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4.1.1 Summary of Section 4

The integration of physiological and molecular responses links an environmental signal to a physiological response. Heat-shock proteins (Hsps) are chaperone proteins that maintain the structure and function of proteins in cells exposed to high temperatures. The initial response of cells to high temperatures is rapid transcription of heat-shock genes, e.g. Hsp70, which is possible because Hsp70 is normally partly transcribed by RNA polymerase II. Resumption of transcription requi
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4.1 Heat-shock proteins

Molecular biology provides further insights into the biochemical and physiological responses of vertebrates to extreme temperatures and aridity in the desert environment. Animals living in hot deserts are at risk of overheating, which in turn results in denaturation of enzymes and other essential proteins. Physiologists were puzzled for a long time about how desert reptiles such as the desert iguana (Dipsosaurus dorsalis) function normally at T b = 44–46°C. Such hi
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3.5 Summary of Section 3

Behavioural mechanisms for reducing water loss are integrated with physiology. While Dipodomys rests in a cool burrow, the nasal counter-current heat exchanger cools exhaled air, conserving water vapour evaporated from respiratory surfaces. Long loops of Henle operate as counter-current multipliers, producing highly concentrated urine. Desert foxes use panting for evaporative cooling, but high rates of evaporative water loss cannot be sustained; hence the crucial importance of dens for
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3.4 Integration of anatomical features and biochemical and physiological strategies in endurers

The endurers, large animals with a relatively low surface area: volume ratio, have problems in losing heat from the body when exposed to high T a. Certain large lizard species behave like endurers, but they are evaders and evaporators too, a salutary reminder that we should not apply classification criteria too rigidly.

Dipsosaurus dorsalis, the desert iguana, lives in the Sonoran desert and is found most commonly in dry sandy areas where creosote bushes grow (
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3.3 Integration of anatomical features and biochemical and physiological strategies in evaporators

Birds and larger desert mammals that use evaporative cooling risk dehydration because of the difficulty of finding sufficient drinking water. For mammals, evaporative heat loss includes panting and sweating.

In small mammals and birds the temperature of exhaled air is often lower than T b, resulting in condensation of water on the nasal mucosa. Small desert mammals rely on this mechanism for water conservation, while resting in their cool burrows during the heat of the
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2.5.1 Summary of Section 2

Desert animals are classified in terms of their body size and physiology into three groups: evaders, evaporators and endurers. The logic for this classification is that the smaller the animal, the larger its surface area to volume ratio. Small animals therefore gain and lose heat faster than large animals, warming rapidly when exposed to intense solar radiation, and cooling rapidly at night. Small endothermic evaders, e.g. kangaroo rats, rest in cool microenvironments, e.g. shade or burrows,
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2.5 Behavioural strategies of endurers

Endurers are defined as large desert mammals such as oryx and camel, and large desert birds, ostrich and emu. The term ‘endurers’ suggests that these animals are forced to endure the extreme conditions of the desert climate because they cannot shelter from high T a and intense solar radiation during the day or low T a at night, as they are too large to hide in burrows or dens. Nevertheless, in spite of their size, endurers do take advantage of aspects o
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2.2 How animals interact with the environment is affected by their body size

Willmer et al. (2000) classify desert animals in terms of the range of body sizes and the rate of evaporation (Figure 8).

Figure 8
Willmer, P., Stone, G. and Johnston, I. (2000) Environmental Physiology of Animals
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6.6.1 Melatonin

Syrian hamsters, which display pronounced circadian temperature fluctuations before hibernation, lose these circadian cycles on entry to hibernation, and start to regain them shortly before arousal. Cycles are distorted during the early recovery period, suggesting that the SON oscillator has either been switched off or de-synchronized in hibernation. Another monoamine, melatonin, is involved in making these adjustments. A hormone rather than a neurotransmitter, melatonin is secreted by the pi
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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
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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
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5.6 The importance of size and habitat

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
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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
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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
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4.3 Cellular changes

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
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3.7 Summary

The physiological details of deep or seasonal hibernation vary widely between species. However, the general pattern is similar, involving controlled entry to torpor, with or without ‘test drops’, and periodic arousals. The intervals between these arousals depend on size, T b and other factors. The frequency of the arousals falls off during the deepest part of the hibernation. Entry to hibernation may be triggered by temperature, daylength and shortage of food, especially
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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
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3.5 Arousal (continued)

Question 9

What alternatives to shivering might act as a source of heat?

Answer

BMR is maintained mainly by a number of tissues with high metabolic activity. One of
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3.4.2 Periodic arousal

All mammalian hibernators arouse periodically. The frequency of the arousal and the length of the euthermic periods between bouts of hibernation vary widely with species, among individuals, and with the time of year (e.g. in deep hibernators, the larger species seem to have longer periods of wakefulness than the smaller ones). The arctic marmot (Marmosa caligata), whose heart rate recording is shown in Author(s): The Open University

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
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