Mechanical independence of wingbeat and breathing in starlings.

The pectoral muscles in birds comprise up to a third of the body weight and provide the principal drive to the wing. Their attachment to the sternum suggests that they could compress the thorax and assist ventilation during flight. Most, but not all, birds have an integer ratio relationship between wingbeat and breathing frequency, but no measurements of the respiratory flow associated with the act of wingbeat are available. We recorded respiratory flow and wing timing in three starlings that flew at 22 knots (11 m.s-1) for up to 5 min in a wind tunnel. Triggering on wingbeat, we ensemble averaged flow records for many wingbeats in each flight. Because wingbeats occurred throughout the respiratory cycle, breathing flow tended to average to zero, and a small flow event related to wingbeat emerged. The volume change associated with wingbeat ranged from 3 to 11% of tidal volume, and this is probably an overestimate. We conclude that wingbeat and breathing in starlings are essentially mechanically independent, despite the direct attachment of the locomotor muscles to the thorax.

An aerodynamic valve in the avian primary bronchus.

The segmentum accelerans in geese is a constriction in the caudal end of the primary bronchus. Experimental evidence suggests that this part of the airway functions as an inspiratory aerodynamic valve, accelerating the incoming airstream past the ventrobronchial openings. The luminal diameter of the segmentum accelerans dilates in the presence of elevated CO2 levels, probably through relaxation of smooth muscle. Physiological control of the segmentum accelerans may permit inspiratory aerodynamic valving to be maintained throughout a wide range of ventilatory flows.

Pressure profiles show features essential to aerodynamic valving in geese.

Inspiratory airflow in the avian lung completely bypasses the most cranial secondary bronchi (the ventrobronchi), and instead enters bronchi arising more caudally (the dorsobronchi). Dotterweich (1936) proposed that 'aerodynamic valves' prevented entry into the ventrobronchi. We have recently provided evidence that inspiratory aerodynamic valving in avian lungs depends on convective inertia in the primary bronchus (Banzett et al., 1987). Theoretical and physical models (Butler et al., 1988; Wang et al., 1988) showed that convective inertia could effect valving, but the effectiveness of valving at resting flows was less than that observed in the bird. This leads us to hypothesize that a segment of the primary bronchus is constricted, accelerating the gas and enhancing the convective inertia. To test this hypothesis in the present work we measured pressures throughout the airways and air sacs in anesthetized, pump-ventilated geese at different flow rates and gas densities. Our data show: (1) there is a large pressure drop in the primary bronchus close to the ventrobronchial junction, indicating the presence of a constriction; (2) this pressure drop increases with gas density and flow; (3) the convective inertia at this site is more than 10 times downstream opposing pressures. We conclude that the primary bronchus just cranial to the first ventrobronchus forms a constriction which accelerates inspired air. Furthermore, we conclude that the convective inertia of gas leaving this segment is sufficient to achieve inspiratory valving.

Bracing arms increases the capacity for sustained hyperpnea.

Patients with severe chronic obstructive pulmonary disease (COPD) frequently lean forward, bracing their arms. We wondered whether the resulting shoulder girdle support improves the function of the ventilatory pump. We tested this possibility in 4 normal men by measuring the maximal ventilation that they could voluntarily sustain for 4 min while seated with their elbows braced firmly on a table and while seated with their elbows held just above the table. Bracing the arms increased ventilatory capacity significantly in all subjects, but the magnitude of the change was small (8%). We attribute the change to improved function of the accessory muscles that expand the rib cage. We speculate that this effect assumes greater importance in patients with COPD, whose diaphragms are flattened and ineffective, because such patients depend more on the inspiratory muscles of the rib cage.

Inspiratory aerodynamic valving in goose lungs depends on gas density and velocity.

The non-reversing gas flow pattern in the avian lung has been attributed to 'aerodynamic valves'. A fundamental property of all aerodynamic valves is their dependence on inertial forces in the gas stream: sufficient reduction of inertial forces will cause aerodynamic valves to fail. If valving in the avian lung is aerodynamic, it should fail when gas stream momentum is reduced. We tested the dependence of the inspiratory valves in the goose lung on gas density and gas flow velocity. A bolus of tracer gas was placed in the tracheal cannula during an end-expiratory pause. Tracer gas appearance in a cranial air sac during the following inspiration and pause was used to deduce failure of the 'inspiratory valve' in cyclically ventilated geese. Little or no tracer entered the sac under control conditions, which approximated resting breathing, indicating highly effective valving. Lower flow rate or lower gas density caused increased tracer appearance, indicating valve failure. These results demonstrate the importance of gas inertial forces to normal valve function, and are direct evidence for the aerodynamic nature of the avian inspiratory valve.