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     

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     

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     

Leaf Photosynthesis,Dark Respiration and Fluorescence as Influenced by Leaf Age in an Evergreen Tree,Prosopis Juliflora

P. juliflora trees produce leaves during two growth periods. The first cohort of leaves is produced during spring in cool conditions, while the second cohort is produced during monsoon under warm conditions. I studied photosynthetic characteristics of young, mature, and old leaves of the previous season (monsoon) in the spring season. Maximum net photosynthetic rate of a young leaf was lower than that of the mature and old leaves. The total CO₂ fixed per day by the young leaves was just 36 % of that in the mature leaves while the old leaves fixed 76 % of that of the mature leaf. The total transpiration rate and water use efficiency (WUE) were similar in the mature and old leaves, while they were much lower in the young leaves. Dark respiration rate was maximal in the young leaves as compared to the mature and old leaves. About 92 % of the total CO₂ fixed per day were respired by the young leaves. The diurnal fluorescence characteristics (F/F_m, q _p, and q _N) of the young, mature, and old leaves showed that photochemical efficiency of photosystem 2 during midday decreased more in the young and old leaves than in the mature ones. However, the fluorescence characteristics showed that in all the three leaf types there was complete recovery of the photochemical efficiency at sunset from the midday depression. F_v/F_m in the young and mature leaves also confirmed this. Hence the young and old leaves were photosynthetically less efficient than mature leaves, but they were well adapted to withstand the harsh environmental conditions.     

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     

Photosynthesis of White Clover (Trifolium Repens L.) Germplasms with Contrasting Leaf Size

In a growth chamber experiment, we determined net photosynthetic rate (PN) and leaf developmental characteristics of cultivars of a relatively small-, intermediate-, and a large-leaf genotype grown under irradiance of 450-500 µmol(photon) m-2 s-1 (HI), shade [140-160 µmol(photon) m-2 s-1] (LI), and after a shade-to-irradiation (LI »HI) transfer. Differences in physiological responses of the genotypes were more pronounced in HI and LI»HI plants than in LI plants. The small- and intermediate-leaf sizes had greater PN in the first measured leaf than the large-leaf type by 70 and 63 % in HI plants, and by 23 and 18 % in LI»HI plants, respectively. Similar relationships were observed in the next developed leaf. The LI plants did not differ significantly in PN. Greater PN in the small- and intermediate-leaf size genotypes were not associated with greater total dry matter of the plant. Under irradiation, the large-leaf genotype accumulated more total nonstructural saccharides (TNS) and starch than the small- or intermediate-leaf size plants. TNS and starch concentrations in LI plants were about one-half those of HI and LI»HI plants. These results should help to develop management practices that capitalize upon the competitive features of white clover in mixed-species swards.     

Relationships among Isoprene Emission Rate, Photosynthesis, and Isoprene Synthase Activity as Influenced by Temperature

Isoprene emissions from the leaves of velvet bean (Mucuna pruriens L. var utilis) plants exhibited temperature response patterns that were dependent on the plant's growth temperature. Plants grown in a warm regimen (34/28°C, day/night) exhibited a temperature optimum for emissions of 45°C, whereas those grown in a cooler regimen (26/20°C, day/night) exhibited an optimum of 40°C. Several previous studies have provided evidence of a linkage between isoprene emissions and photosynthesis, and more recent studies have demonstrated that isoprene emissions are linked to the activity of isoprene synthase in plant leaves. To further explore this linkage within the context of the temperature dependence of isoprene emissions, we determined the relative temperature dependencies of photosynthetic electron transport, CO₂ assimilation, and isoprene synthase activity. When measured over a broad range of temperatures, the temperature dependence of isoprene emission rate was not closely correlated with either the electron transport rate or the CO₂ assimilation rate. The temperature optima for electron transport rate and CO₂ assimilation rate were 5 to 10°C lower than that for the isoprene emission rate. The dependence of isoprene emissions on photon flux density was also affected by measurement temperature in a pattern independent of those exhibited for electron transport rate and CO₂ assimilation rate. Thus, despite no change in the electron transport rate or CO₂ assimilation rate at 26 and 34°C, the isoprene emission rate changed markedly. The quantum yield of isoprene emissions was stimulated by a temperature increase from 26 to 34°C, whereas the quantum yield for CO₂ assimilation was inhibited. In greenhouse-grown aspen leaves (Populus tremuloides Michaux.), the high temperature threshold for inhibition of isoprene emissions was closely correlated with the high temperature-induced decrease in the in vitro activity of isoprene synthase. When taken together, the results indicate that although there may be a linkage between isoprene emission rate and photosynthesis, the temperature dependence of isoprene emission is not determined solely by the rates of CO₂ assimilation or electron transport. Rather, we propose that regulation is accomplished primarily through the enzyme isoprene synthase.     

Development of Ramularia Leaf Spot on Sugar Beet as Influenced by Temperature and the Age of the Host Plant

A method of inoculating sugar beet plants (Beta vulgaris L.) with Ramularia beticola Faut. & Lamb, is described. Following inoculation, disease development in relation to temperature and plant age was studied for more than a month. The incubation period was 18 days at 10°C compared to 14 days at 17°C. At 25°C no symptoms appeared. Both temperature and plant age significantly influenced disease level and rate of disease development. Plants incubated at 17°C were more severely diseased 33 days after inoculation than plants incubated at 10°C. Young plants (3 weeks at inoculation) Were more susceptible than older plants (5 and 8 weeks at inoculatson) under growth chamber conditions. In the field, symptoms of Ramularia leaf spot appear relatively late in the season and young leaves are rarely attacked. The inconsistency of these observations is discussed.     

Defoliation in White Clover: Regrowth, Photosynthesis and N2 Fixation

Single plants of white clover, grown in a controlled environmentand dependent for nitrogen on fixation in their root nodules,were defoliated once by removing approximately half their shoottissue. Their regrowth was compared with the growth of comparableundefoliated plants. Two similar experiments were carried out:in the first, plants were defoliated at 2.5 g, and in the secondat 1.2 g total plant d. wt. Defoliation reduced rate of N₂ fixation by > 70 per cent,rate of photosynthesis by 83–96 per cent, and rate ofplant respiration by 30–40 per cent. Nodule weights initiallydeclined following defoliation as a result of loss of carbohydratesand other unidentified components. No immediate shedding ofnodules was observed but nodules on the most severely defoliatedplants exhibited accelerated senescence. The original rates of N₂ fixation were re-attained after 5–6or 9 d regrowth, with increase in plant size at defoliation.In general, the rate of recovery of N₂ fixation was relatedto the re-establishment and increase of the plant's photosyntheticcapacity. Throughout the growth of both defoliated and undefoliatedplants nodule respiration (metabolism) accounted for at least23 ± 2 per cent of gross photosynthesis. The unit ‘cost’of fixing N₂ in root nodules, in terms of photosynthate, appearedto be unaffected by defoliation, except perhaps for plants veryrecently defoliated. Similarly, the percentage nitrogen contentsof shoot, root and nodules of defoliated plants became adaptedwithin a few days to those characteristic of undefoliated plants. Trifolium repens, white clover, N₂ fixation, defoliation, photosynthesis, respiration     

Effects of Elevated CO2 and High Temperature on Single Leaf and Canopy Photosynthesis of Rice

The increase of atmospheric CO2 concentration is indisputable. In such condition, photosynthetic response of leaf is relatively well studied, while the comparison of that between single leaf and whole canopy is less emphasized. The stimulation of elevated CO2 on canopy photosynthesis may be different from that on single leaf level. In this study, leaf and canopy photosynthesis of rice ( Oryza sativa L. ) were studied throughout the growing season. High CO2 and temperature had a synergetic stimulation on single leaf photosynthetic rate until grain filling. Photosynthesis of leaf was stimulated by high CO2, although the stimulation was decreased by higher temperature at grain filling stage. On the other hand, the simulation of elevated CO2 on canopy photosynthesis leveled off with time. Stimulation at canopy level disappeared by grain filling stage in beth temperature treatments. Green leaf area index was not significantly affected by CO2 at maturity, but greater in plants grown at higher temperature. Leaf nitrogen content decreased with the increase of CO2 concentration although it was not statistically significant at maturity. Canopy respiration rate increased at flowering stage indicating higher carbon loss. Shading effect caused by leaf development reached maximum at flowering stage. The CO2 stimulation on photosynthesis was greater in single leaf than in canopy. Since enhanced CO2 significantly increased biomass of rice stems and panicles, increase in canopy respiration caused diminishment of CO2 stimulation in canopy net photosynthesis, keaf nitrogen in the canopy level decreased with CO2 concentration and may eventually hasten CO2 stimulation on canopy photosynthesis. Early senescence of canopy leaves in high CO2 is also a possible cause.     

Temperature and CO2Responses of Leaf and Canopy Photosynthesis: a Clarification using the Non-rectangular Hyperbola Model of Photosynthesis

The responses of C₃leaf and canopy gross photosynthesis to increasingtemperature and CO₂can be readily understood in terms of thetemperature and CO₂dependencies of quantum yield (     

SHORT COMMUNICATION Growth and Development of White Clover (Trifolium repensL.) as Influenced by P and K Nutrition

The effects of varying P and K supplies on the growth and morphologyof white clover were investigated in a pot experiment. Plantswere treated with three rates of potassium (K) and four ratesof phosphorus (P) in factorial combination, and five harvestswere taken between August 1993 and February 1994. During establishment,shoot and leaf development were more adversely affected by Pthan by K deficiency but, by the final growth period, K deficiencywas the more limiting to growth, causing dramatic reductionsin leaf area index (LAI), numbers of growing points, and stolonand root masses. The adverse affects of K deficiency on shootdevelopment were attributed to the curtailment of petiole andlamina expansion which have major requirements for K. It wasnoted that, under K deficiency, proportionately less assimilatewas partitioned to leaf production than to stolon and root growth;this may be an adaptation enabling K-deficient plants to exploitnutrient supplies in adjacent localities. The results suggestthat, provided white clover can withstand ‘moderate’P stress during establishment, its persistence in swards willprobably be limited more by K deficiency than by low or inadequateP supplies. However, because the experiment was conducted outsidethe normal growing season and with only a single white clovercultivar, it is inappropriate to make any generalizations regardingthe effects of P and K nutrition on white clover in the field.Copyright1998 Annals of Botany Company Mineral nutrition, morphology, phosphorus, potassium,Trifolium repensL.     

群体中叶片光合能力的分布及其对群体光合作用的影响

利用数学变分原理分析了群体中叶片光合能力对环境适应和有限氮资源利用的最优分布。叶片光合能力呈现与光强相同的负指数衰减分布时,“群体的光合速率和对氮的利用率最高;叶片对环境光强适应的优越性随群体消光系数和叶面积指数增加而增加。由此推导了叶片光合能力最优分布下的群体光合模型。     

Single Leaf and Canopy Photosynthesis in a Ryegrass Sward

Swards of S24 perennial ryegrass (Lolium perenne L.) were grownin the field and harvested by repeated cutting. The rate ofphotosynthesis of the swards and of single leaves of plantstaken from them was measured throughout the growing season.Leaf area and light interception of the sward were also measured. In each growth period, sward photosynthesis increased with increasingleaf area and then declined. In the first growth period, duringwhich the crop flowered, the maximum rate of sward photosynthesiswas higher and was reached later than in succeeding vegetativegrowth periods. The photosynthetic capacity of successive youngest expandedleaves was high throughout the first growth period, and onlydeclined due to leaf age when no further leaves were producedafter flowering. However, in the succeeding vegetative growthperiods, although the first leaves to expand had high photosyntheticcapacities, later leaves had progressively lower capacities. The extent to which changes in the photosynthetic capacity ofsingle leaves accounts for the changes in sward photosynthesisobserved in the field is examined using a mathematical model.     

Effect of Leaf Age and Nodal Position on Receptivity of Rice Leaves to Infection by Pyricularia grisea

The effect of leaf age and nodal position on leaf receptivity to rice blast, caused by Pyricularia grisea , was studied by inoculating potted Rosemont plants with blast pathogen race IC-17, isolate 92T107. at different growth stages. Regressioti equations were used to describe the effect ofleaf age (degree days > 10 C) and nodal position on lesion density, the relative frequency of lesion appearance, and the lesion area distribtition. The number of sporulating lesions per cm² of inoculated leaf area was highest for leaves inoculated early in their development and progressively lower for leaves inoculated later in their development. The average lesion density for less than 1-day-old leaves at nodal position 6 was about four times the density for 10-day-old leaves of the same nodal position. Lesion density was similar for leaves of less than 1-day-old from the 4th to the 7th nodal position, but dropped sharply from the 7th to the 11 th nodal position. The flag leaf was the least susceptible of all nodal positions. The average lesion density for young leaves of about 1-day-old at nodal position 6 was about 27 times the density for young flag leaves of the same age. The incubation period increased with leaf age. but was not obviously affected by leaf nodal position. Lesion area increased linearly with time, but was not affected by leaf age and nodal position. The results are of use for epidemiological investigations which rely on detailed quantitative leaf susceptibility data for accurate blast forecasts.     

Influence of Temperature and O2 Concentration on Photosynthesis and Light Activation of Ribulosebisphosphate Carboxylase Oxygenase in Intact Leaves of White Clover (Trifolium repens L.)

Detached leaves of white clover (Trifolium repens L.) were keptfor 1 h under various conditions of temperature, oxygen concentrationand light intensity. Rates of photosynthesis were measured whereappropriate and then ribulosebisphosphate carboxylase oxygenase(RuBPCO) was extracted rapidly and its initial activity measuredimmediately. The extracted activity increased with increased intensity ofillumination of the leaves. Where leaves were pretreated atlow light intensity, the lower the temperature of the leavesthe higher the extracted activity of RuBPCO. At high light intensitytemperature did not affect the activity of subsequently extractedRuBPCO but the light intensity which was necessary for maximumactivity increased with temperature. Activity of RuBPCO fromleaves pretreated in the dark was least when CO₂ was low andtemperature high. Leaves, pretreated at low temperatures andhigh light intensity in 20% O₂, yielded higher activity in extractsthan leaves pretreated under similar conditions but in 2% O₂.A relatively weak temperature response of photosynthesis atlow irradiances was associated with a decrease in extractableRuBPCO activity with increasing temperature. A strong temperaturedependence of the oxygen inhibition of photosynthesis was associatedwith lower extractable RuBPCO activity in leaves pretreatedat low oxygen concentration at low temperatures. With leavesfrom plants grown at low temperatures prior to treatment ofleaves, oxygen inhibition of photosynthesis was less temperaturedependent and activity of RuBPCO in extracts was not decreasedby low O₂ at low temperatures. Differences in the activationof RuBPCO appear to influence photosynthesis and account foran absence of oxygen inhibition of photosynthesis at low temperaturesin plants grown in warm conditions. Key words: Ribulosebisphosphate carboxylase oxygenase activation, Photosynthesis, Temperature, O₂ effect, White clover     

Microclimate, Canopy Structure and Photosynthesis in Canopies of Three Contrasting Temperate Forage Grasses: III. Canopy Photosynthesis, Individual Leaf Photosynthesis and the Distribution of Current Assimilate

The rates of canopy and individual leaf photosynthesis and ¹⁴Cdistribution for three temperate forage grasses Lolium perennecv. S24, L. perenne cv. Reveille and Festuc'a arundinacea cv.SI70 were determined in the field during a summer growth period.Canopy photosynthesis declined as the growth period progressed,reflecting a decline in the photosynthetic capacity of successiveyoungest fully expanded leaves. The decline in the maximum photosyntheticcapacity of the canopies was correlated with a decline in theirquantum efficiencies at low irradiance. Changes in canopy structureresulted in changes in canopy net photosynthesis and dark respiration.No clear relationships between changes in the environment andchanges in canopy net photosynthesis and dark respiration wereestablished. The relative distributions of ¹⁴C in the shootsof the varieties gave a good indication of the amount of drymatter per ground area in the varieties.     

Influence of Carbon Dioxide Concentration on Growth, Carbohydrate Content, Translocation and Photosynthesis of White Clover

White clover ramets were grown at various carbon dioxide concentrations(200, 350 and 1000 µl 1^–1), defoliated and regrownat the same concentrations. Morphological characteristics, dryweights and non-structural carbohydrate contents of plant organs,diurnal variation of sugar and starch content of leaves, translocationof assimilates and photosynthesis were determined. Carbon dioxide concentration influenced the dry weights, butnot the number and size of the plant organs. However, defoliationof plants at low carbon dioxide concentration resulted in decreasedleaf size and stolon length. Carbon dioxide concentration influencedthe content and diurnal variation of starch and sugar in theleaves. Starch was accumulated at medium carbon dioxide concentrationand sugar at a higher concentration when the storage capacityfor starch seemed to be exceeded. Starch was preferentiallyaccumulated in the first and sugar in the second half of thelight period. Translocation was decreased during the periodsof accumulation. Sugar accumulation in the leaves seemed tobe a consequence of the imbalance between sink and source, whereasstarch accumulation seemed to follow an in-built diurnal pattern.Accumulation of both starch and sugar during the photoperiodwas followed by degradation and export during the dark period.Decreased dark export occurred at low carbon dioxide concentrationwhen neither starch nor sugar was accumulated during the photoperiod. Carbon dioxide, white clover, Trifolium repens L., growth, carbohydrates, starch, sugar, translocation, photosynthesis     

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