A thermodynamic analysis of the feasibility of water secretion into xylem vessels against a water potential gradient.

A series of recent publications has launched a debate on trans-membrane water secretion into root xylem vessels against a water potential gradient, energised by a cotransport with salts (e.g. KCl) that follow their chemical potential gradient. Cation-chloride-cotransporter -type transporters that function in this way in mammalian epithelia were detected in root stelar cells bordering on xylem vessels. Using literature data on barley (Hordeum vulgare L.) seedlings, one study confirmed that K+ and Cl- gradients across stelar cell membranes favour salt efflux. Moreover, the energetic costs of putative water secretion into the xylem (required for maintaining ionic gradients) would amount to just 0.12% of the energy captured by photosynthetic C assimilation if transpirational water flow relied exclusively on this mechanism. Here, a detailed thermodynamic analysis of water secretion into xylem vessels is undertaken, including an approach that exploits its analogy to a desalinisation process. Water backflow due to the passive hydraulic conductivity of stelar cell membranes is also considered. By comparing free energy consumption by putative water secretion with (i) the free energy pool provided by root respiration and (ii) stelar ATPase activity, the feasibility of this mechanism is confirmed but is shown to depend critically on the plant's energy status.