define and use, or recognize definitions and applications of, each of the bold terms;

give definitions of the terms ‘hibernation’, ‘torpor’ and ‘adaptive hypothermia’, and the three physiological processes that underlie them;

give examples of the diversity of the major groups of mammals and birds that contain hibernating species;

describe the physiological changes occur]]> By the end of this unit you should be able to:

• define and use, or recognize definitions and applications of, each of the bold terms;

• give definitions of the terms ‘hibernation’, ‘torpor’ and ‘adaptive hypothermia’, and the three physiological processes that underlie them;

• give examples of the diversity of the major groups of mammals and birds that contain hibernating species;

• describe the physiological changes occur]]> This unit examines hibernation, a special form of adaptation that animals can make to the ecological demands of remaining in a chosen habitat in winter. Hibernation is a state which enables energy-efficient survival when ambient temperatures are so low that foraging or simply maintaining normal core body temperature and basal metabolic rate are either energetically too costly or impossible.

  Polar endotherms can maintain a high T _b even when living actively at sub-zero]]> This unit examines hibernation, a special form of adaptation that animals can make to the ecological demands of remaining in a chosen habitat in winter. Hibernation is a state which enables energy-efficient survival when ambient temperatures are so low that foraging or simply maintaining normal core body temperature and basal metabolic rate are either energetically too costly or impossible.

  Polar endotherms can maintain a high T _b even when living actively at sub-zero]]> Adaptive hypothermia occurs in at least six distantly related mammalian orders (Table 1) and in several orders of birds. There is a spectrum running from those species which can tolerate a drop in T _b by 2° C for a few hours, to the seasonal deep hibernators which maintain a T _b as low as 4° C for weeks on end.

  Table 1 Groups o]]> Adaptive hypothermia occurs in at least six distantly related mammalian orders (Table 1) and in several orders of birds. There is a spectrum running from those species which can tolerate a drop in T _b by 2° C for a few hours, to the seasonal deep hibernators which maintain a T _b as low as 4° C for weeks on end.

  Table 1 Groups o]]> Among the birds, torpor occurs in a number of species in the orders Apodiformes (hummingbirds and swifts), Caprimulgiformes (nightjars, nighthawks, goatsuckers and poor wills) and Coliiformes (mousebirds). In all of the hummingbirds (family Trochilidae) studied to date, torpor, if it occurs, takes place on a daily (or more usually nightly) basis. They are able to re-warm themselves independently of T _a and show an increased thermogenesis if T _a falls below ]]> Among the birds, torpor occurs in a number of species in the orders Apodiformes (hummingbirds and swifts), Caprimulgiformes (nightjars, nighthawks, goatsuckers and poor wills) and Coliiformes (mousebirds). In all of the hummingbirds (family Trochilidae) studied to date, torpor, if it occurs, takes place on a daily (or more usually nightly) basis. They are able to re-warm themselves independently of T _a and show an increased thermogenesis if T _a falls below ]]> Hibernating endotherms are not the easiest animals to study. Thus, until the late 1960s many biologists believed that mammalian hibernation was a process in which thermoregulation was simply ‘switched off’, following the receipt of a set of ‘cues’. These cues included a declining T _a, a shortening daylength, the extent of body fat and a lack of food etc. With this model, the hibernator essentially becomes an ectotherm whose T _b follows the T ]]> Hibernating endotherms are not the easiest animals to study. Thus, until the late 1960s many biologists believed that mammalian hibernation was a process in which thermoregulation was simply ‘switched off’, following the receipt of a set of ‘cues’. These cues included a declining T _a, a shortening daylength, the extent of body fat and a lack of food etc. With this model, the hibernator essentially becomes an ectotherm whose T _b follows the T ]]> Emergence can be viewed as the final step in the series of periodic arousals. Instead of re-entering hibernation, the animal maintains the euthermic condition. The cue for maintaining this final arousal is probably not temperature, as some species emerge when T _a is well below zero. It is also difficult to see how arousal could be affected by daylength, since the hibernating animal is usually underground in a cavity or a burrow. Perhaps fat or food stores reach a minimum lev]]> Emergence can be viewed as the final step in the series of periodic arousals. Instead of re-entering hibernation, the animal maintains the euthermic condition. The cue for maintaining this final arousal is probably not temperature, as some species emerge when T _a is well below zero. It is also difficult to see how arousal could be affected by daylength, since the hibernating animal is usually underground in a cavity or a burrow. Perhaps fat or food stores reach a minimum lev]]> Even after many years of research, the phenomenon of hibernation continues to be a mystery to scientists. Despite coming nearer to an understanding of how and why it happens, some fundamental questions remain unanswered. Is there a genetic basis underlying the evolutionary predisposition of animals to hibernate, given its occurrence in many groups of vertebrates and invertebrates? Is the problem of metabolic adaptation in cells separate from thermal regulation which occurs throughout the orga]]> Even after many years of research, the phenomenon of hibernation continues to be a mystery to scientists. Despite coming nearer to an understanding of how and why it happens, some fundamental questions remain unanswered. Is there a genetic basis underlying the evolutionary predisposition of animals to hibernate, given its occurrence in many groups of vertebrates and invertebrates? Is the problem of metabolic adaptation in cells separate from thermal regulation which occurs throughout the orga]]> Hibernation can result in the deposition of fat in adipose tissue. In tissues of finite size which are important sources of energy and sites for fuel metabolism, changes in cell structure (redistribution of organelles involved in energy metabolism and protein synthesis) are the most likely adaptation to a state of torpor. Liver hepatocytes of the hibernating dormouse (Muscardinus avellanarius), are visibly different from those of arousing and euthermic dormice when viewed in thin secti]]> Hibernation can result in the deposition of fat in adipose tissue. In tissues of finite size which are important sources of energy and sites for fuel metabolism, changes in cell structure (redistribution of organelles involved in energy metabolism and protein synthesis) are the most likely adaptation to a state of torpor. Liver hepatocytes of the hibernating dormouse (Muscardinus avellanarius), are visibly different from those of arousing and euthermic dormice when viewed in thin secti]]> The supply of oxygen to tissues such as the heart, liver and WAT is, under euthermic conditions, invariably linked to and dependent upon local blood flow and pulmonary function. However, as we have already seen, under conditions in which blood flow is reduced to a trickle, the control of energy supply switches to local adaptations in the capillaries and tissue cells, including the oxygen affinity of erythrocyte haemoglobin, the supply and metabolism of respiratory fuels and the rate of protei]]> The supply of oxygen to tissues such as the heart, liver and WAT is, under euthermic conditions, invariably linked to and dependent upon local blood flow and pulmonary function. However, as we have already seen, under conditions in which blood flow is reduced to a trickle, the control of energy supply switches to local adaptations in the capillaries and tissue cells, including the oxygen affinity of erythrocyte haemoglobin, the supply and metabolism of respiratory fuels and the rate of protei]]> The use of hibernation to gain energetic advantage must be weighed against a number of considerations, particularly animal size and behaviour, biogeographic distribution and habitat. Small animals, which can carry less fat and have a higher surface area to volume ratio and BMR, are more likely to lose energy as heat and in maintaining life functions if they do not use hypothermic strategies in winter. Few hibernating mammals have a total body mass greater than 5 kg. Indeed, in large animals t]]> The use of hibernation to gain energetic advantage must be weighed against a number of considerations, particularly animal size and behaviour, biogeographic distribution and habitat. Small animals, which can carry less fat and have a higher surface area to volume ratio and BMR, are more likely to lose energy as heat and in maintaining life functions if they do not use hypothermic strategies in winter. Few hibernating mammals have a total body mass greater than 5 kg. Indeed, in large animals t]]> Measurements of thermoregulation, respiration and metabolic depression in the edible dormouse (Myoxus glis) during the early stages of torpor, hibernation and aestivation, indicate remarkable similarities in the profile of physiological changes for all three adaptive phenomena, suggesting that they are controlled by essentially the same mechanism. The capacity for adaptive hypothermia in animals is clearly determined genetically and is manifested in cells from many different tissues. N]]> Measurements of thermoregulation, respiration and metabolic depression in the edible dormouse (Myoxus glis) during the early stages of torpor, hibernation and aestivation, indicate remarkable similarities in the profile of physiological changes for all three adaptive phenomena, suggesting that they are controlled by essentially the same mechanism. The capacity for adaptive hypothermia in animals is clearly determined genetically and is manifested in cells from many different tissues. N]]> Reactive changes in the brain are usually marked by changes in neuronal electrical activity. If these changes are to be of long duration, adjustments in neuronal electrical behaviour may be made through changes in gene expression. Rapid-response genes (sometimes called ‘immediate-early’ genes) are activated within minutes of the onset of such sustained electrical activity. These genes are master controls, acting as a gateway to a series of linked events: alteration of electrical firing pa]]> Reactive changes in the brain are usually marked by changes in neuronal electrical activity. If these changes are to be of long duration, adjustments in neuronal electrical behaviour may be made through changes in gene expression. Rapid-response genes (sometimes called ‘immediate-early’ genes) are activated within minutes of the onset of such sustained electrical activity. These genes are master controls, acting as a gateway to a series of linked events: alteration of electrical firing pa]]> If you have visited a desert you will have noticed the sparse plant cover, or in certain sandy deserts, the almost complete absence of plant life. The low productivity of deserts derives from their defining feature, which is aridity. Scarcity of water restricts the diversity and amount of plant cover, and in turn the diversity and abundance of animals. However, if you were visiting one of the American deserts after rains, you would be rewarded by the sight of the desert ‘in bloom’, as vas]]> If you have visited a desert you will have noticed the sparse plant cover, or in certain sandy deserts, the almost complete absence of plant life. The low productivity of deserts derives from their defining feature, which is aridity. Scarcity of water restricts the diversity and amount of plant cover, and in turn the diversity and abundance of animals. However, if you were visiting one of the American deserts after rains, you would be rewarded by the sight of the desert ‘in bloom’, as vas]]> The unique climate and topography of each desert links to the unique and characteristic flora and fauna found there. From the brief description of deserts provided in Section 1, you can appreciate that a desert provides a variety of niches for animals and plants. The term ‘niche’ applied to animals describes its role in a particular environment, and includes a number of characteristics such as habitat range, how the animal feeds, its diet, its environmental requirements and also its preda]]> The unique climate and topography of each desert links to the unique and characteristic flora and fauna found there. From the brief description of deserts provided in Section 1, you can appreciate that a desert provides a variety of niches for animals and plants. The term ‘niche’ applied to animals describes its role in a particular environment, and includes a number of characteristics such as habitat range, how the animal feeds, its diet, its environmental requirements and also its preda]]>