The alveolar gas equation: do lessons from WW2 research into high flying pilots provide insights into the adaptation to high altitude flight in birds?

Abstract

Daltons law of partial pressures applies equally to birds and mammals so, as gas moves from the nostrils to the smallest gas-diffusion airways, the sequential addition of water vapour and CO2, steadily reduce the partial pressure of O2 (PO2) within the gas mixture. The PO2, at the point of gas exchange, at sea level, will be about 60 mm Hg less than the original PO2 within atmospheric air. As a result, the inspired PO2 is an inaccurate starting point for any model of oxygen transport. In humans, the interactions of gases at the point of diffusion, is described and quantified by the Alveolar Gas Equation (AGE). Its development during WW2, provided valuable insights into human gas exchange and also into the responses to high altitude flight in pilots but, except for an early study of hypoxia in pigeons, the AGE is not mentioned in the avian literature. Even detailed models of oxygen transport in birds omit the effect of CO2 clearance on pulmonary oxygen transfer. This paper develops two related arguments concerning the application of the AGE to birds. The first is that avian blood gas predictions, based on the theory of multicapillary serial arterialization (MSA), are inaccurate because they do not account for the added partial pressure of diffused CO2. The second is that the primary adaptation to hypobaric hypoxia is the same for both classes and consists of defending PaO2 by reducing PaCO2 through increasing hyperventilation. Support for the first is demonstrated by comparing PaO2 predictions made using the AGE, with published values from avian studies and also against values predicted by the theory of MSA. The second is illustrated by comparing the results of high altitude studies of both birds and humans. The application of the AGE to avian respiratory physiology would improve the predictive accuracy of models of the O2 cascade and would also provide better insights into the primary adaptation to high altitude flight.