532:, body temperature, physical activity, ambient temperature, and snow depth were measured to identify the drivers of the start and end of hibernation for bears. This study built the first chronology of both ecological and physiological events from before the start to the end of hibernation in the field. This research found that bears would enter their den when snow arrived and ambient temperature dropped to 0 °C. However, physical activity, heart rate, and body temperature started to drop slowly even several weeks before this. Once in their dens, the bears' heart rate variability dropped dramatically, indirectly suggesting metabolic suppression is related to their hibernation. Two months before the end of hibernation, the bears' body temperature starts to rise, unrelated to heart rate variability but rather driven by the ambient temperature. The heart rate variability only increases around three weeks before arousal and the bears only leave their den once outside temperatures are at their lower critical temperature. These findings suggest that bears are thermoconforming and bear hibernation is driven by environmental cues, but arousal is driven by physiological cues.
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terrestrial environments as a mechanism to keep energy costs low, particularly in harsher than normal environments, as well as being a good way to time the active or reproductive periods in arthropods. It is thought to have originally evolved in three stages. The first is development of neuroendocrine control over bodily functions, the second is pairing of that to environmental changesâin this case metabolic rates decreasing in response to colder temperaturesâand the third is the pairing of these controls with reliable seasonal indicators within the arthropod, like biological timers. From these steps, arthropods developed a seasonal diapause, where many of their biological functions end up paired with a seasonal rhythm within the organism. This is a very similar mechanism to the evolution of insect migration, where instead of bodily functions like metabolism getting paired with seasonal indicators, movement patterns would be paired with seasonal indicators.
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hypothesis that hibernation evolved after endothermy in response to the emergence of colder habitats. Body size also had an effect on the evolution of hibernation, as endotherms which grow large enough tend to lose their ability to be selectively heterothermic, with bears being one of very few exceptions. After torpor and hibernation diverged from a common proto-hibernating ancestor of birds and mammals, the ability to hibernate or go through torpor would have been lost in most larger mammals and birds. Hibernation would be less favored in larger animals because as animals increase in size, the surface area to volume ratio decreases, and it takes less energy to keep a high internal body temperature, and thus hibernation becomes unnecessary.
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environments. As hibernation is a seasonal response, the movement of the ancestor of birds and mammals onto land introduced them to seasonal pressures that would eventually become hibernation. This is true for all clades of animals that undergo winter dormancy; the more prominent the seasons are, the longer the dormant period tends to be on average. Hibernation of endothermic animals has likely evolved multiple times, at least once in mammalsâthough it is debated whether or not it evolved more than once in mammalsâand at least once in birds.
45:
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785:) and for fish that undergo winter dormancy in hypoxic conditions, they do suppress their metabolism like other animals that are dormant in the winter. The mechanism for evolution of metabolic suppression in fish is unknown. Most fish that are dormant in the winters save enough energy by being still and so there is not a strong selective pressure to develop a metabolic suppression mechanism like that which is necessary in hypoxic conditions.
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aging. Reptiles evolved to exploit their ectothermy to deliberately cool their internal body temperatures. As opposed to mammals or birds, which will prepare for their hibernation but not directly cause it through their behavior, reptiles will trigger their own hibernation through their behavior. Reptiles seek out colder temperatures based on a periodic internal clock, which is likely triggered by cooler outside temperatures, as shown in the
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mammals that produce valuable proteins. Protein sequencing of HIT proteins, such as α 1-glycoprotein-like 88 kDa hibernation-related protein HRP, contributes to this research pool. A study in 2014 utilizes recombinant technology to construct, express, purify, and isolate animal proteins (HP-20, HP-25, and HP-27) outside of the animal to study key hibernation proteins (HP).
777:, do not. Instead, they do not actively depress their base metabolic rate, but instead they simply reduce their activity level. Fish that undergo winter dormancy in oxygenated water survive via inactivity paired with the colder temperature, which decreases energy consumption, but not the base metabolic rate that their bodies consume. But for the
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As the ancestors of birds and mammals colonized land, leaving the relatively stable marine environments, more intense terrestrial seasons began playing a larger role in animals' lives. Some marine animals do go through periods of dormancy, but the effect is stronger and more widespread in terrestrial
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opioid and hibernation induction trigger (HIT) proteins were not able to increase the recovery rate of heart tissue during ischemia. While unable to increase recovery rates at the time of ischemia, the protein precursors were identified to play a role in the preservation of veterinary organ function.
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interrupted by periodic, euthermic arousals, during which body temperatures and heart rates are restored to more typical levels. The cause and purpose of these arousals are still not clear; the question of why hibernators may return periodically to normal body temperatures has plagued researchers for
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conservation in the genetic mechanism for diapause. Particularly the timing and extent of the seasonal diapause seem particularly variable, currently evolving as a response to climate change. As typical with hibernation, it evolved after the increased influence of seasonality as arthropods colonized
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As reptiles are ectothermic, having no system to deal with cold temperatures would be deadly in many environments. Reptilian winter dormancy, or brumation, likely evolved to help reptiles survive colder conditions. Reptiles that are dormant in the winter tend to have higher survival rates and slower
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go into hibernation during the winter months in order to give birth to their offspring. The pregnant mothers significantly increase their body mass prior to hibernation, and this increase is further reflected in the weight of the offspring. The fat accumulation enables them to provide a sufficiently
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Hibernation functions to conserve energy when sufficient food is not available. To achieve this energy saving, an endothermic animal decreases its metabolic rate and thereby its body temperature. Hibernation may last days, weeks, or monthsâdepending on the species, ambient temperature, time of year,
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have been documented to be able to survive for three hours without oxygen and at temperatures of up to 26 °C (79 °F) as a means to survive in their shoreline habitat, where water and oxygen levels vary with the tide. Other animals able to survive long periods with very little or no oxygen
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truly hibernate, since they experience only a modest decline in body temperature (3â5 °C) compared with the much larger decreases (often 32 °C or more) seen in other hibernators. Many researchers thought that their deep sleep was not comparable with true, deep hibernation, but this theory
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With the exception of some tropical species, seed germination is typically postponed for some period that is determined by characteristics of seeds and the environmental conditions they are exposed to . seeds may remain ungerminated and persist in the soil for many years. In this case, latency is
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of typical hibernators to three different bear species with respect to season, activity and dormancy, and found that the reduced relaxation (QT) interval of small hibernators was the same for the three bear species. They also found the QT interval changed for both typical hibernators and the bears
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Recent advances in recombinant protein technology make it possible for scientists to manufacture hibernation induction trigger (HIT) proteins in the laboratory without the need for animal euthanasia. Bioengineering of proteins can aid in the protection of vulnerable populations of bears and other
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that is produced in sufficient quantities to satisfy the water needs of the bear. They also do not eat or drink while hibernating, but live off their stored fat. Despite long-term inactivity and lack of food intake, hibernating bears are believed to maintain their bone mass and do not suffer from
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The hibernation of this lemur is strongly dependent on the thermal behaviour of its tree hole: If the hole is poorly insulated, the lemur's body temperature fluctuates widely, passively following the ambient temperature; if well insulated, the body temperature stays fairly constant and the animal
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hibernation. The last two sections point out in particular primates, none of whom were thought to hibernate until recently, and bears, whose winter torpor had been contested as not being "true hibernation" during the late 20th century, since it is dissimilar from hibernation seen in rodents.
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periods. In order to conserve energy, the ancestors of birds and mammals would likely have experienced an early form of torpor or hibernation when they were not using their thermoregulatory abilities during the transition from ectothermy to endothermy. This is opposed to the previously dominant
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Researchers have studied how to induce hibernation in humans. The ability to hibernate would be useful for a number of reasons, such as saving the lives of seriously ill or injured people by temporarily putting them in a state of hibernation until treatment can be given. For space travel, human
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There is evidence that hibernation evolved separately in marsupials and placental mammals, though it is not settled. That evidence stems from development, where as soon as young marsupials from hibernating species are able to regulate their own heat, they have the capability to hibernate. In
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Hibernating Arctic ground squirrels may exhibit abdominal temperatures as low as â2.9 °C (26.8 °F), maintaining sub-zero abdominal temperatures for more than three weeks at a time, although the temperatures at the head and neck remain at 0 °C (32 °F) or above.
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Facultative hibernators enter hibernation only when either cold-stressed, food-deprived, or both, unlike obligate hibernators, who enter hibernation based on seasonal timing cues rather than as a response to stressors from the environment.
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winter dormancy, seems to be plastic and quickly evolving, with large genetic variation and strong effects of natural selection present as well as having evolved many times across many clades of arthropods. As such, there is very little
709:, an ancestor of mammals that lived roughly 252 million years ago. The evolution of endothermy allowed animals to have greater levels of activity and better incubation of embryos, among other benefits for animals in the
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and the individual's body-condition. Before entering hibernation, animals need to store enough energy to last through the duration of their dormant period, possibly as long as an entire winter. Larger species become
365:. These species undergo what has been traditionally called "hibernation": a physiological state wherein the body temperature drops to near ambient temperature, and heart and respiration rates slow drastically.
588:(including fish, reptiles, and amphibians) cannot engage in obligate or facultative hibernation. They can experience decreased metabolic rates associated with colder environments or low oxygen availability (
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726:, only developing the ability to hibernate at a later point. This difference in development is evidence, though inconclusive, that they evolved by slightly different mechanisms and thus at different times.
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Ultsch, Gordon R. 1989. Ecology and
Physiology of Hibernation and Overwintering Among Freshwater Fishes, Turtles, and Snakes. Biological Reviews 64(4), pp. 435-515. doi:10.1111/j.1469-185X.1989.tb00683.x
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and became dormant, but research by David Sims in 2003 dispelled this hypothesis, showing that the sharks traveled long distances throughout the seasons, tracking the areas with the highest quantity of
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induced by external environmental conditions, so these seeds are effectively in hibernation. Because of seed hibernation and dormancy, many plant populations consist of adult individuals as well as
747:. Hypercapnic acidosis evolved as a mechanism to slow metabolism and also interfere with oxygen transport so that oxygen is not used up and can still reach tissues in low oxygen periods of dormancy.
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Horton, Noel D; Kaftani, Dimitra J; Bruce, David S; Bailey, Evans C; Krober, Alan S; Jones, Jeffrey R; Turker, Mitchell; Khattar, Nada; Su, Tsung-Ping; Bolling, Steven F; Oeltgen, Peter R (1998).
743:). One mechanism that reptiles use to survive hibernation, hypercapnic acidosis (the buildup of carbon dioxide in the blood), is also present in mammal hibernation. This is likely an example of
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Obligate hibernators are animals that spontaneously, and annually, enter hibernation regardless of ambient temperature and access to food. Obligate hibernators include many species of
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381:. Other theories postulate that brief periods of high body temperature during hibernation allow the animal to restore its available energy sources or to initiate an immune response.
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There is a variety of definitions for terms that describe hibernation in mammals, and different mammal clades hibernate differently. The following subsections discuss the terms
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Prendergast, B.J.; Freeman, D.A.; Zucker, I.; Nelson, R.J. (2002). "Periodic arousal from hibernation is necessary for initiation of immune responses in ground squirrels".
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warm and nurturing environment for their newborns. During hibernation, they subsequently lose 15â27% of their pre-hibernation weight by using their stored fats for energy.
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Toien, Oivind; Black, J.; Edgar, D. M.; Grahn, D. A.; Heller, H. C.; Barnes, B. M. (February 2011). "Black Bears: Independence of
Metabolic Suppression from temperature".
120:, eating a large amount of food and storing the energy in their bodies in the form of fat deposits. In many small species, food caching replaces eating and becoming fat.
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Lohuis, T. D.; Harlow, H. J.; Beck, T. D. I. (2007). "Hibernating black bears (Ursus americanus) experience skeletal muscle protein balance during winter anorexia".
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in the muscle, as well as regulate the transcription of a suite of genes that limit muscle wasting. A study by G. Edgar Folk, Jill M. Hunt and Mary A. Folk compared
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to describe winter dormancy of reptiles, but the more general term hibernation is believed adequate to refer to any winter dormancy. Many insects, such as the wasp
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that indicated the belief was still current in his time. It is now understood that the vast majority of bird species typically do not hibernate, instead utilizing
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and is now applied based on active metabolic suppression rather than any absolute decline in body temperature. Many experts believe that the processes of daily
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decades, and while there is still no clear-cut explanation, there are multiple hypotheses on the topic. One favored hypothesis is that hibernators build a "
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Harlow, H.J.; Frank, C.L. (2001). "The role of dietary fatty acids in the evolution of spontaneous and facultative hibernation patterns in prairie dogs".
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Carey, H.V.; Andrews, M.T.; Martin, S.L. (2003). "Mammalian hibernation: cellular and molecular responses to depressed metabolism and low temperature".
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Hibernation induction trigger (HIT) proteins isolated from mammals have been used in the study of organ recovery rates. One study in 1997 found that
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457:
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Watts PD, Oritsland NA, Jonkel C, Ronald K (1981). "Mammalian hibernation and the oxygen consumption of a denning black bear (Ursus americanus)".
3291:"The benefit of being still: energy savings during winter dormancy in fish come from inactivity and the cold, not from metabolic rate depression"
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winter temperatures sometimes rise to over 30 °C (86 °F), so hibernation is not exclusively an adaptation to low ambient temperatures.
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2006:"Proteomic and Transcriptomic Changes in Hibernating Grizzly Bears Reveal Metabolic and Signaling Pathways that Protect against Muscle Atrophy"
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Lundberg, D. A.; Nelson, R. A.; Wahner, H. W.; Jones, J. D. (1976). "Protein metabolism in the black bear before and during hibernation".
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Humphries, M. M.; Thomas, D.W.; Kramer, D.L. (2003). "The role of energy availability in mammalian hibernation: A cost-benefit approach".
2239:) which I found in a state of profound torpidity in the winter of 1946â47 in the Chuckawalla Mountains of the Colorado Desert, California
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In both cases, hibernation likely evolved simultaneously with endothermy, with the earliest suggested instance of hibernation being in
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Seldin, Marcus; Byerly, Mardi; Petersen, Pia; Swason, Roy; Balkema-Buschmann, Anne; Groschup, Martin; Wong, G. William (2014).
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Evans, Alina L.; Singh, N.J; Friebe, A.; Arnemo, J.M.; Laske, T.G.; Frobert, O.; Swensen, J.E.; Blanc, S. (11 February 2016).
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2555:"Seasonal oscillation of liver-derived hibernation protein complex in the central nervous system of non-hibernating mammals"
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Hibernating bears are able to recycle their proteins and urine, allowing them to stop urinating for months and to avoid
2762:"Torpor in the Patagonian opossum (Lestodelphys halli): implications for the evolution of daily torpor and hibernation"
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Dausmann, K.H.; Glos, J.; Ganzhorn, J.U.; Heldmaier, G. (June 2004). "Physiology: Hibernation in a tropical primate".
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Dausmann, K.H.; Glos, J.; Ganzhorn, J.U.; Heldmaier, G. (2005). "Hibernation in the tropics: Lessons from a primate".
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can naturally freeze and then "wake" up in the spring. These species have evolved freeze tolerance mechanism such as
2381:"Breathless: A shark with an amazing party trick is teaching doctors how to protect the brains of stroke patients"
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Galster, W.; Morrison, P. R. (1975). "Gluconeogenesis in arctic ground squirrels between periods of hibernation".
139:, which in many invertebrates is referred to as diapause. Some researchers and members of the public use the term
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from summer to winter. This 1977 study was one of the first evidences used to show that bears are hibernators.
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377:" during hibernation, and so must occasionally warm up to sleep. This has been supported by evidence in the
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Folk, G. Edgar; Hunt, Jill M.; Folk, Mary A. (February 1977). "Further
Evidence for Hibernation of Bears".
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Fuster, Gemma; Busquets, SĂlvia; Almendro, Vanessa; LĂłpez-Soriano, Francisco J.; ArgilĂ©s, Josep M. (2007).
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Geiser, Fritz (2004). "Metabolic Rate and Body
Temperature Reduction During Hibernation and Daily Torpor".
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155:, exhibit periods of dormancy which have often been referred to as hibernation, despite their ectothermy.
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Hoekstra, Luke; Schwartz, Tonia; Sparkman, Amanda; Miller, David; Bronikowski, Anne (September 9, 2019).
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While most animals that go through winter dormancy lower their metabolic rates, some fish, such as the
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1737:"Antiproteolytic effects of plasma from hibernating bears: A new approach for muscle wasting therapy?"
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young, which are born either while the mother hibernates or shortly afterwards. For example, female
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Campbell, Hamish; Fraser, Keiron; Bishop, Charles; Peck, Lloyd; Egginton, Stuart (March 5, 2008).
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McKechnie, Andrew W.; Ashdown, Robert A. M., Christian, Murray B. & Brigham, R. Mark (2007).
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3340:"Hypoxia Tolerance in Reptiles, Amphibians, and Fishes: Life with Variable Oxygen Availability"
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Bolling, S.F.; Tramontini, N.L., Kilgore, K.S., Su, T-P., Oeltgen, P.R., Harlow, H.H. (1997).
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2516:"Isolation and partial characterization of an opioid-like 88 kDa hibernation-related protein"
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and hibernation form a continuum and utilise similar mechanisms. The equivalent during the
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2976:"The untapped potential of reptile biodiversity for understanding how and why animals age"
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3138:"Evolutionary and functional genetics of insect diapause: a call for greater integration"
2876:"Evolution of daily torpor and hibernation in birds and mammals: importance of body size"
1374:"Freeze Avoidance in a Mammal: Body Temperatures Below 0 °C in an Arctic Hibernator"
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A good example of the differences between these two types of hibernation can be seen in
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Biochemistry and Physiology Part A: Molecular & Integrative Physiology
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Comparative
Biochemistry and Physiology Part B: Biochemistry and Molecular Biology
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Comparative
Biochemistry and Physiology Part B: Biochemistry and Molecular Biology
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American
Journal of Physiology. Regulatory, Integrative and Comparative Physiology
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The typical winter season for obligate hibernators is characterized by periods of
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Freeze avoidance in a Mammal: Body
Temperatures Below 0°C in an Arctic Hibernator
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Wilsterman, Kathryn; Ballinger, Mallory; Williams, Caroline (November 11, 2021).
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Folk, Edgar; Larson, Anna; Folk, Mary (1976). "Physiology of
Hibernating Bears".
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Earlier I gave an account (Condor, 50, 1948:45) of the behavior of a Poor-will (
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1230:"Warming up for sleep? Ground squirrels sleep during arousals from hibernation"
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Hibernation induction trigger (HIT) protein and recombinant protein technology
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characterized by low body-temperature, slow breathing and heart-rate, and low
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Harvested Human Lung Preservation With the Use of Hibernation Trigger Factors
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2207:(MayâJune 1949). "Further Observations on the Hibernation of the Poor-will".
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in hibernating animals is not necessarily coupled with low body temperature.
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Anthropologists are also studying whether hibernation was possible in early
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of Madagascar, which hibernates in tree holes for seven months of the year.
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Physiological state of dormant inactivity in order to pass the winter season
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2463:"Use of "Natural" Hibernation Induction Triggers for Myocardial Protection"
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Speers-Roesch, Ben; Norin, Tommy; Driedzic, William (September 5, 2018).
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2312:"Seasonal movements and behavior of basking sharks from archival tagging"
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Nelson, R. A. (1980). "Protein and fat metabolism in hibernating bears".
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Some animals can literally survive winter by freezing. For example, some
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3203:"Genetic shift in photoperiodic response correlated with global warming"
2664:"Hibernation in hominins from Atapuerca, Spain half a million years ago"
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that may be composed of seeds produced over a number of growing seasons.
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Sims, DW; Southall, EJ; Richardson, AJ; Reid, PC; Metcalfe, JD (2003).
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depression undergone by some animal species. Hibernation is a seasonal
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2930:"Ontogeny and phylogeny of endothermy and torpor in mammals and birds"
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592:) and exhibit dormancy (known as brumation). It was once thought that
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Ectothermic animals also undergo periods of metabolic suppression and
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Blanco, M.B.; Dausmann, K.; Ranaivoarisoa, J.F.; Yoder, A.D. (2013).
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3136:
Ragland, Gregory; Armbruster, Peter; Meuti, Megan (December 2019).
1734:
1149:
Vierek (Hymenoptera: Vespidae) Hibernate Inside Muddauber Nests?".
956:
794:
714:
611:
602:
468:
438:
230:
183:
175:
136:
520:
In a 2016 study, wildlife veterinarian and associate professor at
2829:"A phenology of the evolution of endothermy in birds and mammals"
942:
710:
685:
638:
541:
444:
was known to hibernate until the discovery of hibernation in the
434:
342:
93:
1522:
2415:
Herpetology: an introductory biology of amphibians and reptiles
800:
774:
555:
441:
426:
369:
330:
193:
156:
105:
101:
92:
Although traditionally reserved for "deep" hibernators such as
82:
52:
37:"Hibernate" redirects here. For the Java database library, see
803: â State of decreased physiological activity in an animal
2713:"A unifying, eco-physiological framework for animal dormancy"
1081:"Predicting climate change impacts on polar bear litter size"
480:
3261:. New York City: Oxford University Press. pp. 219â264.
3201:
Bradshaw, William; Holzapfel, Christina (November 6, 2001).
1900:"Hibernating bear serum hinders osteoclastogenesis in-vitro"
1078:
3255:
Tauber, Maurice; Tauber, Christine; Masaki, ShinzĆ (1986).
3194:
797: â State of minimized physical activity of an organism
630:
508:. They also increase the availability of certain essential
97:
2552:
2389:. Vol. 177, no. 2385. p. 46. Archived from
2309:
869:
159:
may use the term "seed hibernation" to refer to a form of
2253:"Torpor in an African caprimulgid, the freckled nightjar
722:
contrast, placental mammals that hibernate first develop
456:
undergoes regular spells of arousal. Dausmann found that
96:, the term has been redefined to include animals such as
2513:
3375:
3373:
3295:
Proceedings of the Royal Society B: Biological Sciences
2662:
Antonis Bartsiokas, Juan-Luis Arsuaga (December 2020).
1436:
1145:
GonzĂĄlez, Jorge M.; Vinson, S. Bradleigh (2007). "Does
2645:"Hibernating astronauts would need smaller spacecraft"
2095:
1285:
1079:
Molnar PK, Derocher AE, Kianjscek T, Lewis MA (2011).
573:
3382:"Hibernation in an Antarctic Fish: On Ice for Winter"
3331:
3250:
3248:
2880:
Clinical and Experimental Pharmacology and Physiology
2760:
Geiser, Fritz; Martin, Gabriel (September 18, 2023).
895:
893:
677:
hibernation is also under consideration, such as for
3475:. Vol. 1â20 (11th ed.). McGraw-Hill. 2012.
3370:
2967:
1227:
580:
Cryopreservation § Freeze tolerance in animals
30:For the power-management process in computing, see
3473:McGraw-Hill Encyclopedia of Science and Technology
3441:
3282:
3245:
2706:
2704:
2655:
1964:
1190:. New York: Oxford University Press. p. 146.
1034:DeMaster, Douglas P.; Stirling, Ian (1981-05-08).
890:
827: â Cluster of honey bees in cold temperatures
2822:
2820:
2753:
566:), for which hibernation was first documented by
3515:
3338:Bickler, Philip; Buck, Leslie (March 17, 2007).
3200:
2604:"New Hibernation Technique might work on humans"
1827:
1821:
1033:
809: â Annual state for some plants and animals
3207:Proceedings of the National Academy of Sciences
2701:
548:documented anecdotal evidence in his 1789 book
2817:
2623:"Race to be first to 'hibernate' human beings"
1862:
1288:American Journal of Physiology. Legacy Content
1144:
863:
3021:
3019:
2601:
277:
3129:
2620:
2499:: CS1 maint: multiple names: authors list (
2411:
2295:: CS1 maint: multiple names: authors list (
2060:
522:Inland Norway University of Applied Sciences
2921:
2867:
2412:Vitt, Laurie J.; Caldwell, Janalee (2014).
2378:
691:
484:was refuted by research in 2011 on captive
479:Historically it was unclear whether or not
425:While hibernation has long been studied in
81:. It is most commonly used to pass through
3337:
3067:
3016:
2446:: CS1 maint: location missing publisher (
2098:"Drivers of hibernation in the brown bear"
1635:"Drivers of hibernation in the brown bear"
1516:
768:
388:
284:
270:
3415:
3405:
3379:
3314:
3228:
3218:
3169:
3043:
2999:
2973:
2785:
2736:
2578:
2330:
2172:
2123:
2113:
2037:
1933:
1923:
1660:
1650:
1558:
1187:Evolutionary Biology: A Plant Perspective
1112:
1047:
872:Comparative Biochemistry and Physiology A
750:
696:
3356:10.1146/annurev.physiol.69.031905.162529
2357:(2385): 46. 8 March 2003. Archived from
1228:Dan S, Barnes BM, Strijkstra AM (1991).
985:
914:10.1146/annurev.physiol.66.032102.115105
729:
467:
317:
123:Some species of mammals hibernate while
57:
43:
3288:
3254:
2003:
1897:
313:
14:
3516:
2759:
2203:
2146:
1800:
1525:"Underground Hibernation in a Primate"
1371:
1183:
899:
821: â Swedish hibernator (1861â1950)
409:hibernator, while the closely related
2347:"A Shark With an Amazing Party Trick"
1632:
945:Physiological and Biochemical Zoology
2874:Geiser, Fritz (September 25, 1998).
2627:Times Online (from The Sunday Times)
988:"Physiology of Hibernation in Bears"
3076:"Voluntary hypothermia in reptiles"
3032:Integrative and Comparative Biology
2826:
2063:Bears: Their Biology and Management
2004:Mugahid, Douaa (27 December 2019).
1971:Bears: Their Biology and Management
1865:Bears: Their Biology and Management
1482:Journal of Comparative Physiology B
574:Dormancy and freezing in ectotherms
69:is a state of minimal activity and
24:
3435:
3026:Malan, André (February 28, 2014).
2892:10.1111/j.1440-1681.1998.tb02287.x
25:
3550:
3480:
3142:Current Opinion in Insect Science
3135:
1633:Evans, Alina (11 February 2016).
1372:Barnes, Brian M. (30 June 1989).
62:Bats hibernating in a silver mine
3074:Regal, Philip (March 24, 1967).
2927:
2873:
2827:Lovegrove, Barry (May 7, 2016).
2602:Britt, Robert (April 21, 2005).
2276:10.1111/j.2007.0908-8857.04116.x
1967:"Bone Metabolism in Black Bears"
1300:10.1152/ajplegacy.1975.228.1.325
488:and again in 2016 in a study on
182:
3258:Seasonal Adaptations of Insects
3073:
3025:
2710:
2637:
2614:
2595:
2559:Journal of Experimental Biology
2546:
2507:
2454:
2405:
2372:
2339:
2303:
2244:
2197:
2140:
2089:
2054:
1997:
1958:
1891:
1856:
1794:
1767:
1677:
1626:
1575:
1473:
1430:
1365:
1322:
1279:
1221:
1177:
551:The Natural History of Selborne
2467:The Annals of Thoracic Surgery
2319:Marine Ecology Progress Series
1138:
1129:
1072:
1027:
979:
936:
837:
544:hibernated, and ornithologist
499:. They stay hydrated with the
333:in preparation for hibernation
13:
1:
3100:10.1126/science.155.3769.1551
2621:Harlow, John (May 27, 2007).
2532:10.1016/S0305-0491(98)00056-X
2479:10.1016/s0003-4975(97)00631-0
2379:Douglas Fox (March 8, 2003).
1898:Nasoori; et al. (2020).
831:
779:Antarctic yellowbelly rockcod
558:. One known exception is the
540:Ancient people believed that
3407:10.1371/journal.pone.0001743
2680:10.1016/j.anthro.2020.102797
2147:Geiser, Fritz (2013-03-04).
1925:10.1371/journal.pone.0238132
1246:10.1016/0304-3940(91)90276-Y
1184:Cruzan, Mitchell B. (2018).
884:10.1016/0300-9629(81)90645-9
665:
596:settled to the floor of the
7:
3344:Annual Review of Physiology
2928:Fritz, Geiser (June 2008).
2418:(4th ed.). Amsterdam.
1965:Floyd T, Nelson RA (1990).
902:Annual Review of Physiology
845:"Do Black Bears Hibernate?"
788:
420:
10:
3555:
3456:10.1152/physrev.00008.2003
3162:10.1016/j.cois.2019.08.003
2946:10.1016/j.cbpa.2007.02.041
2237:Phalaenoptilus nuttallinii
2030:10.1038/s41598-019-56007-8
1842:10.1016/j.cbpb.2006.12.020
1753:10.1016/j.clnu.2007.07.003
1343:10.1152/ajpregu.00562.2001
986:Hellgren, Eric C. (1998).
849:North American Bear Center
669:
577:
296:
36:
29:
3487:Do Black Bears Hibernate?
2778:10.1007/s00114-013-1098-2
2174:10.1016/j.cub.2013.01.062
2115:10.1186/s12983-016-0140-6
1652:10.1186/s12983-016-0140-6
1494:10.1007/s00360-004-0470-0
1163:10.3958/0147-1724-32.1.69
1151:Southwestern Entomologist
815: â Branch of biology
475:mother and cubs "denning"
2264:Journal of Avian Biology
692:Evolution of hibernation
564:Phalaenoptilus nuttallii
535:
463:
411:black-tailed prairie dog
403:white-tailed prairie dog
248:Critical thermal maximum
3504:Potential medical usage
2992:10.1111/1365-2435.13450
2738:10.1111/1365-2435.13718
1776:Mayo Clinic Proceedings
1706:10.1126/science.1199435
1401:10.1126/science.2740905
769:Winter dormancy in fish
389:Facultative hibernation
32:Hibernation (computing)
3307:10.1098/rspb.2018.1593
3220:10.1073/pnas.241391498
755:Seasonal diapause, or
751:Diapause in arthropods
697:In endothermic animals
530:heart rate variability
476:
446:fat-tailed dwarf lemur
379:Arctic ground squirrel
334:
329:material for its warm
63:
55:
3444:Physiological Reviews
2649:European Space Agency
2255:Caprimulgus tristigma
1451:10.1007/s003600000148
730:Brumation in reptiles
471:
321:
258:Developmental biology
61:
47:
39:Hibernate (framework)
2651:. November 18, 2019.
2393:on February 29, 2012
2102:Frontiers in Zoology
1639:Frontiers in Zoology
1234:Neuroscience Letters
783:Notothenia coriiceps
745:convergent evolution
314:Obligate hibernation
3398:2008PLoSO...3.1743C
3213:(25): 14509â14511.
3154:2019COIS...36...74R
3092:1967Sci...155.1551R
3086:(3769): 1551â1553.
2766:Naturwissenschaften
2729:2021FuEco..35...11W
2165:2013CBio...23.R188G
2022:2019NatSR...919976M
1916:2020PLoSO..1538132N
1698:2011Sci...331..906T
1596:2004Natur.429..825D
1543:2013NatSR...3E1768B
1439:J. Comp. Physiol. B
1418:on 16 December 2008
1393:1989Sci...244.1593B
1387:(4912): 1593â1595.
1097:2011NatCo...2..186M
741:Phrynosoma cornutum
737:Texas horned lizard
672:Suspended animation
643:antifreeze proteins
586:ectothermic animals
169:Part of a series on
3497:2021-04-29 at the
3045:10.1093/icb/icu002
2980:Functional Ecology
2833:Biological Reviews
2717:Functional Ecology
2633:on August 7, 2008.
2571:10.1242/jeb.095976
2332:10.3354/meps248187
2010:Scientific Reports
1741:Clinical Nutrition
1530:Scientific Reports
1337:(4): R1054âR1062.
1147:Polistes exclamans
1105:10.1038/ncomms1183
477:
351:European hedgehogs
335:
236:Embryonic diapause
146:Polistes exclamans
64:
56:
3539:Animal metabolism
2845:10.1111/brv.12280
2565:(15): 2667â2679.
2425:978-0-12-386919-7
2205:Jaeger, Edmund C.
1809:(12): 2955â2958.
1692:(6019): 906â909.
1590:(6994): 825â826.
1551:10.1038/srep01768
1040:Mammalian Species
1036:"Ursus maritimus"
616:red-eared sliders
294:
293:
16:(Redirected from
3546:
3529:Sleep physiology
3476:
3467:
3450:(4): 1153â1181.
3430:
3429:
3419:
3409:
3377:
3368:
3367:
3335:
3329:
3328:
3318:
3286:
3280:
3279:
3277:
3275:
3252:
3243:
3242:
3232:
3222:
3198:
3192:
3191:
3173:
3133:
3127:
3126:
3124:
3122:
3071:
3065:
3064:
3062:
3060:
3047:
3023:
3014:
3013:
3003:
2971:
2965:
2964:
2962:
2960:
2925:
2919:
2918:
2916:
2914:
2871:
2865:
2864:
2839:(2): 1213â1240.
2824:
2815:
2814:
2812:
2810:
2789:
2757:
2751:
2750:
2740:
2708:
2699:
2698:
2696:
2694:
2659:
2653:
2652:
2641:
2635:
2634:
2629:. Archived from
2618:
2612:
2611:
2599:
2593:
2592:
2582:
2550:
2544:
2543:
2511:
2505:
2504:
2498:
2490:
2458:
2452:
2451:
2445:
2437:
2409:
2403:
2402:
2400:
2398:
2376:
2370:
2369:
2367:
2366:
2361:on 26 April 2003
2343:
2337:
2336:
2334:
2316:
2307:
2301:
2300:
2294:
2286:
2284:
2278:. Archived from
2261:
2248:
2242:
2241:
2201:
2195:
2194:
2176:
2159:(5): R188âR193.
2144:
2138:
2137:
2127:
2117:
2093:
2087:
2086:
2058:
2052:
2051:
2041:
2001:
1995:
1994:
1962:
1956:
1955:
1937:
1927:
1895:
1889:
1888:
1860:
1854:
1853:
1825:
1819:
1818:
1798:
1792:
1791:
1771:
1765:
1764:
1732:
1726:
1725:
1681:
1675:
1674:
1664:
1654:
1630:
1624:
1623:
1579:
1573:
1572:
1562:
1520:
1514:
1513:
1477:
1471:
1470:
1434:
1428:
1427:
1425:
1423:
1417:
1411:. Archived from
1378:
1369:
1363:
1362:
1326:
1320:
1319:
1283:
1277:
1276:
1274:
1273:
1264:. Archived from
1225:
1219:
1218:
1206:
1204:
1181:
1175:
1174:
1142:
1136:
1133:
1127:
1126:
1116:
1076:
1070:
1069:
1051:
1031:
1025:
1024:
1022:
1021:
1015:
1009:. Archived from
992:
983:
977:
976:
940:
934:
933:
897:
888:
887:
867:
861:
860:
858:
856:
841:
679:missions to Mars
624:bar-headed geese
607:Epaulette sharks
431:ground squirrels
339:ground squirrels
286:
279:
272:
186:
166:
165:
85:months â called
21:
3554:
3553:
3549:
3548:
3547:
3545:
3544:
3543:
3514:
3513:
3499:Wayback Machine
3483:
3471:"Hibernation".
3470:
3438:
3436:Further reading
3433:
3378:
3371:
3336:
3332:
3287:
3283:
3273:
3271:
3269:
3253:
3246:
3199:
3195:
3134:
3130:
3120:
3118:
3072:
3068:
3058:
3056:
3024:
3017:
2972:
2968:
2958:
2956:
2926:
2922:
2912:
2910:
2872:
2868:
2825:
2818:
2808:
2806:
2772:(10): 975â981.
2758:
2754:
2709:
2702:
2692:
2690:
2668:L'Anthropologie
2660:
2656:
2643:
2642:
2638:
2619:
2615:
2600:
2596:
2551:
2547:
2512:
2508:
2492:
2491:
2459:
2455:
2439:
2438:
2426:
2410:
2406:
2396:
2394:
2377:
2373:
2364:
2362:
2345:
2344:
2340:
2314:
2308:
2304:
2288:
2287:
2282:
2259:
2249:
2245:
2223:10.2307/1365104
2202:
2198:
2153:Current Biology
2145:
2141:
2094:
2090:
2075:10.2307/3872841
2059:
2055:
2002:
1998:
1983:10.2307/3872912
1963:
1959:
1910:(8): e0238132.
1896:
1892:
1877:10.2307/3872787
1861:
1857:
1826:
1822:
1799:
1795:
1782:(11): 716â722.
1772:
1768:
1733:
1729:
1682:
1678:
1631:
1627:
1604:10.1038/429825a
1580:
1576:
1521:
1517:
1478:
1474:
1435:
1431:
1421:
1419:
1415:
1376:
1370:
1366:
1327:
1323:
1284:
1280:
1271:
1269:
1226:
1222:
1202:
1200:
1198:
1182:
1178:
1143:
1139:
1134:
1130:
1077:
1073:
1049:10.2307/3503828
1032:
1028:
1019:
1017:
1013:
990:
984:
980:
941:
937:
898:
891:
868:
864:
854:
852:
843:
842:
838:
834:
819:Karolina Olsson
791:
771:
753:
732:
699:
694:
674:
668:
651:
582:
576:
560:common poorwill
538:
501:metabolic water
466:
423:
391:
316:
299:
290:
149:and the beetle
51:hibernating in
42:
35:
28:
23:
22:
15:
12:
11:
5:
3552:
3542:
3541:
3536:
3531:
3526:
3512:
3511:
3506:
3501:
3489:
3482:
3481:External links
3479:
3478:
3477:
3468:
3437:
3434:
3432:
3431:
3369:
3350:(1): 145â170.
3330:
3281:
3267:
3244:
3193:
3128:
3066:
3038:(3): 484â496.
3015:
2966:
2940:(2): 176â180.
2920:
2886:(9): 736â740.
2866:
2816:
2752:
2700:
2654:
2636:
2613:
2594:
2545:
2526:(4): 787â805.
2506:
2473:(3): 623â627.
2453:
2424:
2404:
2371:
2338:
2302:
2285:on 2008-12-17.
2270:(3): 261â266.
2243:
2217:(3): 105â109.
2196:
2139:
2088:
2053:
1996:
1957:
1890:
1871:(1): 373â380.
1855:
1820:
1793:
1766:
1747:(5): 658â661.
1727:
1676:
1625:
1574:
1515:
1488:(3): 147â155.
1472:
1429:
1364:
1321:
1294:(1): 325â330.
1278:
1240:(2): 265â268.
1220:
1196:
1176:
1137:
1128:
1071:
1026:
978:
957:10.1086/367950
951:(2): 165â179.
935:
889:
862:
835:
833:
830:
829:
828:
825:Winter cluster
822:
816:
810:
804:
798:
790:
787:
770:
767:
752:
749:
731:
728:
698:
695:
693:
690:
667:
664:
650:
647:
594:basking sharks
578:Main article:
575:
572:
537:
534:
497:muscle atrophy
465:
462:
458:hypometabolism
422:
419:
390:
387:
315:
312:
298:
295:
292:
291:
289:
288:
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263:
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206:
196:
188:
187:
179:
178:
171:
170:
79:metabolic rate
26:
9:
6:
4:
3:
2:
3551:
3540:
3537:
3535:
3532:
3530:
3527:
3525:
3522:
3521:
3519:
3510:
3507:
3505:
3502:
3500:
3496:
3493:
3490:
3488:
3485:
3484:
3474:
3469:
3465:
3461:
3457:
3453:
3449:
3445:
3440:
3439:
3427:
3423:
3418:
3413:
3408:
3403:
3399:
3395:
3391:
3387:
3383:
3376:
3374:
3365:
3361:
3357:
3353:
3349:
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2149:"Hibernation"
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1268:on 2020-09-28
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1016:on 2016-09-08
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565:
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546:Gilbert White
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3272:. Retrieved
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2769:
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2631:the original
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2391:the original
2384:
2374:
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2359:the original
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2291:cite journal
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1438:
1432:
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1413:the original
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1270:. Retrieved
1266:the original
1237:
1233:
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1208:
1203:21 September
1201:. Retrieved
1186:
1179:
1157:(1): 67â71.
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1011:the original
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994:
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901:
878:(1): 121â3.
875:
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865:
853:. Retrieved
851:. 2008-01-28
848:
839:
782:
772:
762:phylogenetic
754:
740:
733:
720:
704:
700:
683:
675:
660:
654:
652:
628:
583:
563:
549:
539:
519:
506:osteoporosis
494:
478:
454:
424:
417:hibernator.
414:
406:
399:prairie dogs
396:
392:
383:
367:
355:insectivores
347:mouse lemurs
336:
306:
302:
300:
221:Cryptobiosis
204:Hibernaculum
198:
152:Bolitotherus
150:
144:
134:
122:
114:
91:
75:heterothermy
66:
65:
49:Northern bat
2693:21 December
2608:LiveScience
2397:November 9,
2325:: 187â196.
1977:: 135â137.
1422:23 November
1001:: 467â477.
908:: 239â274.
813:Cryobiology
807:Winter rest
724:homeothermy
707:Thrinaxodon
510:amino acids
490:brown bears
486:black bears
415:facultative
307:facultative
243:Winter rest
216:Aestivation
199:Hibernation
129:black bears
118:hyperphagic
110:aestivation
67:Hibernation
3518:Categories
3268:0195036352
2787:11336/3465
2365:2006-10-06
2210:The Condor
1272:2019-12-05
1213:Seed Banks
1085:Nat Commun
1020:2017-01-06
855:12 October
832:References
670:See also:
635:amphibians
620:wood frogs
526:heart rate
473:Black bear
375:sleep debt
363:marsupials
359:monotremes
353:and other
325:gathering
108:months is
3188:202026266
3148:: 74â81.
2804:253639063
2747:228924549
2688:229399008
2442:cite book
2434:839312807
2183:0960-9822
1952:221357509
1058:0076-3519
757:arthropod
688:species.
666:In humans
598:North Sea
323:Groundhog
226:Brumation
157:Botanists
125:gestating
71:metabolic
18:Hibernate
3534:Ethology
3495:Archived
3464:14506303
3426:18320061
3386:PLOS ONE
3364:17037980
3325:30185640
3301:(1886).
3274:25 April
3239:11698659
3180:31539788
3121:25 April
3116:85053896
3059:25 April
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2809:25 April
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1171:84040488
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965:12794670
930:22397415
922:14977403
795:Dormancy
789:See also
715:Triassic
639:reptiles
612:goldfish
610:include
603:plankton
542:swallows
450:Malagasy
439:tropical
429:(namely
421:Primates
407:obligate
341:, other
303:obligate
231:Diapause
176:dormancy
137:dormancy
3417:2254195
3394:Bibcode
3316:6158517
3171:7212789
3150:Bibcode
3108:6020475
3088:Bibcode
3080:Science
3001:7480806
2900:9750966
2861:3488097
2725:Bibcode
2580:4117459
2540:9787770
2487:9307448
2231:1365104
2161:Bibcode
2125:4750243
2083:3872841
2039:6934745
2018:Bibcode
1991:3872912
1935:7451522
1912:Bibcode
1885:3872787
1815:6998737
1803:FASEB J
1694:Bibcode
1686:Science
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1620:4366123
1592:Bibcode
1560:3641607
1539:Bibcode
1409:2740905
1389:Bibcode
1381:Science
1359:8967165
1316:1125482
1308:1147024
1254:1945046
1114:3105343
1093:Bibcode
1091:: 186.
1066:3503828
1007:3873159
711:Permian
686:hominid
655:delta 2
590:hypoxia
435:primate
427:rodents
343:rodents
327:nesting
297:Mammals
174:Animal
141:brumate
94:rodents
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1848:
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1788:825685
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1759:
1720:
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1584:Nature
1567:
1557:
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1500:
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963:
928:
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801:Torpor
775:cunner
637:, and
622:, and
556:torpor
442:mammal
405:is an
401:. The
370:torpor
361:, and
331:burrow
194:Torpor
106:summer
102:torpor
83:winter
53:Norway
3524:Sleep
3230:64712
3184:S2CID
3112:S2CID
2904:S2CID
2857:S2CID
2800:S2CID
2743:S2CID
2684:S2CID
2315:(PDF)
2283:(PDF)
2260:(PDF)
2227:JSTOR
2213:. 3.
2108:: 7.
2079:JSTOR
1987:JSTOR
1948:S2CID
1881:JSTOR
1718:S2CID
1616:S2CID
1506:S2CID
1463:S2CID
1416:(PDF)
1377:(PDF)
1355:S2CID
1312:S2CID
1258:S2CID
1167:S2CID
1062:JSTOR
1014:(PDF)
1003:JSTOR
995:Ursus
991:(PDF)
969:S2CID
926:S2CID
536:Birds
481:bears
464:Bears
433:) no
413:is a
253:Sleep
98:bears
3460:PMID
3422:PMID
3360:PMID
3321:PMID
3276:2023
3263:ISBN
3235:PMID
3176:PMID
3123:2023
3104:PMID
3061:2023
3050:PMID
3006:PMID
2961:2023
2950:PMID
2915:2023
2896:PMID
2849:PMID
2811:2023
2792:PMID
2695:2020
2585:PMID
2536:PMID
2501:link
2483:PMID
2448:link
2430:OCLC
2420:ISBN
2399:2006
2297:link
2187:PMID
2179:ISSN
2130:PMID
2044:PMID
1940:PMID
1846:PMID
1811:PMID
1784:PMID
1757:PMID
1710:PMID
1667:PMID
1608:PMID
1565:PMID
1498:PMID
1455:PMID
1424:2008
1405:PMID
1347:PMID
1304:PMID
1250:PMID
1205:2019
1192:ISBN
1119:PMID
1054:ISSN
961:PMID
918:PMID
857:2019
713:and
631:fish
305:and
3452:doi
3412:PMC
3402:doi
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3311:PMC
3303:doi
3299:285
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3215:doi
3166:PMC
3158:doi
3096:doi
3084:155
3040:doi
2996:PMC
2988:doi
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2938:150
2888:doi
2841:doi
2782:hdl
2774:doi
2770:100
2733:doi
2676:doi
2575:PMC
2567:doi
2563:217
2528:doi
2524:119
2475:doi
2355:177
2327:doi
2323:248
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