Birds are a diverse group of vertebrates, with over 10,000 extant species. Diversification into volant, aquatic and terrestrial environmental niches has precipitated a remarkable morphological diversity between species. Birds have a unique respiratory system consisting of a rigid lung connected to an air sac system. Air is pumped into the respiratory system via movements of the ribcage and sternum. Previous research identified the uncinate processes, ossified projections extending from the vertebral ribs, as critical respiratory and locomotor structures. Uncinate processes facilitate inspiration and expiration through associated muscles that displace the ribs and therefore sternum. External intercostal muscles project from the processes and function during locomotion to stabilise body roll. Therefore uncinate processes provide a link between breathing and locomotion in birds. The objective of my PhD is to extend beyond this basic research on uncinate processes to investigate how diversity in avian body morphology relates to the fundamental functions of breathing and locomotion.While the function of uncinate processes in respiration has been identified, the mechanism whereby ventilatory movements are elicited is not known. Therefore I present a model that demonstrates how respiratory movements of the skeleton are facilitated by the lever action of uncinate processes. Furthermore, variation in process and sternal morphology is driven by adaptation to different forms of locomotion. Therefore fundamental differences in breathing mechanics may be associated with specialisation to locomotor behaviour. Detailed developmental studies of the uncinate processes in birds are almost nonexistent. I provide the first detailed description of developmental changes in the uncinate processes in the turkey. Ossification of the uncinate processes begins around the time of hatch. However, the base is cartilaginous upon hatching and so the lever action of the processes may be compromised in the chick. I provide further evidence for a functional link between process length and respiratory physiology, since elongated processes support an elevated resting metabolic rate in birds. This link was further explored in physiological experiments where the energetic cost of walking in the barnacle goose was manipulated by load carrying. Carrying extra mass on the sternum is more energetically costly than an equivalent back load indicating that the cost of breathing increased. A directly proportional relationship exists between increasing mass of back load and metabolic rate, while sternal loads were approximately twice as expensive to carry during locomotion. Leg loads incurred the greatest increase in metabolism. Finally, I demonstrate how uncinate processes functioned as respiratory structures in basal avian species and a theropod ancestor of modern birds. Development of the uncinate processes may have been an important step in the evolution of the avian lung - air sac system.The principal findings of the five first author research articles presented in this PhD thesis shed important new light on the ventilatory mechanics in birds and highlight interactions between breathing and locomotion. Diversity in avian body morphology driven by adaptation to various locomotor behaviours has resulted in modification of the respiratory system.
|Date of Award||31 Dec 2010|
- The University of Manchester
|Supervisor||Jonathan Codd (Supervisor)|
- uncinate process