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     

Influence of the Form of Nitrogen Supply on Root Uptake and Translocation of Cations in the Xylem Exudate of Maize (Zea mays L)

Concentrations of inorganic cations are often lower in plantssupplied with NH₄⁺ as compared with NO₃^–. To examine whetherthis is attributable to impaired root uptake of cations or lowerinternal demand, the rates of uptake and translocation of K,Mg, and Ca were compared in maize plants (Zea mays L.) withdifferent growth-related nutrient demands. Plants were grownin nutrient solution with either 1·0 mol m^–3 NO₃^–or NH₄⁺ and the shoot growth rate per unit weight of roots wasmodified by varying the temperature of the shoot base (SBT)including the apical shoot meristem. The shoot growth rate per unit weight of roots, which was takenas the parameter for the nutrient demand imposed on the rootsystem, was markedly lower at 12°C than at 24°C SBT.As a consequence of the lower nutrient demand at 12°C SBT,uptake rates of NO₃^– and NH₄⁺ declined by more than 50%Compared with NO₃^– supply, NH₄⁺ nutrition depressed theconcentrations of K and particularly of Ca in the shoot, bothin plants with high and with low nutrient demand. This indicatesa control of cation concentration by internal demand ratherthan by uptake capacity of the roots. Translocation rates of K, Mg and Ca in the xylem exudate werelower in NH₄⁺- than in NO₃^–-fed plants. Net accumulationrates of Ca in the shoot were also decreased, whereas net accumulationrates of K in the shoot were even higher in NH₄⁺-fed plants.It is concluded that reduced cation concentrations in the xylemsap of plants supplied with NH₄⁺ are due to the lower demandof cations for charge balance. The lower K translocation tothe shoot is compensated by reduced retranslocation to the roots.For Ca, in contrast, decreased translocation rates in NH₄⁺-fedplants result in lower shoot concentration. Key words: Nitrogen form, cation nutrition, charge balance, xylem exudate, recirculation     

Plant Growth in Relation to the Supply and Uptake of NO3-: A Comparison Between Relative Addition Rate and External Concentration as Driving Variables

Two approaches to quantifying relationships between nutrientsupply and plant growth were compared with respect to growth,partitioning, uptake and assimilation of NO₃^– by non-nodulatedpea (Pisum sativum L. cv. Marma). Plants grown in flowing solutionculture were supplied with NO₃^– at relative addition rates(RAR) of 0·03, 0·06, 0·12, and 0·18d^–1, or constant external concentrations ([NO₃^–)of 3, 10, 20, and 100 mmol m^–3 over 19 d. Following acclimation,relative growth rates (RGR)approached the corresponding RARbetween 0·03–0.12 d^-1, although growth was notlimited by N supply at RAR =0.18 d^-1. Growth rates showed littlechange with [NO₃–] between 10–100 mmol m^–3(RGR=0·15 –0·16 d^-1). The absence of growthlimitation over this range was suggested by high unit absorptionrates of NO₃^–, accumulation of NO₃^– in tissues andprogressive increases in shoot: root ratio. Rates of net uptakeof NO₃^– from 1 mol m^–3 solutions were assessed relativeto the growth-related requirement for NO₃^–, showing thatthe relative uptake capacity increased with RGR between 0·03–0·06d^–1 , but decreased thereafter to a theoretical minimumvalue at RGR     

Compensatory Roles of Nitrogen Uptake and Photosynthetic N-use Efficiency in Determining Plant Growth Response to Elevated CO2: Evaluation Using a Functional Balance Model

We used a modified functional balance (FB) model to predictgrowth response of Helianthus annuus L. to elevated CO₂. Modelpredictions were evaluated against measurements obtained twiceduring the experiment. There was a good agreement between modelpredictions of relative growth rate (RGR) responses to elevatedCO₂and observations, particularly at the second harvest. Themodel was then used to compare the relative effects of biomassallocation to roots, nitrogen (N) uptake and photosyntheticN-use efficiency (PNUE) in determining plant growth responseto elevated CO₂. The model predicted that a rather substantialincrease in biomass allocation to root growth had little effecton whole plant growth response to elevated CO₂, suggesting thatplasticity in root allocation is relatively unimportant in determininggrowth response. Average N uptake rate at elevated comparedto ambient CO₂was decreased by 21–29%. In contrast, elevatedCO₂increased PNUE by approx. 50% due to a corresponding risein the CO₂-saturation factor for carboxylation at elevated CO₂.The model predicted that the decreased N uptake rate at elevatedCO₂lowered RGR modestly, but this effect was counterbalancedby an increase in PNUE resulting in a positive CO₂effect ongrowth. Increased PNUE may also explain why in many experimentselevated CO₂enhances biomass accumulation despite a significantdrop in tissue nitrogen concentration. The formulation of theFB model as presented here successfully predicted plant growthresponses to elevated CO₂. It also proved effective in resolvingwhich plant properties had the greatest leverage on such responses.Copyright 2000 Annals of Botany Company Elevated CO₂, functional balance model, Helianthus annuus L., N uptake, photosynthetic nitrogen use efficiency, root:shoot ratio     

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     

Effect of CO2 Concentration on Growth, Carbon Distribution and Loss of Carbon from the Roots of Maize

The effects of three ranges of CO₂ concentration on growth,carbon distribution and loss of carbon from the roots of maizegrown for 14 d and 28 d with shoots in constant specific activity¹⁴CO₂ are described. Increasing concentrations of CO₂ led toenhancement of plant growth with the relative growth rate (RGR)of the roots affected more than the RGR of the shoots. Between16% and 21% of total net fixed carbon (defined as ¹⁴C retainedin the plant plus ¹⁴C lost from the root) was lost from theroots at all CO₂ concentrations at all times but the amountsof carbon lost per unit weight of plant decreased with time.Possible mechanisms to account for these observations are discussed. Key words: Growth, Roots, Carbon loss, [CO₂]     

Effects and Interactions of Ozone and the Anti-ozonant EDU at Different Stages of Radish {Raphanus sativus L.) Development

The use of the anti-ozonant ethylenediurea (EDU) to assess croplosses due to atmospheric ozone (O₃), is controversial becauseof incomplete knowledge on the optimal dosage, the extent ofprotection provided and potential effects on plant performanceof EDU per se. In a greenhouse experiment, radish (Raphanussativus L.) cv. ‘Cherry Belle’ plants were treatedwith EDU as a soil drench (150 mg1¹, 60 ml plant¹) at threestages of development: early, maximum and late hypocotyl thickening.Two days later the plants were exposed to O₃, mimicking an ambientpattern (70 nl 1¹ O₃, 7 h d¹, 5 d, and up to 120 nl 1¹ O₃ on2 d), or remained in filtered air. Assessment of leaf injury showed complete protection from ozone-inducedleaf damage at all stages of growth due to EDU. EDU did causeslight leaf margin necrosis and hyponastic leaf deformationirrespective of the atmospheric treatment, but biomass and relativegrowth rate (RGR) of the shoot and leaf number duration werenot altered and leaf area duration was increased when comparedto control plants. EDU did not affect dry weight accumulationnor RGR of the hypocotyl. Ozone-induced reductions in growthof below-ground organs (hypocotyl and root), which were mostprominent in the intermediate and early stage of growth, werelargely suppressed by the EDU-treatment. Decreases of hypocotylbiomass due to O₃ were transient and significant increases inRGR of the hypocotyl during the post-exposure period indicateda substantial capability of compensation during the period ofrecovery. Key words: Radish (Raphanus sativus), ozone, anti-ozonant, ethylenediurea (EDU), development, phytotoxicity     

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     

Growth of Actinorhizal Plants as Influenced by the Form of N with Special Reference to Myrica gale and Alnus incana

Growth of two actinorhizal species was studied in relation tothe form of N supply in water culture. Non-nodulated bog myrtle(Myrica gale) and grey alder (Alnus incana) were grown withNH₄⁺, NH₄NO₃^– or NO₃^– (4 mol m^–3 N). A nodulatedseries of bog myrtle was also cultivated in N-free medium. Relative growth rate (RGR), utilization rate of N, and shoot/rootratio were highest for the two species with the N completelysupplied as NH₄⁺. In both species, nitrate was largely reducedin the roots and the presence of NO₃^– in combined-N supplyalways affected the RGR and N utilization rate negatively. BothN₂ fixation and complete NO₃^– nutrition represented conditionsof relative N-deficiency resulting in relatively low tissue-Nconcentrations and a greater allocation of dry mass to the roots.The physiological N status of nodulated M. gale plants was comparativelygood, as indicated by a normal nodule weight ratio and a relativelyhigh N₂-fixing rate per unit nodule mass. However, whole-plantN₂-fixing capacity remained relatively low in comparison withacquisition rates of N in combined-N plants. The anion charge from the nitrate reduction was largely directlyexcreted as an OH^– efflux. H ⁺ /N ratios generally agreedwith the theory. In comparison with NH₄⁺ nutrition, carboxylateconcentrations were higher in N₂-fixing M. gale plants and theH ⁺ /N ratio in nodulated plants was less than unity below thevalue for ammonium plants as previously found for other actinorhizalspecies. Therefore, NH₄⁺ should be an energetically more attractiveN source for actinorhizal plants than N₂. The results agree with commonly accepted views on energeticsof N uptake and assimilation in higher plants and support theconcept of a basically similar physiological behaviour betweennon-legumes and legumes. Key words: Actinorhizal symbioses, ammonium, H⁺/OH^– efflux, nitrate, N₂ fixation, NRA     

Effects of nitrogen nutrition on salt-stressed Nicotiana tabacum var. Samsun in vitro plantlets

Tobacco shoots were grown in vitro for 35 d, in MS culture mediummodified to include various sources (nitrate-N, ammonium-N ora mixture) and levels (0–120 mM) of N, and in the presenceof 0–180 mM NaCI or iso-osmotic concentrations of mannitol.Growth of control plantlets was significantly inhibited whenNH₄⁺-N was the sole N source, and at high (120 mM) NO^–₃-N supply. Under conditions of salt stress (90 and 180 mM NaCI)growth was repressed, with roots being more severely affectedthan shoots. Salinity also inhibited root emergence in vitro.The only alleviation of the salt stress by nitrate nutritionobserved in this study was on shoot growth parameters of plantletsgrown on 60 mM NO^–₃-N and 90 mM NaCI. Although both weresignificantly inhibited by NaCI, nitrate reduc-tase activitywas more severely affected than nitrate uptake. When mannitolreplaced NaCI in the culture medium, similar Inhibition of growth,nutrient uptake and enzyme activity were recorded. These observations,together with the relatively low recorded values for Na⁺ andCI^– uptake, indicate that under in vitro salt stress conditionsthe negative effects of NaCI are primarily osmotic. Key words: Growth, nitrogen metabolism, osmotic stress, salinity     

Nutrient Relations of Groundsel (Senecio vulgaris) Infected by Rust (Puccinia lagenophorae) at a Range of Nutrient Concentrations II. Uptake of N, P and K and Shoot-Root Interactions

Groundsel (Senecio vulgaris), healthy or infected with rust,Puccinia lagenophorae, was grown at a range of nutrient concentrationsin sand culture. Specific absorption rates calculated on thebasis of root dry weight (SAR_W) were greater in rusted thancontrol groundsel for nitrogen, potassium and phosphorus. Whilethe magnitudes of these stimulations varied, they occurred acrossthe whole range of nutrient concentrations. By contrast, specificabsorption rate on the basis of root length (SAR_L) were littlechanged by rust at any external nutrient concentration; SAR_Lfor phosphate and potassium were slightly reduced when nutrientswere freely available. Water flux per unit dry root weight and length was stimulatedby rust because transpiration per unit leaf area was more rapidin infected plants after fungal sporulation. However, water-fluxand the rate of uptake of nutrients were correlated only whenexpressed on the basis of root weight and increased transpirationdid not appear to be the mechanism underlying increased rootactivity. Rather, increased SAR_W for N, P and K could very largelybe attributed to increased shoot demand per unit root, whichresulted from the higher shoot: root (S: R) ratios of infectedindividuals. Changes in S: R accounted for 92, 81 and 57% oftotal variation in SAR_W for K, P and N respectively. Greatervalues for SAR_W were possible because specific root length (SRL)increased, producing more functional root per unit root weight.The lack of stimulation in SAR_L in response to rust could beexplained since the higher SRL of infected plants resulted instable values of shoot weight per unit root length, i.e. shootdemand was not increased by infection on this basis. Senecio vulgaris, Puccinia lagenophorae, rust infection, nutrient uptake, water uptake, shoot: root interactions     

Regulation of K+ Influx in Barley: Evidence for a Direct Control of Influx by K+ Concentration of Root Cells

Siddiqi, M. Y. and Glass, A. D. M. 1987. Regulation of K⁺ influxin barley: Evidence for a direct control of influx by K⁺ concentrationof root cells.—J. exp. Bot. 38: 935–947. The kinetics of K⁺ (⁸⁶Rb⁺) influx into intact roots of barley(Hordeum vulgare L. cv. Fergus) seedlings having different combinationsof root and shoot [K⁺], different growth rates and differentroot:shoot weight ratios were studied. K⁺ influx was stronglycorrelated with root [K⁺]; shoot [K⁺], growth rates, and root:shoot ratios appeared to have little effect on K⁺ influx. Adetailed study showed that both V_max and K_m for K⁺ influx wereaffected by root [K⁺] but not by shoot [K⁺]. We have suggestedthat factors such as growth rates and root: shoot ratio mayaffect K⁺ influx indirectly primarily via their influence onroot factors such as root [K⁺]. We have reiterated that othertypes of kinetic control, e.g. increased or decreased synthesisof ‘carrier systems’, may operate in addition todirect (allosteric?) control of K⁺ influx by root [K⁺]. Thenegative feedback signal from root [K⁺] appeared to be the primeeffector in the regulation of K⁺ influx. Key words: Barley, K⁺ influx     

Concurrent Rates of N2 Fixation, Nitrate and Ammonium Uptake by White Clover in Response to Growth at Different Root Temperatures

Nodulated white clover plants (Trifolium repens L. cv. Huia)were grown for 71 d in flowing nutrient solutions containingN as 10 mmol m_–3 NH₄NO₃, under artificial illumination,with shoots at 20/15°C day/night temperatures and root temperaturereduced decrementally from 20 to 5°C. Root temperatureswere then changed to 3, 7, 9, 11, 13, 17 or 25°C, and theacquisition of N by N₂ fixation, NH₄+ and NO₃^– uptakewas measured over 14 d. Shoot specific growth rates (d. wt)doubled with increasing temperature between 7 and 17°C,whilst root specific growth rates showed little response; shoot:root ratios increased with root temperature, and over time at11°C. Net uptake of total N per plant (N₂ fixation + NH₄++ NO₃^–) over 14 d increased three-fold between 3 and 17°C.The proportion contributed by N₂ fixation decreased with increasingtemperature from 51% at 5°C to 18% at 25°C. Uptake ofNH₄+ as a proportion of NH₄+ + NO₃^– uptake over 14 d variedlittle (55–62%) with root temperature between 3 and 25°C,although it increased with time at most temperatures. Mean ratesof total N uptake per unit shoot f. wt over 14 d changed littlebetween 9 and 25°C, but decreased progressively with temperaturebelow 9°C, due to the decline in the rates of NH₄+ and NO₃^–uptake, even though N₂ fixation increased. The results suggestthat N₂ fixation in the presence of sustained low concentrationsof NH₄+ and NO₄^– is less sensitive to low root temperaturethan are either NH₄+ or NO₃^– uptake systems. White clover, Trifolium repens L. cv. Huia, root temperature, nitrogen fixation, ammonium, nitrate     

Growth and Preferences for Ammonium or Nitrate Uptake by Barley in Relation to Root Termperature

Barley plants (Hordewn vulgare L. cv. Atem) were grown fromseed for 28 d in flowing solution culture, during which timeroot temperature was lowered decrementally to 5?C. Plants werethen subjected to root temperatures of 3, 5, 7, 9, 11, 13, 17or 25 ?C, with common air temperature of 25/15 ?C (day/night).Changes in growth, plant total N, and NO₃^– levels, andnet uptake of NH₄⁺ and NO₃^– from a maintained concentrationof 10 mmol m^–3 NH₄NO₃ were measured over 14 d. Dry matterproduction increased 6-fold with increasing root temperaturebetween 3–25 ?C. The growth response was biphasic followingan increase in root temperature. Phase I, lasting about 5 d,was characterized by high root specific growth rates relativeto those of the shoot, particularly on a fresh weight basis.During Phase I the shoot dry weight specific growth rates wereinversely related to root temperature between 3–13 ?C.Phase 2, from 5–14 d, was characterized by the approachtowards, and/or attainment of, balanced exponential growth betweenshoots and roots. Concentrations of total N in plant dry matterincreased with root temperature between 3–25 ?C, moreso in the shoots than roots and most acutely in the youngestfully expanded leaf (2?l–6?9% N). When N contents wereexpressed on a tissue fresh weight basis the variation withtemperature lessened and the highest concentration in the shootwas at 11 ?C. Uptake of N increased with root temperature, andat all temperatures uptake of NH₄⁺, exceeded that of NO₃^–,irrespective of time. The proportions of total N uptake over14 d absorbed in the form of NH₄⁺ were (%): 86, 91, 75, 77,76, 73, 77, and 80, respectively, at 3, 5, 7, 9, Il, 13, 17,and 25 ?C. At all temperatures the preference for NH₄⁺ overNO₃^– uptake increased with time. An inverse relationshipbetween root temperature (3–11 ?C) and the uptake of NH₄⁺as a proportion of total N uptake was apparent during PhaseI. The possible mechanisms by which root temperature limitsgrowth and influences N uptake are discussed. Key words: Hordeum vulgare, root temperature, ammonium, nitrate, ion uptake, growth rate     

H+/OH- Excretion and Nutrient Uptake in Upper and Lower Parts of Lupin (Lupinus angustifolius L.) Root Systems

The cultivation of narrow-leafed lupins (Lupinus angustifoliusL.) increase rates of subsoil acidification, and this is thoughtto be partly related to their pattern of nutrient uptake andH⁺/OH^- excretion. The main hypothesis of this study was thatH⁺ and OH^- excretion is not distributed evenly over the entirelength of the root system but is limited to zones where excesscation or anion uptake occur. Seedlings of nodulated lupinswere grown in solution culture using vertically split pots thatallowed the upper and lower zones of the root system to be suppliedwith varying concentrations of K⁺ and NO^-₃. Net H⁺/OH^- excretionwas equated to the addition of NaOH/HCl required to maintaina constant pH in the nutrient solution during a 4-d treatmentperiod and nutrient uptake was measured by depletion from solutionin each zone of the split pots. The excess of cation over anion uptake was positively correlatedwith H⁺ excretion in each rooting zone. In zones where K⁺ wassupplied at 1200 µM, cation uptake was dominated by K⁺and up to twice as much H⁺ was excreted than in zones whereK⁺ was absent. In zones where NO^-₃ was supplied at 750 µM,the anion/cation uptake was balanced, however H⁺ excretion continuedto occur in the zone. When NO^-₃ was supplied at 5000 µM,anion uptake exceeded cation uptake but there was no OH^- excretion.Organic acid anions may be excreted by lupins to maintain theirinternal electroneutrality when anion uptake exceeds cationuptake. Rhizosphere pH would not increase unless the pK_a ofthe excreted organic anions was greater than the external pH.Copyright1993, 1999 Academic Press Lupinus angustifolius L., H⁺/OH^- excretion, nutrient uptake, cation-anion balance, vertical split root     

Effects of elevated carbon dioxide on three montane grass species: I. Growth and dry matter partitioning

Upland grasslands are a major component of natural vegetationwithin the UK. Such grasslands support slow growing relativelystable plant communities. The response of native montane grassspecies to elevated atmospheric carbon dioxide concentrationshas received little attention to date. Of such studies, mosthave only focused on short-term (days to weeks) responses, oftenunder favourable controlled environment conditions. In thisstudy Agrostis caplllaris L.⁵, Festuca vivipara L. and Poa alpinaL. were grown under semi-natural conditions in outdoor open-topchambers at either ambient (340µmol mol^–1) or elevated(680µmol mol^–1) concentrations of atmospheric carbondioxide (CO₂ for periods from 79 to 189 d, with a nutrient availabilitysimilar to that of montane Agrostis-Fescue grassland in Snowdonia,N. Wales. Whole plant dry weight was increased for A. capillarisand P. alpina, but decreased for F. vivipara, at elevated CO₂.Major components of relative growth rate (RGR) contributingto this change at elevated CO₂ were transient changes in specificleaf area (SLA) and leaf area ratio (LAR). Despite changes ingrowth rate at 680 µmol mol^–1 CO₂, partitioningof dry weight between shoot and root in plants of A. capillarisand P. alpina was unaltered. There was a significant decreasein shoot relative to root growth at elevated CO₂ in F. viviparawhich also showed marked discoloration of the leaves and increasedsenescence of the foliage. Key words: Allometry, growth analysis, elevated CO₂, grasses     

Sugar Accumulation in Barley and Rice Grown in Solutions with low Concentrations of Oxygen

Barley and rice, at the early tillering stage, were grown inaerated nutrient solutions (> 7 mg O₂ l^–1) and transferredto solutions of low O₂ concentrations (< 0.5 mg l ^–1). For barley, low O₂ concentrations during the first 5 days severelyinhibited growth of seminal roots had less effect on nodal roots,and did not reduce shoot growth. Longer exposure to low O₂ concentrationsreduced shoot as well as root growth. Sugar concentrations inroots and shoots increased within 7 h after transfer of plantsto low O₂ concentrations. After 5 days at low O₂ concentrationssugar concentrations were very high in fast growing nodal rootsand in shoots, as well as in the slower growing seminal roots. In rice, low O₂ concentrations increased sugar levels of rootsduring summer, but not during winter. In summer, the highersugar levels at low O₂ concentrations persisted throughout adiurnal cycle. In root apices, sugar concentrations were increasedby low O₂ concentrations, even though the experiment was donein winter and the bulk of the root system showed no differencein sugar levels. The data indicate that sugar accumulation, at low O₂ concentrations,is caused by reduced growth and also that even apices of rootsgrown at low O₂ concentrations have sufficient substrates forrespiration. Hordeum vulgare L, barley, Oryza sativa L, rice, sugar accumulation, oxygen concentration     

The Effect of Root Temperature on Growth and Uptake of Ammonium and Nitrate by Brassica napus L. cv. Bien venu in Flowing Solution Culture: II. UPTAKE FROM SOLUTIONS CONTAINING NH4NO3

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. CV. Bien venu in flowing solution culture.II. Uptake from solutions containing NH₄NO₃.—J. exp. Bot.38: 53–66 The effects of root temperature on uptake and assimilation ofNH₄⁺ and NO₃^– by oilseed rape (Brassica napus L. CV. Bienvenu) were examined. Plants were grown for 49 d in flowing nutrientsolution at pH 6?0 with root temperature decrementally reducedfrom 20?C to 5?C; and then exposed to different root temperatures(3, 5, 7, 9, 11, 13, 17 or 25?C) held constant for 14 d. Theair temperature was 20/15?C day/night and nitrogen was suppliedautomatically to maintain 10 mmol m^–3 NH₄NO₃ in solution.Total uptake of nitrogen over 14 d increased threefold between3–13?C but was constant above 13?C. Net uptake of NH₄⁺exceeded that of NO₃^– at all temperatures except 17?C,and represented 47–65% of the total uptake of nitrogen.Unit absorption rates of NH₄⁺ and of 1?5–2?7 for NO₃^–suggested that NO₃^– absorption was more sensitive thanNH₄⁺ absorption to temperature. Rates of absorption were relativelystable at 3?C and 5?C compared with those at 17?C and 25?C whichincreased sharply after 10 d. Tissue concentration of N in theshoot, expressed on a fresh weight basis, was independent ofroot temperature throughout, but doubled between 3–25?Cwhen expressed on a dry weight basis. The apparent proportionof net uptake of NO₃^– that was assimilated was inverselyrelated to root temperature. The results are used to examinethe relation between unit absorption rate adn shoot:root ratioin the context of short and long term responses to change ofroot temperature Key words: Brassica napus, oilseed rape, root temperature, nitrogen uptake     

Root Distribution, Growth, Respiration, and Hydraulic Conductivity for Two Highly Productive Agaves

Cultivated Agave mapisaga and A. salmiana can have an extremelyhigh above-ground dry-weight productivity of 40 Mg ha^–1yr^–1. To help understand the below-ground capabilitiesthat support the high above-ground productivity of these Crassulaceanacid metabolism plants, roots were studied in the laboratoryand in plantations near Mexico City. For approximately 15-year-oldplants, the lateral spread of roots from the plant base averaged1.3 m and the maximal root depth was 0.8 m, both considerablygreater than for desert succulents of the same age. Root andshoot growth occurred all year, although the increase in shootgrowth at the beginning of the wet season preceded the increasein growth of main roots. New lateral roots branching from themain roots were more common at the beginning of the wet season,which favoured water uptake with a minimal biomass investment,whereas growth of new main roots occurred later in the growingseason. The root: shoot dry weight ratio was extremely low,less than 0.07 for 6-year-old plants of both species, and decreasedwith plant age. The elongation rates of main roots and lateralroots were 10 to 17 mm d^–1, higher than for various desertsucculents but similar to elongation rates for roots of highlyproductive C₃ and C₄ agronomic species. The respiration rateof attached main roots was 32 µmol CO₂ evolved kg^–1dry weight s^–1 at 4 weeks of age, that of lateral rootswas about 70% higher, and both rates decreased with root age.Such respiration rates are 4- to 5-fold higher than for Agavedeserti, but similar to rates for C₃ and C₄ agronomic species.The root hydraulic conductivity had a maximal value of 3 x 10^–7ms^–1 MPa^–1 at 4 weeks of age, similar to A. deserti.The radial hydraulic conductivity from the root surface to thexylem decreased and the axial conductivity along the xylem increasedwith root age, again similar to A. deserti. Thus, although rootsof A. mapisaga and A. salmiana had hydraulic properties perunit length similar to those of a desert agave, their highergrowth rates, their higher respiration rates, and the greatersoil volume explored by their roots than for various desertsucculents apparently helped support their high above-groundbiomass productivity Key words: Crassulacean acid metabolism, productivity, root elongation rate, root system, water uptake     

Respiration and Phloem Translocation in the Roots of Chickpea (Cicer arietinum)

Root growth in chickpea (Cicer arietinum) has been studied fromthe early vegetative phase to the reproductive stage in orderto elucidate its growth and maintenance respiration and to quantifythe translocation of assimilates from shoot to root. A carbonbalance has been drawn for this purpose using the growth andrespiration data. The increase in the sieve tube cross-sectionalarea was also followed simultaneously. Plants growing in a nutrient culture medium were studied todetermine the relative growth rate (RGR) 5–60 d aftergermination. RGR declined from 113 to 41 mg d^–1 g^–1during the measurement period. Simultaneous with the RGR analysis,respiration rate was also measured using an oxygen electrode.The respiration rate declined as the plants aged and a drasticreduction was recorded following anthesis. The relationshipbetween RGR and respiration rate was used to extrapolate themaintenance respiration (m) and growth respiration (1/Y_EG).The respiration quotient (r.q.) of the roots was 1.2 and theQ₁₀ in the range 20–25 °C was 2·2. A carbon balance for the roots was constructed by subtractingthe carbon lost during respiration from that gained during growth.The roots were found to respire no less than 80% of the carbontranslocated. The increase in the cross-sectional area composed of sieve tubeswas measured near the root-shoot junction as the plants grew.Chickpea has storied sieve plates which simplifies these measurements.Their cross-sectional area increased during growth mainly becauseof an increase in sieve tube number. The diameter of individualsieve tubes remained constant. Specific mass transfer (SMT) values for seive tubes into theroots have been computed during various stages of growth. SMTvalues were relatively constant before anthesis (approx. 6·5g h^–1 cm^–2), but decreased following anthesis. Wedid not evaluate possible retranslocation from roots: any suchretranslocation would have the effect of increasing our SMTvalues. Chickpea, Cicer arietinum, legume, root, respiration, phloem, translocation, carbon balance, specific mass transfer, sieve-tube dimensions