Incomplete gas mixing in air sacs of the duck

During normal breathing, the CO2 concentration in caudal air sacs of birds is higher, and the O2 concentration lower, than expected on the basis of the known air flow pattern. We have experimentally tested two hypotheses which could explain this finding: (1) Preferential shunting of re-inspired dead space gas into caudal air sacs; (2) Incomplete mixing of inspired and residential air sac gas. - Different portions of the inspired air in anesthetized ducks were labeled by injecting a small bolus of argon (Ar) into the trachea. The resulting Ar concentration was recorded continuously in the caudal thoracic air sac at the ostium and in deeper regions.-The amount of Ar entering the sac was found to be independent of the volume inspired prior to injection of the label, and hypothesis (1) thus dismissed. However, during inspiration and subsequent expiration the Ar bolus was found to be neither perfectly mixed within the inspired gas nor with the air sac residential gas. More than 10 sec of breath-hold were necessary for air sac gas to approach an equilibrium value. Gas layering (stratification) in caudal air sacs gas is proposed to cause the high CO2 and low O2 levels during steady state breathing, as air sac residential gas equilibrates with a layer of dead space gas that enters the air sac on each breath and contains a higher CO2 and lower O2 concentration than the mixed inspirate.