Ammonium Nutrition in Ricinus communis: Its Effect on Plant Growth and the Chemical Composition of the Whole Plant, Xylem and Phloem Saps

Allen, S. and Smith, J A. C. 1986. Ammonium nutrition in Ricinuscommunis: its effect on plantgrowth and the chemical compositionof the whole plant, xylem and phloem saps.—J. exp. Bot.37: 1599–1610. The growth and chemical composition of Ricinus communis cultivatedhydroponically on 12 mol m ^{– 3} NO₃^–-N were comparedwith plants raised on a range of NH₄⁺-N concentrations. At NH₄⁺-Nconcentrations between 0·5 and 4·0 mol m^–3,fresh- and dry-weight yields of 62-d-old plants were not significantlydifferent from those of the NO₃^–-N controls. Growth wasreduced at 0·2 mol m^–3 NH₄⁺-N and was associatedwith increased root. shoot and C: organic N ratios, suggestingthat the plants were N-limited. At 8·0 mol m^–3NH₄⁺-N, growth was greatly restricted and the plants exhibitedsymptoms of severe ‘NH₄⁺ toxicity’. Plants growingon NH₄⁺-N showed marked acidification of the rooting medium,this effect being greatest on media supporting the highest growthrates. Shoot carboxylate content per unit dry weight was lower at mostNH₄⁺-N concentrations than in the NO₃^–-N controls, althoughit increased at the lowest NH₄⁺-N levels. Root carboxylate contentwas comparable on the two N sources, but also increased substantiallyat the lowest NH₄⁺-N levels. N source had little effect on inorganic-cationcontent at the whole-plant level, while NO₃^– and carboxylatewere replaced by Cl as the dominant anion in the NH₄⁺-N plants.This was reflected in the ionic composition of the xylem andleaf-cell saps, the latter containing about 100 mol m^–3Cl^– in plants on 8·0 mol m^–3 NH₄⁺. Xylem-saporganic-N concentration increased more than threefold with NH₄⁺-N(with glutamine being the dominant compound irrespective ofN source) while in leaf-cell sap it increased more than 12-foldon NH₄⁺-N media (with arginine becoming the dominant species).In the phloem, N source had little or no effect on inorganic-cation,sucrose or organic-N concentrations or sap pH, but sap fromNH₄⁺-N plants contained high levels of Cl^– and serine. Collectively, the results suggested that the toxic effects ofhigh NH₄⁺ concentrations were not the result of medium acidification,reduced inorganic-cation or carboxylate levels, or restrictedcarbohydrate availability, as is commonly supposed. Rather,NH₄⁺ toxicity in R. communis is probably the result of changesin protein N turnover and impairment of the photorespiratoryN cycle. Key words: Ricinus, ammonium nutrition, nitrate, whole-plant composition, xylem, Phloem, amino acids, carboxylate