Summary

In vivo measurements of pectoralis muscle force during different modes of free flight (takeoff, level flapping, landing, vertical ascending and near vertical descending flight) were obtained using a strain gauge attached to the dorsal surface of the delto-pectoral crest (DPC) of the humerus in four trained pigeons (Columba livia). In one bird, a rosette strain gauge was attached to the DPC to determine the principal axis of strain produced by tension of the pectoralis. Strain signals recorded during flight were calibrated to force based on in situ measurements of tetanic force and on direct tension applied to the muscle's insertion at the DPC. Rosette strain recordings showed that at maximal force the orientation of tensile principal strain was −15° (proximo-anterior) to the perpendicular axis of the DPC (or +75° to the longitudinal axis of the humerus), ranging from +15 to −25° to the DPC axis during the downstroke. The consistency of tensile principal strain orientation in the DPC confirms the more general use of single-element strain gauges as being a reliable method for determining in vivo pectoralis force generation. Our strain recordings show that the pectoralis begins to develop force as it is being lengthened, during the final one-third of the upstroke, and attains maximum force output while shortening during the first one-third of the downstroke. Force is sustained throughout the entire downstroke, even after the onset of the upstroke for certain flight conditions. Mean peak forces developed by the pectoralis based on measurements from 40 wingbeats for each bird (160 total) were: 24.9+/−3.1 N during takeoff, 19.7+/−2.0 N during level flight (at speeds of about 6–9 m s-1 and a wingbeat frequency of 8.6+/−0.3 Hz), 18.7+/−2.5 N during landing, 23.7+/−2.7 N during near-vertical descent, and 26.0+/−1.8 N during vertical ascending flight. These forces are considerably lower than the maximum isometric force (67 N, P0) of the muscle, ranging from 28 % (landing) to 39 % (vertical ascending) of P0. Based on estimates of muscle fiber length change determined from high- speed (200 frames s-1) light cine films taken of the animals, we calculate the mass-specific power output of the pigeon pectoralis to be 51 W kg-1 during level flight (approximately 8 m s-1), and 119 W kg-1 during takeoff from the ground. When the birds were harnessed with weighted backpacks (50 % and 100 % of body weight), the forces generated by the pectoralis did not significantly exceed those observed in unloaded birds executing vertical ascending flight. These data suggest that the range of force production by the pectoralis under these differing conditions is constrained by the force- velocity properties of the muscle operating at fairly rapid rates of shortening (4.4 fiber lengths s-1 during level flight and 6.7 fiber lengths s-1 during takeoff).