40.

5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior

We will examine how animal form and function work together in regulating the internal environmentspecifically, the regulation of body temperature. Thermoregulation is the process by which animals maintain an internal temperature within a tolerable range. This ability is critical to survival because most biochemical and physiological processes are very sensitive to changes in body temperature. The rates of most enzymemediated reactions increase two to threefold for every 10C temperature increase until temperature is high enough to begin to denature proteins. The properties of membranes also change with temperature. These thermal effects dramatically influence animal functioning. Although different species of animals are adapted to different environmental temperatures, each species has an optimal temperature range. Thermoregulation helps keep body temperature within that optimal range, enabling cells to function most effectively, even as the external temperature fluctuates. Section Vocabulary Thermoregulation - The maintenance of internal body temperature within a tolerable range. Ectotherm - An animal, such as a reptile (other than birds), fish, or amphibian, that must use environmental energy and behavioral adaptations to regulate its body temperature. Endotherm - An animal, such as a bird or mammal, that uses metabolic heat to regulate body temperature. Integumentary System - The outer covering of a mammals body, including skin, hair, and nails. Vasodilation - An increase in the diameter of superficial blood vessels triggered by nerve signals that relax the muscles of the vessel walls. Vasoconstriction - A decrease in the diameter of superficial blood vessels triggered by nerve signals that contract the muscles of the vessel walls. Countercurrent Heat Exchanger - An arrangement of blood vessels that helps trap heat in the body core and is important in reducing heat loss in many endotherms. Nonshivering Thermogenesis (NST) - The increased production of heat in some mammals by the action of certain hormones that cause mitochondria to increase their metabolic activity and produce heat instead of ATP. Brown Fat - A tissue in some mammals, located in the neck and between the shoulders, that is specialized for rapid heat production. Acclimatization - Physiological adjustment to a change in an environmental factor. StressInduced Proteins - Molecules, including heatshock proteins, that are produced within cells in response to exposure to marked increases in temperature and to other forms of severe stress, such as toxins, rapid pH changes, and viral infections. HeatShock Protein - A protein that helps protect other proteins during heat stress. Heatshock proteins are found in plants, animals, and microorganisms. Torpor - In animals, a physiological state that conserves energy by slowing down metabolism. Hibernation - A physiological state that allows survival during long periods of cold temperatures and reduced food supplies, in which metabolism decreases, the heart and respiratory system slow down, and body temperature is maintained at a lower level than normal. Estivation - Summer torpor; a physiological state that is characterized by slow metabolism and inactivity and that permits survival during long periods of elevated temperature and diminished water supplies. Daily Torpor - A daily decrease in metabolic activity and body temperature during times of inactivity for some small mammals and birds. There are important differences in how various species manage their heat budgets. One way to classify the thermal characteristics of animals is to emphasize the role of metabolic heat in determining body temperature. As you learned earlier, ectotherms gain most of their heat from the environment. An ectotherm has such a low metabolic rate that the amount of heat it generates is too small to have much effect on body temperature. In contrast, endotherms can use metabolic heat to regulate their body temperature. In a cold environment, an endotherms high metabolic rate generates enough heat to keep its body substantially warmer than its surroundings. Many endotherms, including humans, maintain high and very stable internal temperatures even as the temperature of their surroundings fluctuates. Many ectotherms can thermoregulate by behavioral means, such as basking in the sun or seeking out shade. But in general, ectotherms tolerate greater variation in internal temperature than do endotherms (Figure 40.12). Most invertebrates, fishes, amphibians, lizards, snakes, and turtles are

Ectotherms and Endotherms

40.5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior
ectotherms. Mammals, birds and a few other reptiles, some fishes, and numerous insect species are endotherms. It is important to note that animals are not classified as ectotherms or endotherms based on whether they have variable or constant body temperatures, a common misconception. As mentioned earlier, it is the source of heat used to maintain body temperature that distinguishes ectotherms from endotherms. A different set of terms is used to imply variable or constant body temperatures. The term poikilotherm refers to animals whose internal temperatures vary widely, and the term homeotherm refers to animals that maintain relatively stable internal temperatures. However, as scientists have gained more knowledge of animal thermoregulatory mechanisms, these terms have largely fallen out of use. Many marine fishes and invertebrates, classified as poikilotherms, inhabit water with such stable temperatures that their body temperature varies less than that of humans and other mammals. Furthermore, some mammals that were classified as homeotherms experience great variation in internal temperature. For example, a chipmunk sustains a high body temperature while it is active, but its temperature drops as hibernation begins. Because of such exceptions, the terms ectotherm and endotherm are generally preferred. Another common misconception is the idea that ectotherms are coldblooded and endotherms are warmblooded. Ectotherms do not necessarily have low body temperatures. In fact, when sitting in the sun, many ectothermic lizards have higher body temperatures than mammals. Thus, most biologists avoid the familiar terms coldblooded and warmblooded because they are so often misleading. It is also important to note that ectothermy and endothermy are not mutually exclusive thermoregulatory strategies. For example, a bird is an endotherm, but it may warm itself in the sun on a cold morning, much as an ectothermic lizard does. Endothermy has several important advantages. Being able to generate a large amount of heat metabolically, along with other biochemical and physiological adaptations associated with endothermy (such as elaborate circulatory and respiratory systems), enables endotherms to perform vigorous activity for much longer than is possible for most ectotherms (Figure 40.9). Sustained intense activity, such as longdistance running or powered (flapping) flight, is usually only feasible for animals with an endothermic way of life. Endothermy also solves certain thermal problems of living on land, enabling terrestrial animals to maintain stable body temperatures in the face of environmental temperature fluctuations that are generally more severe than in aquatic habitats. For example, no ectotherm can be active in the belowfreezing weather that prevails during winter over much of Earths surface, but many endotherms function very well in these conditions. Most of the time, endothermic vertebratesbirds and mammalsare warmer than their surroundings, but these animals also have mechanisms for cooling the body in a hot environment, which enables them to withstand heat loads that are intolerable for most ectotherms. Endotherms are better buffered against external temperature fluctuations compared to ectotherms, but keep in mind that ectotherms can usually tolerate larger fluctuations in their internal temperatures. Being endothermic is liberating, but it is also energetically expensive. For example, at 20C, a human at rest has a metabolic rate of 1,300 to 1,800 kcal per day (BMR). In contrast, a resting ectotherm of similar weight, such as an American alligator, has a metabolic rate of only about 60 kcal per day at 20C (SMR). Thus, endotherms generally need to consume much more food than ectotherms of equivalent sizea serious disadvantage for endotherms if food supplies are limited. For this and other reasons, ectothermy is an extremely effective and successful strategy in most of Earths environments, as shown by the abundance and diversity of ectothermic animals. Modes of Heat Exchange Whether it is an ectotherm or an endotherm, an organism, like any object, exchanges heat by four physical processes: conduction, convection, radiation, and evaporation. Figure 40.13 distinguishes these processes, which account for the flow of heat within an organism and between an organism and its external environment. Note that heat is always transferred from an object of higher temperature to one of lower temperature.

40.5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior
Balancing Heat Loss and Gain For endotherms and for those ectotherms that thermoregulate, the essence of thermoregulation is managing the heat budget so that rates of heat gain are equal to rates of heat loss. If the heat budget is unbalanced, the animal becomes either warmer or colder. Five general categories of adaptations help animals thermoregulate. A major thermoregulatory adaptation in mammals and birds is insulation (hair, feathers, or fat layers), which reduces the flow of heat between an animal and its environment and lowers the energy cost of keeping warm. In mammals, the insulating material is associated with the integumentary system, the outer covering of the body, consisting of the skin, hair, and nails (claws or hooves in some species). Skin is a key organ of the integumentary system. In addition to functioning as a thermoregulatory organ by housing nerves, sweat glands, blood vessels, and hair follicles, the skin protects internal body parts from mechanical injury, infection, and drying out. The skin consists of two layers, the epidermis and the dermis, underlain by a tissue layer called the hypodermis (Figure 40.14). The epidermis is the outermost layer of skin and is composed mostly of dead epithelial cells that continually flake and fall off. New cells pushing up from lower layers replace the cells that are lost. The dermis supports the epidermis and contains hair follicles, oil and sweat glands, muscles, nerves, and blood vessels. The hypodermis contains adipose tissue, which includes fatstoring cells and blood vessels. Adipose tissue provides varying degrees of insulation, depending on the species. The insulating power of a layer of fur or feathers mainly depends on how much still air the layer traps. (Hair loses most of its insulating power when wet.) Most land mammals and birds react to cold by raising their fur or feathers, thereby trapping a thicker layer of air. Humans rely more on a layer of fat just beneath the skin as insulation (Figure 40.14); goose bumps are a vestige of hair raising inherited from our furry ancestors. Marine mammals, such as whales and seals, have a very thick layer of insulating fat called blubber just under their skin. Marine mammals swim in water colder than their body core temperature, and many species spend at least part of the year in nearly freezing polar seas. The transfer of heat to water occurs 50 to 100 times more rapidly than heat transfer to air, and the skin temperature of a marine mammal is close to water temperature. Even so, the blubber insulation is so effective that marine mammals maintain body core temperatures of about 36 38C, with metabolic rates about the same as those of land mammals of similar size. Many endotherms and some ectotherms can alter the amount of blood (and hence heat) flowing between the body core and the skin. Elevated blood flow in the skin normally results from vasodilation, an increase in the diameter of superficial blood vessels (those near the body surface) triggered by nerve signals that relax the muscles of the vessel walls. In endotherms, vasodilation usually warms the skin, increasing the transfer of body heat to a cool environment by radiation, conduction, and convection (see Figure 40.13). The reverse process, vasoconstriction, reduces blood flow and heat transfer by decreasing the diameter of superficial vessels. Another circulatory adaptation is an arrangement of blood vessels called a countercurrent heat exchanger that is important for reducing heat loss in many endotherms, including marine mammals and birds. Figure 40.15 explores two examples of countercurrent heat exchangers. In some species, blood can either go through the heat exchanger or bypass it by way of other blood vessels. In this way, the relative amount of blood that flows through the two different paths may vary, adjusting the rate of heat loss as an animals physiological state or environment changes. Unlike most fishes, which are thermoconformers with internal body temperatures usually within 12C of the surrounding water temperature, some specialized endothermic bony fishes and sharks have circulatory adaptations that retain metabolic heat in the body. These include large, powerful swimmers such as bluefin tuna and swordfish, as well as the great white shark. Large arteries convey most of the cold blood from the gills to tissues just under the skin. Branches deliver blood to the deep muscles, where the small vessels are arranged into a countercurrent heat exchanger (Figure 40.16). Endothermy enables the vigorous, sustained activity that is characteristic of these animals by keeping the main swimming muscles several degrees warmer

Insulation

Circulatory Adaptations

40.5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior
than the tissues near the animals surface, which are about the same temperature as the surrounding water. Some reptiles also have physiological adaptations that regulate heat loss. For example, in the marine iguana, which inhabits the Galpagos Islands (Figure 22.1), body heat is conserved by vasoconstriction of superficial blood vessels, routing more blood to the central core of the body when the animal is swimming in the cold ocean. Many endothermic insects (bumblebees, honeybees, and some moths) have a countercurrent heat exchanger that helps maintain a high temperature in the thorax, where the flight muscles are located. For example, the heat exchanger keeps the thorax of certain winteractive moths at about 30C during flight, even on cold, snowy nights when the external temperature may be subfreezing (Figure 40.17). In contrast, insects flying in hot weather run the risk of overheating because of the large amount of heat produced by working flight muscles. In some species, the countercurrent mechanism can be shut down, allowing muscleproduced heat to be lost from the thorax to the abdomen, and from there to the environment. A bumblebee queen incubates her eggs this way: She generates heat by shivering her flight muscles and then transfers the heat to her abdomen, which she presses against her eggs. Cooling by Evaporative Heat Loss Many mammals and birds live in places where thermoregulation requires cooling as well as warming. If environmental temperature is above body temperature, animals gain heat from the environment as well as from metabolism, and evaporation is the only way to keep body temperature from rising rapidly. Terrestrial animals lose water by evaporation across the skin and when they breathe. Water absorbs considerable heat when it evaporates; it is 50 to 100 times more effective than air in transferring heat. Some animals have adaptations that can greatly augment this cooling effect. Panting is important in birds and many mammals. Some birds have a pouch richly supplied with blood vessels in the floor of the mouth; fluttering the pouch increases evaporation. Pigeons, for example, can use evaporative cooling to keep body temperature close to 40C in air temperatures as high as 60C, as long as they have sufficient water. Sweating or bathing moistens the skin and enhances evaporative cooling (Figure 40.18). Many terrestrial mammals have sweat glands controlled by the nervous system (see Figure 40.14). Other mechanisms that promote evaporative cooling include spreading saliva on body surfaces, an adaptation of some kangaroos and rodents for combating severe heat stress. Some species of amphibians, such as bullfrogs, can vary the amount of mucus they secrete from their surface, a response that regulates evaporative cooling. Both endotherms and ectotherms use behavioral responses to control body temperature. Many ectotherms can maintain a very constant body temperature through relatively simple behaviors. More extreme behavioral adaptations in some animals include hibernation or migration to a more suitable climate. All amphibians and most reptiles other than birds are ectothermic. Therefore, these organisms control body temperature mainly by behavior. The optimal temperature range for amphibians varies substantially with the species. For example, certain closely related species of salamanders have average body temperatures ranging from 7 to 25C. When exposed to air, most amphibians lose heat rapidly by evaporation from their moist body surfaces, making it difficult to keep sufficiently warm. However, by moving to a location where solar heat is available, an amphibian can maintain a satisfactory body temperature. And when the surroundings are too warm, amphibians seek cooler microenvironments, such as shaded areas. Like amphibians, reptiles other than birds thermoregulate mainly by behavior. When cold, they seek warm places, orienting themselves toward heat sources and expanding the body surface exposed to a heat source. When hot, they move to cool areas or turn in another direction. Many reptiles keep their body temperatures very stable over the course of a day by shuttling back and forth between warm and cool spots. Many terrestrial invertebrates can adjust internal temperature by the same behavioral mechanisms used by vertebrate ectotherms. The desert locust, for example, must reach a certain temperature to become active, and on cold days it orients in a direction that maximizes the absorption of sunlight. Other terrestrial invertebrates have certain postures that enable them to maximize or minimize their absorption of heat from the sun (Figure 40.19).

Behavioral Responses

40.5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior
Honeybees use a thermoregulatory mechanism that depends on social behavior. In cold weather, they increase heat production and huddle together, thereby retaining heat. They maintain a relatively constant temperature by changing the density of the huddling. Individuals move between the cooler outer edges of the cluster and the warmer center, thus circulating and distributing the heat. Even when huddling, honeybees must expend considerable energy to keep warm during long periods of cold weather, and this is the main function of storing large quantities of fuel in the hive in the form of honey. Honeybees also control the temperature of their hive by transporting water to it in hot weather and fanning with their wings, which promotes evaporation and convection. Thus, a honeybee colony uses many of the mechanisms of thermoregulation seen in single organisms. Adjusting Metabolic Heat Production Because endotherms generally maintain body temperatures considerably warmer than the environment, they must counteract constant heat loss. Endotherms can vary heat production to match changing rates of heat loss. For example, heat production is increased by such muscle activity as moving or shivering. In some mammals, certain hormones can cause mitochondria to increase their metabolic activity and produce heat instead of ATP. This nonshivering thermogenesis (NST) takes place throughout the body, but some mammals also have a tissue called brown fat in the neck and between the shoulders that is specialized for rapid heat production. Through shivering and NST, mammals and birds in cold environments can increase their metabolic heat production by as much as five to ten times the minimal levels that occur in warm conditions. For example, small birds called chickadees, which weigh only 20 g, can remain active and hold body temperature nearly constant at 40C in environmental temperatures as low as 40Cas long as they have enough food to supply the large amount of energy necessary for heat production. A few large reptiles become endothermic in particular circumstances. For example, female pythons that are incubating eggs increase their metabolic rate by shivering, generating enough heat to keep their body (and egg) temperatures 57C warmer than the surrounding air for weeks at a time. This temporary endothermy consumes considerable energy. Researchers continue to debate whether certain groups of dinosaurs were endothermic. As mentioned earlier, many species of flying insects, such as bees and moths, are endothermic the smallest of all endotherms. The capacity of such endothermic insects to elevate body temperature depends on powerful flight muscles, which generate large amounts of heat when operating. Many endothermic insects use shivering to warm up before taking off. They contract their flight muscles in synchrony, so that only slight wing movements occur but considerable heat is produced. Chemical reactions, and hence cellular respiration, speed up in the warmedup flight motors, enabling these insects to fly even on cold days or at night (Figure 40.20). The regulation of body temperature in humans and other mammals is a complex system facilitated by feedback mechanisms (Figure 40.11). Nerve cells that control thermoregulation, as well as those that control many other aspects of homeostasis, are concentrated in a region of the brain called the hypothalamus. The hypothalamus contains a group of nerve cells that functions as a thermostat, responding to changes in body temperature above or below a set point (actually above or below a normal range) by activating mechanisms that promote heat loss or gain (Figure 40.21). Nerve cells that sense temperature are in the skin, in the hypothalamus itself, and in several other body regions. Warm receptors signal the hypothalamic thermostat when temperatures increase; cold receptors signal temperature decrease. At body temperatures below the normal range, the thermostat inhibits heat loss mechanisms and activates heatsaving ones such as vasoconstriction of superficial vessels and erection of fur, while stimulating heat generating mechanisms (shivering and nonshivering thermogenesis). In response to elevated body temperature, the thermostat shuts down heat retention mechanisms and promotes body cooling by vasodilation, sweating, or panting. The thermostat can also respond to external temperature (sensed as skin temperature) even without changes in body core temperature. Many animals can adjust to a new range of environmental temperatures over a period of days or weeks, a physiological response called acclimatization. Both ectotherms and endotherms

Feedback Mechanisms in Thermoregulation

Adjustment to Changing Temperatures

40.5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior
acclimatize, but in different ways. In birds and mammals, acclimatization often includes adjusting the amount of insulationby growing a thicker coat of fur in the winter and shedding it in the summer, for exampleand sometimes varying the capacity for metabolic heat production in different seasons. These changes help endotherms keep a constant body temperature in both warm and cold seasons. Acclimatization in ectotherms involves compensating for changes in temperature. These adjustments can strongly affect physiology and temperature tolerance. For example, summeracclimatized bullhead catfish can survive water temperatures up to 36C but cannot function in cold water; after winter acclimatization, they can easily tolerate cold water, but a temperature above 28C is lethal. Acclimatization responses in ectotherms often include adjustments at the cellular level. Cells may increase the production of certain enzymes or produce variants of enzymes that have the same function but different optimal temperatures. Membranes may also change the proportions of saturated and unsaturated lipids they contain, which helps keep membranes fluid at different temperatures. Some ectotherms that experience subzero body temperatures protect themselves by producing antifreeze compounds (cryoprotectants) that prevent ice formation in the cells. In arctic regions or on cold mountain peaks, cryoprotectants in the body fluids let overwintering ectotherms, such as some frogs and many arthropods and their eggs, withstand body temperatures considerably below zero. Cryoprotectants are also found in certain species of fishes from arctic and antarctic seas, where water temperatures can be as cold as 1.8C, well below the freezing point of unprotected body fluids (about 0.7C). Cells can often make rapid adjustments to temperature changes. For example, mammalian cells grown in laboratory cultures respond to a marked increase in temperature and to other forms of severe stress by accumulating molecules called stressinduced proteins, including heatshock proteins. Within minutes of being shocked by a rapid change in temperature from 37C to about 43C, cultured mammalian cells begin synthesizing heatshock proteins. These molecules help maintain the integrity of other proteins that would be denatured by severe heat. Found in bacteria, yeasts, and plant cells as well as in animals, stressinduced proteins help prevent cell death when an organism is challenged by severe changes in the cellular environment. Torpor and Energy Conservation Despite their many adaptations for homeostasis, animals may occasionally encounter conditions that severely challenge their abilities to balance heat, energy, and materials budgets. For example, at certain seasons of the year (or certain times of day), temperatures may be extremely hot or cold, or food may be unavailable. An adaptation that enables animals to save energy while avoiding difficult and dangerous conditions is torpor, a physiological state in which activity is low and metabolism decreases. Hibernation is longterm torpor that is an adaptation to winter cold and food scarcity. When vertebrate endotherms (birds and mammals) enter torpor or hibernation, their body temperatures declinein effect, their bodys thermostat is turned down. The temperature reduction may be dramatic: Some hibernating mammals cool to as low as 12C, and a few even drop slightly below 0C in a supercooled (unfrozen) state. The resulting energy savings due to lower metabolic rate and less heat production are huge; metabolic rates during hibernation can be several hundred times lower than if the animal attempted to maintain normal body temperatures of 3638C. This allows hibernators to survive for very long periods on limited supplies of energy stored in the body tissues or as food cached in a burrow. Certain ground squirrels are favorite research models for biologists studying the physiology of hibernation (Figure 40.22). For example, a Beldings ground squirrel (Spermophilus beldingi ) living in the high mountains of California is active only during spring and summer, when it maintains a body temperature of about 37C and a metabolic rate of about 85 kcal per day. In September, the squirrel retreats to a safe burrow where it spends the next eight months hibernating. For most of the hibernation season, the squirrels body temperature is only slightly above burrow temperature (which may be close to freezing), and its metabolic rate is extremely

40.5 Thermoregulation contributes to homeostasis and involves anatomy, physiology, and behavior
low (Figure 40.22). Every week or two it arouses for a few hours, using metabolic heat to warm up to about 37C (these periodic arousals may be needed for maintenance functions that require high body temperature). In late spring, when outside temperature is climbing, the squirrel resumes normal endothermy. By hibernating, Beldings ground squirrels avoid severe cold and greatly reduce the amount of energy they need to survive the winter, when their normal food of grasses and seeds is not available. Instead of having to spend 150 kcal per day to maintain normal body temperatures in winter weather, a squirrel in its burrow spends an average of only 58 kcal per day and can live on stored fatwithout eatingfor the entire hibernation season. Estivation, or summer torpor, also characterized by slow metabolism and inactivity, enables animals to survive long periods of high temperatures and scarce water supplies. Hibernation and estivation are often triggered by seasonal changes in the length of daylight. As the days shorten, some hibernators prepare for winter by storing food in their burrows; other species eat huge quantities of food. For example, ground squirrels double their weight in a month of gorging. Many small mammals and birds exhibit a daily torpor that seems to be adapted to their feeding patterns. For instance, most bats and shrews feed at night and go into torpor during daylight hours. Chickadees and hummingbirds feed during the day and often go into torpor on cold nights; the body temperature of chickadees drops as much as 10C at night, and that of hummingbirds can fall 2530C. All endotherms that use daily torpor are relatively small; when active, they have high metabolic rates and thus very high rates of energy consumption. During hours when they cannot feed, torpor enables them to survive on stored energy. An animals daily cycle of activity and torpor appears to be a builtin rhythm controlled by its biological clock (see Chapter 48). Even if food is made available to a shrew all day, it still goes through its daily torpor. The need for sleep in humans and the slight drop in body temperature that accompanies it may be an evolutionary remnant of a more pronounced daily torpor in our early mammalian ancestors. 1. Can ectotherms have stable body temperatures? Explain. a. Yes, ectotherms in the deep sea and in constanttemperature freshwater springs have constant body temperatures. And terrestrial ectotherms can maintain relatively constant body temperatures by behavioral means.

2.

What mode of heat exchange is involved in wind chill, when the air feels colder than the actual temperature? a. Heat loss through convection

3.

Some birds in tropical dry forests periodically go into torpor, especially in the dry season. Explain. a. Food and water supplies may be short during the dry season, and torpor enables animals to survive at a much lower metabolic rate.