A Comparison of the Uptake and Translocation of Some Organic Herbicides and a Systemic Fungicide by Barley: II. RELATIONSHIP BETWEEN UPTAKE BY ROOTS AND TRANSLOCATION TO SHOOTS

The relationship between uptake by barley roots and translocationto shoots has been examined under different conditions for sixherbicides and a systemic fungicide (four triazines, diuron,2,4-dichlorophenoxyacetic acid and ethirimol). For all thesecompounds a large proportion of the material taken up by rootsdoes not appear to move readily to the shoots. Elution withwater of the roots of intact plants which had previously beenplaced in labelled solutions of these compounds enabled thematerial released to be separated into three fractions differingin their rates of diffusion out of the roots. There was reasonablygood correlation between the concentration of the most readilydiffusible fraction in the roots and the concentration in thetranspiration stream. The results obtained were consistent withthe postulate that lipophilic compounds can diffuse into thevacuoles of the cortical cells where they are available fortransport to the shoots, whereas for lipophobic compounds materialreaching the shoots originates largely from the free space inthe roots.     

A Comparison of the Uptake and Translocation of Some Organic Herbicides and a Systemic Fungicide by Barley: I. ABSORPTION IN EELATION TO PHYSICO-CHEMICAL PROPERTIES

A comparative study has been made of the uptake by and translocationfrom roots of intact barley plants of six herbicides and a systemicfungicide (four triazines, diuron, 2,4-dichloro-phenoxyaceticacid (2,4-D) and ethirimol). Relationships between uptake andtranspiration rate are discussed in the light of the physico-chemicalproperties of these compounds, notably their partition coefficientsin oil/water systems and their dissociation constants. Apartfrom 2,4-D, sorption of these compounds appears to be a passiveprocess. At pH4 the uptake of 2,4-D seems to be influenced bymetabolism; not only may the concentration of this compoundin the transpiration stream be considerably greater than thatin the medium surrounding the roots but absorption by rootsis markedly reduced at low temperatures and by sodium azide. The initial rate of uptake of these compounds correlates reasonablywell with their partition coefficients in olive oil/water orn-dodecane/water systems; likewise the concentration in thetranspiration stream is greater for lipophilic than for lipophobicsubstances. Whereas the hydrogen ion and calcium concentrations of the ambientmedium appear to have no effect on the uptake of compounds withlow pK's, the uptake of those substances which protonate betweenpH4 and pH6 is affected by them. These findings are discussedfrom the viewpoint that the pathways of transport of lipophilicand lipophobic compounds across the roots may differ. Although there is some evidence that retention by roots canlimit transport to shoots, there is no simple inverse correlationbetween the total concentration of the different substancesin the roots and that in the transpiration stream. This questionis discussed in a subsequent paper.     

Origins of the Electrical Potential Difference between the Xylem Sap of Maize Roots and the External Solution

The effects of 2,4-dinitrophenol, of changes in the temperatureand concentration of the ambient solution and of variationsin salt status on the electrical potential difference betweenthe xylem exudate of maize roots and the ambient solution havebeen examined. The results are discussed in the light of someof the factors which could give rise to a potential differencebetween the sap and the solution. The rapid response of thepotential difference to dinitrophenol and to changes in temperaturesuggest that, at least in part, it arises directly from metabolicprocesses. Rapid changes in the potential difference broughtabout by addition of salts may be attributed to differentialrates of movement of anions and cations in the initial uptakeprocess. Over longer periods the potential difference appearsto be dependent on the concentration, but not the compositionof the ambient solution, and on the salt status of the roots.The salt status influences the relative rates at which anionsand cations are transported to the xylem sap, and a correlationhas been found between the potential difference and the ratioof the rates of movement of chloride or sulphate to potassiumto the sap. The implications of these findings on the elucidationof the pathways whereby ions are transported to the sap arediscussed.     

The Initial Uptake of Ions by Barley Roots: I. UPTAKE OF ANIONS

Experiments have been carried out to examine the applicabilityof the Goüy–Chapman electrical double-layer theoryto the initial entry of anions into excised barley roots. Anattempt was made to separate the total quantity of labellediodide or sulphate ions which had entered the roots under varyingconditions into three fractions: the two fractions which couldbe successively eluted with water and extracted with an exchangingsolution and that which remained in the roots after these treatments.Except at a low temperature and a low hydrion concentration,the three fractions could not clearly be distinguished, andthere were indications of the presence of metabolically maintainedpositive sites. However, the effect of varying pH on the rateat which anions were eluted with water was consistent with thepresence in the roots of a system of pores, the walls of whichcarried a varying number of negative charges depending on thehydrion concentration. The effect of calcium ions on the magnitudeof the water extractable fraction also accorded qualitativelywith the double-layer theory.     

The Initial Uptake of Ions by Barley Roots: II. APPLICATION OF MEASUREMENTS ON SORPTION OF ANIONS TO ELUCIDATE THE STRUCTURE OF THE FREE SPACE

In previous experiments on excised barley roots, results werepresented showing the effect of varying the concentration ofthe uptake solution on the amount of anion which could subsequentlybe extracted in water. It was concluded that the system couldbe more satisfactorily analysed by means of the Goüy–Chapmanelectrical double-layer theory than by the Donnan theory. Onthe basis that ions diffuse into a system of negatively chargedpores within the root, the relationship between the quantityof water-extractable anion and its concentration in the uptakesolution has been examined in the light of the electrical double-layertheory. For potassium iodide and sulphate the experimental resultsconformed reasonably well with a structural model for the FreeSpace of parallel, charged surfaces separated by a mean distanceof about 200 A. The internal surface area of the free spacewas estimated as 2x10⁶ cm²/g dry weight. The uptake of iodidefrom calcium iodide was somewhat greater than that predictedby the theory; this may be ascribed to interaction between theanions and divalent cations within the double layer.     

Measurement of free space and sorption of large molecules by cereal roots

Abstract. Large molecular weight solutes that do not penetrate the root have been used to correct for the surface film in measurements with mannitol of the volume of the Apparent Free Space (FS) in bailey roots. The results are compared with those obtained using other correction techniques for elimination of the surface film. Large molecules seem to be adsorbed on the root surface and the kinetics of adsorption differ between the polyhydric alcohol mannitol or the polysaccharide dextran on the one hand, and the polyether polyethylene glycol (PEG-4000) on the other. The significance of this difference in kinetics is discussed in relation to the use of PEG as an osmoticum in studies on root water relations and its effect on ion uptake. Although smaller molecular weight PEG's penetrate the FS and diminish sodium uptake from 10 mol m⁻³ NaCl, more dilute solutions of mannitol and larger PEG polymers are unlikely to affect ion uptake from dilute nutrient solutions. Use of these substances along with labelled nutrients in kinetic studies of the compartmentation of ions in roots can help to distinguish between ions associated with the surface film, those in the FS and those that have crossed the cell membranes into the protoplast.     

Effect of Sodium Chloride Stress and Nitrogen Source on Respiration, Growth and Photosynthesis in Lucerne (Medicago sativa L.)

The effects of salinity at different light intensities on freshweight growth and on carbon dioxide influx and efflux were examinedin young plants of lucerne (Medicago sativa L.) that had beengrown in solution culture with nitrogen supplied either as nitrateor by a symbiotic rhizobium. Although the inoculated plantsgrew more slowly than those supplied with nitrate, NaCl at alevel equivalent to an osmotic stress of –0.3 MPa didnot reduce the growth rate of either type of plant under a 12h day-length in a growth chamber. With a day-length of 5 h saltstress (0.0 to –0.6 MPa) did not greatly affect grossphotosynthesis of plants grown on nitrate but respiration ratereached a maximum at –0.3 MPa and declined at larger saltconcentrations. Salt diminished both gross and net photosynthesisin the inoculated plants at a 5 h day-length without stimulatingrespiration. The relationship between photosynthesis and respiration as thephoton flux density was successively decreased was used to inferthe effect of salt on maintenance respiration of the plantssupplied with nitrate. Growth and maintenance components ofrespiration could not clearly be separated in the inoculatedplants suggesting that these were unable to maintain themselvesunder the combined stresses of salt and low light intensity.This view was supported by chemical analysis of the plant material.We conclude that the failure of the inoculated plants to adaptto these conditions could be attributed to the greater demandfor assimilates by the rhizobium. Key words: Medicago sativa, Nitrogen source, Salt stress     

The Absorption of Silica from Aqueous Solutions by Plants

Silica dissolves in water to a small extent and below pH7 existsin solution essentially in a non-polar form. In this respectit contrasts with nutrient cations and anions and a study ofits uptake by plants was thought to be of interest. It was foundthat in common with the absorption of ionized solutes the entryof silica into plants appears to require the expenditure ofmetabolic energy since the process is sensitive to both metabolicinhibitors and variations in temperature. Furthermore, exceptunder conditions of very low humidity, silica enters barleyplants at a relatively greater rate than the water lost in transpiration,and its concentration in the xylem sap of bean plants may begreater than that in the external solution.     

Electrochemical Relations in the Transfer of Ions to the Xylem Sap of Maize Roots

The difference in electrical potential between the xylem exudateof isolated roots of 1-week-old maize seedlings and the culturesolution surrounding the roots has been determined togetherwith the concentrations of potassium, calcium, chloride, andsulphate in the xylem sap. The difference in electro-chemicalpotential (µ) for these ions has been calculated fromthe measurements. The effects on µ of varying the saltstatus of the roots, the composition of the culture solutionand of 2-4-dinitrophenol have been examined. µ for potassiumand chloride was always positive, implying that movement ofthese ions to the xylem sap was under metabolic constraint.However, pretreatment of the maize seedlings with potassiumchloride and increasing levels of dinitrophenol in the culturesolution over the exudation period caused little or no significantchange in µ for potassium although the rate of movementof ions to the xylem was substantially reduced. For calcium,µ was negative with roots in dilute culture solutionsin the absence of dinitrophenol, implying that calcium couldenter the xylem by passive diffusion. Addition of dinitrophenolchanged the sign of µ and thus brought about an apparentlyactive transport of calcium.Only in the absence of competinganions was the rate of entry of chloride significantly correlatedwith µ, and no such relationship was found for potassiumor calcium individually. These results encourage doubt as towhether measurements of µ provide a valid basis for decidingto what extent the movement of individual ions depends on specific‘active’ or ‘passive’ transport processes.Thedifference in electrical potential appears to be a characteristicof the living root and to depend on the total concentrationof ions in the external solution and on their rate of transferinto the xylem sap.     

Factors Affecting Absorption and Translocation of Simazine by Barley

The factors affecting the absorption and translocation of simazineby young barley plants in short-term experiments in water culturehave been investigated. Chromatographic examination of the xylemsap indicated no extensive breakdown of the herbicide in thetranspiration stream. Under varying conditions of humidity,light intensity, temperature, and in the presence of metabolicinhibitors, the concentration of simazine in the transpirationstream relative to that in the uptake medium was always lessthan unity. This could, in part, be attributed to retentionof simazine at a higher concentration on a fresh weight basisin the root tissues than in the ambient medium. There was littleevidence that the absorption and translocation of simazine wasinfluenced by metabolism except in so far as this affected movementof water, and micro-organisms present on the plant roots atambient laboratory levels had no effect on the uptake and transportof the herbicide. Simazine brings about a decrease in the rate of transpirationand a reduction in the total uptake of rubidium and phosphateat relatively high concentrations. However, when the phosphateconcentration in the ambient medium was below the level at whichtranslocation of this ion is affected by the rate of transpiration,the herbicide did not reduce uptake of phosphate. There is thereforeno evidence that in short-term experiments simazine has an effecton active transport processes.