temperature regulation The human body is a heat-generating object. Even at complete rest at a comfortable temperature, the vital functions of the body generate heat. When it is at a minimum this is called basal metabolic heat. Heat production rises with activity: the more vigorous the activity the greater the quantity of heat produced. Excess heat has to be lost. Considering this variation in internal heat production, together with the wide variety of thermal environments inhabited by humans, it is remarkable that, except during exercise or fever, the body temperature can be kept within such narrow limits. This is achieved partly by the physiological mechanisms that control the rate of heat loss and heat production to achieve a balance. If the rate of heat loss exceeds the rate of heat production, the body temperature will fall, leading ultimately to hypothermia. Conversely, if the rate of heat loss is less than the rate of heat production, or is defected by external heating the body temperature will rise, leading eventually to heatstroke. Death may be the end result of either.

Body temperature is controlled through a central mechanism in the brain which, while it acts in a manner similar to a thermostat, is not a simple on/off device but is more akin to a ‘black box’ with a complex system of neurons cross-linking the sensory input and the effector output. The thermostat is activated by impulses from central receptors, which detect changes in the temperature of the blood, and from peripheral receptors mainly in the skin. The thermostat regulates the temperature of the body by adjusting heat production and heat loss, but the setting of the thermostat itself may be altered. During sleep the cerebral thermostat is reset to a new low level, skin blood vessels relax, causing an immediate rise in skin temperature and heat loss, and the metabolic rate is reduced.

There are racial variations in the response to heat and cold, and at the extremes of age there is an increased risk of hypothermia and heatstroke. Many medical disorders and a range of drugs predispose to temperature disorders.

Heat is lost, and gained, through convection, radiation, and conduction, while evaporation contributes only to heat loss. Even for humans, the standard thermodynamic laws of physics apply. The rate at which the body loses heat depends on the temperature difference between the skin and the environment. In the cold, therefore, the aim is to reduce the skin temperature as much as possible, whereas in the heat the skin temperature should be as high as possible.

Conductive

heat transfer depends on the surface area in contact with a substance, the temperature of the substance, and its thermal conductivity. Air is a poor conductor of heat, and therefore most insulating systems, including clothing, act by trapping air. Conductive transfer is high in water. Wet clothing not only has decreased insulating properties, but it rapidly conducts heat from the body to the colder surface of the clothing. In an emergency in the cold, conductive heat loss may be forgotten, with the casualty being well covered with blankets but continuing to lie with nothing between him and the ground.

Convective

heat loss is increased by wind, hence the ‘wind chill’ index, which gives the total heat loss under any particular combination of cold and wind speed, and relates it to the still air temperature which would produce the same rate of heat loss. The rate of heat loss at –10°C in still air is almost exactly the same as that at +10°C with a 10 mph wind. Convective heat loss is also increased by movement, not only by the person physically moving forward but also through the ‘pendulum’ movement of the limbs, and through the clothing producing a bellows effect and blowing warm air out to be replaced by cold air, or water, at the next movement. Increasing convective loss is dangerous in the cold but beneficial in the heat.

Radiant

heat exchange is often overlooked. Standing near a glacier causes a feeling of chill even though there is no contact and no wind. Similarly, people may gain heat radiating from hot walls, concrete, or sand in a hot environment, as well as from fires or central heating radiators in the cold.

Evaporation

is an effective way of losing heat because of the high latent heat of evaporation. However, the rate of evaporation depends on the surface area of fluid and its vapour pressure immediately above the surface. Evaporative heat loss is greatest when the air is dry and the moisture vapour is constantly being removed from the body surface, and least when there is high humidity and no air movement. Any factor which increases convective heat loss also increases evaporative heat loss. In man evaporative heat loss occurs through sweating and when clothing is wet. Wet clothing is used to reduce the temperature in heatstroke.

In animals with lungs, heat is also lost by warming and humidifying the inspired air during breathing, with the greater proportion being through humidification. Even in a mist in winter, with an ambient relative humidity of 100%, when the inspired air is warmed to core body temperature the relative humidity falls and the body expends heat in raising the relative humidity to 100% before it reaches the alveolar surfaces of the lung. Although some of the heat and moisture is retrieved from the air on its way out again, in temperate conditions 10–15% of the total heat loss from the body is from the respiratory tract, and the proportion increases in colder or drier air.

In the cold

The body responds by constriction of the superficial blood vessels, mainly via the sympathetic nervous system but also through direct action of cold on the blood vessels. Vasoconstriction is very effective in reducing heat loss by limiting blood flow to the periphery, thus increasing the depth of ‘shell’ insulation and reducing the temperature differential between the skin and the environment (see Figure). In fact vasoconstriction can result in the outermost inch of the body having a thermal conductivity equivalent to that of cork. To reduce the surface area exposed to cold the person may adopt a ball-like fetal position. There is also a counter-current exchange of heat between the arteries and veins in the distal half of the limbs. Below a limb temperature of 10–12°C the peripheral vasoconstriction fails and alternating vasodilatation and vasoconstriction occurs, though there may actually be very little increase in the volume of blood circulating in the skin during the vasodilatation, which would preserve the insulating effect of the vasoconstriction. Though there is some vasoconstriction in the face, it is minimal in the scalp, and this plus the lack of tissue depth makes the regulation of heat loss from the head inefficient; the rate of heat loss by this route increases in a linear manner between ambient temperatures of +32°C and –20°C, and indeed may be equal to half the total body heat production at rest at –4°C. Mental stress even of as mild a degree as mental arithmetic increases this heat loss, as also do nausea, vomiting, fainting, trauma, and haemorrhage.

Heat production rises with muscle activity, either deliberate activity (10–15-fold during hard physical exercise) or shivering. In severe cold stress catecholamine secretion increases and stimulates increased heat production. The less fit a person is, the less the degree of cold stress at which the catecholamine ‘overdrive’ occurs. Any increase in heat production is always accompanied by a rise in oxygen consumption: in shivering it may double or treble. Activity and shivering are not economical in thermoregulation, because they are accompanied by an increased blood supply to the muscles and this in turn raises the surface temperature and increases heat loss. In fact only 48% of the extra heat generated is retained in the body.

The greatest rate of heat production which an individual can achieve depends on his maximal ability for muscular work, linked to his maximal rate of oxygen usage. This in turn depends on the supply of oxygen and therefore on the greatest rate at which the heart can pump blood around the body, and on the efficiency with which the muscles can utilize the oxygen. These all increase with improving fitness. For any level of exercise the oxygen consumption and cardiac output need to be higher in a cold environment than in a warm one; this explains, for example, why angina may develop during a particular level of activity in the cold but not at normal temperatures. Also, during sleep in the cold, unfit people are repeatedly awakened by shivering, whereas the greater heat generating ability of ‘fit’ muscles allows sleep undisturbed by shivering.

In conditions of very severe cold, if the person has to undertake very vigorous exercise, the maximal oxygen uptake may be insufficient to provide for the high demand of both the exercise and the severe cold stress, and a person can develop unexpected and unsuspected hypothermia despite vigorous muscular activity. If hypoxia is also present, as at high altitude, this decreases the total possible oxygen uptake, and therefore heat production and shivering may be inhibited. Finally, if the person is exhausted or suffering from malnutrition he cannot increase heat production because of the lack of substrate (fuel) for metabolism.

Alcohol produces a number of effects which increase the risk of hypothermia; the greatest danger is decreasing the awareness of cold and increasing bravado, while impairing the ability to assess risks.

In the heat

The skin and superficial blood vessels open up and the skin temperature rises, thus increasing the possibility of heat loss. Sweating increases and, in a dry, hot environment, evaporation can become the major means of losing heat. Unfortunately in a hot and humid environment there is less scope for evaporation, and sweating becomes less effective. The person tends to adopt an ‘open’ posture e.g. lying spread-eagled to maximize the surface area available. This is usually done for suntanning but also works for heat loss. By contrast with the cold, there is little scope for adjusting the production of heat except by reducing activity.

In fever the thermostat in the brain is set at a higher level. Therefore the person shivers to raise the temperature to the new, higher setting, and, when the fever breaks and the thermostat setting drops back to normal, the person sweats to dispose of the (now) excess heat.

Clothing and shelter

Clothes are the obvious protection from the cold, especially those that trap air. It is more efficient to have several thin layers than one thick. Cold weather clothing effectively reduces the temperature of the outermost surface and therefore reduces heat loss. A windproof layer stops or reduces convective loss. In the heat, clothing should be thin and loose to cause the least barrier to heat loss by conduction and convection. White or light-coloured clothing reduces the heat load by reflecting radiant heat.

Shelter may take many forms, from simply sheltering behind a boulder or digging a snow hole to the high technology of modern housing. In cold climates houses tend to have wooden floors with carpets, fires or central heating, attic insulation, and double glazing, and everything is done to reduce draughts. In a hot climate many houses have stone floors, the windows and doors are placed opposite each other to encourage through draughts, and the buildings are often painted white to reflect radiant heat from the sun.

Comfort

Humans are very sensitive to differences and changes in temperature, especially changes downwards. The sensation of cold is related to the lowered average skin temperature, and the sensation of local cold becomes more intense with increasing duration of exposure. A hot or cold stimulus is considered pleasant when it tends to restore temperature towards normal, and unpleasant if it has the reverse tendency. Thus cold applied to the skin is pleasant if the core temperature is raised but unpleasant if the core temperature is lowered. A person may feel warm on moving from the cold into the warmth, even if the body temperature is still low; this suggests that the discomfort caused by cold may be the result of the superficial vasoconstriction. Subjective discomfort is greater if certain areas, such as the forehead or feet, are cold, and the discomfort is increased by shivering. Because cold is a common human experience, the fact that it can cause injury, illness, and death is often overlooked.

Symptomless cooling

The central thermostat responds to the rate of firing of temperature-sensitive neurons, and the rate of firing is partially dependent on the rate of change of the stimulus. If the peripheral and central temperatures change slowly enough, the body may fail to activate the mechanisms for reducing heat loss and increasing heat production. During experimental cooling (in a water bath) it is possible to abolish shivering and the sensation of cold without interrupting the progressive reduction in body temperature, by slightly raising the temperature of the water, and therefore of the skin, at the onset of shivering; the perceived warmth is increased although the water temperature remains cold enough to cause continued cooling.

Perceived warmth has practical and clinical implications:(i) Deep divers have hot water flowing over the surface of the body, which gives a sensation of warmth whilst it may not be enough to balance the large quantities of heat they lose through breathing cold oxyhelium. They may therefore become hypothermic while feeling warm.(ii) Coming out of a refrigerated room at –30°C into a cold room at 0°C gives a feeling of warmth and comfort but the risk of hypothermia is still present, especially if the person is moving frequently between the two chambers.(iii) For the elderly a single bar fire in a large cold room gives a sensation of warmth which prevents vasoconstriction and therefore allows an increased heat loss for which the heat from the small fire cannot compensate; so they can become hypothermic without being aware of discomfort. Inadequate heating may therefore be more dangerous than no heating.

It is salutary to realize that humans can live in and explore very hostile areas of the world, not because of the physiology of thermoregulation, but because of man's technological ability to manipulate his thermal environment by the use of clothing, heating, and shelter. The proof of this is the fact that the temperature of the skin of the abdominal wall is the same in the Arctic as in the tropics.

Evan L. Lloyd

Bibliography

Lloyd, E. L. (1986). Hypothermia and cold stress. Croom Helm, London.
Maclean, D. and and Emslie-Smith, D. (1977). Accidental hypothermia. Blackwell, Oxford.

Temperature Regulation

Externally heated animals

Internally heated animals

Resources

Temperature regulation, or thermoregulation, is the ability of an organism to maintain its body temperature when other temperatures surround it. An organism maintains homeostasis, a steady internal state, only if its body temperature stays within prescribed limits. Cellular activities require an optimum amount of heat.

They depend on enzyme action, and enzymes function within a narrow range of temperature. For this reason, living things can only tolerate a limited rise or drop in temperature from the optimum. Mechanisms exist that regulate body temperature to stay within its survival limits.

Cells require a source of heat for their activities. The heat source can be external or internal. The sun is an external source of heat for living things. Cellular metabolism is an internal heat source. During cellular respiration, the chemical energy in food is converted to high-energy phosphate groups in adenosine triphosphate, ATP. In the process, cells lose about 60% of the released energy as heat. Plants and most animals give up this heat to the environment. Birds, mammals, and some fish, however, make use of some of the heat that metabolism yields.

Organisms exchange heat with the environment by four major physical processes. First, there is conduction, the direct transfer of heat from a warmer to a cooler object. Humans cool off during a swim because the heat of the body is conducted to the cool water. During another process called convection, a breeze passes over a surface and brings about heat loss. This is why a fan cools people on a hot day. Radiation, the emission of electromagnetic waves, transfers heat between objects that are not in direct contact. Animals absorb heat that the sun radiates. Finally, evaporation, the physical change of liquid to gas removes heat from a surface. Sweating and panting are cooling processes. Perspiration evaporates from the skin, and panting increases evaporation from the respiratory system. Both procedures take heat away from the body.

Externally heated animals

Ectotherms are animals that warm their bodies by absorbing heat from their surroundings. In most ectotherms, the body temperature fluctuates with changes in the surrounding temperature. The body temperature of snakes, for example, cools in cold weather and warms up in hot weather. Most marine fish and invertebrates, however, live in water that stays the same temperature. Their body temperature, therefore, does not change. In everyday language, scientists say that these animals are cold-blooded. Amphibians, reptiles, most fish, and invertebrates are considered cold-blooded. This term is misleading, however, because many lizards have high body temperatures. External temperature plays a major role in the activity rate of ectothermic animals. When the weather is warm, they become active. They slow down when the temperature drops.

Certain ectotherm behaviors help regulate body temperature. To warm up, reptiles find sunny places, and stretch out for maximum exposure. If it gets too warm, lizards alternate between sun and shade. Amphibians warm up by moving into the sun or diving into warm water. They cool off by entering the shade. In cold weather, honeybees huddle together to retain heat. Bees and large moths build up heat before takeoff by contracting their flight muscles without moving their wings.

In addition to behaviors, physiological adaptations help ectotherms regulate temperature. Diving reptiles conserve heat because their blood circulates inward toward the body core during a dive. The skin of bullfrogs secretes more mucus when it is hot, allowing more cooling by evaporation. Many ectotherms exist at a lower temperature during torpor, a state of slowed metabolism. This helps them survive a food shortage. If the food supply increases, they come out of torpor in a few hours.

KEY TERMS

Cooling center —Thermoregulating area of the hypothalamus whose impulses result in temperature-lowering activities of various parts of the body.

Ectotherm —A cold-blooded animal, whose internal body temperature is similar to that of its environment. Ectotherms produce little body heat, and are dependent on external sources (such as the sun) to keep their body temperature high enough to function efficiently.

Endotherm —An animal that uses its metabolism as a primary source of body heat and uses physiological mechanisms to hold its body temperature nearly constant.

Heating center —Thermoregulating area of the hypothalamus whose impulses result in temperature-raising activities of various parts of the body.

Hibernation —A type of torpor in which an animal’s body temperature drops, thereby enabling it to withstand prolonged cold and little food.

Internally heated animals

Endotherms are animals that warm their bodies mainly from their own metabolism. Mostly birds and mammals, they maintain a constant body temperature regardless of changes in the surrounding temperature. Scientists commonly call them warm-blooded, because the internal temperature they generate is usually higher than the environmental temperature. Endotherms have certain advantages. Their consistent higher temperature allows them to be active at all times. It also gives them the freedom to live in varied habitats. It helps them survive on land, where the air temperature changes more drastically than water temperature.

Although most fish are ectothermic, mackerals, tuna, swordfish, marlins, and some sharks are endotherms; although in swordfish and some mackeral, warming occurs only in the central nervous system and the retina of the eye. Endothermic fish are more closely related to their ectothermic relatives than they are to each other. Thus, endothermy evolved independently in these different groups, as an adaptation that allowed the animals to expand their habitats into waters of varying temperatures.

Certain adaptations accompany the maintenance of a constant body temperature. A high metabolic rate helps supply heat. An efficient circulatory system conducts and distributes heat around the body. Fat layers, fur, and feathers insulate the body and retain heat. Shivering muscles contract to increase body heat, and hormones such as epinephrine and thyroxin increase the rate of metabolism. Under unfavorable conditions some endotherms experience torpor, for example, hibernation, in which the body temperature drops. Hibernation enables animals to survive long periods of cold and lack of food.

Receptors in various parts of the body, such as the skin, deliver information to the hypothalamus of the brain about the body’s temperature. The hypothalamus acts as a thermostat. It contains two thermoregulating areas that stimulate nerve impulses. One area, the heating center, sends out impulses that raise the body temperature. It causes blood vessels near the surface of the body to contract, thus preventing heat loss. It also causes fur to stand erect and become more insulating. It causes shivering by muscles that produce heat when they contract, and it stimulates hormonal production. The other area, the cooling center, sends out impulses that bring about a temperature drop. It causes surface blood vessels to expand thereby releasing heat. It brings about sweating or panting.

Resources

BOOKS

Burshtein, A.I. Introduction to Thermodynamics and Kinetic Theory of Matter. Weinheim, Germany: Wiley-VCH, 2005.

Germann, William J. Principles of Human Physiology. San Francisco, CA: Pearson Benjamin Cummings, 2005.

Hall, John E., ed. Guyton & Hall Physiology Review and Problems of Human Anatomy and Physiology. Philadelphia, PA: Elsevier Saunders, 2006.

Roy, Bimalendu Naravan. Fundamentals of Classical and Statistical Thermodynamics. West Sussex, UK, and New York: John Wiley, 2002.

Turns, Stephen R. Thermodynamics: Concepts and Applications. New York: Cambridge University Press, 2006.

Van De Graaff, Kent M., and R. Ward Rhees, eds. Human Anatomy and Physiology: Based on Schaum’s Outline of Theory and Problems of Human Anatomy and Physiology. New York: McGraw-Hill, 2001.

Bernice Essenfeld

Temperature regulation

An organism maintains homeostasis , a steady internal state, only if its body temperature stays within prescribed limits. Cellular activities require an optimum amount of heat . They depend on enzyme action, and enzymes function within a narrow range of temperature. For this reason, living things can only tolerate a limited rise or drop in temperature from the optimum. Mechanisms exist that regulate body temperature to stay within its survival limits.

Cells require a source of heat for their activities. The heat source can be external or internal. The Sun is an external source of heat for living things. Cellular metabolism is an internal heat source. During cellular respiration, the chemical energy in food is converted to high-energy phosphate groups in adenosine triphosphate , ATP. In the process, cells lose about 60% of the released energy as heat. Plants and most animals give up this heat to the environment. Birds , mammals , and some fish , however, make use of some of the heat that metabolism yields.

Organisms exchange heat with the environment by four major physical processes. First, there is conduction, the direct transfer of heat from a warmer to a cooler object. We cool off during a swim because the heat of the body is conducted to the cool water . During another process called convection , a breeze passes over a surface and brings about heat loss. This is why a fan cools us on a hot day. Radiation , the emission of electromagnetic waves, transfers heat between objects that are not in direct contact. Animals absorb heat that the sun radiates. Finally, evaporation , the physical change of liquid to gas removes heat from a surface. Sweating and panting are cooling processes. Perspiration evaporates from the skin, and panting increases evaporation from the respiratory system . Both procedures take heat away from the body.

Externally heated animals

Ectotherms are animals that warm their bodies by absorbing heat from their surroundings. In most ectotherms, the body temperature fluctuates with changes in the surrounding temperature. The body temperature of snakes , for example, cools in cold weather and warms up in hot weather. Most marine fish and invertebrates , however, live in water that stays the same temperature. Their body temperature, therefore, does not change. In everyday language, we say that these animals are cold-blooded. Amphibians , reptiles , most fish, and invertebrates are considered cold-blooded. This term is misleading, however, because many lizards have high body temperatures. External temperature plays a major role in the activity rate of ectothermic animals. When the weather is warm, they become active. They slow down when the temperature drops.

Certain ectotherm behaviors help regulate body temperature. To warm up, reptiles find sunny places, and stretch out for maximum exposure. If it gets too warm, lizards alternate between sun and shade. Amphibians warm up by moving into the sun or diving into warm water. They cool off by entering the shade. In cold weather, honeybees huddle together to retain heat. Bees and large moths build up heat before takeoff by contracting their flight muscles without moving their wings.

In addition to behaviors, physiological adaptations help ectotherms regulate temperature. Diving reptiles conserve heat because their blood circulates inward toward the body core during a dive. The skin of bullfrogs secretes more mucus when it is hot, allowing more cooling by evaporation. Many ectotherms exist at a lower temperature during torpor, a state of slowed metabolism. This helps them survive a food shortage. If the food supply increases, they come out of torpor in a few hours.

Internally heated animals

Endotherms are animals that warm their bodies mainly from their own metabolism. Mostly birds and mammals, they maintain a constant body temperature regardless of changes in the surrounding temperature. We commonly call them warm-blooded, because the internal temperature they generate is usually higher than the environmental temperature. Endotherms have certain advantages. Their consistent higher temperature allows them to be active at all times. It also gives them the freedom to live in varied habitats. It helps them survive on land, where the air temperature changes more drastically than water temperature.

Although most fish are ectothermic, mackerals, tuna , swordfish , marlins , and some sharks are endotherms; although in swordfish and some mackeral, warming occurs only in the central nervous system and the retina of the eye . Endothermic fish are more closely related to their ectothermic relatives than they are to each other. Thus, endothermy evolved independently in these different groups, as an adaptation that allowed the animals to expand their habitats into waters of varying temperatures.

Certain adaptations accompany the maintenance of a constant body temperature. A high metabolic rate helps supply heat. An efficient circulatory system conducts and distributes heat around the body. Fat layers, fur, and feathers insulate the body and retain heat. Shivering muscles contract to increase body heat, and hormones such as epinephrine and thyroxin increase the rate of metabolism. Under unfavorable conditions some endotherms experience torpor, for example, hibernation , in which the body temperature drops. Hibernation enables animals to survive long periods of cold and lack of food.

Receptors in various parts of the body, such as the skin, deliver information to the hypothalamus of the brain about the body's temperature. The hypothalamus acts as a thermostat . It contains two thermoregulating areas that stimulate nerve impulses. One area, the heating center, sends out impulses that raise the body temperature. It causes blood vessels near the surface of the body to contract, thus preventing heat loss. It also causes fur to stand erect and become more insulating. It causes shivering by muscles that produce heat when they contract, and it stimulates hormonal production. The other area, the cooling center, sends out impulses that bring about a temperature drop. It causes surface blood vessels to expand thereby releasing heat. It brings about sweating or panting.

Resources

books

Coping with Change. Films for the Humanities and Sciences. Princeton, 1994-95.

Guyton & Hall. Textbook of Medical Physiology. 10th ed. New York: W. B. Saunders Company, 2000.

periodicals

Abass, Mageed, and C.B. Rajashekar. "Abscisic Acid Accumulation in Leaves and Cultured Cells During Heat Acclimation in Grapes." HortScience (January 1993).

Bookspan, Jolie. "Exposing the Myths of Heat Loss and Gain for Divers." Underwater USA (July 1993).

Comis, Don. "Resetting a Plant's Thermostat." AgriculturalResearch (July 1992).

Heinrich, Bernd. "Kinglets' Realm of Cold." Natural History (February 1993).

Luck-Baker. "Taking the Temperature of T. Rex." New Scientist (July 23, 1994).

Monastersky, Richard. "The Pulse of T. Rex." Science News (May 14, 1994).

Bernice Essenfeld

KEY TERMS

Cooling center

—Thermoregulating area of the hypothalamus whose impulses result in temperature-lowering activities of various parts of the body.

Ectotherm

—A cold-blooded animal, whose internal body temperature is similar to that of its environment. Ectotherms produce little body heat, and are dependent on external sources (such as the sun) to keep their body temperature high enough to function efficiently.

Endotherm

—An animal that uses its metabolism as a primary source of body heat and uses physiological mechanisms to hold its body temperature nearly constant.

Heating center

—Thermoregulating area of the hypothalamus whose impulses result in temperature-raising activities of various parts of the body.

Hibernation

—A type of torpor in which an ani mal's body temperature drops, thereby enabling it to withstand prolonged cold and little food.

Temperature Regulation

Humans and other mammals are homeothermic, able to maintain a relatively constant body temperature despite widely ranging environmental temperatures. Although the average human body temperature is 36.7 degrees Celsius (98.2 degrees Fahrenheit), this temperature varies depending on individual differences, time of day, the stage of sleep, and the ovulatory cycle in women. Temperature regulation, or thermoregulation, is the balance between heat production mechanisms and heat loss mechanisms that occur to maintain a constant body temperature.

Heat flows from higher temperature to lower temperature. Conduction is the transfer of heat between objects that are in direct contact with each other. For instance, if a person sits on the cold ground, heat moves from the body to the cold ground. Convection is the transfer of heat by the movement of air or liquid moving past the body. This explains why a breeze across the skin may cool one down, whereas trapping air inside clothing keeps the body warm.

A lizard sunning itself on a rock on a warm summer day illustrates radiation: the transfer of heat energy via electromagnetic waves. Whereas conduction, convection, and radiation can cause both heat loss and heat gain to the body, evaporation is a mechanism of heat loss only, in which a liquid is converted to a gas. Perspiration evaporating off the skin is an example of this heat loss mechanism.

When the body is too hot, it decreases heat production and increases heat loss. One way of increasing heat loss is through peripheral vasodilation, the dilation of blood vessels in the skin. When these vessels dilate, large quantities of warmed blood from the core of the body are carried to the skin, where heat loss may occur via radiation, convection, and conduction. Evaporation of fluids from the body also causes heat loss. Humans constantly lose fluids from the skin and in exhaled air. The unconscious loss of fluid is called insensible perspiration.

Although the body has no active control over insensible perspiration, the sympathetic nervous system controls the process of sweating and can stimulate secretion up to 4 liters (4.22 liquid quarts) of sweat per hour. In order for the sweat to evaporate and cool the body, the environmental air must have a relatively low humidity.

When the body is too cold, it increases heat production and decreases heat loss. Vasoconstriction, the constriction of the vessels of the skin, helps prevent heat loss. Shivering, which is a rhythmic contraction of skeletal muscles, produces heat. Heat can also be produced by nonshivering thermogenesis, an increase in metabolic heat production.

Hormones such as epinephrine, norepinephrine, and thyroid hormone increase the metabolic rate by stimulating the breakdown of fat. Humans also change posture, activity, clothing, or shelter to adjust for fluctuations in temperature. The goose bumps that arise on the skin in the cold are another sign the body is trying to prevent heat loss. They are due to piloerection, the erection of the hair follicles on the skin. This is a vestige of the time when humans were covered in hair: piloerection would trap air and retain heat.

Body temperature is regulated by a system of sensors and controllers across the body. The brain receives signals regarding body temperature from the nerves in the skin and the blood. These signals go to the hypothalamus, which coordinates thermoregulation in the body. Signals from the hypothalamus control the sympathetic nervous system, which affects vasoconstriction, metabolism , shivering, sweating, and hormonal controls over temperature. In general, the posterior hypothalamus controls responses to cold, and the anterior hypothalamus controls responses to heat.

Hypothermia, or low body temperature, is a result of prolonged exposure to cold. With a decrease in body temperature, all metabolic processes begin to slow. Hypothermia can be life-threatening.

Hyperthermia describes a body temperature that is higher than normal. One example of hyperthermia is fever. A fever is generally considered to be a body temperature over 38 degrees Celsius (100.4 degrees Fahrenheit). A fever is the body's natural defense to an infection by a bacterium or virus. Fevers are one of the body's mechanisms for eliminating an invading organism. Fevers may even make the immune system work more effectively. Heat exhaustion and heatstroke are other examples of hyperthermia. These occur when heat production exceeds the evaporative capabilities of the environment. Heatstroke may be fatal if untreated.

see also Hormones; Hypothalamus; Metabolism, Human; Nervous Systems; Skin; Thyroid Gland

Martha S. Rosenthal

Bibliography

Marieb, Elaine. "Nutrition, Metabolism, and Body Temperature Regulation." In Human Anatomy and Physiology, 5th ed. New York: Benjamin Cummings, 2001.

Sherwood, Laurelee. "Energy Balance and Temperature Regulation." In Fundamentals of Physiology: A Human Perspective, 2nd ed. New York: West Publishing Company, 1995.