Anatomy and histochemistry of spread-wing posture in birds. 3. Immunohistochemistry of flight muscles and the "shoulder lock" in albatrosses.

As a postural behavior, gliding and soaring flight in birds requires less energy than flapping flight. Slow tonic and slow twitch muscle fibers are specialized for sustained contraction with high fatigue resistance and are typically found in muscles associated with posture. Albatrosses are the elite of avian gliders; as such, we wanted to learn how their musculoskeletal system enables them to maintain spread-wing posture for prolonged gliding bouts. We used dissection and immunohistochemistry to evaluate muscle function for gliding flight in Laysan and Black-footed albatrosses. Albatrosses possess a locking mechanism at the shoulder composed of a tendinous sheet that extends from origin to insertion throughout the length of the deep layer of the pectoralis muscle. This fascial "strut" passively maintains horizontal wing orientation during gliding and soaring flight. A number of muscles, which likely facilitate gliding posture, are composed exclusively of slow fibers. These include Mm. coracobrachialis cranialis, extensor metacarpi radialis dorsalis, and deep pectoralis. In addition, a number of other muscles, including triceps scapularis, triceps humeralis, supracoracoideus, and extensor metacarpi radialis ventralis, were found to have populations of slow fibers. We believe that this extensive suite of uniformly slow muscles is associated with sustained gliding and is unique to birds that glide and soar for extended periods. These findings suggest that albatrosses utilize a combination of slow muscle fibers and a rigid limiting tendon for maintaining a prolonged, gliding posture.

Effects of mass distribution on the mechanics of level trotting in dogs.

The antero-posterior mass distribution of quadrupeds varies substantially amongst species, yet the functional implications of this design characteristic remain poorly understood. During trotting, the forelimb exerts a net braking force while the hindlimb exerts a net propulsive force. Steady speed locomotion requires that braking and propulsion of the stance limbs be equal in magnitude. We predicted that changes in body mass distribution would alter individual limb braking-propulsive force patterns and we tested this hypothesis by adding 10% body mass near the center of mass, at the pectoral girdle, or at the pelvic girdle of trotting dogs. Two force platforms in series recorded fore- and hindlimb ground reaction forces independently. Vertical and fore-aft impulses were calculated by integrating individual force-time curves and Fourier analysis was used to quantify the braking-propulsive (b-p) bias of the fore-aft force curve. We predicted that experimental manipulation of antero-posterior mass distribution would (1) change the fore-hind distribution of vertical impulse when the limb girdles are loaded, (2) decrease the b-p bias of the experimentally loaded limb and (3) increase relative contact time of the experimentally loaded limb, while (4) the individual limb mean fore-aft forces (normalized to body weight + added weight) would be unaffected. All four of these results were observed when mass was added at the pelvic girdle, but only 1, 3 and 4 were observed when mass was added at the pectoral girdle. We propose that the observed relationship between antero-posterior mass distribution and individual limb function may be broadly applicable to quadrupeds with different body types. In addition to the predicted results, our data show that the mechanical effects of adding mass to the trunk are much more complex than would be predicted from mass distribution alone. Effects of trunk moments due to loading were evident when mass was added at the center of mass or at the pelvic girdle. These results suggest a functional link between appendicular and axial mechanics via action of the limbs as levers.