Long Distance Transport of Abscisic Acid in NaCI-Treated Intact Plants of Lupinus albus

Plants of Lupinus albus were grown for 51 d under control (1.1mol m^–3 NaCl) and saline (40 mol m^–3 NaCl) conditions.Plants were harvested and changes of carbon, nitrogen and abscisicacid (ABA) contents of individual organs were determined 41d and 51 d after germination. In the period between the twoharvests xylem and phloem saps were collected and respirationand photosynthesis of individual organs were measured. Usingflows of carbon, C/ABA ratios and increments of ABA flows ofABA in phloem and xylem and rates of biosynthesis and degradationof ABA were calculated. Both under control and saline conditionsnet biosynthesis occurred in the root, the basal strata of leavesand in the inflorescence. Metabolic degradation of ABA tookplace in the stem internodes and apical leaf strata. Salt stress increased xylem transport of ABA up to 10-fold andphloem transport to the root up to 5-fold relative to that ofthe controls. A considerable amount of ABA in the xylem saporiginated from biosynthesis in the roots, i.e. 55% in salt-treatedand smaller than 28% in control plants. The remaining part ofABA in the xylem sap originated from the shoot: it was translocatedin the phloem from fully differentiated leaves towards the rootand from there it was recirculated back to the aerial partsof the plant. The data suggest that ABA may serve as a hormonalstress signal from the root system. Key words: Lupinus albus, salt stress, abscisic acid, long distance transport     

The uptake and flow of C, N and ions between roots and shoots in Ricinus communis L. III. Long-distance transport of abscisic acid depending on nitrogen nutrition and salt stress

Seedlings of Ricinus communis L. were cultivated in quartz sandand supplied with media which contained either different concentrationsof nitrate or ammonium nitrogen and were treated with a lowsalt stress. The concentration of ABA was determined in tissuesand in xylem and phloem saps. Between 41 and 51 day after sowing,abscisic acid (ABA) flows between roots and shoots were modelled.Long-distance transport of ABA was not stimulated under conditionsof nitrate deficiency (0.2 mol m^–3). However, when ammoniumwas given as the only N source (1.0 mol m^–3), ABA transportin both xylem and phloem was increased significantly. Mild saltstress (40 mol m^–3 NaCl) increased ABA transport in nitrate-fedplants, but not in ammonium-fed plants. The leaf conductancewas lowered by salt treatment with both nitrogen sources, butit was always lower in ammonium-fed compared to nitrate-fedplants. A negative correlation of leaf conductance to ABA levelsin leaves or flow in xylem was found only in comparison of ammonium-fedto nitrate-fed plants. Key words: Abscisic acid, ammonium, Ricinus communis, phloem, xylem, transport, nitrate, nitrogen nutrition     

The Role of the Mesocotyl in Sodium Exclusion from the Shoot of Zea mays L. (cv. Pioneer 3906)

Drew, M. C. and Lauchli, A. 1986. The role of the mesocotylin sodium exclusion from the shoot of Zea mays L. (cv. Pioneer3906).—J. exp. Bot. 38: 409–418. The mesocotyl, located between the root and shoot, can stronglyaccumulate Na⁺ from the ascending transpiration stream, therebypotentially acting as a sink to protect the shoot from excessNa⁺. To determine the quantitative importance of the mesocotylas a Na⁺ sink, we grew plants with either short (9·0mm) or long(21 mm) mesocotyls, the latter resembling the sizefound in field-grown plants. At 13 d, plants were transferredfrom Na ⁺ -free nutrient solution to a ²²Na⁺ labelled solutionin which the concentration of NaCl was (mol m^–3) 1·0,10 or 100. The concentration of Na⁺ accumulated in the mesocotylin 24 h (g^–1 fr. wt.) exceeded that in the roots thatwere directly exposed to the nutrient solution. The amountsof ²²Na⁺ retained in the long mesocotyl were about double thatin the short ones and increased with time of exposure and NaClconcentration. At 1·0 and 10 mol m^–3 NaCl, theamounts of ²²Na⁺ retained in the mesocotyl were 6–19%of those reaching the shoot in 24 h, but with 100 mol m^–3NaCl, a damaging concentration for maize, this declined to 3–8%.The mesocotyl, even as a fully elongated structure is, therefore,unlikely to provide an appreciable alternative sink for Na⁺when NaCl reaches injurious concentrations. Key words: Ion transport, potassium, roots, salinity     

The Effects of Partially Drying Part of the Root System of Helianthus annuus on the Abscisic Acid Content of the Roots, Xylem Sap and Leaves

Plants of Helianthus annuus were grown in soil in pots suchthat approximately 30% of the root system protruded throughthe base of the pot. After 7 d further growth in aerated nutrientsolution, the attached, protruding roots were air-dried for10–15 min and thereafter surrounded with moist still air,in the dark, for 49 h, whilst the soil was kept at field capacity.The roots of the control plants remained in the nutrient solutionthroughout the experiment. This treatment rapidly reduced the water content of protrudingroots from 20.5 to 17.8 g g^–1 dry mass (DM), which remainedless than that of the control roots for the rest of the experiment.This treatment also reduced root turgor and water potential.The abscisic acid (ABA) concentrations in the protruding roots,xylem sap and leaves of the treated plants increased significantly,compared to values recorded for control plants. In treated roots, the ABA concentration was significantly increased4 h after treatment, with a maximum of 4.4+0.1 nmol g^–1(DM) after 25 h. The ABA concentration in the xylem sap of thetreated plants was significantly greater than in the controls25 h, 30 h, and 49 h after the partial drying of the roots,with a maximum concentration of approximately 970 pmol ABA cm-3at 49 h. Initially, the ABA concentration in the leaves was0.45 nmol g^–1 (DM) which increased significantly to 1.1±0.1 nmol g^–1 at 25 h, to 1.7±0.3 nmol g^–1at 49 h. Leaf conductance was significantly less in plants with air-driedroots than in the controls 8 h after the start of the treatmentand thereafter. The water relations of the leaves of the treatedplants did not differ from those of the control plants. These results confirm previous reports that ABA is rapidly generatedin partially-dried and attached root systems and demonstratesa concomitant large increase in the ABA content of the xylemsap. It is suggested that partial dehydration of some of theroots of Helianthus annuus, increases ABA concentration in thetranspiration stream and decreases leaf conductance in the absenceof changes in leaf water status. As these responses were initiatedin free-growing roots the stimulus is independent of any increasesin soil shear strength that are associated with soil drying. Key words: Soil drying, roots, ABA, leaf conductance, water relations     

Delivery rates of abscisic acid in xylem sap of Ricinus communis L. plants subjected to part-drying of the soil

Abscisic acid (ABA) moving from roots to shoots in the transpirationstream is a potential hormonal message integrating perceptionof a root stress with adaptive changes in the shoot. A twinroot system was used to study ways of estimating effects ofdroughting the upper roots of Ricinus communis L. on abscisicacid (ABA) transport to the shoot in the transpiration stream.Droughted plants transpired more slowly than controls. Droughtingalso increased concentrations of ABA up to I I-fold in sap inducedto flow from the roots of freshly decapitated plants at ratesof whole plant transpiration. However, because of dilution effectsarising from the different sap flows in control and droughtedplants, these changes in ABA concentration in the xylem sapdid not accurately reflect amounts of ABA transported. To overcomethis problem, delivery rates were calculated by multiplyingconcentration with sap flow rate to generate ABA delivery interms of µmol s–1 per plant. Droughting for 24 hor more increased ABA delivery from roots to shoots by 5-fold.Since droughting can alter the relative sizes of the roots andshoots and also the root:shoot ratio these delivery rates wererefined in several ways to reflect both the amount of root generatingthe ABA message and the size of the recipient shoot system. Key words: Abscisic acid, Ricinus communis L., soil drying, xylem sap     

Growth, Nitrogen Uptake and Flow in Maize Plants Affected by Root Growth Restriction

The objective of the present study was to investigate the influence of a reduced maize root-system size on root growth and nitrogen (N) uptake and flow within plants. Restriction of shoot-borne root growth caused a strong decrease in the absorption of root: shoot dry weight ratio and a reduction in shoot growth. On the other hand, compensatory growth and an increased N uptake rate in the remaining roots were observed. Despite the limited long-distance transport pathway in the mesocotyl with restriction of shoot-borne root growth, N cycling within these plants was higher than those in control plants, implying that xylem and phloem flow velocities via the mesocotyl were considerably higher than in plants with an intact root system. The removal of the seminal roots in addition to restricting shoot-borne root development did not affect whole plant growth and N uptake, except for the stronger compensatory growth of the primary roots. Our results suggest that an adequate N supply to maize plant is maintained by compensatory growth of the remaining roots, increased N uptake rate and flow velocities within the xylem and phloem via the mesocotyl, and reduction in the shoot growth rate.     

Concentrations of abscisic acid and other solutes in xylem sap from root systems of tomato and castor-oil plants are distorted by wounding and variable sap flow rates

We investigated if concentrations of abscisic acid (ABA) andother solutes measured in the first few droplets of xylem sapfrom detopped root systems, are good estimates of those in thetranspiration stream as it enters the shoot-base of whole plants.Xylem sap from root systems of pot-grown tomato plants (Lycopersiconesculentum Mill., cv. Ailsa Craig), at the seven-leaf stage,was obtained by placing root systems in chambers pressurizedto 0.3 MPa with air. The first sample was taken from the cut-surfaceof the hypo-cotyl stump within 30 s of removing the shoot. ABA,sucrose and other osmolytes were more concentrated in the initial100–200 mm³ of xylem sap than in subsequent samples. Thissuggested the sap was contaminated and not unchanged transpirationfluid. The effect was reproduced on the same plant, severaltimes, by recutting the hypocotyl prior to reassembling thesap collecting set-up and repressurizing. Similar results werefound with castor-oil plants (Ricinus communis L., cv. Gibsonii).However, neither release of ABA from the cut surface of thehypocotyl stump, nor the effects of pressure to the roots causedthe contamination. Instead, small radial pressures exerted bya rubber sleeve attached to the hypocotyl stump, for collectingthe sap, were responsible. The effect was reproduced by lightlysqueezing the hypocotyl by hand. The possibility was examined that reliable estimates of ABAconcentrations in transpiration stream fluid may be obtainedfrom sap samples taken immediately after rejecting the initial,contaminated 200 mm³. However, ABA concentrations in these latersamples were also unsatisfactory since they changed with rateof sap flow. The problem may be overcome by analysing sap inducedto flow through detached root systems at rates close to thoseof whole-plant transpiration. Key words: Tomato, Lycopersicon esculentum Mill., Castor-oil plant, Ricinus communis L., roots, root to shoot communication, xylem sap, abscisic acid, sucrose, transpiration stream     

Hormonal responses to partial drying of the root system of Helianthus annuus

Plants of Helianthus annuus were pot-grown in soil, with approximately30% of the root system protruding through the base. After 7d, the upper part of the root system of half of the plants wasexposed to drought (internal roots) while the lower part waskept in aerated nutrient solution (protruding root). The treatmentrapidly reduced the internal roots' water content from 26.1to 21.9 g g^–1 dry weight (DW), while in protruding rootsof stressed plants it slowly and continuously decreased from31.9 to 25.2 g g^–1 DW. Leaf water content rapidly decreasedin treated plants from 7.4 to 6.4 g g^–1 DW in the first2d and then reached a plateau. In stressed plants leaf stomatalresistance was significantly higher in the first 3 d while leafwater potential was lower only on the last day. Abscisic acid (ABA) concentration in treated plants increasedsignificantly compared to the controls. In treated internalroots, ABA rose from the first day, reaching a maximum of 1.48±0.49nmol g^–1 DW after 3 d. In treated protruding roots a maximumof 0.99±0.09 nmol g^–1 DW was reached after 1 d.ABA concentration in the xylem sap increased 2 d and 3 d afterthe start of soil drying, with a maximum of 113±12nmoll^–1 during the third day. The ABA rise in the leaves oftreated plants was less significant. Indol-3yl-acetic acid (IAA) concentration in internal rootsof treated plants reached a maximum of 22.54±3.34 nmolg^–1 DW on the third day, then decreased dramatically.The protruding root system of control plants showed a maximumvalue of 16.05±1.77 nmol g^–1 DW on the sixth day. Little difference in cytokinin content of xylem sap was notedbetween control and treated plants. Hormonal variations in different parts of the plant are discussedin relation to drought stress. Key words: Soil drying, roots, ABA, IAA, cytokinins     

Release of Guttation Fluid from Passive Hydathodes of Intact Barley Plants. II. The Effects of Abscisic Acid and Cytokinins

Guttation was used as a non-destructive way to study the flowof water and mineral ions from the roots and compared with parallelmeasurements of root exudation. Guttation of the leaves of barley seedlings depends on age andon the culture solution. Best rates of guttation were obtainedwith the primary leaves of 6- to 7-day-old seedlings grown onfull mineral nutrient solution. The growing leaf tissue becomessaturated with K⁺ below 1.5 mM K⁺ in the medium, whereas K⁺concentration in the guttated fluid still increases furtheras K⁺ concentration in the medium is raised. At 3 mM K⁺ averagevalues of guttation were 1.4–2.4 mm³ h^–1 per plantwith a K⁺ concentration of 10–20 mM; for exuding plantsthe flow was 4.2–7.6 mm³ h^–1 per plant and K⁺ concentration35–55 mM. Abscisic acid (ABA) at 10^–6 to 10^–4 M 0–2h after addition to the root medium increased volume flow ofguttation and exudation and the amount of K⁺ exported. Threeh after addition of ABA both volume and amount of K⁺ were reduced.There was an ABA-dependent increase in water permeability (Lp)of exuding roots shortly after ABA addition. Later Lp was decreasedby 35 per cent and salt export by 60 per cent suggesting aneffect of ABA on salt transport to the xylem apart from itseffect on Lp. Benzyladenine (5 x 10^–8 to 10^–5 M)and kinetin (5 x 10^–6 M) progressively reduced volumeflow and K⁺ export in guttation and exudation and reduced Lp. Guttation showed a qualitatively similar response to phytohormonesas found here and elsewhere using exuding roots. Hordeum vulgare L., barley, guttation, abscisic acid, cytokinins, benzyl adenine, kinetin     

Sodium Exclusion from the Shoots by Roots of Zea mays (cv. LG 11) and its Breakdown with Oxygen Deficiency

The effects of sodium chloride salinity and root oxygen deficiency(anoxia) were studied in 11-12d old maize plants (Zea mays L.cv. LG 11) in nutrient solution culture. Transport of ²²Na bythe roots to the shoot in 24 h was markedly increased by anoxiawhen the external concentration of NaCl was in the range 0·1-10·9mol m^–3. Anoxia severely inhibited uptake of ⁴²K by rootsand its transport to the shoot, so that the ratio of Na⁺/K⁺moving into the shoot was increased by a factor of approximately10. When the external concentration of NaCl was increased to2.4 mol m^–3, the roots showed much less ability to excludeNa⁺ under aerobic conditions, and anoxia caused no further increasein the movement of Na⁺ to the shoot. It is concluded that atthe higher concentration the ability of the roots to excludeNa⁺, presumably through an active mechanism in the xylem parenchymacells or in the root cortex and transporting Na⁺ to the outersolution, is saturated by excessive inward diffusion of Na⁺.The ratio of Na⁺/K⁺ transported to the shoot increased by afactor of 600 when the concentration of NaCl was increased from2·4 mol m^–3 to 40 mol m^–3 and roots weremade anoxic. Such imbalances in the supply of cations to theshoot, particularly when roots are oxygen-deficient, may contributeto salinity damage. Key words: Anaerobic, Anoxic, Oxygen deficiency, Roots, Salinity, Salt stress, Sodium chloride, Zea mays     

Does Xylem Sap ABA Control the Stomatal Behaviour of Water-Stressed Sycamore (Acer pseudoplatanus L.) Seedlings?

Sycamore seedlings were grown with their root systems dividedequally between two containers. Water was withheld from onecontainer while the other container was kept well-watered. Effectsof soil drying on stomatal behaviour, shoot water status, andabscisic acid (ABA) concentration in roots, xylem sap and leaveswere evaluated. At 3 d, root ABA in the drying container increased significantly,while the root ABA in the unstressed container of the same plantsdid not differ from that of the control. The increase in rootABA was associated with the increase in xylem sap ABA and withthe decrease in stomatal conductance without any significantperturbation in shoot water status. At 7 d, despite the continuous increase in root ABA concentration,xylem sap ABA showed a marked decline when soil water contentwas depleted below 013 g g^–1. This reduction in xylemsap ABA coincided with a partial recovery of stomatal conductance.The results indicate that xylem sap ABA is a function of rootABA as well as the flow rate of water from roots to shoots,and that this ABA can be a sensitive indicator to the shootof the effect of soil drying. Key words: Acer pseudoplatanus L., soil drying, stomatal behaviour, xylem sap ABA     

The Uptake and Flow of C, N and lons between Roots and Shoots in Ricinus communis L.

Seedlings of Ricinus communis L. cultivated in quartz sand weresupplied with a nutrient solution containing either 1 mol m^–3NO^–₃ or 1 mol m^–3 NH⁺₄ as the nitrogen source. Duringthe period between 41 and 51 d after sowing, the flows of N,C and inorganic ions between root and shoot were modelled andexpressed on a fresh weight basis. Plant growth was clearlyinhibited in the presence of NH⁺₄. In the xylem sap the majornitrogenous solutes were nitrate (74%) or glutamine (78%) innitrate or ammonium-fed plants, respectively. The pattern ofamino acids was not markedly influenced by nitrogen nutrition;glutamine was the dominant compound in both cases. NH⁺₄ wasnot transported in significant amounts in both treatments. Inthe phloem, nitrogen was transported almost exclusively in organicform, glutamine being the dominant nitrogenous solute, but theN-source affected the amino acids transported. Uptake of nitrogenand carbon per unit fresh weight was only slightly decreasedby ammonium. The partitioning of nitrogen was independent ofthe form of N-nutrition, although the flow of nitrogen and carbonin the phloem was enhanced in ammonium-fed plants. Cation uptakerates were halved in the presence of ammonium and lower quantitiesof K⁺, Na⁺ and Ca²⁺ but not of Mg²⁺ were transported to theshoot. As NH⁺₄ was balanced by a 30-fold increase in chloride in thesolution, chloride uptake was increased 6-fold under ammoniumnutrition. We concluded that ammonium was predominantly assimilated inthe root. Nitrate reduction and assimilation occurred in bothshoot and root. The assimilation of ammonium in roots of ammonium-fedplants was associated with a higher respiration rate. Key words: Ricinus communis, nitrogen nutrition (nitrate/ammonium), phloem, xylem, transport, partitioning, nitrogen, carbon, potassium, sodium, magnesium, calcium, chloride     

Ion Transport to the Xylem in Aerenchymatous Roots of Zea mays L.

Aerenchyma was induced in the developing nodal (adventitious)roots of maize (Zea mays L. cv. LG11) in solution culture bytreatment with an atmosphere of 5% (v/v) oxygen (oxygen partialpressure 5.0 k Pa). Ion uptake by aerenchymatous and ordinary(non-aerenchymatous) roots was then compared to determine theeffect of degeneration of much of the cortex in aerenchymatousroots on radial ion movement to the xylem. Rates of uptake andtransport of phosphate, potassium and chloride were followedby labelling 4–12mm length segments of intact roots usingradioactive tracers. In some experiments, enhanced rates ofphosphate transport were induced by means of a divided rootsystem and nutrient deprivation. For all three nutrient ions,and over a range of concentration for phosphate (10–100mmol m^–3 and potassium (0·25–5·0 molm^–3 and with enhanced rates of transport, aerenchymatousroots were at least as effective as ordinary roots (per mm³of root) and sometimes more effective (per mm root length).These findings are discussed in relation to hypotheses concerningthe pathways for radial ion movement across the cortex in ordinaryand aerenchymatous roots. Correspondence to: Long Ashton Research Station, Long Ashton,Bristol BS18 9AF, U.K. Key words: Aerenchyma, ion transport, mineral nutrition, oxygen deficiency, radio-isotopic tracers, roots, structure, Zea mays     

The Vascular Pattern of Italian Ryegrass (Lolium multiflorum Lam.) 2. Development of the Seedling Axis to Maturity

The vascular system present in a grass seedling axis persistsin a functional state at the base of a maturing plant, but undergoesa number of modifications. Two strands of phloem, accompanied by some internal xylem, differentiatein association with the bicollateral mesocotyl trace at rightangles to the existing phloem, resulting in a tetrarch bundle.Lateral seminal roots are themselves tetrarch and the vascularinsertion of a seminal root on to the mesocotyl is a distinctivethree-dimensional feature. At the base of the mesocotyl thetetrarch bundle merges with the tetrarch bundle of the primaryseminal root via a transition zone. The four phloem poles uniteand then diverge again; the central xylem strand splits intothree and then reunites, the two tissues being intimately interlockedby this rearrangement. The additional vascular tissue of the mesocotyl extends up intothe coleoptilar node and becomes involved in the vascular attachmentof nodal roots at this point. Additional vascular tissue continuesto differentiate in the periphery of the maturing stem and ishere termed the ‘peripheral plexus’. In the seedling, the xylem of the ‘bridge’ linkingthe mesocotyl trace with the scutellar trace is associated withxylem transfer cells and also contains tracheids with distinctive,thin-barred scalariform thickening. These transfer cells disappearas the plant matures but numerous tracheids with thin-barredscalariform thickening are then to be found. The possible significanceof transfer cells in the coleoptilar node is discussed.     

Cation and Chloride Partitioning through Xylem and Phloem within the Whole Plant of Ricinus communis L. under Conditions of Salt Stress

Uptake and partitioning through the xylem and phloem of K⁺,Na⁺, Mg²⁺ , Ca²⁺ and Cl^– were studied over a 9 d intervalduring late vegetative growth of castor bean (Ricinus communisL.) plants exposed to a mean salinity stress of 128 mol m^–3NaCl. Empirically based models of flow and utilization of eachion within the whole plant were constructed using informationon ion increments of plant parts, molar ratios of ions to carbonin phloem sap sampled from petioles and stem internodes andpreviously derived information on carbon flow between plantsparts in xylem and phloem in identical plant material. Salientfeatures of the plant budget for K⁺ were prominent depositionin leaves, high mobility of K⁺ in phloem, high rates of cyclingthrough leaves and downward translocation of K⁺ providing theroot with a large excess of K⁺ . Corresponding data for Na⁺showed marked retention in the root, lateral uptake from xylemby hypocotyl, stem internodes and petioles leading to low intakeby young leaf laminae and substantial cycling from older leavesback to the root. The partitioning of the anionic componentof NaCl salinity, Cl^–, contrasted to that of Na⁺ in thatit was not substantially retained in the root, but depositedmore or less uniformly in stem, petiole and leaf lamina tissues.The flow pattern for Mg²⁺ showed relatively even depositionthrough the plant but some preferential uptake by young leaves,generally lesser export than import by leaf laminae, and a returnflow of Mg²⁺ from shoot to root considerably less than the recordedincrement of the root. Ca²⁺ partitioning contrasted with thatof the other ions in showing extremely poor phloem mobility,leading to progressive preferential accumulation in leaf laminaeand negligible cycling of the element through leaves or root.Features of the response of Ricinus to salinity shown in thepresent study were discussed with data from similar modellingstudies on white lupin (Lupinus albus L.) and barley (Hordeumvulgare L.) Key words: Ricinus communis L, potassium, sodium, chloride, calcium, magnesium, phloem, xylem, transport, partitioning, salinity     

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     

Regulation of Sulphate Transport in a Tropical Legume, Macroptilium atropurpureum, cv. Siratro

When young plants of Macroptilium atropurpureum, cv. Siratrowere deprived of external sulphate (-S plants) growth of shootsand roots continued at rates comparable to those in plants wellsupplied with sulphate (control) for 3 d and 5 d respectively.Dilution of internal sulphur therefore took place and redistributionof sulphur occurred between inorganic and organic forms andbetween roots and younger leaves. Even when S-deficiency limitedgrowth, plants contained 16% of their total sulphur as sulphate,but most of this was retained in old leaves and redistributedslowly to growing zones. The capacity for sulphate uptake increased in roots of –Splants very soon after they were deprived of external sulphate;within 24 h the absorption from 0.25 mol m^–3 SO₄^2–was more than five times that of control roots. Maximum increasedcapacity was reached after 2–3 d stress when the V_maxof system 1 was 1948 nmol h^–1g^–1root fr. wt. in–S plants and 337 nmol h^–1g^–1root fr. wt.in controls. The K^mfor system 1 did not change significantlywith S-stress being between 5–8 µM in both setsof plants. Absorption of L-cysteine was not stimulated by S-stress. There was a close, positive relationship between plant growthrate and the rate at which sulphate uptake capacity was enhancedby withholding sulphate from culture solutions. When –S plants were replaced in sulphate-containing solutiontheir capacity for SO₄^2– declined to the control levelwithin 24 h. Very marked repression of capacity was also foundwhen –S plants were treated with L-cysteine, but therewas no immediate effect with methionine. Roots of this species appear to have a very active system fordegrading L-cysteine to sulphate, 30% of the label in ³⁵S-cysteineabsorbed by roots was recovered in ³⁵SO₄^2– after 20 minor 2 h incubation. By contrast, roots had a very weak abilityto reduce sulphate. When part of the root system was in solution lacking sulphatethere was enhanced uptake of sulphate by other parts which themselveswere amply supplied with sulphate. This is seen as an exampleof compensatory absorption. The response to S-stress is specific and there were no positiveinteractions between S-stress and the absorption of phosphate,or P-stress and the uptake of sulphate. The results are discussed in relation to the close control ofsulphate uptake by internal sulphate concentration, redistributionof forms of sulphur during stress and mobility of sulphate inthe phloem. Key words: Kinetics, Amino-S, Sulpholipid, Repression;, Deficiency     

Modelling of the Partitioning, Assimilation and Storage of Nitrate within Root and Shoot Organs of Castor Bean (Ricinus communis L.)

An experimentally-based modelling technique was developed todescribe quantitatively the uptake, flow, storage and utilizationof NO₃-N over a 9 d period in mid-vegetative growth of sandcultured castor bean (Ricinus communis L.) fed 12 mol m^–3nitrate and exposed to a mean salinity stress of 128 mol m^–3NaCl. Model construction used information on increments or lossesof NO₃-N or total reduced N in plant parts over the study periodand concentration data for NO₃-N and reduced (amino acid) Nin phloem sap and pressure-induced xylem exudates obtained fromstem, petiole and leaf lamina tissue at various levels up ashoot. The resulting models indicated that the bulk (87%) of incomingnitrate was reduced, 51% of this in the root, the remainderprincipally in the laminae of leaves. The shoot was 60% autotrophicfor N through its own nitrate assimilation, but was oversuppliedwith surplus reduced N generated by the root and fed to theshoot through the xylem. The equivalent of over half (53%) ofthis N returned to the root as phloem translocate and, mostly,then cycled back to the shoot via xylem. Nitrate comprised almosthalf of the N of most xylem samples, but less than 1% of phloemsap N. Laminae of leaves of different age varied greatly inN balance. The fully grown lower three leaves generated a surplusof reduced N by nitrate assimilation and this, accompanied byreduced N cycling by xylem to phloem exchange, was exportedfrom the leaf. Leaf 4 was gauged to be just self-sufficientin terms of nitrate reduction, while also cycling reduced N.The three upper leaves (5–7) met their N balance to varyingextents by xylem import, phloem import (leaves 6 and 7 only)and assimilation of nitrate. Petioles and stem tissue generallyshowed low reductase activities, but obtained most of theirN by abstraction from xylem and phloem streams. The models predictedthat nodal tissue of lower parts of the stem abstracted reducedN from the departing leaf traces and transferred this, but notnitrate, to xylem streams passing further up the shoot. As aresult, xylem sap was predicted to become more concentratedin N as it passed up the shoot, and to decrease the ratio ofNO₃-N to reduced N from 0·45 to 0·21 from thebase to the top of the shoot. These changes were reflected inthe measured N values for pressure-induced xylem exudates fromdifferent sites on the shoot. Transfer cells, observed in thexylem of leaf traces exiting from nodal tissue, were suggestedto be involved in the abstraction process. Key words: Ricinus communis, nitrogen, nitrate, nitrate reduction, partitioning, phloem, xylem, flow models     

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     

The influence of salinity on the utilization of root anaplerotic carbon and nitrogen metabolism in tomato seedlings

In hydroponically grown Lycopersicon esculentum (L.) Mill. cv.F144 the site of NO₃^– reduction and assimilation withinthe plant was shifted from the shoot to the root by salinity.Uptake of NO₃^– from the root solution was strongly inhibitedby salinization. Consequently, NO₃^– concentrations inthe leaf, stem and root tissues as well as the nitrate reductaseactivities of the leaves were lower in salinized than in controlplants. Lower NO₃^–, but higher reduced-N, concentrationswere observed in the xylem sap as a result of the enhanced participationof the root in NO₃^– reduction in salinized plants. Lowerstem K⁺ concentrations and leaf malate concentrations were foundin salinized compared to control plants which indicates reducedfunctioning of the K⁺–shuttle in the salinized plants. Incorporation of inorganic carbon by the root was determinedby supplying a pulse of NaH¹⁴CO₃ followed by extraction andseparation of the labelled products on ion exchange resins.The rate of H¹⁴CO₃^– incorporation was c. 2-fold higherin control than in salinized plants. In salinized plants theproducts of H¹⁴CO₃^– incorporation within the roots werediverted into amino acids, while the control plants divertedrelatively more ¹⁴C into organic acids. Products of inorganiccarbon incorporation in the roots of salinized plants providean anaplerotic source of carbon for assimilation of reducedNO₃^– into amino acids, while in control plants the productswere predominantly organic acids as part of mechanisms to maintainionic balance in the cells and in the xylem sap. Key words: Tomato, nitrate, PEPc, respiration, salinity