Relative Nitrogen Limitation at Steady-state Nutrition as a Determinant of Plasticity in Five Grassland Plant Species

Partitioning of biomass between roots and different shoot partshas often been used to explain the response of plants to variationsin resource availability. There are still many uncertaintiesin the importance of this trait for plant performance, and clearguidelines on how partitioning should be quantified in relationto growth rate and resource supply are of fundamental importancefor such an understanding. This paper reports an attempt toshow how plant nitrogen status relates to root:shoot partitioningand other plastic responses, in a manner that can be used forquantitative predictions. The reactions to nitrogen limitationof five grassland plant species, with different ecological demands,were compared. The species used were the forbs Polygala vulgarisand Crepis praemorsa, and the grasses Danthonia decumbens, Agrostiscapillaris and Dactylis glomerata. The experiment was conductedin a climate chamber where the plants were grown hydroponically(1) under non-limiting nutrient conditions and (2) at a steady-statenitrogen limitation, which enabled the plants to express halfof their growth potential. The relative growth rate (RGR) ofthe species was strongly related to plant nitrogen concentration(PNC) and leaf area ratio (LAR), whereas the effects on netassimilation rate (NAR) were very small. Despite large differencesin maximum relative growth rate, the species showed remarkablesimilarities in dry matter partitioning between root and shoot.It is concluded that root:shoot partitioning can be treatedas a direct function of the relative resource limitation ofthe plant. The difficulty of attaining well-defined levels ofresource limitation in soil, other solid substrates and manyhydroponic systems may be the most important reason for thedivergent results in earlier studies. Better knowledge of soil-rootinteractions, and plant responses to the whole span of resource-supplylevels, is required for a thorough understanding of how nutrientslimit growth. Copyright 1999 Annals of Botany Company Growth rate, plant strategies, plasticity, partitioning, biomass, nitrogen, nutrient limitation, grassland.     

Nutrient-limited growth rates: quantitative benefits of stress responses and some aspects of regulation

Young sunflower plants (Helianthus annuus L.) under stress oflow nitrate or phosphate availability exhibited increases inroot: shoot ratio and in kinetic parameters for uptake. Theyshowed no significant changes in photosynthetic utilizationof either nutrient. Increases in root: shoot ratio were achievedby early and persistent suppression of shoot growth, but notroot growth. Affinity for phosphate uptake, 1/K_m(P), increasedwith phosphate stress, as did affinity for nitrate uptake, 1/K_m(N),with nitrate stress. Maximal uptake rate, V_max, for phosphateuptake increased with phosphorus stress; V_max for nitrate didnot increase with nitrogen stress. Phosphate V_max was relatedstrongly to root nutrient status. Decreases in V_max with plantage were not well explained by changes in age structure of roots.Estimated benefits of acclimatory changes in root: shoot ratioand uptake kinetics ranged up to 2-fold increases in relativegrowth rate, RGR. The relation of RGR to uptake physiology followedpredictions of functional balance moderately well, with somesystematic deviations. Analyses of RGR using growth models implyno significant growth benefit from regulating Vmax, specifically,not from down-regulating it at high nutrient availability. Quantitativebenefits of increases in root: shoot ratio and uptake parametersare predicted to be quite small under common conditions whereinnutrient concentrations are significantly depleted by uptake.The root: shoot response is estimated to confer the smallestbenefit under non-depleting conditions and the largest benefitunder depleting conditions. Even then, the absolute benefitis predicted to be small, possibly excepting the case of heterogeneoussoils. Depleting and non-depleting conditions are addressedwith very different experimental techniques. We note that atheoretical framework is lacking that spans both these cases,other than purely numerical formulations that are not readilyinterpreted. Key words: Nutrient stress, nutrient uptake, nutrient use efficiency, relative growth rate, Helianthus annuus     

The Effect of Root Temperature on Growth and Uptake of Ammonium and Nitrate by Brassica napus L. in Flowing Solution Culture: I. GROWTH

Macduff, J. H., Hopper, M. J. and Wild, A. 1987. The effectof root temperature on growth and uptake of ammonium and nitrateby Brassica napus L. in flowing solution culture. I. Growth.—J.exp. Bot. 38: 42–52 Oilseed rape (Brassica napus L. cv. Bien venu) was grown for49 d in flowing nutrient solution at pH 6?0 with root temperaturedecrementally reduced from 20?C to 5?C; and then exposed todifferent root temperatures (3, 5, 7, 9, 11, 13,17 or 25?C)held constant for 14 d. The air temperature was 20/15?C day/nightand nitrogen was supplied automatically to maintain 10 mmolm^–3 NH₄NO₃ in solution. Total dry matter production wasexponential with time and similar at all root temperatures givinga specific growth rate of 0?0784 g g^–1 d^–1. Partitioningof dry matter was influenced by root temperature; shoot: rootratios increased during treatment at 17?C and 25?C but decreasedafter 5 d at 3?C and 5?C. The ratio of shoot specific growthrate: root specific growth rate increased with the ratio ofwater soluble carbohydrates (shoot: root). Concentrations ofwater soluble carbohydrates in shoot and root were inverselyrelated to root temperature; at 3, 5 and 7?C they increasedin stem + petioles throughout treatment, coinciding with a decreasein the weight of tissue water per unit dry matter. These resultssuggest that the accumulation of soluble carbohydrates at lowtemperature is the result of metabolic imbalance and of osmoticadjustment to water stress. Key words: Brassica napus, oilseed rape, root temperature, specific growth rate     

Determination of a Critical Nitrogen Dilution Curve for Winter Oilseed Rape

Several controlled environmental and field experiments werecarried out to define the critical nitrogen dilution curve forwinter oilseed rape, cultivar Goeland. This curve is describedby the following power equation:N=4.48 W^-0.25,whereNis the totalnitrogen concentration in the shoot biomass andWthe shoot biomass.This curve has been validated over the range of shoot dry matterof 0.88 to 6.3 t ha^-1. For lower shoot biomasses this equationoverestimated the critical nitrogen concentration; we proposea constant value of 4.63 (Nis expressed in reduced N, whichis a more stable N fraction in the shoot at these stages ofdevelopment). These results have been validated in several pedoclimaticconditions in France on a single variety in 1994 and 1995. Thehigher position of this curve relative to the C₃species referencecurve (Greenwoodet al.,Annals of Botany67: 181–190, 1990)can be explained by the experimental conditions obtained byGreenwoodet al. (1990); therefore, all their rape data are ratherclose to the critical curve that we propose. The differencesfound between wheat and winter oilseed rape critical N dilutioncurves correspond to their respective leaf:stem dry matter ratioand the specific leaf loss phenomenon occuring in rape. Winteroilseed rape has a higher capacity of N accumulation in itsshoot than wheat for the same aerial dry matter. The proportionof nitrate in shoots rises with the nitrogen nutrition index(N.N.I.) and is more important for rapeseed than for wheat forthe same N.N.I. This difference is especially high at the beginningof flowering when the shade provided by the canopy of rapeseedflowers decreases nitrate reductase activity.Copyright 1998Annals of Botany Company Winter oilseed rape;Brassica napusL.; plant N concentration; nitrate; reduced N; shoot biomass; critical nitrogen concentration; dilution curve; N productivity.     

Relationships Between Nitrogen and Water Concentration in Shoot Tissue ofMolinia caeruleaDuring Shoot Development

Plants ofMolinia caeruleawere supplied with either a low (0.2mol m^-3) or high (10 mol m^-3) supply of nitrogen over two growingseasons. A total of 14 destructive plant harvests were made:when plants were in an over-wintering state prior to the secondseason; immediately following bud burst; and on 12 further occasionsthroughout the second season. The relationships between shootnitrogen concentration on a dry mass basis, shoot water contentand plant developmental stage were investigated. Shoot nitrogenconcentration on a dry mass basis fell as the growing seasonprogressed. In contrast, the concentration of nitrogen in tissuewater after bud burst showed only a slight reduction. The concentrationof nitrogen both on a dry mass basis and in tissue water wasgreater for plants receiving the higher supply of nitrogen.Shoot water content was highest immediately following bud burstthen declined as the season progressed, with plants receivingthe low nitrogen supply having slightly greater shoot watercontents. It was concluded that the decline in shoot nitrogenconcentration ofM. caeruleaon a dry mass basis as the mass increasedwas mainly explained by changes in shoot water content. Theobserved increase in the rate of decline of both shoot nitrogenconcentration and water content with increased shoot mass coincidedwith the cessation of leaf tissue production and was thereforedue to a switch from the production of leaves to other tissues.Copyright1999 Annals of Botany Company Molinia caerulea(L.), purple moor grass, nitrogen, water content, shoot development.     

The Uptake of Phosphate by Plants from Flowing Nutrient Solution: IV. EFFECT OF PHOSPHATE CONCENTRATION ON THE GROWTH OF TRIFOLIUM REPENS L. SUPPLIED WITH NITRATE, OR DEPENDENT UPON SYMBIOTICALLY FIXED NITROGEN

Breeze, V. G. and Hopper, M. J. 1987. The uptake of phosphateby plants from flowing nutrient solution. IV. Effect of phosphateconcentration on the growth of Trifolium repens L. suppliedwith nitrate, or dependent upon symbiotically fixed nitrogen.—J.exp. Bot. 38: 618–630. Nodulated white clover plants were subjected to a range of phosphateconcentrations in flowing solution culture (0.32 to 8.0 mmolm^–3 P) at 41 d from sowing, either supplied with nitrateor dependent on symbiotically-fixed nitrogen. No effect of phosphateconcentration in solution on dry matter production, relativegrowth rate, root/shoot ratio, or water soluble carbohydrateconcentration of the plant tissue was observed after 24 d fromthe start of the experiment, although the plants supplied withnitrate yielded more than the others. Phosphate uptake throughoutthe experimental period was related to the solution concentration,but the source of nitrogen did not affect the phosphorus concentrationsof the shoots. However, the roots of the plants dependent onsymbiotically-fixed nitrogen had higher concentrations of phosphorusthan those supplied with nitrate, but this did not appear tobe due to an increased phosphorus requirement for nitrogen fixation,because the amount fixed was unaffected by the phosphate concentrationin solution. The cation-anion balance showed that plants dependenton nitrogen fixation had no larger requirement for calcium thanplants supplied with nitrate, but a requirement for hydroxylions equivalent to over 130 kg lime per tonne of dry shoot.It is suggested that the enhanced phosphate uptake by plantsdependent on nitrogen fixation is due to this need for a cation-chargebalancing anion. Key words: Phosphate uptake, nitrogen fixation, Trifolium repens L., repens L., cation-anion balance, flowing solution culture     

A Mechanistic Analysis of Self-thinning in Terms of the Carbon Balance of Trees

The self-thinning behavior of a monoculture forest is examinedfrom a mechanistic viewpoint, in terms of the carbon balanceof trees at the stand level. Two approaches are described: thefirst is based on simple assumptions concerning the balancebetween growth, photosynthesis and respiration; the second usesthe process-based I.T.E. EDINBURGH FOREST model, extended toinclude tree birth and death, to examined the assumptions ofthe first approach at a more mechanistic level and to interpretthe observed variation in the responses to shading and soilfertility in terms of a single model. The first approach leads to a power law relation m n ^-1/y betweenmean biomass per tree (m) and the number of trees per unit groundarea (n), where y is the exponent characterising the rate atwhich respiration (r) scales with biomass (i.e. r m^y). Forvalues of y less than 1, m and n are predicted to increase anddecrease, respectively, as powers of time (t) such that thebiomass per unit ground area (M) increases linearly with t forany value of y. When y equals 1, m and n vary exponentiallywith t and M remains constant. When r is assumed to be proportional to stem cambium surfacearea, the value of y is shown to lie in the range 0·5to 1 depending on the rate of stem taper. This leads to slopeson a log m vs. log n self-thinning plot in range -1 to -2 andtypically around -1·5 for rates of taper based on mechanicalrequirements. For the second approach, the I.T.E. EDINBURGH FOREST model,a previously published mechanistic model of plantation growthat the stand level (Thornley, 1991), is extended to naturalstands by introducing average stem birth and death rates thatare functions of the carbon and nitrogen substrate status oftrees. Growth of even-aged and mixed-age forest over about 500years is simulated under a variety of environmental and physiologicalconditions. In general, the forest model predicts a curved log m vs. logn thinning line, but has a reasonably well-defined linear sectionunder most conditions. The slope of the linear portion flattensas the rate at which cambium surface area scales with structuralbiomass increases, in qualitative agreement with the analyticalprediction of the first approach. Quantitative differences betweenthe two approaches are interpreted in terms of the process representedin the forest model. In general, the height of the thinningline increases with irradiance, but is relatively insensitiveto soil nitrogen, with no change in slope in either case. However,at very low irradiance or soil nitrogen levels, the slope flattenscontinuously and the entire thinning line becomes markedly non-linear.Even-aged stands and mixed-age stands have different thinningslopes.Copyright 1993, 1999 Academic Press Self-thinning, carbon balance, mechanistic model, forest growth, respiration     

Nitrogen Uptake and Plant Growth I. Effect of Nitrogen Removal on Growth of Polygonum cuspidatum

Polygonun cuspidatum was grown hydroponically to examine theeffect of nitrogen removal from the nutrient solution upon plantgrowth and the partitioning of dry matter and nitrogen amongorgans. Nitrogen removal reduced the growth rate mainly dueto the reduced growth of leaf area. Accelerated root growthwas observed only in plants which earlier had received highlevels of nitrogen. Nitrogen removal caused almost exclusiveallocation of available nitrogen to root growth. Nitrogen fluxfrom the shoot to the root occurred in plants which had receivedlow nitrogen. Not only was net assimilation rate (NAR) littleaffected by nitrogen removal, but it also was not correlatedwith the concentration of leaf nitrogen on an area basis. Light-saturatedCO₂ exchange rate (CER) was highly correlated with the concentrationof leaf nitrogen. Nitrogen use efficiency (NUE) in CER (CERdivided by leaf nitrogen) remained constant against leaf nitrogen,indicating efficient use of nitrogen under light saturation,while NUE in terms of NAR decreased with higher concentrationof leaf nitrogen. Polygonum cuspidatum Sieb. et Zuce., CO₂ exchange rate, growth analysis, leaf nitrogen, net assimilation rate, nitrogen use efficiency, partitioning of dry matter and nitrogen     

Nitrogen Uptake and Plant Growth II. An Empirical Model of Vegetative Growth and Partitioning

An empirical model of vegetative plant growth is presented.The model is based on experimental data on Polygonum cuspidatum,which showed (1) that the partitioning of dry matter and nitrogenamong organs was linearly related to the nitrogen concentrationof the whole plant and (2) that leaf thickness was negativelycorrelated with leaf nitrogen concentration. The model properlydescribes the behaviour of plants. Steady-state solutions ofthe model give the relative growth rate, specific leaf weight,and partitioning of dry matter and nitrogen among organs withthe net assimilation rate and the specific absorption rate asenvironmental variables. The effect of nitrogen removal on drymatter and nitrogen partitioning was examined as non-steady-statedynamic solutions of the model. The model predicted not onlyreduced leaf growth and enhanced root growth but also a fluxof nitrogen from the leaf to the root, which agreed with theexperimental results. Mathematical model, partitioning of dry matter and nitrogen, plant nitrogen, relative growth rate, shoot: root ratio, specific leaf weight     

Determination of a Critical Nitrogen Dilution Curve for Winter Wheat Crops

A set of N-fertilization field experiments was used to determinethe 'critical nitrogen concentration', i.e, the minimal concentrationof total N in shoots that produced the maximum aerial dry matter,at a given time and field situation. A unique 'critical nitrogendilution curve' was obtained by plotting these concentrationsN_ct (% DM) vs. accumulated shoot biomass DM (t ha^-1). It couldbe described by the equation: N_ct = 5·35DM^-0·442 when shoot biomass was between 1·55 and 12 t ha^-1. Anexcellent fit was obtained between model and data (r² = 0·98,15 d.f.). A very close relationship was found using reducedN instead of total N, because the nitrate concentrations inshoots corresponding to critical points were small. The criticalcurve was rather close to those reported by Greenwood et al.(1990) for C₃ plants. However, this equation did not apply whenshoot biomass was less than 1·55 t ha^-1. In this case,the critical N concentration was independent of shoot biomass:the constant critical value N_ct = 4·4% is suggested forreduced-N. The model was validated in all the experimental situations,in spite of large differences in growth rate, cultivar, soiland climatic conditions; shoot biomass varying from 0·2to 14 t ha^-1. Plant N concentration was found to vary by a factor of fourat a given shoot biomass level. In the heavily fertilized treatments,shoot N concentration could be 60% higher than the criticalconcentration. Most (on average 80%) of the extra N accumulatedwas in the form of reduced N. The proportion of nitrate to totalN in shoot mainly depended on the crop stage of development.It was independent of the nitrogen nutrition level.Copyright1994, 1999 Academic Press Winter wheat, Triticum aestivum, arable crops, plant N concentration, aerial biomass, critical nitrogen, dilution curve, fertilization, reduced N, nitrate     

Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

Assimilate Partitioning in Red and White Clover Either Dependent on N2 Fixation in Root Nodules or Utilizing Nitrate Nitrogen

During vegetative growth in controlled environments, the patternof distribution of ¹⁴C-labelled assimilates to shoot and root,and to the meristems of the shoot, was measured in red and whiteclover plants either wholly dependent on N₂ fixation in rootnodules or receiving abundant nitrate nitrogen but lacking nodules. In experiments where single leaves on the primary shoot wereexposed to ¹⁴CO₂, nodulated plants of both clovers generallyexported more of their labelled assimilates to root (+nodules),than equivalent plants utilizing nitrate nitrogen, and thiswas offset by reduced export to branches (red clover) or stolons(white clover). The intensity of these effects varied with experiment.The export of labelled assimilate to growing leaves at the terminalmeristem of the donor shoot was not influenced by source ofnitrogen. Internode elongation in the donor shoot utilized nolabelled assimilate. Whole plants of white clover exposed to ¹⁴CO₂ on seven occasionsover 32 days exhibited the same effect on export to root (+nodules),which increased slightly in intensity with increasing plantage. Nodulated plants had larger root: shoot ratios than theirequivalents utilizing nitrate nitrogen. Trifolium repens, Trifolium pratense, red clover, white clover, nitrogen fixation, nitrate utilization, assimilate partitioning     

Differences in Nitrogen Metabolism During Growth of Plants from Aged Potato (Solanum tuberosum L.) Meristems

The effect of advanced meristem age on growth and accumulationof plant nitrogen (N) in potato (Solanum tuberosum L.) was studied.Etiolated plantlets, excised from sprouted, single-eye-containingcores from 7 and 19-month-old seed-tubers, were transplantedinto aerated nutrient culture. Rates of shoot and root dry matterand shoot soluble-N (which included nitrate-N) accumulationwere similar for plants from both meristem ages over a 30 dinterval of log-linear growth. The rate at which nitrate-N accumulatedwas consistently 17 per cent higher in shoots from 19-month-oldcompared to those from 7-month-old meristems. However, accumulationof free amino-N and soluble protein-N were 21 and 15 per centlower, respectively in shoots from 19-month-old meristems. Abuild-up of shoot nitrate, along with lower rates of accumulationof amino-N and soluble protein-N, suggests a lower capacityfor nitrate reduction during early growth of plants from oldermeristems. Furthermore, these effects can be attributed to age-inducedchanges in the meristem or bud tissue as the plants were separatedfrom the tuber tissue initially in the study. Long-term ageingof seed-potatoes apparently affects changes within meristemsthat translate into a lower capacity to accumulate reduced formsof nitrogen during early plant growth. Potatoes (Solanum tuberosum L.), meristem age, nitrogen metabolism, plant growth potential     

Effects of Low Temperature on Growth and Nutrient Accumulation in Rye (Secale cereale) and Wheat (Triticum aestivum)

Rye (Secale cereale cv. Rheidol) and wheat (Triticum aestivumcv. Mardler) were grown at shoot/root temperatures of 20/20°C (warm grown, WG plants), 8/8 °C (cold grown, CG plants)and 20/8 °C (differential grown, DG plants). Plants fromcontrasting growth temperature regimes were standardized andcompared using a developmental timescale based on accumulatedthermal time (°C d) at the shoot meristem. Accumulationof dry matter, nitrogen and potassium were exponential overthe time period studied (150–550 °C d). In rye, therates of plant dry matter and f. wt accumulation were linearlyrelated to the temperature of the shoot meristem. However, inwheat, although the rates of plant dry matter and f. wt accumulationwere temperature dependent, the linear relationship with shootmeristem temperature was weaker than in rye. The shoot/rootratio of rye was stable irrespective of growth temperature treatment,but the shoot/root ratio of wheat varied with growth temperaturetreatment. The shoot/root ratio of DG wheat was 50% greaterthan WG wheat. In both cereals, nutrient concentrations anddry matter content tended to be greater in organs exposed directlyto low temperatures. The mean specific absorption rates of nutrientswere calculated for the whole period studied for each species/temperaturecombination and were positively correlated with both plant shoot/rootratio and relative growth rate. The data suggest that nutrientuptake rates were influenced primarily by plant demand, withno indication of specific nutrient limitations at low temperatures. Nutrient accumulation, relative growth rate (RGR), rye, Secale cereale cv. Rheidol, temperature, thermal time, Triticum aestivum cv. Mardler, wheat     

Optimization of Tissue Nitrogen and Root-Shoot Allocation

A model is explored which describes the joint optimization oftissue nitrogen and root-shoot allocation in response to variationin nitrogen availability in the environment. The model plantis composed of root and shoot biomass and tissue nitrogen, andrelative growth rate is the parameter maximized. A 'source' (uptake limited) model of joint tissue nitrogen androot-shoot allocation is reviewed (Hilbert, 1990) that optimallybalances the uptake of carbon and nitrogen. Modifications includingroot respiratory costs and fixed root tissue nitrogen are explored.Then the role of tissue nitrogen in regulating 'sink' strengthis combined with the balanced source model, and modified byassuming separate tissue nitrogen involved in source vs. sink.Generally, the results indicate that as available nitrogen increases,optimal root allocation declines and tissue nitrogen increases.These results appear to be robust for more complicated versionsof the model provided that various internal nitrogen compartments'compete' for the same nitrogen.Copyright 1993, 1999 AcademicPress Plants growth, RGR, leaf nitrogen, nitrogen productivity, allocation, partitioning, root-shoot ratio     

Effect of elevated CO2 and nutrient status on growth, dry matter partitioning and nutrient content of Poa alpina var. vivipara L.

Poa alpina var. vivipara L. was grown in an atmosphere containingeither 340 or 680 µmol CO₂ mol^–1 within controlledenvironment chambers. The available nutrient regime was variedby altering the supply of nitrogen and phosphorus within a completenutrient solution. At a high, but not low, N and P supply regime,elevated CO₂ markedly increased growth. Differences betweennutrient supply, but not atmospheric CO₂ concentration, alteredthe allometric relations between root and shoot. Net photosynthesisof mature leaf blades and leaf N and P concentration were reducedin plants grown at the elevated CO₂ concentration. The question was asked: is it possible to ascribe all of theseeffects to elevated CO₂ or are some due to nutrient deficiencycaused by dilution with excess carbon? Several criteria, includingthe nutrient content of sink tissue, root:shoot allometry andthe use of divalent cations to estimate integrated water flowsare suggested in order to make this distinction. It is concludedthat only at a low supply of N and P₁ and elevated CO₂ concentration,was low leaf N concentration due to induced nutrient deficiency.The data are consistent with a model where the capacity of sinksto use photosynthetically assimilated carbon sets both the rateof import into those sinks (and thus rate of export from sourceleaves) and the rate of photosynthesis of source leaves themselves. Key words: Poa alpina L., growth, photosynthesis, carbohydrate, export, nitrogen, phosphorus     

The Rate of Growth and Partitioning of Assimilates in Young Grass Plants: A Mathematical Model

A model describing the increase in weight with time of younggrass plants is formulated. The parameters are the relativegrowth rates of the root and shoot systems; k, the ratio ofthe relative growth rate of the root system relative to thatof the shoot system; b, the weight of the root system when thatof the shoot system is unity, and u the rate of increase inweight of the whole plant per unit of shoot system per unitof time, k and b are the constants in the allometric formula,r = bs^k where r and are the weights of the root and shoot systems.The model enables the effect of changes in the distributionof assimilates between the root and shoot systems upon the rateof growth of the plant to be assessed. Data from a number ofexperiments are analysed in this manner and the significanceof the results discussed.     

The influence of the parasitic angiosperm Striga gesnerioides on the growth and photosynthesis of its host, Vigna unguiculata

Cowpea plants (Vigna unguiculata) infected with the root hemiparasiticangiosperm Striga gesnerioides accumulated less biomass thanuninfected plants over a growth period of 60 d. The allometricrelationship between shoot and root dry weight was similar inparasitized plants relative to control plants, as was the proportionof dry matter partitioned into leaf, stem and root tissue. However,infected plants failed to make any significant investment ofdry matter in pods. The rate of photosynthesis of the youngestfully expanded leaf of parasitized plants was significantlylower than for control plants. The lower rates of photosynthesiswere not attributable to stomatal limitation, a loss of chlorophyllor to an accumulation of carbohydrate. The depression of photosynthesisin the young leaves was transient. As control leaves aged, photosynthesisdeclined. This also occurred in Striga infected plants, butto a lesser extent resulting in higher rates of photosynthesisin mature leaves when compared to those of uninfected plants.The foliar nitrogen content of parasitized plants was higherthan control plants consistent with the slower rate of photosyntheticdecline of older leaves. The data are discussed with respectto the influence of parasitic weeds on host growth and photosynthesis. Key words: Cowpea, hemiparasite, allometry, nitrogen     

Growth and partitioning of C and fixed N in the shrub legume Acacia littorea in the presence or absence of the root hemiparasite Olax phyllanthi

Nodulated plants of Acacia littorea were pot cultured singlyin minus nitrogen sand culture in the presence or absence ofa transplanted seedling of the root hemiparasite Olax phyllanthiand harvests of cultures made 4 and 8 months after introducingthe parasite. Parasitism decreased host shoot growth while increasingroot growth to a similar extent. Final shoot:root dry weightratio was 2.2 for parasitized versus 4.3 for unparasitized Acacia.Partitioning of fixed N showed 4-fold larger N increments inshoots than roots of unparasitized plants, whereas parasitizedplants lost a small amount of shoot N, made a root gain of Ndouble that of unparasitized plants and lost over half of theirN to Olax. The increment of fixed N in the host:parasite associationwas similar to that of unparasitized Acacia. Data on dry mattergain per unit foliage area and mean CO₂ assimilation rates pershoot of Olax and Acacia (parasitized or unparasitized) werediscussed in relation to an estimated heterotrophic gain ofxylem C from the host equivalent to 40% of the increment ofdry matter C made by the parasite. Growth of Olax was accompaniedby large increases in numbers of haustoria, 9% of which wereattached to root nodules as opposed to roots. Structural andnutritional features of direct parasitism of nodules are described.Models of flow and utilization of C and N in the Acacia:Olaxassociation and unparasitized Acacia are discussed in relationto published data for other host:parasite associations. Key words: Olax phyllanthi, host-parasite relationships, C and N partitioning, Acacia, N₂ fixation     

The Acquisition and Utilization of Carbon in Early Spring by Kiwifruit Shoots

Measurements of net photosynthetic rate (at 1450µ molm^-2s^-1photosynthetically active radiation) of leaves, of leafand stem respiration, and of shoot growth of potentially-fruitinglaterals on kiwifruit (Actinidia deliciosa ) were used to estimateweekly shoot carbon balances over the first 10 weeks of shootgrowth (budburst to anthesis). Consistent differences in therate of shoot elongation, of internode expansion and of increasein basal diameter were found among shoots. Faster-growing (long)shoots acquired carbon by photosynthesis at a faster rate evenin the first few weeks after budburst, but the amount of carbonrequired to sustain this growth resulted in shoot carbon deficitswhich were approx. seven times greater than those of the slower-growing(short) shoots. It was estimated that the transition from shootcarbon deficit to carbon surplus occurred 3–4 weeks afterbudburst, irrespective of shoot growth rate. As a result ofsubsequent rapid increases in shoot photosynthetic rate, longshoots had a shoot carbon surplus of 4.4 g C week^-1in the weekbefore anthesis, approx. three times that of the short shoots.Defoliation (66%) of shoots 1 week after budburst, and subsequentremoval of later-emerging leaves to maintain the level of defoliation,had the effect of slowing shoot growth in the carbon deficitperiod, particularly for the long shoots. However, the durationof shoot expansion in the defoliated shoots was longer, resultingultimately in shoots which were longer than the control shoots.Linkages among early carbon balance dynamics of shoots, shootlength at anthesis, and fruit growth are discussed. Actinidia deliciosa ; kiwifruit; shoot growth; carbon acquisition; respiration; photosynthesis