RESUMEN
Root growth in higher plants is sensitive to excess ammonium (NH(4)(+)). Our study shows that contact of NH(4)(+) with the primary root tip is both necessary and sufficient to the development of arrested root growth under NH(4)(+) nutrition in Arabidopsis. We show that cell elongation and not cell division is the principal target in the NH(4)(+) inhibition of primary root growth. Mutant and expression analyses using DR5:GUS revealed that the growth inhibition is furthermore independent of auxin and ethylene signalling. NH(4)(+) fluxes along the primary root, measured using the Scanning Ion-selective Electrode Technique, revealed a significant stimulation of NH(4)(+) efflux at the elongation zone following treatment with elevated NH(4)(+), coincident with the inhibition of root elongation. Stimulation of NH(4)(+) efflux and inhibition of cell expansion were significantly more pronounced in the NH(4)(+)-hypersensitive mutant vtc1-1, deficient in the enzyme GDP-mannose pyrophosphorylase (GMPase). We conclude that both restricted transmembrane NH(4)(+) fluxes and proper functioning of GMPase in roots are critical to minimizing the severity of the NH(4)(+) toxicity response in Arabidopsis.