Cardiovascular function in large to small hibernators: bears to ground squirrels
Mammalian hibernation has intrigued scientists due to extreme variations in normal seasonal physiological homeostasis. Numerous species manifest a hibernation phenotype although the characteristics of the hypometabolic state can be quite different. Ground squirrels (e.g., Sciuridae) are often considered the prototypical hibernator as individuals in this genus transition from an active, euthermic state (37 °C) to a nonresponsive hibernating state where torpid body temperature commonly falls to 3–5 °C. However, the hibernating state is not continuous as periodic warming and arousals occur. In contrast, the larger hibernators of genus Ursus are less hypothermic (body temperatures decline from approximately 37°–33 °C), are more reactive, and cyclical arousals do not occur. Both species dramatically reduce cardiac output during hibernation from the active state (bears ~75 % reduction in bears and ~97 % reduction in ground squirrels), and both species demonstrate hypokinetic atrial chamber activity. However, there are several important differences in cardiac function between the two groups during hibernation. Left ventricular diastolic filling volumes and stroke volumes do not differ in bears between seasons, but increased diastolic and stroke volumes during hibernation are important contributors to cardiac output in ground squirrels. Decreased cardiac muscle mass and increased ventricular stiffness have been found in bears, whereas ground squirrels have increased cardiac muscle mass and decreased ventricular stiffness during hibernation. Molecular pathways of cardiac muscle plasticity reveal differences between the species in the modification of sarcomeric proteins such as titin and α myosin heavy chain during hibernation. The differences in hibernation character are likely to account for the alternative cardiac phenotypes and functional strategies manifested by the two species. Molecular investigation coupled with better knowledge of seasonal physiological alterations is dramatically advancing our understanding of small and large hibernators and their evolutionary differences.