Control of water uptake by rice (Oryza sativa L.):: role of the outer part of the root

A new pressure-perfusion technique was used to measure hydraulic and osmotic properties of the outer part of roots (OPR) of 30-day-old rice plants (lowland cultivar: IR64, and upland cultivar: Azucena). The OPR comprised rhizodermis, exodermis, sclerenchyma and one cortical cell layer. The technique involved perfusion of aerenchyma of segments from two different root zones (20-50 mm and 50-100 mm from the tip) at precise rates using aerated nutrient solution. The hydraulic conductivity of the OPR (Lp(OPR)=1.2x10(-6) m s(-1) MPa-1) was larger by a factor of 30 than the overall hydraulic conductivity (Lp(r)=4x10(-8) m s(-1) MPa-1) as measured by pressure chamber and root pressure probe. Low reflection coefficients were obtained for mannitol and NaCl for the OPR (sigma(sOPR)=0.14 and 0.09, respectively). The diffusional water permeability (P-dOPR) estimated from isobaric flow of heavy water was smaller by three orders of magnitude than the hydraulic conductivity (Lp(OPR)/P-fOPR). Although detailed root anatomy showed well-defined Casparian bands and suberin lamellae in the exodermis, the findings strongly indicate a predominantly apoplastic water flow in the OPR. The Lp(OPR) of heat-killed root segments increased by a factor of only 2, which is in line with the conclusion of a dominating apoplastic water flow. The hydraulic resistance of the OPR was not limiting the passage of water across the root cylinder. Estimations of the hydraulic properties of aerenchyma suggested that the endodermis was rate-limiting the water flow, although the aerenchyma may contribute to the overall resistance. The resistance of the aerenchyma was relatively low, because mono-layered cortical septa crossing the aerenchyma ('spokes') short-circuited the air space between the stele and the OPR. Spokes form hydraulic bridges that act like wicks. Low diffusional water permeabilities of the OPR suggest that radial oxygen losses from aerenchyma to medium are also low. It is concluded that in rice roots, water uptake and oxygen retention are optimized in such a way that hydraulic water flow can be kept high in the presence of a low efflux of oxygen which is diffusional in nature.