The problem of living in the sea: the uptake of inorganic carbon and nutrients in Posidonia oceanica (L.) Delile

Abstract

The genus Posidonia exhibits a peculiar geographical distribution. It is composed by nine species, eight of which are distributed along the Australian coasts and only one, Posidonia oceanica (L.) Delile, is a Mediterranean endemism. Like other angiosperms, P. oceanica has adapted secondarily to the marine environment, and has developed anew mechanisms to face a liquid and alkaline medium (pH 8.2) that contains a high salt concentration (0.5 M NaCl). The liquid environment limits the diffusive flow of CO2 and nutrients and, furthermore, CO2 dissolves in water and forms HCO3-, the more abundant chemical species of inorganic carbon at pH 8.2. Like other green plants P. oceanica uses CO2 for photosynthesis. In addition, this species shows a transport system in the plasma membrane for the direct uptake of HCO3-, that uses H+ as the driving ion. The addition of HCO3- provokes a transient hyperpolarization of the plasma membrane followed by a depolarization; at the same time, the cytosolic pH (pHc) becomes transiently acidic and next it gets alkaline, and remains alkaline throughout the HCO3- pulse. The alkalinization of the pHc is due to the cytosolic accumulation of HCO3- and OH- and it is sensitive to the addition of ethoxyzolamide, an inhibitor of the internal carbonic anhydrase. The increase of negative charges in the cytosol triggers the release of Cl- to recover the values of the resting membrane potential. The plasmalemma of P. oceanica exhibits a reduced Na+ permeability and shows a H+/Na+ antiporter activity that keeps low and relatively constant the cytosolic Na+ concentration (17 mM Na+). The inside negative membrane potential (-178 mV) and the low [Na+]c generate a tremendous Na+-motive force that this plant uses for the high affinity transport of NO3- (Km= 21 µM), and of the amino acids alanine (Km= 37 µM) and cysteine (Km= 10 µM). The uptake of these compounds shows a strict dependence on the presence of Na+ in the medium. Moreover, the addition of micromolar concentrations of NO3-, alanine or cysteine gives rise to millimolar increments of [Na+]c. Experiments with external LIX pH mini-electrodes show that the uptake of glucose is not Na+ but H+ dependent. Thus, the model for the ion transport energization in this species seems to be mixed, with a H+-ATPase as the primary pump and a series of carriers that use H+ (HCO3-, Na+, glucose) or Na+ (NO3-, amino acids) as the driving ion.

Project Funding: CTM 2011-30356 (MEC)