An Analysis of the Photosynthesis and Productivity of Vegetative Crops in the United Kingdom

The rate of total dry matter production of a vegetative crop,under optimal water and nutrient regimens is related to someleaf and canopy photosynthetic characteristics. Three leaf photosyntheticcharacteristics are examined in detail: the light utilizationefficiency at normal ambient CO₂ and O₂ concentrations, a, therate of light saturated photosynthesis per unit leaf area, F_max,and the ratio of the rates of photorespira tion and gross photosynthesis.The genetic variability in each of these characteristics issought from published data on a wide range of C₃ and C₄ planttypes. Within C₃ and C₄ plant types there are significant genetic differencesonly in F_max,, although differences exist between C₃ and C₄plants in the other two characteristics. The effects of thesedifferences on the rate total dry matter production are estimated,and it is concluded that there is no compelling evidence toindicate that improvements in total dry matter production rates,in the U.K., are likely to result from genetic manipulationof these characteristics in the existing range of plant material.     

Dry Matter Production in a Tomato Crop: Measurements and Simulation

Simulation of dry matter production by the explanatory glasshousecrop growth model SUKAM (Gijzen, 1992, Simulation Monographs),based on SUCROS87 (Spitters, Van Keulen and Van Kraalingen,1989, Simulation and systems management in crop protection),was validated for tomato. In the model, assimilation rates arecalculated separately for shaded and sunlit leaf area at differentcumulative leaf area in the canopy, taking into account thedifferent interception of direct and diffuse components of light.Daily crop gross assimilation rate (P_gd) is computed by integrationof these rates over total crop leaf area and over the day. Leafphotochemical efficiency and potential gross assimilation rateat saturating light depend on temperature and CO₂ concentrationand are approximated as being identical in the whole canopy.Crop growth results from P_gd minus maintenance respiration rate(R_m; dependent on temperature and crop dry weight), multipliedby the conversion efficiency (carbohydrates to structural drymatter; C_f). Growth experiments (periodic destructive harvest) with differentplanting dates and plant densities and two data-sets from commerciallygrown crops, were used for model validation. Hourly averagesfor global radiation outside the glasshouse, glasshouse temperatureand CO₂ concentration, together with measured leaf area index,dry matter distribution (for calculation of C_f) and organ dryweights (for calculation of R_m) were the inputs to the model. Dry matter production (both level and dynamic behaviour) wassimulated reasonably well for most experiments, but final drymatter production was under-estimated by about 27% for the commerciallygrown crops. At low irradiance and with large crop dry weight,growth rate was under-estimated, probably as a result of over-estimationof R_m. This could almost completely explain the large under-estimationfor the commercially grown crops, which had large dry weight.Final dry matter production was over-estimated by 7-11% if dailyaverages instead of hourly input of climatic data were used. It is concluded that SUKAM is a reliable model for simulatingdry matter production in a tomato crop, except for those situationswhere R_m has a large influence on crop growth rate (low irradianceand large crop dry weight). An improved estimate of R_m wouldtake into account the influence of metabolic activity. A preliminaryattempt to relate maintenance costs to relative growth rate(a measure for metabolic activity), showed promising results.Copyright1995, 1999 Academic Press Crop growth, dry matter production, glasshouse, maintenance respiration, metabolic activity, model, relative growth rate, respiration, simulation, tomato, model validation     

The Seasonal Pattern of Growth and Production of a Temperate C4 Species, Cyperus longus

Growth and production of the temperate C₄ species Cyperus longusL. was measured throughout a growing season in an establishedplot in Eastern Ireland. The maximum standing live biomass reachedwas 2·5 kg m^–2. Estimates of unit leaf rate (ULR)and leaf area index (LAI) were made. The product of these quantitiesgave the crop growth rate (CGR) each week. C. longus was foundto maintain high values of LAI throughout the summer, with amaximum value of about 13 in early August. CGR reached a peakin early July. The optimum LAI was 11·6. Temperaturesat five levels in the plant canopy, and the amount of solarradiation intercepted by the canopy were measured continuouslyduring the summer. The mean daily rate of leaf extension waspositively correlated with the mean daily air temperature abovethe canopy but the temperature coefficient of the process waslow compared with other temperate species. The percentage ofsolar radiation intercepted by the canopy increased rapidlyin early summer, and canopy closure had occurred by mid-June.Rates of net photosynthesis were measured on young and old leafmaterial in situ at the time of peak LAI. In young leaves themaximum rates of net photosynthesis were higher than those publishedfor a range of temperate C₃ species, but similar to those foundin another temperate C₄ species, Spartina townsendii. Key words: C₄ photosynthesis, leaf growth, productivity     

Use of Interactive Computer Graphics for Simulation of Radiation Interception and Photosynthesis for Canopies of Kiwifruit Vines with Heterogeneous surface Shape and Leaf Area Distribution

A method incorporating interactive computer graphics to simulatespatially variable interception and canopy photosynthesis isdescribed. The method presents a graphical interface to a conventionalmodel of radiation interception and canopy photosynthesis. Includedis the capacity to consider a large number of positions withinthe canopy, thus providing a rapid and convenient representationof the dynamics of photosynthesis while also overcoming limitationsof one-dimensional models applied to complex plant canopies.The method was applied to examine spatial variability of photosynthesiswithin canopies of kiwifruit (Actinidia deliciosa) vines growingon two trellis types. The diurnal integral of simulated canopyphotosynthesis, assuming sunny conditions, for a vine trainedon a horizontal 'Pergola' trellis was 14% higher than that fora vine with similar leaf area distribution trained on a 'T-bar'trellis with inclined surfaces. Simulations of photosynthesisfor vines on a T-bar trellis, assuming spatially variable leafarea distributions as measured under filed conditions, indicateddisproportionate contributions from different regions of thecanopy. Canopy regions inclined to the east or the west wereusually the major sites for photosynthesis immediately aftersunrise and before sunset respectively, while regions near thecordon were the most important overall. For any day, the maximumsimulated photosynthetic rate generally declined with distancefrom the cordon and, at any distance from the cordon, increasedwith leaf index. For a vine with an average leaf area indexof 2·7, diurnal integrals of photosynthesis on a sunnyday in late summer ranged from 1·0 mol CO₂ m^-2 near thecordon to 0·5 mol CO₂ m^-2 at 1·5 m from the cordon.Within-canopy shading was more important on sunny days thanon cloudy days, while the spatial distribution of leaf areawas especially important on cloudy days. Comparison of simulationswith direct measurements of canopy photosynthesis indicatedthat a numerical integral of simulated photosynthesis, basedon a large number of canopy positions, provided a reasonableestimate of total canopy photosynthesis.Copyright 1993, 1999Academic Press Actinidia deliciosa, kiwifruit, interactive computer graphics, mathematical modelling, photosynthesis, radiation interception, spatial heterogeneity     

Nitrogen Deficiency in Small Closed Communities of S24 Ryegrass. I. Photosynthesis, Respiration, Dry Matter Production and Partition

Small communities of S24 ryegrass were grown under supplementarylights in a glasshouse at 20°C, and abundantly suppliedwith a complete nutrient solution containing 300 p.p.m. of nitrogen,until they had a leaf area index of 5 and fully interceptedthe light. Half were then given a solution containing only 3p.p.m. of nitrogen (LN) while the rest were kept at 300 p.p.m.(HN). The LN plants had a rate of single leaf photosynthesis lowerthan that of the HN plants at all but the lowest light intensities(33 per cent lower at the saturating irradiance of 170 W m^–2).Similarly, the LN communities had rates of canopy gross photosynthesis(P_sc) markedly lower than those of the HN communities. A comparisonof the observed rates of P_sc with those predicted by a mathematicalmodel of canopy photosynthesis indicated that it was the effectof nitrogen on single leaf photosynthesis, rather than differencesbetween the communities in leaf area, which led to the observeddifferences in P_sc. The superiority of the HN communities in terms of P_sc was partlyoffset by a higher rate of respiration so that they only exceededthe LN communities in terms of canopy net photosynthesis atirradiances in excess of 180 W m_–2, and produced only15 per cent more total dry matter. Nevertheless, the HN plantsdirected less of that dry matter into root and more into topsso that they came to possess twice the weight of live laminae,and the HN communities twice the leaf area, of their nitrogendeficient counterparts. Lolium perenne, S24 ryegrass, photosynthesis, respiration, dry matter production and partition, nitrogen dekieacy     

Canopy Photosynthesis and Crop Growth Rate of Eight Temperate Forage Grasses

The rates of canopy and individual leaf photosynthesis, ratesof growth of shoots and roots, and the extinction coefficientfor light of eight temperate forage grasses were determinedin the field during early autumn. Canopy gross photosynthesiswas calculated as net photosynthesis plus dark respiration adjustedfor temperature using a Q₁₀ = 2. The relationships between canopygross photosynthesis and light intensity were hyperbolic, andthe initial slopes of these curves indicated that light wasbeing utilized efficiently at low light intensities. The initialslope depended on the distribution of light in the canopy andthe quantum efficiency of the individual leaves. The maximumrate of canopy gross photosynthesis reflected the maximum rateof individual leaf photosynthesis. Although the maximum rateof canopy gross photosynthesis was correlated with crop growthrate, there was no significant relationship between daily grossphotosynthesis and crop growth rate. Indeed, daily gross photosynthesisvaried by only 22 per cent, whereas the daily growth of shootsand roots varied by 120 per cent. This poor correlation is influencedby a negative correlation (P < 0.01) between the maximumrate of canopy gross photosynthesis and the initial slope ofthe curve relating canopy gross photosynthesis and light intensity.Difficulties in the interpretation of measurements of dark respirationappeared to confound attempts to relate daily net photosynthesisto crop growth rate, individual leaf photosynthesis, and theextinction coefficient for light.     

Evaluation of a Dynamic Simulation Model for Tomato Crop Growth and Development

A dynamic simulation model for tomato crop growth and development,TOMSIM, is evaluated. Potential crop growth and daily crop grossassimilation rate (P_gc,d) is computed by integration of leafassimilation rates over total crop leaf area throughout theday. Crop growth results fromP_gc,dminus maintenance respirationrate (R_m), multiplied by the conversion efficiency. Dry matterdistribution is simulated, based on the sink strength of theplant organs, which is quantified by their potential growthrate. Within the plant, individual fruit trusses and vegetativeunits (three leaves and stem internodes between two trusses)are distinguished. Sink strength of a truss or a vegetativeunit is described as a function of its developmental stage.In this paper, emphasis is on the interactions between the twosubmodels of, respectively, dry matter production and dry matterdistribution. Sensitivity analysis showed that global radiation,CO₂concentration, specific leaf area (SLA) and the developmentalstage of a vegetative unit at leaf pruning had a large influenceon crop growth rate, whereas temperature, number of fruits pertruss, sink strength of a vegetative unit and plant densitywere less important. Leaf area index (LAI) was very sensitiveto SLA and the developmental stage of a vegetative unit at leafpruning. Temperature did not influence the simulated R_m, asincreased respiration rate per unit of biomass at higher temperatureswas compensated by a decrease in biomass. The model was validatedfor four glasshouse experiments with plant density and fruitpruning treatments, and on data from two commercially growncrops. In general, measured and simulated crop growth ratesfrom 1 month after planting onwards agreed reasonably well,average overestimation being 12%. However, crop growth ratesin the first month after planting were overestimated by 52%on average. Final crop dry mass was overestimated by 0–31%,due to inaccurate simulation of LAI, resulting partly from inaccurateSLA prediction, which is especially important at low plant densityand in a young crop.Copyright 1999 Annals of Botany Company Crop growth, dry matter production, glasshouse, leaf area,Lycopersicon esculentum, partitioning, simulation model, tomato, TOMSIM.     

The Effects of Nitrogen Fertilization and the Growing Season on Photosynthesis of Field-grown Tall Fescue (Festuca arundinacea Schreb.) Canopies

Canopy gross photosynthesis of tall fescue receiving three tofour rates of N fertilization was studied under field conditionsduring three contrasting growing seasons. Under non-limitingN growing conditions, the growing seasons did not have a significanteffect on the maximum canopy gross photosynthesis (canopy grossphotosynthesis at saturating PAR) and the maximum light yield(quantum efficiency of the canopy at low PAR). In the absenceof N fertilization and for a similar LAI, the values of themaximum canopy gross photosynthesis were approximately equalto 70% of those obtained under non-limiting N conditions. Thisresponse of the tall fescue canopy to N concentration is muchsmaller than that reported at the leaf level. The reductionin canopy photosynthetic capacity with no N applied comparedto non-limiting N conditions is much less than the reductionobserved previously in above-ground dry matter accumulation.The effect of N fertilization on above-ground dry matter accumulationis due primarily to changes in C partitioning and the resultingfaster leaf area development and greater light interceptionrather than the effect of N on the canopy photosynthetic capacityper se .Copyright 1993, 1999 Academic Press Festuca arundinacea Schreb., photosynthesis, nitrogen, grass, carbon     

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     

Growth Response of a Chrysanthemum Crop to the Environment. II. A Mathematical Analysis Relating Photosynthesis and Growth

Vegetative crops of chrysanthemum were grown for 5 weeks inthree replicate daylit assimilation chambers. Weekly harvestswere made from each crop for growth analysis, and on seven occasionsduring the 5-week period continuous measurements of the netCO₂ exchange rate of each crop were made over a 24 h period.A semi-empirical model for canopy photosynthesis was fittedto these data. The photosynthesis model was then incorporatedinto a simple, dynamic growth model. Using fitted values ofthe canopy photosynthesis parameters, the daily total radiationintegrals, and the experimentally observed relationship betweenthe leaf area index and crop dry matter per unit ground area,the crop growth model was used to simulate growth over the 5-weekperiod. The predicted and measured crop dry weights were inclose agreement over the whole period.     

Slow Leaf Development of Evergreen Broad-leaved Tree Species in Japanese Warm Temperate Forests

Rates of light-saturated net photosynthesis (P_Nmax) and darkrespiration (Rd) on a leaf area basis, leaf dry mass per area(LMA), leaf nitrogen content on a leaf area basis (LNa) andinstantaneous nitrogen use efficiency (NUE=P_Nmax/LNa) were followedduring leaf development in six evergreen broad-leaved tree speciestypical of warm-temperate forests in Japan. These species wereCastanopsissieboldii, Quercus myrsinaefolia, Quercus glauca, Machilus thunbergii,Cinnamomum japonicumandNeolitsea sericea.When expansion of leafarea was complete, P_Nmax was about one third of its peak valueand increased for another 15 to 44 d. Rd at full leaf expansionwas about 1.5 to 3.5-times greater than steady-state rates.These facts suggest that leaf development was still underwayat the time of full leaf area expansion. Low P_Nmax at full leafexpansion was caused both by low leaf nitrogen content and lowNUE. P_Nmax increased with the increase in LMA during leaf developmentin all six species; data from the literature for other specieswith different life forms also indicated a similar tendency.The steady-state LMA varied markedly among species. Becauseleaves with larger steady-state LMAs need more resources fortheir construction, they will also need longer periods for maturation.We hypothesized that the period required for the attainmentof peak P_Nmax, the ‘leaf maturation period’, dependson the steady-state LMA. Plotting data from the present studytogether with those from literature for other plants acrossseveral life forms showed a strong positive relationship betweenleaf maturation period and steady-state LMA, supporting thehypothesis.Copyright 1998 Annals of Botany Company. Castanopsis sieboldii, Cinnamomum japonicum,delayed period, expansion period, full leaf expansion,Machilus thunbergii,maturation period,Neolitsea sericea, Quercus glauca, Quercus myrsinaefolia,steady-state LMA.     

Dynamics of Vertical Leaf Nitrogen Distribution in a Vegetative Wheat Canopy. Impact on Canopy Photosynthesis

The development of vertical canopy gradients of leaf N has beenregarded as an adaptation to the light gradient that helps tomaximize canopy photosynthesis. In this study we report thedynamics of vertical leaf N distribution during vegetative growthof wheat in response to changes in N availability and sowingdensity. The question of to what extent the observed verticalleaf N distribution maximized canopy photosynthesis was addressedwith a leaf layer model of canopy photosynthesis that integratesN-dependent leaf photosynthesis according to the canopy lightand leaf N distribution. Plants were grown hydroponically attwo amounts of N, supplied in proportion to calculated growthrates. Photosynthesis at light saturation correlated with leafN. The vertical leaf N distribution was associated with thegradient of absorbed light. The leaf N profile changed duringcrop development and was responsive to N availability. At highN supply, the leaf N profiles were constant during crop development.At low N supply, the leaf N profiles fluctuated between moreuniform and steep distributions. These changes were associatedwith reduced leaf area expansion and increasing N remobilizationfrom lower leaf layers. The distribution of leaf N with respectto the gradient of absorbed irradiance was close to the theoreticaloptimum maximizing canopy photosynthesis. Sensitivity analysisof the photosynthesis model suggested that plants maintain anoptimal vertical leaf N distribution by balancing the capacityfor photosynthesis at high and low light. Copyright 2000 Annalsof Botany Company Canopy photosynthesis, leaf nitrogen distribution, nitrogen, Triticum aestivum L, wheat     

Effects of Different CO2 Environments on the Photosynthesis-Yield Relationship and the Carbon and Water Balance of a White Clover (Trifolium repens L. cv. Blanca) Sward

Effects of atmospheric CO₂ enrichment to a level above 600 parts10^–6 on leaf and canopy gas exchange characteristics wereinvestigated in Trifolium repens, using an open system for gasexchange measurement. The cuvettes of the system served as growthchambers, allowing continuous measurement in a semi-controlledenvironment of ±350 and ±600 parts 10^–6CO₂, respectively. Carbon balance data were compared with cropyield and effects on the canopy level were compared with measuredleaf responses of photosynthesis and stomatal behaviour. Photosyntheticstimulation by high CO₂ was stronger at the canopy level (103%on average) than for leaves (90% in full light), as a consequenceof accelerated foliage area development. The latter increasedabsolute water consumption by 16%, despite strong stomatal closure.The overall result was a 63% improvement in canopy water useefficiency (WUE), while leaf WVE increased almost 3-fold insaturating light. The stomatal response was such that, whilethe internal CO₂ concentration in the leaf, ch increased withrising atmospherical CO₂ concentration, c_a, ci/c_a was somewhatdecreased. Total canopy resistance, R_c, was generally lowerat high CO₂ levels, despite higher leaf resistance. Higher canopyCO₂ loss at night and faster light extinction in a larger-sizedhigh CO₂ canopy were major drawbacks which prevented a furtherincrease in dry matter production (the harvest index was increasedby a factor 1.83). Key words: CO₂ enrichment, canopy CO₂ exchange, carbon balance, water use efficiency, leaf and canopy resistance     

Growth of Lettuce, Onion, and Red Beet. 1. Growth Analysis, Light Interception, and Radiation Use Efficiency

A field experiment was carried out to analyse the growth oflettuce, onion and red beet in terms of: (a) canopy architecture,radiation interception and absorption; (b) efficiency of conversionof absorbed radiation into biomass; and (c) dry matter partitioning.Growth analysis, total solar radiation interception, PAR interceptionand absorption by the crop canopy, ground cover, maintenancerespiration of onion bulbs and red beet storage roots were measured.Models for different leaf angle distribution and ground coverwere used to simulate light transmission by the crop canopy. The three crops are shown to have contrasting growth patternsfrom both a morphological and a physiological point of view.Lettuce showed very high light interception and growth afterthe early growth stages but, throughout the growth cycle, thisleafy crop showed the lowest radiation use efficiency due tothe respirational cost of the high leaf area. Onion showed alower early relative growth rate than lettuce and red beet.This was due partly to the low light interception per unit leafarea in the later stages of growth and partly to the low initialradiation use efficiency compared with the other two crops.On the other hand, thanks to more uniform distribution of theradiation inside the canopy, to the earlier termination of leafdevelopment and to the very low level of bulb respiration, onionshowed high radiation use efficiency and was able to producea large amount of dry matter. Red beet leaf posture and canopystructure resulted in high light interception and absorption.Its radiation use efficiency was lower than that of onion, partlyperhaps because of the more adverse distribution of the interceptedradiation fluxes within the canopy and partly because of thehigh respiration cost of a continuous dry-matter allocationto the leaves. However, this crop can accumulate a very largeamount of dry matter as leaf blade development and storage rootgrowth can both continue almost indefinitely, providing continuouslyavailable sinks. Ground cover gave a good estimate of the PAR interception onlyat low values of light interception but, in general, it underestimatedPAR interception in all three crops. Ratios between attenuationcoefficients established by considering PAR or total solar radiationand LAI or ground cover were calculated. Lettuce,Lactuca sativa L. var.crispa ; onion,Allium cepa L.; red beet; Beta vulgaris L. var.conditiva ; growth analysis; light interception and absorption; canopy architecture; ground cover; radiation use efficiency; maintenance respiration rate; dry matter distribution     

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

Photosynthesis of Stands of Tomato, Cucumber and Sweet Pepper Measured in Greenhouses under Various CO2-concentrations

The rate of net photosynthesis (P) of whole plant stands oftomato (Lycopersicon esculentum Mill.), cucumber (Cucumis sativusL.) and sweet pepper (Capsicum annuum L.) was measured in sixlong-term experiments in large greenhouses under normal operatingconditions and CO₂-concentrations between 200 and 1200 µmolmol^-1. The objective was to quantify the responses to lightand carbon dioxide and to obtain data sets for testing simulationmodels. The method of measuring canopy photosynthesis involvedan accurate estimation of the greenhouse CO₂ balance, usingnitrous oxide (N₂O) as tracer gas to determine, on-line, theexchange rate between greenhouse and outside air. The estimatedrelative error in the observed P was about ± 10%, exceptthat higher relative errors could occur under particular conditions. A regression equation relating P to the photosynthetically activeradiation, the CO₂ concentration and the leaf area index explained83-91% of the variance. The main canopy photosynthesis characteristicscalculated with the fitted regression equations were: canopyP_max 5-9 g m^-2 h^-1 CO₂ uptake; ratio P_max/LAI 1·5-3 gm^-2 h^-1; light compensation point 32-86 µmol s^-1 m^-2;light use efficiency (quantum yield) at low light 0·06-0·10µmol µmol^-1 and CO₂ compensation point 18-54 µmolmol^-1. The results were related to the prevailing conditions.Copyright1994, 1999 Academic Press Canopy photosynthesis, Capsicum annuum L., carbon dioxide, CO₂, CO₂ balance, CO₂ use efficiency, cucumber, _{Cucumis sativus} L., glasshouse, greenhouse, light use efficiency, Lycopersicon esculentum Mill., sweet pepper, tomato, tracer gas     

Flag Leaf Anatomy of Triticum and Aegilops Species in Relation to Photosynthetic Rate

Quantitative anatomical and other measurements were made onfully expanded flag leaves of a series of diploid, tetraploidand hexaploid Triticum and Aegilops species, and photosyntheticrates per unit leaf area were measured at light saturation (P_max). Diploids had the highest P_max, hexaploids the lowest with tetraploidsbeing intermediate. The anatomical features of tetraploids andhexaploids were generally similar, but different from the diploids.The diploids had thinner leaves with less dry matter and chlorophyllper unit area. The surface area of the mesophyll cells per unitvolume of mesophyll tissue was similar for all ploidy levels,as was the ratio mesophyil cell surface area per unit leaf area.It is argued that while these anatomical features are unlikelyto account for the observed variation in Pmax, it is possiblethat other structural factors with which they are correlatedmay causally influence P_max. One such feature is the averagediffusion path length from the plasmalemma at the cell surfaceto the sites of carboxylation. Anatomy, photosynthesis, mesophyll, cell size, Triticum, Aegilops, polyploidy     

Dynamic Model of Leaf Photosynthesis with Acclimation to Light and Nitrogen

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

A Generic Equation for Nitrogen-limited Leaf Area Index and its Application in Crop Growth Models for Predicting Leaf Senescence

Appropriate quantification of leaf area index (LAI) is importantfor accurate prediction of photosynthetic productivity by cropgrowth models. Estimation of LAI requires accurate modellingof leaf senescence. Many models use empirical turnover coefficients,the relative leaf-death rate determined from frequent fieldsamplings, to describe senescence during growth. In this paper,we first derive a generic equation for nitrogen-determined photosyntheticallyactive LAI (LAI_N), and then describe a method of using thisequation in crop growth models to predict leaf senescence. Basedon the theory that leaf-nitrogen at different horizons of acanopy declines exponentially, LAI_N, which is counted from thetop of the canopy to the depth at which leaf-nitrogen equalsthe minimum value for leaf photosynthesis, is calculated analyticallyas a function of canopy leaf-nitrogen content. At each time-stepof crop growth modelling, LAI_Nis compared to an independentcalculation of the non-nitrogen-limited LAI assuming no leafdeath during that time-step (LAI_NLD). In early stages, LAI_Nishigher than LAI_NLD; but with the advancement of crop growth,LAI_Nwill become smaller than LAI_NLD. The difference betweenLAI_NLDand LAI_N, whenever LAI_Nis smaller than LAI_NLD, gives theestimate of leaf area senesced at the time-step; the senescedleaf area divided by specific leaf area (SLA) gives the estimateof senesced leaf mass. The method was incorporated into twocrop models and the models adequately accounted for the LAIobserved in field experiments for rice and barley. The novelfeatures of the approach are that: (1) it suggests a coherent,biologically reasonable picture of leaf senescence based onthe link with photosynthesis and leaf nitrogen content; (2)it avoids the use of empirical leaf-turnover coefficients; (3)it avoids over-sensitivity of LAI prediction to SLA; and (4)it is presumably of sufficient generality as to be applicableto plant types other than crops. The method can be applied tomodels where leaf-nitrogen is used as an input variable or issimulated explicitly. Copyright 2000 Annals of Botany Company Leaf area index, leaf senescence, canopy nitrogen, modelling