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 Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions II. Nitrogen Fixation and Nitrate Uptake

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and harvested at intervals over and88 d growht period. The swards received a nutrient solutiondaily, which was either High (220 mg l^–1) or Low (10 mgl^–1) in nitrate N. The nitrate was labelled with the ¹⁵Nisotope. An acetylene reduction assay was carried out on eachsward just prior to harvest. Rates of acetylene reduction agreed qualitatively with the ^l5Nanalyses but absolute values did not match (assuming a 4:1 C₂H₄:N₂ratio) and errors in the acetylene assay are discussed. In theLow-N swards clover relied almost entirely on symbioticallyfixed N₂, fixing more than ten times as much as the High-N cloverplants. In the Low-N treatment the grass was N-deficient despiteobtaining much more nitrate per unit root dry weight than clover.In the High-N swards, however, clover took up more nitrate perunit root weight than grass. The High-N clover plants also fixedsome N₂ and maintained a higher total-N content than grass throughoutthe period. There was no evidence of transfer of symbioticallyfixed N from the clover to the grass in either treatment. Trifolium repens, Lolium perenne, nitrate, nitrogen fixation, ¹⁵N, acetylene reduction     

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     

The Use of a Closed System Flow-Through Enclosure Apparatus for Studying the Effects of Partial Pressure of Dinitrogen in the Atmosphere on Growth of Trifolium repens L. and Lolium perenne L.

A closed system flow-through enclosure apparatus was constructedand used to enclose mixtures ofLolium perenne L. (perennialryegrass cv. Trani) and nodulatedTrifolium repens L. (whiteclover cv. Blanca) growing in soil in pots. There were no significantdifferences between the shoot growth, in terms of dry matteraccumulation and nitrogen content, of mixtures in the systemcompared to that of mixtures grown in a standard growth cabinet.This demonstrated that stable ambient conditions could be achievedby the closed system and its control circuits and that therewere no apparent side-effects of the recirculatory gases.Reducingthe partial pressure of dinitrogen in the atmosphere affectedwhite clover, but not perennial ryegrass. A fairly rapid effectwas observed 4 h after reduction in dinitrogen partial pressureas some of the clover leaves folded downwards along the petiole.These same effects were observed at two different partial pressuresof dinitrogen (22% and 39%) and with two different replacementgases (argon and helium). In the longer term (11 to 16 d) drymatter accumulation and nitrogen content of the clover shootswere significantly reduced. These effects of reduced partialpressure of dinitrogen were observed in both nodulated (NOD⁺)and nitrate-dependent (NOD^–) clover. Possible reasonsfor these effects were discussed with particular reference toimpurities in the gases used, stomatal responses and plant waterrelations. It was concluded that the closed system flow-throughapparatus provides a useful tool for studying whole plant-soilsystems and, in particular, the cycling of nitrogen. However,the use of a replacement gas to reduce the cost of labelleddinitrogen was obviously not a viable proposition. Key words: Dinitrogen, flow-through system, Lolium perenne, partial pressure, Trifolium repens     

The Effect of Shade During Leaf Expansion on Photosynthesis by White Clover Leaves

In three experiments measurements of photosynthesis were madeon single leaves of white clover (Trifolium repens L.) on threecultivars grown in a controlled environment. Plants which had grown under an irradiance of 30 J m^–2s^–1, or in shade within a simulated mixed sward, producedleaves with photosynthetic capacities some 30 per cent lowerthan did plants grown at 120 J m^–2 s^–1 without shade.There were no differences between treatments either in photosynthesismeasured at 30 J m^–2 s^–1, or in respiration ratesper unit leaf dry weight. Respiration per unit leaf area washigher in the plants grown at 120 J m^–2 s^–1, reflectingthe lower specific leaf area of these leaves. There were nodifferences between the three cultivars examined. Leaves which were removed from the shade of a simulated swardshortly after becoming half expanded achieved photosyntheticcapacities as high as those which were in full light throughouttheir development. It is suggested that it is this characteristicwhich enables clover plants growing in an increasingly densemixed sward to produce a succession of leaves of high photosyntheticcapacity, even though each lamina only reaches the top of thesward at a relatively late stage in its development. Trifolium repens L., white clover, photosynthesis, leaf expansion, shade, specific leaf area, stomatal conductance     

Effect of Previous Defoliation Regime and Mineral Nitrogen on Regrowth in White Clover Swards: Photosynthesis, Respiration, Nitrogenase Activity and Growth

Small swards of white clover (Trifolium repens L.) cv. Haifawere grown in solution culture in a controlled environment at24 °C day/18 °C night and receiving 500 µE m^-2S^–1 PAR during a 14-h photoperiod. The swards were cuteither frequently (10-d regrowth periods) or infrequently (40-dregrowth) over 40 d before being cut to 2 cm in height. Halfof the swards received high levels of nitrate (2–6 mMN in solution every 2 d) after defoliation while the othersreceived none. Changes in d. wt, leaf area and growing pointnumbers were recorded over the following 10 d. CO₂ exchangewas measured independently on shoots and roots and nitrogenase-linkedrespiration was estimated by measuring nodulated root respirationat 21% and 3% oxygen in the root atmosphere. There was a general pattern in all treatments consisting ofan initial d. wt loss from roots and stubble and reallocationto new leaves, followed by a period of total d. wt gain andrecovery, to a greater or lesser extent, of weight in non-photosyntheticparts. Frequently cut swards had a smaller proportion of theirshoot d. wt. removed by cutting and had a greater shoot d. wt,growing point number and leaf area at the start of the regrowthperiod. As a result of these differences, and also because ofdifferences in relative growth rates, frequently cut swardsmade more regrowth than infrequently cut. Initial photosyntheticrates were higher in frequently cut swards, although the laminaarea index was very low, and it was concluded that stolons andcut petioles made a significant contribution to carbon uptakeduring the first few d. Infrequently cut swards continued toallocate carbon to new and thinner leaves at the expense ofroots and stubble for longer than frequently cut swards andas a result achieved a similar lamina area index after 10 d. Nitrogenase-linked respiration was low in all treatments immediatelyafter cutting: frequently cut swards receiving no nitrate maintainedhigh nitrogenase activity, whereas recovery took at least 5d in infrequently cut swards. Swards which received nitrateafter cutting maintained only low rates of nitrogenase-linkedrespiration and their total nodulated root respiration overthe period was lower than those receiving no nitrogen: greaterregrowth in nitrate fed swards over the 10 d compared to N₂-fixingswards was in proportion to this lower respiratory burden. White clover (Trifolium repens L.), defoliation, regrowth, nitrogen, photosynthesis, respiration, nitrogenase-activity     

Effects of Temperature on Growth and Nitrogen Fixation by Swards of Subterranean Clover

Subterranean clover (Trifolium subterraneum L.) cv. ‘Woogenellup’ swards were grown at 10, 15, 20 and 25 Cwith a 12 h photoperiod of 500 or 1000 µmol m^–2s^–1 [low and high photosynthetic photon flux density (PPFD)].Nitrogen-fixing swards received nutrient solution lacking combinednitrogen while control swards received a complete nutrient solution.Growth was measured by infra-red analysis of carbon dioxideexchange and by accumulation of dry matter. Swards were harvestedat intervals between 95 and 570 g d. wt m^–2 for estimationof nitrogenase activity by acetylene reduction and hydrogenevolution assays. Nitrogen fixation was also measured by increasein organic nitrogen. The growth rate was highest at 10 C at low PPFD, and at 10–15C at high PPFD. Nitrogen-fixing swards grew slower than thosereceiving combined nitrogen. Nitrogen fixation measured by increasein organic nitrogen responded similarly to the growth rate,as did acetylene reduction between 10 and 20 C. At 25 C therelationship between acetylene reduction and nitrogen fixationwas distrupted. The difference between the rates of acetylenereduction and hydrogen evolution, theoretically proportionalto nitrogen fixation, was not a reliable indicator of nitrogenfixation because hydrogen uptake developed. Trifolium subterraneum L, subterranean clover, growth, nitrogen fixation, temperature, acetylene reduction     

The Sensitivity of Net Photosynthesis in Several Plant Species to Short-term Fumigation with Sulphur Dioxide

The effects of exposure of up to 2 h with sulphur dioxide ona range of plant species was observed by measuring changes inthe rate of net photosynthesis under closely controlled environmentalconditions. Ryegrass, Lolium perenne ‘S23’ was thespecies most sensitive to SO₂; significant inhibition was detectedat 200 nl l^–1. Fumigations at 300 nl l^–1 also inhibitedphotosynthesis in field bean (Vicia faba cv. ‘Three FoldWhite’ and ‘Blaze’) and in barley (Hordeumvulgare cv. ‘Sonja’). No effect was detected inwheat (Triticum aestivum cv. ‘Virtue’) at concentrationsup to 600 nl l^–1 SO₂, or in oil-seed rape (Brassica napuscv. ‘Rafal’) except at 800 nl l^–1 SO₂). Recoverycommenced immediately after the fumigation was terminated andwas complete within 2 h when inhibition had not exceeded 20%during the SO₂ treatment. Key words: Sulphur dioxide, short-term fumigation, photosynthesis     

The Effect of Nitrogenous Fertilizer on the Photosynthesis and Growth of White Clover/Perennial Ryegrass Swards

The application of nitrogenous fertilizer in March to a whiteclover (cv. Blanca) and perennial ryegrass (cv. S23) sward resultedin a rapid suppression of the clover, relative to clover ina treatment given no added nitrogen. Thereafter, the cloverin both treatments grew more rapidly than the grass and itsproportion of the total leaf area in the mixture increased,as the leaf area index rose to 8. After a second applicationof N in early July, clover was not suppressed to the same extentas in the first growth period. Overall, the photosynthetic capacities of newly expanded cloverlaminae were similar in the two treatments. Clover laminae hadhigher photosynthetic capacities than grass, even in the grass-dominant+ N treatment. Lamina area, petiole length, and the number of live leaves perstolon were similar in the two treatments, indicating that thedifferences in total leaf area were due to the presence of fewerstolon growing points in the + N treatment. Trifolium repens L., white clover, Lolium perenne L., perennial ryegrass, nitrogen, leaf area index, photosynthesis, growth     

Radiation Interception, Partitioning and Use in Grass -Clover Mixtures

Mixed swards of perennial ryegrass /white clover were grownin competition under controlled environmental conditions, attwo temperatures and with different inorganic nitrogen supplies.The swards were studied after canopy closure, from 800 to 1200°C d cumulative temperatures. Clover contents did not varysignificantly during the period. A simulation model of lightinterception was used to calculate light partitioning coefficientsand radiation use efficiencies for both components of the mixturein this controlled environment experiment. Additionally, thissame radiative transfer model was applied to the field datafrom Woledge (1988) (Annals of Applied Biology112: 175 –186)and from Woledge, Davidson and Dennis (1992) (Grass and ForageScience47: 230 –238). The measured and simulated valuesof light transmission, at different depths in the mixed canopy,were highly correlated (P<0.001) with more than 80% of thetotal variance explained. The daily average of photosyntheticallyactive radiation (PAR) interception in a natural environmentwas estimated from simulations, for the field and controlledenvironment data. Under these conditions, white clover capturedsignificantly more light per unit leaf area than perennial ryegrassat low, but not at high, nitrogen supply. In the controlled environment experiment, the radiation useefficiency of the legume was lower than that of its companiongrass. For both species, radiation use efficiency was negativelycorrelated with the mean irradiance of the leaf. The role ofa compensation between light interception and light use forstabilizing the botanical composition of dense grass –cloverswards is discussed. Light interception; radiation transfer model; growth analysis; radiation use efficiency; white clover; perennial ryegrass; Trifolium repensL.; Lolium perenneL.; grassland     

Comparisons of the Respiratory Effluxes of Nodules and Roots in Six Temperate Legumes

The specific respiration rates of nodulated root systems, ofnodules and of roots were determined during active nitrogenfixation in soya bean, navy bean, pea, lucerne, red clover andwhite clover, by measurements on whole plants before and afterthe removal of nodule populations. Similar measurements weremade on comparable populations of the six legumes, lacking nodulesbut receiving abundant nitrate-nitrogen, to determine the specificrespiration of their roots. All plants were grown in a controlled-environmentclimate which fostered rapid growth. The specific respiration rates of nodulated root systems ofthe three grain and three forage legumes during a 7–14-dayperiod of vegetative growth varied between 10 and 17 mg CO₂g^–1 (dry weight) h^–1. This mean value consistedof two components: a specific root respiration rate of 6–9mg CO₂ g^–1 h^–1 and a specific nodule respirationrate of 22–46 mg CO₂ g^–1 h^–1. Nodule respirationaccounted for 42–70 per cent of nodulated root respiration;nodule weight accounted for 12–40 per cent of nodulatedroot weight. The specific respiration rates of roots lackingnodules and utilizing nitrate nitrogen were generally 20–30per cent greater than the equivalent rates of roots from nodulatedplants. The measured respiratory effluxes are discussed in thecontext of nitrogen nitrogen fixation, nitrate assimilation. Glycine max, Phaseolus vulgaris, Pisum sativum, Medicago sativa, Trifolium pratense, Trifolium repens, soya bean, navy bean, pea, lucerne, red clover, white clover, nodule respiration, root respiration, fixation, nitrate assimilation     

Environment-induced Modifications to Root Longevity in Lolium perenne and Trifolium repens

Understanding how environmental factors affect the longevityof roots is essential if root mortality linked nutrient cyclingprocesses within ecosystems are to be understood, and the impactof natural and anthropogenic climate change properly evaluated.In this study the longevity of roots at two geographically andclimatically distinct sites were compared to identify the scaleof change that can occur due to environmental differences. Minirhizotrontubes were inserted into swards sown with the same variety ofLolium perenne and Trifolium repens at sites in the UK and Italy.Roots were viewed using a video camera and digitized imagesof roots generated at intervals. From these images the lifehistory of individual roots was determined and compared. Therewere few differences in patterns of longevity between differentspecies at the same site. Major differences, however, were observedbetween roots of the same species at different sites. For example,73% of Trifolium repens roots failed to survive for 21 d inItaly compared to only 29% at the UK site. Similarly, over 84%of Lolium perenne roots failed to survive for more than 21 din Italy compared to 38% in the UK. These data suggest thatenvironmental factors can have a major impact on root longevity.Copyright 2000 Annals of Botany Company Climate change, Lolium perenne L., minirhizotron, perennial ryegrass, root longevity, Trifolium repens L., white clover     

Photosynthesis by White Clover Leaves in Mixed Clover/Ryegrass Swards

Measurements of rates of net photosynthesis were made on singleBlanca white clover leaves on plants taken from a field-grown,mixed clover/perennial ryegrass sward during two regrowth periods. Net photosynthesis fell by 20 per cent in the first measurementperiod as leaf area index increased and the grass componentof the crop flowered, but did not change significantly in thesecond measurement period during which the grass remained vegetative. Leaves which had been artificially protected from shading inthe sward did not have significantly different photosyntheticcapacities from leaves in the undisturbed sward, even in thefirst measurement period. As leaf area index and sward height increased, successive cloverpetioles were longer, keeping the newly expanded leaves nearthe top of the sward where they received full light. It is suggestedthat it is this which allows successive clover leaves, unlikethose of vegetative grasses, to attain a high photosyntheticcapacity throughout a growth period. Trifolium repens, Lolium perenne, Photosynthetic capacity, shading, growth     

Effect of Contrasting Patterns of Nitrate Application on the Nitrate Uptake, N2-Fixation, Nodulation and Growth of White Clover

White clover (Trifolium repens L.) plants were grown from seedin perlite, inoculated with effective rhizobia and exposed tothe same ‘concentration x days’ of ¹⁵N-labellednitrate in four contrasting patterns of doses. Acetylene reductionwas measured at intervals using an open, continuous-flow sytem.Mean dry weight per nodule and rates of acetylene reductionfell rapidly (2–3 d) during periods of exposure to highnitrate concentrations (> 7 mM N) and rose again, equallyrapidly, when nitrate was withdrawn or substantially reduced.The fall in mean dry weight per nodule (50–66 per cent)was almost certainly too large to be accounted for by loss ofsoluble or storage carbohydrate only. No new nodules were formedduring periods of high nitrate availability. When nitrate wassupplied continuously at a moderate concentration (5.7 mM N)nodule numbers stabilised although existing nodules increasedin dry weight by almost four-fold over the 30 d measurementperiod. Treatment had no effect on the percentage nitrogen in planttissues although there were large differences in the proportionsderived from nitrate and N₂-fixation. Plants exposed continuouslyor frequently to small doses of nitrate took up more nitrate,and hence relied less heavily on N₂-fixation, than those exposedto larger doses less often. Increased reliance on nitrate broughtwith it increased total dry weight and shoot: root ratios. Possiblemechanisms involved in bringing about these differences in nitrogennutrition and growth are discussed. White clover, Trifolium repens, nitrate, N₂-fixation, nodule, acetylene reduction, ¹⁵N     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.): 3. Partition of Assimilates between Root and Shoot during the Transition from Vegetative to Reproductive Growth

The partition of ¹⁴C labelled current assimilates to root insimulated swards of Lolium perenne cv. S24 was measured duringthe transition from vegetative growth in autumn to reproductivegrowth in spring under close to natural conditions of lightand temperature. Assimilate partitioning was also measured in‘established’ swards cut three times during thegrowing season and in vegetative ‘seedling’ swardsgrowing in autumn and in spring. All measurements were madewhen the swards had achieved more than 90 per cent light interception,and all swards were abundantly supplied with water and mineralnutrients. During autumn there was a gradual decrease in the proportionof assimilates partitioned to the roots in both the ‘established’and the ‘seedling’ swards. In the established swards,partition to roots was low over winter, increased during earlyspring, but decreased dramatically, later in the spring, whenstem elongation began. In contrast, in the unvernalized vegatativeseedling swards in spring, partition to roots remained high. The seasonal pattern of assimilate partitioning is consideredin relation to changes in the natural environment and the rateat which the crop fixed carbon in photosynthesis. A decreasein the proportion of assimilates partitioned to roots duringlate spring was significant in increasing the production ofshoot at that time but seasonal differences in partition contributedvery little to the marked differences in shoot growth betweenthe spring and autumn crop. Lolium perenne L., perennial ryegrass, partition of assimilates, flowering     

Effects of Soil Nutrients and Moisture on Root/Shoot Ratios in Lolium perenne L. and Trifolium repens L

The dry weights of foliage and roots (excluding stembases) werecompared in a 3³x2² factorial experiment with Lolium perenneL. and Trifolium repens L. grown in pots with three nitrogen(0–300 ppm N), phosphorus (0–300 ppm P), and soilmoisture (pF 3.2, pF 2.4, and waterlogged) treatments, eachduplicated. Root/shoot ratios were influenced significantly (P < 0.001)by N, P, and W (soil moisture), but these main effects werequalified by highly significant first-order interactions andsecond-order interactions which included N and P. The lowestR/S ratios occurred where N, P, and W were all non-limiting,and the highest where there was an imbalance of these factors. The explanation suggested is that a decrease in the availabilityof N, P, or W causes an increase in the relative weight of roots,but in the presence of N, P, or W deficiency an increase inany one of these factors causes a further increase in the R/Sratio by increasing the relative efficiency of the foliage.     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.). 1. Response of Leaf Extension to Temperature during the Transition from Vegetative to Reproductive Growth

The potential for leaf extension of plants of Lolium perennecv S24 growing in small artificial communities under naturalconditions was measured as the plants progressed from the vegetativeto the reproductive state In two consecutive years, 1975 and 1976, ‘simulated swards’were sown in autumn and overwintered in an open, unheated glasshouseIndividual swards from the batch sown in 1975 were brought into a growth room on two occasions in spring 1976 to measuretheir potential for leaf extension at a range of temperatures(5–20 °C) Individual swards from the batch sown inautumn 1976 were brought in to the growth room on 15 occasionsbetween November 1976 and May 1977 and their potential for leafextension was measured at a single temperature of 15 °CFrequent dissections were made in both years to describe changesin the developing apex. The potential for leaf extension at 15 °C decreased fromc 17 mm day^–1 in November to c 10 mm day^–1 in mid-winter.In January, the potential rapidly increased threefold to reach30mm day^–1 by mid February The increase began coincidentwith the earliest stages of floral initiation and was completedby the time of double-ridge formation Spring-grown vegetativeplants, however, showed potential rates of < 20 mm day^–1at 15 °C The results are discussed in relation to reproductive developmentand to changes in the carbohydrate strategy of the plants inearly spring Lolium perenne L perennial ryegrass, leaf extension, temperature response     

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     

Rates of Hydrogen Ion Efflux by Nodulated Legumes Grown in Flowing Solution Culture with Continuous pH Monitoring and Adjustment

The net efflux of H⁺ from lucerne (Medicago saliva L.), redclover (Trifolium pratense L.) and white clover (Trifolium repensL.) growing in flowing solution culture and dependent upon symbioticfixation of atmospheric N, was measured over a 75 d experimentalperiod. Considerable and rapid increases in acidity of the nutrientsolution of up to 1.45 pH units were recorded when the pH wasriot held constant over a 30 h period. There was little differencein H⁺ efflux when solution pH was held constant at 4.75, 5.75or 6.75, but there was an immediate cessation when it was adjustedto 3.75. Differences in the daily net efflux of H⁺ closely followedthe pattern of daily differences in incoming radiation, andthere was also evidence of a diurnal pattern of H⁺ efflux. Althoughthere were initially distinct differences between the speciesin the calculated rate of net H⁺ efflux (µg H⁺ g^–1dry shoot d^–), by day 75 these had diminished. In allspecies, however, the maximum rate of efflux per unit of shootsoccurred during the earlier rapid phases of growth. The measuredefflux of H⁺ was well equated with the plant content of excesscations (as measured by ash alkalinity) and, on average, theratio of acidity produced to N assimilated (expressed as anequivalent) was 0-24. Medicago sativa L., Trifolium pratense L., Trifolium repens L., lucerne, red clover, white clover, acidification, cation/anion balance, flowing solution culture, H⁺ efflux, nitrogen fixation     

Absorption and Distribution of Copper in Plants with Sufficient or Deficient Supplies

Wheat, red clover and ryegrass were grown in flowing solutionculture with sufficient (+ Cu) and deficient (–Cu) suppliesof copper. The rates of Cu absorption (µg g^–1 dryroot day^–1) did not differ greatly between species ineither treatment. Wheat plants, when transferred from the –Cu to the +Cu treatment, absorbed Cu at a much slower rate thanthose which had remained throughout in the + Cu treatment. Inall plants considerable proportions of the absorbed Cu wereretained in the roots, even when the plants were Cu-deficient,and the concentration in roots usually exceeded that in anypart of the shoots in both treatments. Transferring wheat plantsfrom the +Cu to the –Cu treatment decreased the concentrationin all plant parts except old leaves; similarly, transferringfrom the –Cu to +Cu treatment increased the concentrationin all parts of the shoots, execept old leaves, and in the roots. Lolium perenne, Trifolium pratense, Triticum aestivum, ryegrass, red clover, wheat, absorption, copper, flowing solution culture