Keri Massie and Emily Williamson Biology 425, Spring 2009

Introduction  many theories about the evolution of hibernation (Kemp, 2005)  None have extra- ordinary evidence supporting them (Kemp, 2005) We’re going to explore theories specifically related to the evolution of hibernation in mammals

What is hibernation?!?! Reduced activity in order to battle extreme environmental conditions.

Who hibernates Mammals Birds Insects Reptiles

Causes in mammals Physiologically Decreases in body temperature, heart rate, and metabolism resulting in long-term dormancy Interrupted by brief “awake” periods and re- warming (Yuan et al., 2007). Genetically Different gene expression patterns during winter – PRL-2 (Yuan et al., 2007) – Genes for Pancreatic lipase – Genes for PDK-4

Genetically controlled hibernation (Geiser, 2002) PDK-4 helps control cardiac physiology allowing a prolonged dormant state Fig. 1: Expression of PDK-4 during the hibernation season.

Genetically controlled hibernation PRL-2 found in bats, highly conserved in many mammals Has different expression pattern during active times and hibernating times shown below

The Evolution of Hibernation in Mammals: the Genetic Effect Mutation in hemoglobin causing higher affinity for oxygen lead to changes needed for increased O 2 absorption and endothermy (Kemp, 2005) Genes that are highly conserved in species may be selected for PRL-2 (Yuan et al., 2007) Nonsynonymous and synonymous mutations both had less than 2% chance of becoming fixed over populations PRL-2 is thus passed down with small variability from generation to generation Physiological traits required for hibernation happen in invertebrates (Geiser, 1998)

The Evolution of Hibernation in Mammals: from Endotherms (Kemp, 2005)  Many believe hibernation has evolved from early endotherms  Endothermy allows animals to survive varying temperatures and supports higher degree of organization (Kemp, 2005) ○ have higher aerobic activity potential ○ higher breathing capacity

Ectotherm to Endotherm Mechanism First, evolved control over blood flow and shelter seeking at night. Insulation, more shelter seeking Controlled heat production, shelter seeking postponed until late in evening. Climate: Cool: significant controlled heat production, no need to seek shelter in active season, seek retreat in cold season. Warm: energetic needs favor year round activity. Cold: seek retreat in winter Various: year round endothermy Pattern Morning warm-up, stable Tb during day, cooling to air temp. at night. Tb maintained warm well into evening, active season extended. Tb maintained much later into evening, season extended further. Climate: Regular/occasional torpor, facultative winter hibernation. climate too warm for torpor Entry to prolonged hibernation in winter Tb essentially stable all day and night throughout year.

The Evolution of Hibernation in Mammals : Polyphyletically (Geiser, 1998) Some believe hibernation and torpor evolved separately in different mammals and in birds Geiser analyzed phylogenetic tree and compared torpor in bird and mammal species

The Evolution of Hibernation in Mammals: Plesiomorphically Many believe hibernation has evolved from a common ancestor, and has only evolved once from daily torpor Evidence: Small ancestral mammals are thought to be heterothermic and have a different pattern of torpor than in modern mammals with control of body temperature regulation. (Geiser, 2002) Hibernation was shown to be plesiomorphic in Eutheria, Marsupialia and Monotremata. Evolved from a common pattern of circadian heterothermy, ectothermic at first then facilitated by endothermic thermogenisis. (Grigg, 2000)

The Evolution of Hibernation in Mammals: Size and Diet Hypothesis (Geiser, 1998) Both mammals and birds that hibernate are small and rely on a varying diet Possibly all early animals that were small went through some form of hibernation or torpor. Most hibernated in order to conserve energy and food, necessary in order to survive extreme conditions in the winter.

How does this effect the evolution of hibernation? Theory of evolution of hibernation still debated over Researchers believe it is not one evolutionary process acting on mammals, but has evolved in small steps over many generations(Kemp, 2005)

Literature Cited Geiser, F Evolution of daily torpor and hibernation in birds and mammals: importance of body size. Clinical and Experimental Pharmacology and Physiology 25: Geiser, F., N. Goodship, C. Pavey Was basking important in the evolution of mammalian endothermy. Physiological Zoology 89: Grigg, G., L. Beard Hibernation by Echidnas in mild climates: Hints about the evolution of endothermy. In: Heldermeier, G. Klingenspor M (eds) Life in the Cold: Eleventh International Hibernation Symposium. Springer, Berlin Heidelberg New York, pp Kemp, T.S The origin of mammalian endothermy: a paradigm for the evolution of complex biological structure. Zoological Journal of the Linnean Society 147: Yuan, L., J. Chen, B. Lin, J. Zhung, and S. Zhang Differential expression and functional constraint of PRL-2 in hibernating bat. Comparative Biochemistry and Physiology, Part b 148:

Photo sources: National Museum of Natural History Microsoft Office 2007, clip art Google images k_hibernating.jpg k_hibernating.jpg