Introduction This unit is the first in a series of three on Animals at the extreme. It is concerned with the integration of behaviour anatomy, physiology and biochemistry in diverse vertebrates that live in deserts. Once you have completed this unit, you will be all the more able to appreciate the linked units that follow, Animals at the extreme: hibernation and torpor and Animals at the extreme: the polar environment. These units build on and develop some of the science you will stud
6.3 Metabolic regulation and the midbrain As you found in the last section, the physiological evidence points to the likelihood that different components of regulation may be regulated separately. The hypothalamus, which appears to be central to the depression and recovery of body temperature during entry to torpor and arousal, is not the only player in the control of metabolic processes underlying non-behavioural thermogenesis. In many respects, the initiation of thermogenesis is the prime event in the reactivation of a cold body: t
6.2 The hypothalamus as central regulator Research in the past 30–40 years has established that the hypothalamus, which lies below the thalamus and above the optic nerve chiasma and the pituitary gland in the brain, fulfils all of the functions listed above, at least in part. The main function of the hypothalamus is homeostasis. Factors such as blood pressure, body temperature, fluid and electrolyte balance, and body weight are held to constant values called the set-points. Although set-points can vary over time, from day to
5.7 Summary BMR is regulated independently of T
b at least in hibernating mammals. Entry into hibernation is characterized by a gradual fall in RQ, which indicates a switch from carbohydrate to lipid metabolism for energy provision (through the phosphorylation of pyruvate dehydrogenase, the inhibitor of mitochondrial fatty acid uptake). There is evidence that some other vertebrates, such as hibernating frogs, may continue to use carbohydrate catabolism or activate gluconeogenic pathways
5.5 Energy budgeting – the benefits of hibernation and torpor Studies performed on ground squirrels in the wild and in the laboratory have allowed estimates to be made of energy expenditure in hibernating and euthermic animals over similar periods (Wang, 1987). The average time spent by Richardson's ground squirrel in a periodic arousal in the wild is about 10 hours and the frequency of arousal decreases during November-March, when animals are spending more than 90% of their time in torpor. Monthly total oxygen consumption in January is about 35% of tha
5.4 Inspiratory drive 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
5.3 Mitochondrial adaptations During the winter months, whilst hibernating vertebrates maintain a very low metabolic rate, major reorganization of mitochondrial metabolism occurs. The phenomenon has been studied in some detail in frogs which, although not hibernators in the true sense, can endure very low water temperatures under the conditions of profound hypoxia that exist when they lie dormant for long periods below the surface. In contrast to normoxic conditions, the muscle mitochondria of dormant frogs depress their
4.4 Cell survival mechanisms Physical damage is not the only danger that faces cells recovering from low temperatures in the absence of oxygen (due to a 90% drop in blood flow to the brain) and energy supplies. A universal sign of recovery from such conditions is the production of reactive oxygen species (ROS) (Box 4). The electron transfer chain that participates in the formation of water from oxygen in mitochondrial respiration can also be used in the production of the free radical superoxide, sometimes called ‘singl
4.2 Arresting protein synthesis The regulation of T
b in hibernators has traditionally been viewed as the fundamental physiological process in hibernation. But recently, questions have been raised about whether thermal changes initiate or simply accompany metabolic depression. Is the metabolic inactivity of animal tissues during bouts of torpor or in hibernation, the cause or the result of hypothermia? A common-sense view is that temperature directly influences metabolism by regulating enzyme activity. Evi
3.6 Length of torpor bouts in hibernation It is obvious that there is a very high energetic cost to arousal, and an even higher one to the periods of euthermic wakefulness prior to re-entering torpor. If an animal could simply enter torpor once, and arouse 2, 4 or 6 months later, depending on the environment, it would represent a huge energy saving. Thus, it has been assumed that either prolonged torpor is physiologically impossible, or there is some strong selective value to the species in regular arousal. In the case of some small
2.3 Hibernators as eutherms 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
2.2 Species showing torpor or deep hibernation 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
2.1 Degrees of torpor 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. 5.4.1 Summary of Section 5 Several anatomical and biochemical adaptations to living in very cold water have evolved in polar fish, particularly those of the southern oceans, which have evolved in isolation for many millions of years. Cold, turbulent water is rich in oxygen. One family of fairly large fish lacks blood pigments but its blood is less viscous and it has additional respiratory surfaces. Many fish have cryoprotectants in the blood and other body fluids, and the muscles of some contain numerous mitochondria a 5.4 Fatty acids as indicators of diet Although polar fish and invertebrates are difficult to study alive for the reasons just described, some information about their diet and habits can be obtained from analysis of the lipid composition of their tissues. At high latitudes, the supply of most kinds of marine food changes with the seasons, just as it does on land, and many fish eat little or nothing for long periods, living off their reserves of triacylglycerols. Lipids are major fuels for polar fish and, although many fish have li 4.3 Humans in polar regions Humans evolved in tropical Africa and gradually colonized colder climates during the Pleistocene ice ages. There have been permanent populations in the Arctic for several thousand years, mostly Inuit (Eskimos) in what are now Canada, Alaska and Greenland, and several groups in northern Europe and Russia, such as the Saami (Lapp) in Scandinavia and the Chukchi in Siberia. Such people do not grow crops and keep only a few domestic animals, mostly for transport (e.g. husky dogs or reindeer), not 3.5 The structure of adipose tissue Since food is only available seasonally or intermittently at high latitudes, many arctic birds and mammals, including polar bears, Svalbard reindeer, arctic foxes, seals and walruses, naturally accumulate large stores of fat. The quantity of energy stored and the metabolic control of its use are finely adjusted to the habits and habitat of the species. This section is concerned with the cellular structure and anatomical organization of adipose tissue in such naturally obese species. Most labo 3.3.1 Dormancy in black and brown bears The dormant state of bears differs from true hibernation in that the body temperature does not fall below 31–35° C and a major disturbance (such as an intruding biologist) can arouse them to full activity in a few minutes. Dormant bears do not eat, drink, urinate or defaecate, the heart rate drops from 50–60 beats min−1 to 8–12 beats min−1, and oxygen consumption is only 32% of that of actively foraging bears. Nonetheless, the rate of protein turnover, as mea 3.1 Introduction It is clear from Sections 1 and 2 that seasonal or irregular periods of fasting are an integral part of living at high latitudes, especially for large animals. When people (and many tropical and temperate-zone mammals) lose weight, either because they are eating less or because they are suffering from a digestive or metabolic disorder, protein is broken down in substantial quantities long before the lipid stores are exhausted. Even frequent and vigorous exercise cannot prevent the breakdown o 2.4.1 Summary of Sections 1 and 2 Large seasonal changes in temperature and sunlight dominate primary plant production and hence the food supply. Food intake is regulated by the endogenous seasonal control of appetite, fattening and activity, as well as by food availability. Energetically demanding activities such as breeding and migration are only feasible during a brief period and must be tightly synchronized to season. Greater accessibility of food suitable for chicks makes long-distance migration to and from high arctic r
Table 1 Groups o
