Flapping of the entire wing occurs primarily through the actions of the chest muscles, the pectoralis and the supracoracoideus. These muscles are highly developed in birds and account for a higher percentage of body mass than in most mammals. These attach to a blade-shaped keel, like that of a boat, located on the sternum. The sternum of birds is larger than that of other vertebrates, which accommodates the large muscles required to generate enough upward force to generate lift with the flapping of the wings. Another skeletal modification found in most birds is the fusion of the two clavicles (collarbones), forming the furcula or wishbone. The furcula is flexible enough to bend and provide support to the shoulder girdle during flapping.
An important requirement of flight is a low body weight. As body weight increases, the muscle output required for flying increases. The largest living bird is the ostrich, and while it is much smaller than the largest mammals, it is flightless. For birds that do fly, reduction in body weight makes flight easier. Several modifications are found in birds to reduce body weight, including pneumatization of bones. Pneumatic bones are bones that are hollow, rather than filled with tissue ( [link] ). They contain air spaces that are sometimes connected to air sacs, and they have struts of bone to provide structural reinforcement. Pneumatic bones are not found in all birds, and they are more extensive in large birds than in small birds. Not all bones of the skeleton are pneumatic, although the skulls of almost all birds are.
Other modifications that reduce weight include the lack of a urinary bladder. Birds possess a cloaca, a structure that allows water to be reabsorbed from waste back into the bloodstream. Uric acid is not expelled as a liquid but is concentrated into urate salts, which are expelled along with fecal matter. In this way, water is not held in the urinary bladder, which would increase body weight. Most bird species only possess one ovary rather than two, further reducing body mass.
The air sacs that extend into bones to form pneumatic bones also join with the lungs and function in respiration. Unlike mammalian lungs in which air flows in two directions, as it is breathed in and out, airflow through bird lungs travels in one direction ( [link] ). Air sacs allow for this unidirectional airflow, which also creates a cross-current exchange system with the blood. In a cross-current or counter-current system, the air flows in one direction and the blood flows in the opposite direction, creating a very efficient means of gas exchange.