How does the study of hummingbird metabolism exemplify one approach to the study of animal movement?

Hummingbirds have the highest mass-specific rate of metabolism, as well as the broadest metabolic range of any known vertebrate. At the peak of physical activity they are capable of utilizing approximately 40 ml of oxygen per gram per hour, and at rest can lower their metabolism to about 3 ml of oxygen per gram per hour (Healy et al. 2006). Such novel metabolic capabilities make the hummingbird an organism of great biological significance. This incredibly efficient metabolism has likely been evolved to maintain the hummingbirds highly specialized form of locomotion, hovering flight. This form of flight enables hummingbirds to move at a faster rate than any other comparably sized avian, terrestrial, or aquatic organism (Healy et al. 2006). To better understand this mode of locomotion is crucial to understanding the other specializations in avian physiology and morphology, as well as broader scopes of bird behavior and evolution. Certainly, locomotion is more energetically expensive than rest; however, some modes of locomotion may be more energetically efficient than others-depending on the organism and the environment that it has adapted to. To understand the kind of strain that different forms of locomotion have on an organism, it is crucial to gain a better understanding of the organisms metabolism. An important aspect of metabolism is the utilization of various fuel types in order to maximize energy available for muscular use. Hummingbirds use fatty acid oxidation serves as the primary energy source in resting birds, while foraging hummingbirds fuel their activity by a rapid shift to carbohydrates (Suarez et al. 1990). This is an optimal system as the oxidation of carbohydrates produces an immediate source of energy which is perfect for quick movements and activities that elicit an immediate response, such as foraging. The oxidation of fats is far more beneficial during rest, as it provides twice as much energy (per carbon atom) as carbohydrates do, allowing longer periods of time before more energy is needed to be consumed.

Hummingbirds have also been found to be capable of tolerating plasma glucose levels so high that such concentrations would cause serious neurological and microvascular pathologies in diabetic humans (Beuchat et al. 1998). This capability of humming bird metabolism, along with some of the highest rates of carrier-mediated glucose transport in the intestine seen in vertebrate species (Diamond et al. 1986), allow hummingbirds to ingest and process more glucose without suffering the deleterious effects that such high sugar concentrations would have in other species. Furthermore, hummingbirds are most typically found at altitudes ranging from 1000 to 2500m and are progressively colonizing higher elevations over the generations (Bleiweiss et al. 1998). Thus, not only are hummingbirds utilizing extremely high levels of oxygen during their daily locomotive activities, but they are doing so at elevations in which the partial pressure of oxygen is significantly lower than that at sea level. Thus, respiratory and cardiovascular performances as displayed by the hummingbird show exciting promises of vertebrate metabolic capacity in the face of hypoxic and energetically challenging conditions (Dudley et al. 2001). So what exactly can the preceding studies on hummingbird metabolism provide us with in terms of better understanding hummingbird locomotion? As I mentioned above, animal structural design has functional implications that determine physiological processes and ultimately the ability to exist under specific physiological constraints. To understand the metabolic processes of an organism is to better understand the mechanism of their evolved, and highly specialized, physiologies. In many organisms, maximal metabolic activity (during aerobic exercise) is primarily limited by oxygen transport and delivery to the working muscles. However, in highly metabolically active organisms (such as the hummingbird), maximal metabolic activity is more likely to be dependent upon the design limitations of muscles and transport systems (Turner et al. 2006). Metabolic rate is a logical indicator of how organisms provide

ATP to the muscles that they use to move themselves through their environment, which in effect, defines the study of locomotion. Bibliography Healy S, Hurly AT, Quick Guide: Hummingbirds. Current Biology 16(11): 392-393, (2006). Suarez RK, Lighton JRB, Moyes CD, Brown GS, Gass CL, Hochachka PW, Fuel Selection in rufous hummingbirds: Ecological implications of metabolic biochemistry. Ecology 87: 9207-9210, (1990). Beuchat CA, Chong CR, Hyperglycemia in hummingbirds and its consequences for hemoglobin glycation. Comparative Biochemistry and Physiology 120(3): 409-416, (1998). Diamond JM, Karasov WH, Phan D, Carpenter FL, Digestive physiology is a determinant of foraging bout frequency in hummingbirds. Nature 320: 6263. (1986). Bleiweiss R, Origin of hummingbird faunas. Biological Journal of the Linnean Society 65: 77-97 (1998). Dudley R, Limits to human locomotor performance: phylogenetic origins and comparative perspectives. The Journal of Experimental Biology 204: 3235-3240 (2001). Turner N, Hulbert AJ, Else PL, Limits to physical performance and metabolism across secies.Current Opinion in Clinical Nutrition and Metabolic Care 9(6): 691-696 (2006).