained a consistently higher Ncontent by virtue of N₂-fixation. At High-N, grass dominated throughout the measurement period.Earlier, when plants grew as spaced individuals, clover grewless well than grass, but once the canopy was closed it hada similar relative growth rate and thus maintained a steadyproportion of total sward dry weight. It is proposed that earlyin the development of the crop, leaf area production is thelimiting factor for growth, and that in this respect cloveris adversely affected by low temperature relative to grass.Later, as the LAI of the crop builds up, and the canopy becomesfully light intercepting, net canopy photosynthesis plays amore dominant role and here the higher photosynthetic rate perunit leaf area of the clover is crucial. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, low temperature, nitrogen, photosynthesis     

Effect of Temperature and Nitrogen Supply on the Growth of Perennial Ryegrass and White Clover. 2. A Comparison of Monocultures and Mixed Swards

White clover (Trifolium repens L.) and Perennial ryegrass (Loliumperenne L.) plants were grown, in Perlite, in simulated swardsas either monocultures or mixtures of equal plant numbers. Theywere supplied with a nutrient solution either high (220 µgg^–1) or low (40 µg g^–1) in ¹⁵N-labelled nitrateand grown to ceiling yield at either high (20°C day/15°Cnight) or low (10°C day/8°C night) temperature. Temperature had little effect on the maximum rates of grosscanopy photosynthesis which were similar in High-N grass andHigh-N and Low-N clover monocultures. However these maxima werereached more slowly in clover than grass, and more slowly atlow rather than high temperature. Nitrogen supply increasedphotosynthesis in grass but not in clover. Clover had higherN contents than grass in all four treatments, although in anygiven treatment its N content was lower, and contribution ofN₂-fixation relative to nitrate uptake higher, in mixture thanin monoculture. Conversely, grass had higher N contents in mixturethan monoculture, because more nitrate was available per plantand not because of transfer of biologically fixed N from clover. Under Low-N, clover outyielded grass in mixture, particularlyat high temperature. The grass plants in the Low-N mixtureshad higher N contents and higher SLA, LAR and shoot: root ratiosthan those in monoculture. It is proposed that competition forlight is the cause of the low relative yield and negative aggressivityof grass in these swards. Under High-N, grass outyielded cloverin monoculture and mixture, at both temperatures but particularlyat low temperature when grass had a high aggressivity. Nitrogenand yield component analyses shed no light on clover's apparentlylow competitive ability and evidence is drawn from the previouspaper to demonstrate that grass grew faster than clover onlyas spaced individuals during non-com petitive growth. The relativemerits of measures of competitive ability based on final harvestdata and physiological data taken over a growth period are discussed. Trifolium repens L., white clover, Lolium perenne, perennial ryegrass, competition, temperature, nitrogen     

Components of Growth and Dark Respiration of Kikuyu (Pennisetum clandestinum Chiov.) at Various Temperatures

Growth and dark respiration were measured in dense, miniatureswards of kikuyu grass grown at constant temperatures of 15,20, 25 and 30 °C. Total respiration over the first 12 hof darkness was very high and CO² efflux per unit surface areavaried from 2.4 to 3.9 g CO₂ m^–2 h^–1 at 15 and 30°C respectively. Such rates were consistent with the correspondinglyhigh net growth rates of 24 and 63 g d. wt m^–2 d^–1and the heavy yields of herbage. When plants were kept in thedark, CO₂ efflux subsequently declined rapidly to a lower, constantrate which was taken to be the maintenance respiration rate.The half-life of the declining phase of respiration averaged10.9 and 6.0 h at 15 and 30 °C respectively, and was curvilinearlyrelated to the specific maintenance respiration rate (m). Therapid decline in respiration was consistent with the low concentrationsof total soluble carbohydrate and starch in the herbage. Valuesof m for lamina and top growth increased with temperature witha Q₁₀ of 2.6 and 1.42 respectively, but m of stems alone wasnot affected by temperature. Using results from this study forkikuyu and from McCree (1974) for sorghum and white clover,it was noted that all three species have similar m when grownat temperatures which are near their respective optimums forgrowth. Kikuyu, Pennisetum clandestinum, growth, respiration, temperature     

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     

Response of Soybean Canopy Photosynthesis to CO2 Concentration, Light, and Temperature

Photosynthetic rates of outdoor-grown soybean (Glycine max L.Merr. cv. Bragg) canopies increased with increasing CO₂ concentrationduring growth, before and after canopy closure (complete lightinterception), when measured over a wide range of solar irradiancevalues. Total canopy leaf area was greater as the CO₂ concentrationduring growth was increased from 160 to 990 mm³ dm^–3.Photosynthetic rates of canopies grown at 330 and 660 mm³ CO₂dm^–3 were similar when measured at the same CO₂ concentrationsand high irradiance. There was no difference in ribulose bisphosphatecarboxylase/oxygenase (rubisco) activity or ribulose 1,5-bisphosphate(RuBP) concentration between plants grown at the two CO₂ concentrations.However, photosynthetic rates averaged 87% greater for the canopiesgrown and measured at 660 mm³ CO₂ dm^–3. A 10°C differencein air temperature during growth resulted in only a 4°Cleaf temperature difference, which was insufficient to changethe photosynthetic rate or rubisco activity in canopies grownand measured at either 330 or 660 mm³ CO₂ dm^–3. RuBP concentrationsdecreased as air temperature during growth was increased atboth CO₂ concentrations. These data indicate that the increasedphotosynthetic rates of soybean canopies at elevated CO₂ aredue to several factors, including: more rapid development ofthe leaf area index; a reduction in substrate CO₂ limitation;and no downward acclimation in photosynthetic capacity, as occurin some other species. Key words: CO₂ concentration, soybean, canopy photosynthesis     

Effect of Elevated CO2 on the Photosynthesis, Respiration and Growth of Perennial Ryegrass

Single, seed-grown plants of ryegrass (Lolium perenne L. cv.Melle) were grown for 49 d from the early seedling stage ingrowth cabinets at a day/night temperature of 20/15 C, witha 12 h photoperiod, and a CO₂ concentration of either 340 or680µI 1^–1 CO₂. Following complete acclimation tothe environmental regimes, leaf and whole plant CO₂ effluxesand influxes were measured using infra-red gas analysis techniques.Elevated CO₂ increased rates of photosynthesis of young, fullyexpanded leaves by 35–46% and of whole plants by morethan 50%. For both leaves and whole plants acclimation to 680µI^–1 CO₂ reduced rates of photosynthesis in bothCO₂ regimes, compared with plants acclimated to 340µll^–1. There was no significant effect of CO₂ regime onrespiration rates of either leaves or whole plants, althoughleaves developed in elevated CO₂ exhibited generally lower ratesthan those developed in 340µI I^–1 CO₂. Initially the seedling plants in elevated CO₂ grew faster thantheir counterparts in 340µI I^–1 CO₂, but this effectquickly petered out and final plant weights differed by onlyc. 10%. Since the total area of expanded and unexpanded laminaewas unaffected by CO₂ regime, specific leaf area was persistently13–40% lower in elevated CO₂ while, similarly, root/shootratio was also reduced throughout the experiment. Elevated CO₂reduced tissue nitrogen contents of expanded leaves, but hadno effect on the nitrogen contents of unexpanded leaves, sheathsor roots. The lack of a pronounced effect of elevated CO₂ on plant growthwas primarily due to the fact that CO₂ concentration did notinfluence tiller (branch) numbers. In the absence of an effecton tiller numbers, any possible weight increment was restrictedto the c. 2.5 leaves of each tiller. The reason for the lackof an effect on tillering is not known. Key words: Lolium perenne, ryegrass, elevated CO₂, photosynthesis, respiration, growth, development     

Effect of Elevated CO2 on Stomatal Size and Distribution in Perennial Ryegrass

Plants of ryegrass (Lolium perenne L. cv. Melle) were grownfrom the early seedling stage in growth cabinets at a day/nighttemperature of 20/15 °C, with a 12-h photoperiod, and aCO₂ concentration of either 340 or 680 ± 15 µl1^–1 CO₂. Young, fully-expanded, acclimated leaves fromprimary branches were sampled for length of stomata, and ofepidermal cells between stomata, numbers of stomata and epidermalcells per unit length of stomatal row, numbers of stomatal rowsacross the leaf and numbers of stomatal rows between adjacentvein ridges. Elevated CO₂ had no significant effect on any ofthe measured parameters. Elevated CO₂, Lolium perenne, ryegrass, stomatal distribution, stomatal size     

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

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

Effects of elevated carbon dioxide on three grass species from montane pasture II. Nutrient uptake, allocation and efficiency of use

Agrostis capillaris L.⁴ Festuca vivipara L. and Poa alpinaL.were grown in outdoor open-top chambers at either ambient (340µmol mol^–1) or elevated (680 µmol^–1)CO² for periods from 79 to 189 d. Under these conditions thereis increased growth of A. caplllarls and P. alpina, but reducedgrowth of F. vivipara. Nutrient use efficiency, nutrient productivity(total plant dry weight gain per unit of nutrient) and nutrientallocation of all three grass species were measured in an attemptto understand their individual growth responses further andto determine whether altered nutrient-use efficiencies and productivitiesenable plants exposed to an elevated atmospheric CO₂ environmentto overcome potential limitations to growth imposed by soilfertility. Total uptake of nutrients was, in general, greater in plantsof A. capillaris and P. alpina (with the exception of N andK in the latter) when grown at 680 µmol mol^–1 CO₂.In F. vivipara, however, uptake was considerably reduced inplants grown at the higher CO₂ concentration. Overall, a doubling of atmospheric CO₂ concentration had littleeffect on the nutrient use efficiency or productivity of A.capillaris. Reductions in tissue nutrient content resulted fromincreased plant growth and not altered nutrient use efficiency.In P. alpina, potassium, magnesium and calcium productivitieswere significantly reduced and photosynthetic nitrogen and phosphorususe efficiencies were doubled at elevated CO₂ with respect toplants grown at ambient CO₂ F. vivipara grown for 189 d showedthe most marked changes in nutrient use efficiency and nutrientproductivity (on an extracted dry weight basis) when grown atelevated CO₂, F. vivipara grown at elevated CO₂ however, showedlarge increases in the ratio of non-structural carbohydrateto nitrogen content of leaves and reproductive tissues, indicatinga substantial imbalance between the production and utilizationof assimilate. Key words: Nutrient, allocation, nutrient use efficiency, grasses, nutrient productivity, elevated CO₂, cliniate change     

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     

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     

Concurrent Rates of N2 Fixation, Nitrate and Ammonium Uptake by White Clover in Response to Growth at Different Root Temperatures

Nodulated white clover plants (Trifolium repens L. cv. Huia)were grown for 71 d in flowing nutrient solutions containingN as 10 mmol m_–3 NH₄NO₃, under artificial illumination,with shoots at 20/15°C day/night temperatures and root temperaturereduced decrementally from 20 to 5°C. Root temperatureswere then changed to 3, 7, 9, 11, 13, 17 or 25°C, and theacquisition of N by N₂ fixation, NH₄+ and NO₃^– uptakewas measured over 14 d. Shoot specific growth rates (d. wt)doubled with increasing temperature between 7 and 17°C,whilst root specific growth rates showed little response; shoot:root ratios increased with root temperature, and over time at11°C. Net uptake of total N per plant (N₂ fixation + NH₄++ NO₃^–) over 14 d increased three-fold between 3 and 17°C.The proportion contributed by N₂ fixation decreased with increasingtemperature from 51% at 5°C to 18% at 25°C. Uptake ofNH₄+ as a proportion of NH₄+ + NO₃^– uptake over 14 d variedlittle (55–62%) with root temperature between 3 and 25°C,although it increased with time at most temperatures. Mean ratesof total N uptake per unit shoot f. wt over 14 d changed littlebetween 9 and 25°C, but decreased progressively with temperaturebelow 9°C, due to the decline in the rates of NH₄+ and NO₃^–uptake, even though N₂ fixation increased. The results suggestthat N₂ fixation in the presence of sustained low concentrationsof NH₄+ and NO₄^– is less sensitive to low root temperaturethan are either NH₄+ or NO₃^– uptake systems. White clover, Trifolium repens L. cv. Huia, root temperature, nitrogen fixation, ammonium, nitrate     

In situ Effects of Elevated Atmospheric CO2on Leaf Freezing Resistance and Carbohydrates in a Native Temperate Grassland

The objectives of this study were to quantify changes in leaffreezing resistance and carbohydrate concentrations caused bylong-term (6 years) exposure to elevated CO₂(ambient: 360 µll^-1, elevated: 600 µl l^-1) in five dominant plant speciesgrowing in situ in a native temperate grassland. Across allfive species tested from three functional groups, the mean temperatureat which all leaves were damaged (T₁₀₀) significantly (P = 0.016)increased from -9.6 to -8.5 °C under elevated CO₂ , anda similar marginally significant (P = 0.079) reduction was observedfor the mean temperature that caused 50% leaf damage (T₅₀),from -6.7 to -6.0 °C. The mean temperature at which initialleaf damage was observed (T₀) was not significantly influencedby elevated CO₂ . Although concentrations of soluble sugars(+25%,P = 0.042), starch (+53%, P < 0.001), and total non-structuralcarbohydrates (TNC, +40%, P < 0.001) were significantly higherunder elevated CO₂ , leaf freezing resistance actually decreasedunder elevated CO₂ . Concentrations of soluble sugars were positivelycorrelated with freezing resistance when viewed across all fivecommunity dominants, but within any individual species, no suchrelationships were found. We also found no evidence for ouroriginal hypothesis that increased concentrations of solublesugars increase freezing resistance. Thus, future atmosphericCO₂levels may instead increase the risk of late spring freezingdamage. Furthermore, the strong differences in freezing resistanceobserved among the species, along with decreased freezing resistance,may increase the risk of losing species that have inherentlyweak freezing resistances from the plant community. Copyright2001 Annals of Botany Company CO₂enrichment, frost hardiness, sugar, starch, total non-structural carbohydrates (TNC)     

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

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

The Growth and Photosynthesis of Seedling Plants of White Clover at Low Temperature

The growth of white clover (Trifolium repens L.) in conditionstypical of April in Southern England (8 °C day/4 °Cnight, 12 h photoperiod of 90 J m^–2 s^–1 visibleradiation) was extremely slow, whether the plants were dependentfor nitrogen on fixation by their root nodules or were suppliedwith abundant nitrate; although growth was slower in the nodulatedplants. The reasons for slow growth were a large root: shootratio and a small leaf area, particularly in the nodulated plants,and a low photosynthetic rate in all plants. The probable effectsof these characteristics on the growth of white clover withgrasses in mixed pastures are discussed. Trifolium repens L, white clover, low temperature, leaf area, photosynthetic rate, nitrogen supply, growth     

Contrasting CO2 and temperature effects on leaf growth of perennial ryegrass in spring and summer

The effects of increased atmospheric carbon dioxide (CO₂) of700 µmol mol^–1 and increased air temperature of+ 4C were examined in Lolium perenne L. cv. Vigor, growingin semi-controlled greenhouses. Leaf growth, segmental elongationrates (SER), water relations, cell wall (tensiometric) extensibility(%P) and epidermal cell lengths (ECL) were measured in expandingleaves in spring and summer. In elevated CO₂, shoot dry weight (SDW) increased in mid-summer.In both seasons, SDW decreased in elevated air temperatureswith this reduction being greater in summer as compared to spring.Specific leaf area (SLA) decreased in elevated CO₂ and in CO₂ temperature in both seasons. In spring, increased leaf extensionand SER in elevated CO₂ were linked with increased ECL, %P andfinal leaf size whilst in summer all were reduced. In high temperature,leaf extension, SER, %P and final leaf size were reduced inboth seasons. In elevated CO₂ temperature, leaf extension,SER, %P, and ECL increased in spring, but final leaf size remainedunaltered, whilst in summer all decreased. Mid-morning waterpotential did not differ with CO₂ or temperature treatments.Leaf turgor pressure increased in elevated CO₂ in spring andremained similar to the control in summer whilst solute potentialdecreased in spring and increased in summer. Contrasting seasonalgrowth responses of L. perenne in response to elevated CO₂ andtemperature suggests pasture management may change in the future.The grazing season may be prolonged, but whole season productivitymay become more variable than today. Key words: Lolium perenne, ryegrass, CO₂ and temperature, leaf extension, cell wall rheology     

Carbon Exchange Rate of Intact Individual Soya Bean Pods. 2. Ontogeny of Temperature Sensitivity

Diurnal temperature fluctuations induced change in soya bean-pod[Glycine max (L.) Merr.] carbon exchange rate (CER, where positiveCER represents CO₂ evolution). CER appeared to depend linearlyon temperature. Linear regressions of CER on temperature interceptedthe temperature axis at 5°C (i.e. zero CER at 5°C).Slopes of these regressions (i.e. temperature sensitivity) changedover the season. The CER-temperature sensitivity coefficient,K, (calculated from observed values of CER. pod temperatureand temperature intercept) rose from less than 0·02 mgCO₂ h^–1 pod^–1 °C^–1 during early pod-flll,peaked at over 0·04 mg CO₂ h^–1 pod^–1 °C^–1at mid pod-fill, and then declined during late pod-fill andmaturation. Glycine max (L.) Merr., Soya bean, carbon exchange rate, temperature