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     

Modelling the Dynamical Components of the Sugar Beet Crop

Many models have been constructed to describe the growth ofthe sugar beet crop up to harvesting. In general, these modelshave a complex physiological basis, requiring a large numberof parameters yet relying on empirical functions with no mechanisticbasis to partition assimilates within the crop. An importantfactor in considering the growth of the crop, both from an economicand environmental point of view, is the response of the cropto varying amounts of available nitrogen in the soil. In thispaper, a model is described for crop growth using soil nitrogencontent and solar radiation as driving functions. The parsimoniousapproach to construction resulted in a 14 parameter model, sevenof which are associated with the driving variables. This issubstantially fewer than for other crop models. The model containsa new dynamical way of describing partitioning of assimilatesbetween shoot, storage root and fibrous roots. The partitioningmodel is derived from observations on the effect of soil nitrogenon crop growth. Interception of light is determined by foliagecover, which makes the model suitable for use with data collectedfrom satellite imaging. The model fits well to three independentdata sets with estimated parameters lying within biologicallyreasonable bounds. The model is used to test the sensitivityof yield to changes in soil nitrogen. Modelling; partitioning; parameter estimation; sugar beet; Beta vulgarisL.; nitrogen; crop growth dynamics     

An Analysis of the Relation between Dry Matter Allocation to the Tuber and Earliness of a Potato Crop

Compared with late cultivars, early potato cultivars allocatea larger part of the available assimilates to the tubers earlyin the growing season, leading to shorter growing periods andlower yields. A dynamic simulation model, integrating effectivetemperature and source –sink relationships of the crop,was used to analyse this relation, using data from experimentsin the Netherlands carried out over 5 years. Dry matter allocationto the tuber in these field experiments was simulated well whenthe tuber was considered as a dominant sink that affects earlinessof a potato crop in two ways: early allocation of assimilatesto the tubers stops foliage growth early in the season and reducesthe longevity of individual leaves. In a sensitivity analysisthe influence of tuber initiation, leaf longevity and the maximumrelative tuber growth rate (R_tb) on assimilate allocation andcrop earliness was evaluated. It was found that the maximumrelative tuber growth rate can influence crop earliness morethan the other two factors, but when conditions for tuber growthare optimal, the leaf longevity is most important. Solanum tuberosumL.; simulation model; source –sink relationships; cultivars     

Dark Respiration of Winter Wheat Crops in Relation to Temperature and Simulated Photosynthesis

The dark respiration of shoots (measured between March and anthesisin mid-June) and of ears (measured between anthesis and maturityat end of July) of winter wheat crops, grown in 1982 and 1985under different nitrogen application and irrigation conditions,was determined in the field. The respiration rate of 126 averagesof four samples was measured hourly for a 12–14-h darkperiod including the night. Respiration (expressed per unitdry mass) generally declined through the season for both shootsand ears. The average rate of respiration obtained on the samenight was greater for fertilized and irrigated crops, comparedwith unfertilized and droughted crops. The relationship betweenthe measured respiration and photosynthesis, simulated usinga modified version of the model developed by Spitters (1986),was analysed. This revealed that: (a) Shoot respiration was less well correlated with photosynthesisfrom the day preceding measurement than with the average ofthe photosynthesis from the two days preceding measurement. (b) The constants relating shoot respiration to total crop photosynthesisper unit crop mass and ear respiration to total crop photosynthesisper unit ear mass had similar values. This suggests that allgrowth respiration takes place in the ears at the end of theseason. (c) Crop growth respiration consumes about 35% of assimilatebefore anthesis, and that growth respiration of the ear consumesabout 40% of assimilate at the end of the season. (d) No significant effect of treatment on the relationship betweenrespiration and photosynthesis was detected, suggesting thatthe observed effect of treatment on respiration is due entirelyto differences in photosynthesis. Triticwn aestivum var. Avalon, winter wheat, dark respiration, growth coefficient, photosynthesis model, nitrogen nutrition, irrigation     

Validation of a Photosynthesis Model through the Use of the CO2Balance of a Greenhouse Tomato Canopy

Several leaf photosynthesis models were developed from wellcontrolled experiments in growth chambers. However, only a fewhave been validated under greenhouse conditions for their quantitativeand qualitative adequacy. In this paper, rates of net photosynthesisfor a tomato crop (Lycopersicon esculentum Mill) were measuredin a semi-commercial greenhouse (615 m³) for a significant timeperiod. Concomitant measurements of climatic conditions andLAI were used for simulation of net photosynthesis using theTOMGRO model which integrates Acock's model for photosynthesiscalculations. From simulations and from sensitivity analysis,the prediction of net photosynthesis appeared to be very sensitiveto the quantum use efficiency. The Acock model with originalparameters underestimated the net photosynthesis rate, but anincrease in the quantum use efficiency by 10% gave a good fit.In an effort to generalize the validity of the model, a residualanalysis was performed and showed a systematic bias relatedto light intensity intercepted by the canopy. The Marquardtalgorithm was used to adjust our data to the model but did noteliminate residual heterogeneity of variance with new parametervalues. On the basis of collected data, the criteria of goodnessof fit used showed that the photosynthesis model is inadequatein describing the CO₂-balance of the greenhouse agrosystem.However, it was determined that it could be used as a submodelwithin a more complex model for predicting growth and development.Copyright 1999 Annals of Botany Company Greenhouse, CO₂-balance, photosynthesis, TOMGRO model, Acock's model, residuals, tomato Lycopersicon esculentum Mill.     

Effects of Low Oxygen on Photosynthesis, Translocation and Growth in Green Pepper (Capsicum annuum)

Green pepper (Capsicum annuum cv. Bell Boy) plants were exposedin chambers to low (2%) oxygen and controlled carbon dioxideconcentrations. Vegetative and fruiting plants showed short-termincreases in net photosynthesis in low oxygen or elevated carbondioxide (up to 900 µl CO₂ l^–1). Photosynthesis ofyoung vegetative plants increased in low oxygen in the short-termbut there was no long-term benefit. Low oxygen enhancement ofphotosynthesis declined with time and after 10 d, leaf areaand root dry weight were less than in plants grown in normalair. Labelled assimilates were translocated from leaves to otherregions at similar rates in low oxygen and normal air. Low oxygenreduced respiratory losses from leaves and reduced the proportionof soluble carbohydrate converted to polysaccharide in all plantparts. Thus, low-oxygen environments decrease the utilisationof assimilates which then may lead to inhibition of photosynthesis. Capsicum annuum, photosynthesis, photorespiration, translocation, utilization of assimilates     

Optimal Nitrogen Content and Photosynthesis in Cauliflower (Brassica oleracea L. botrytis). Scaling up from a Leaf to the Whole Plant

A simple model of photosynthesis is described which is dependenton leaf area, organic nitrogen content and distribution withinthe canopy as well as on the light and temperature environments.The model is parameterized using a cauliflower crop as an example.The optimized protein-N profile within the canopy is calculatedwith respect to daily growth rate. By comparison with measuredprotein-N contents, the amount of super-optimal N-uptake, i.e.the N-uptake which does not increase productivity, is assessedfor two different nitrogen and light treatments. The amountof super-optimal N accumulated in cauliflower depends on N-supplyand can exceed 80 kg N ha^-1. Copyright 2000 Annals of BotanyCompany Brassica oleracea L. botrytis, cauliflower, nitrogen, photosynthesis, respiration, model, optimization     

A Comparison of Two Approaches for Modelling Cassava (Manihot esculenta Crantz.) Crop Growth

Two approaches for modelling the growth and development of cassava,Manihot esculenta Crantz, are described and evaluated. The twomodels differ only in the hypotheses accounting for storageroot growth. In model 1, assimilate allocation to storage rootsis governed by the combined Chanter's (1976: Mathematical modelsin mushroom research and production. PhD Thesis, Universityof Sussex, UK) growth equation; and in model 2 the spill-overhypothesis for assimilate allocation to storage root governsstorage root growth. In both models, canopy photosynthesis generatesthe carbon substrate required for all growth processes. Thegrowth rates of leaves, stems and storage roots are definedby growth equations subject to substrate saturation kinetics.A key feature of both models is that the growth demands of thestem, fibrous roots and storage roots are related to leaf demandrates. Allocation to stems and branches was modelled by meansof a modified logistic growth equation which includes all theparameters and variables (number of nodes, internode lengths,stem density, stem modulus of elasticity and branch tensilestrength) that define the limits of the load bearing capacityof the shoot's supportive structures. The correlation coefficientsfor determination of yield prediction for the models werer =0.898,P =0.0385 (model 1) and r =0.954, P =0.0117 (model 2). For agrowth season of 290 d (after which leaf area index equals zeroand crop growth ceases), both models simulate the sigmoidaltransition from the lag to exponential phase of crop growth.Both models are equally well corroborated by observed data;however, model 1 has greater explanatory power. Copyright 2000Annals of Botany Company Allocation, simulation, model, crop growth, cassava, Manihot esculenta Crantz.     

A Model for the Simulation of the Growth of Beta vulgaris L.

A simple explanatory model for the simulation of the growthof Beta is described. It takes account of photosynthesis, respiration,and the light-intercepting properties of the foliage, but usesthe observed distribution pattern of dry matter. A crop of Beta was grown and, by periodic sampling of plants,growth curves were obtained.These are compared with simulatedgrowth curves and the reasons for the discrepancies are discussed. From simulations with different values of the constants of themodel, the effects on yield of variations in these constantsare predicted     

Modelling the Components of Plant Respiration: Representation and Realism

This paper outlines the different ways in which plant respirationis modelled, with reference to the principles set out in Cannelland Thornley (Annals of Botany85: 55–67, 2000), firstin whole-plant ‘toy’ models, then within ecosystemor crop models using the growth-maintenance paradigm, and finallyrepresenting many component processes within the Hurley Pasture(HPM) and Edinburgh Forest Models (EFM), both of which separateC and N substrates from structure. Whole-plant models can beformulated so that either maintenance or growth respirationtake priority for assimilates, or so that growth respirationis the difference between total respiration and maintenanceassociated with the resynthesis of degraded tissues. All threeschemes can be converted to dynamic models which give similar,reasonable predictions of plant growth and respiration, butall have limiting assumptions and scope. Ecosystem and cropmodels which use the growth-maintenance respiration paradigmwithout separating substrates from structure, implicitly assumethat maintenance respiration is a fixed cost, uncoupled to assimilatesupply, and use fixed rate coefficients chosen from a rangeof measured values. Separation of substrates in the HPM andEFM enables estimates to be made of respiration associated withlocal growth, phloem loading, ammonium and nitrate N uptake,nitrate reduction, N₂fixation and other mineral ion uptake,leaving a ‘residual maintenance’ term. The lattercan be explicitly related to C substrate supply. Simulated changesin grassland respiration over a season and forest respirationover a rotation show that the ratio of total respiration togross canopy photosynthesis varies within the expected limitedrange, that residual maintenance accounts for 46–48% oftotal respiration, growth 36–42%, phloem loading 10–12%and the other components for the small remainder, with the ratiosbetween components varying during a season or forest rotation.It is concluded that the growth-maintenance approach to respiration,extended to represent many of the component processes, has considerablemerit. It can be connected to reality at many points, it givesmore information, it can be examined at the level of assumptionsas well as at the level of predictions, and it is open to modificationas more knowledge emerges. However, currently, there are stillparameters that require adjustment so that the predictions ofthe model are acceptable. Copyright 2000 Annals of Botany Company Respiration, photosynthesis, growth, maintenance, substrate, N uptake, mineral uptake, phloem loading, model.     

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.     

Growth Response of a Chrysanthemum Crop to the Environment. III, Effects of Radiation and Temperature on Dry Matter Partitioning and Photosynthesis

Vegetative crops of chrysanthemum were grown for 5 or 6 weekperiods in daylit assimilation chambers. Crop responses to differentradiation levels and temperatures were analysed into effectson dry matter partitioning, specific leaf area, leaf photosynthesisand canopy light interception. The percentage of newly formed dry matter partitioned to theleaves was almost constant, although with increasing radiationor decreasing temperature, a greater percentage of dry matterwas partitioned to stem tissue at the expense of root tissue.There was a positive correlation between the percentage of drymatter in shoot material and the overall carbon: dry matterratio. Canopy photosynthesis was analysed assuming identical behaviourfor all leaves in the crop. Leaf photochemical efficiency wasonly slightly affected by crop environment. The rate of grossphotosynthesis per unit leaf area at light saturation, P_A (max),increased with increasing radiation integral, but the same parameterexpressed per unit leaf dry matter, P_w (max) was almost unaffectedby growth radiation. In contrast, P_A (max) was hardly affectedby temperature but P_w (max) increased with increasing growthtemperature. This was because specific leaf area decreased withdecreasing temperature and increased with decreasing radiation.There was a positive correlation between canopy respirationintegral and photosynthesis integral, and despite a four-foldchange in crop mass during the experiments, the maintenancecomponent of canopy respiration remained small and constant. Canopy extinction coefficient showed no consistent variationwith radiation integral but was negatively correlated with temperature.This decrease in the efficiency of the canopy at interceptingradiation exactly cancelled the increase in specific carbonassimilation rate that occurred with increasing growth temperature,giving a growth rate depending solely on the incident lightlevel. Chrysanthemum, dry matter partitioning, photosynthesis, specific leaf area     

A Physiological Approach to the Analysis of Crop Growth Data: 3. The Effects of Repeated Short Term Soil Water Deficits on the Growth of Spaced Plants of the Legume, Macroptilium atropurpureum cv. Siratro

A physiological analysis of crop growth, reported in the previouspapers in this series, is extended to deal with the growth ofspaced plants, and applied to the growth data for M. atropurpureumsubject to several successive cycles of water deficit. Differences in growth between stressed and unstressed plantscan be quantitatively attributed to a reduction in the photosyntheticactivity of the leaves of stressed plants induced by increasingtheir leaf water deficits, and to an increase in the rate ofleaf abscission by the stressed plants. Macroptilium atropurpureum, growth analysis, water stress, photosynthesis, leaf abscission, stomatal conductance     

Use of the Expolinear Growth Model to Analyse the Growth of Faba bean, Peas and Lentils at Three Densities: Predictive Use of the Model

The expolinear equation for crop growth (Goudriaan and Monteith,Annalsof Botany66: 695–701, 1990; Goudriaan, 1994. In: MonteithJL, Scott RK, Unsworth MH, eds.Resource capture by crops.Nottingham:Nottingham University Press, 99–110) has the potentialto predict growth under specified environmental conditions.The choice of suitable parameters is discussed and tested usingdata from field experiments with faba bean, peas and lentils(Ishag and Dennett,Annals of Botany82: 497–505. 1998).It is suggested that suitable parameters for predicting cropgrowth are the fraction of solar radiation intercepted at emergence,the extinction coefficient for solar radiation, the maximumfraction of radiation intercepted, the maximum relative growthrate and the maximum crop growth rate for a crop interceptingall the incident radiation. In the experiments considered, thedifferences in growth patterns were due mainly to differencesin the maximum relative growth rate associated with differencesin temperature.Copyright 1998 Annals of Botany Company Expolinear equation, grain legumes, crop growth rate, crop density, relative growth rate, growth modelling, faba bean,Vicia fabaL., peas,Pisum sativumL., lentils,Lens culinarsMedic.     

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     

Modelling Light Absorption and Canopy Net Photosynthesis of Glasshouse Row Crops and Application to Cucumber

A model of light absorption and photosynthesis applicable toglasshouse row crops is constructed and applied to cucumber.Light absorption is calculated using a method suggested fordiscontinuous canopies; photosynthesis is modelled with a non-rectangularhyperbola. The predictions of this model are compared with experimentaldata in the preceding paper. Here the model is used to simulateresponses to light and CO₂ concentration and especially to examinethe effects of varying the parameters of the crop that can becontrolled by the grower. These include the number of plantsin each row, the number and width of the rows, the gap betweenrows, and the height of the crop. For example, it is shown that,for high values of crop net photosynthesis, the number of rowsis more important at high light than at low light, whereas cropheight is more important at low light than at high light. Theimplications of these and other findings are discussed. Key words: Cucumis sativus L., glasshouse crops, cucumber, model, light absorption, photosynthesis, CO₂, row crops, simulation     

The Photosynthesis and Cell Shape Rhythms can be Naturally Uncoupled from the Biological Clock in Euglena gracilis

A naturally occurring population of Euglena (Klebs Strain Z)cells, with unusual biological clock properties, has been isolated.The photosynthesis reactions, which are usually controlled bythe biological clock, are uncoupled from the clock in the newpopulation. The rate of oxygen evolution is influenced predominantlyby the environmental growth parameters instead of the biologicalclock. In addition, the rhythm in cellular shape has a differenttiming from the control population and can be temporarily uncoupledfrom the clock by lowering the light intensity used for growth Key words: Biological clock, photosynthesis, Euglena     

Growth and Photosynthesis of Mutant Pea Seedlings

The recessive of gene, producing tendrils in place of leaves,and the recessive st gene, reducing stipule size, produce phenotypesof pea that are termed leafless (afafstst) and semi-leafless(afafStSt). Photosynthesis and growth of these two types werecompared with the conventional phenotype (AfAfStSt) during thefirst 9 days of post-emergent growth. The conventional seedlingshowed faster net photosynthesis per unit dry weight than theleafless phenotype, whilst the semi-leafless seedlings wereintermediate. Differences in dark respiration were small butleafless seedlings had significantly higher rates soon afteremergence. Where the three phenotypes used were isogenic, except for ofand st, the rates of shoot growth were in the same ranking orderas net CO₂ uptake. With three other genotypes, representingthe three phenotypes, more similar shoot growth was found betweenthe conventional and semi-leafless phenotype, possibly becauseof compensating differences in embryonic axis size. The ratesof growth of roots and the rates of dry weight loss from thecotyledons showed no consistent differences between phenotypes. The results are discussed in relation to the potential for thesemi-leafless phenotype as an alternative to the conventionalphenotype for the dried pea crop. Pea seedling, Pisum sativum, leafless pea, photosynthesis, seedling growth