Respiratory gas exchange at lungs, gills and tissues: mechanisms and adjustments.

(1) A general model for external gas exchange organs of vertebrates is presented, in which the main parameters are the ventilatory, diffusive and perfusive conductances for O2 and CO2. The relevant properties of the external medium (air or water) and of the internal medium (blood) are analysed in terms of capacitance coefficients (effective solubilities) for O2 and CO2. The models for the main types of gas exchange organs (fish gills, amphibian skin, and avian and mammalian lungs) are compared in terms of their intrinsic gas exchange efficacy. The adjustments to increased metabolic rate or to hypoxia are achieved by increasing the conductances. (2) The gas exchange at tissue level is analysed using the Krogh cylinder and a simplified model containing a diffusive and a perfusive conductance. The adjustments to increased load (exercise, hypoxia) consist in both increased local blood flow and in improvement of diffusion conditions (enlargement and recruitment of capillaries). (3) Some particular features of respiration in transitional (unsteady) states, such as occurring at the beginning of exercise and of hypoxia, are examined. The additional physical variables are the O2 (and CO2) stores acting according to their capacitances and partial pressure changes. Delayed increase in O2 uptake at the beginning of exercise is due to the limited speed of physiological adjustments. The ensuing O2 debt is energetically covered by anoxidative energy releasing processes (hydrolysis of high-energy phosphates and anaerobic glycolysis). Finally, the reduction of metabolic rate as adjustment to hypoxia is discussed.