Standing-Gradient Osmotic Flow A mechanism for coupling of water and solute transport in epithelia

At the ultrastructural level, epithelia performing solute-linked water transport possess long, narrow channels open at one end and closed at the other, which may constitute the fluid transport route (e.g., lateral intercellular spaces, basal infoldings, intracellular canaliculi, and brush-border microvilli). Active solute transport into such folded structures would establish standing osmotic gradients, causing a progressive approach to osmotic equilibrium along the channel's length. The behavior of a simple standing-gradient flow system has therefore been analyzed mathematically because of its potential physiological significance. The osmolarity of the fluid emerging from the channel's open end depends upon five parameters: channel length, radius, and water permeability, and solute transport rate and diffusion coefficient. For ranges of values of these parameters encountered experimentally in epithelia, the emergent osmolarity is found by calculation to range from isotonic to a few times isotonic; i.e., the range encountered in epithelial absorbates and secretions. The transported fluid becomes more isotonic as channel radius or solute diffusion coefficient is decreased, or as channel length or water permeability is increased. Given appropriate parameters, a standing-gradient system can yield hypertonic fluids whose osmolarities are virtually independent of transport ratp/'over a wide range, as in distal tubule and avian salt gland. The results suggest ~ that water-to-solute coupling in epithelia is due to the ultrastructural geometry of the transport route. T h e purpose of this paper is to examine whether some distinctive physiological properties of epithelia migh t arise as geometr ical consequences of epithelial ul t rastructure. Mos t epithelia absorb or secrete specific fluids, such as bile, the g lomerular filtrate, gastric juice, aqueous humour , sweat, and cerebrospinal fluid. T h e p r imary t ranspor ted fluid m a y be ei ther isotonic or hyper tonic to p lasma (hypotonic secretions in vertebrates appear to result f rom isotonic secretion