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Controlling the internal environment Controlling the internal environment Most organisms can survive fluctuations in the external environment that are more extreme than any of their individual cells could tolerate.During the day , a human may be exposed to substantial changes in outside temperatures but will die if the internal body temperature goes more than a few degrees above or below an average of about 37C. The spring peeper, a North American tree frog often spends an entire winter with much of its body frozen. Most of its blood and interstitial fluid turns to ice, but its cells and a film of water surrounding them are kept from freezing by specialized proteins and a high level of glucose throughout the frog's body. Without these protectants, ice crystals would rupture the cell membranes, and the animal would die. Homeostatic mechanisms protect an animal's internal environment from harmful fluctuations: an overview: In most organism, the majority of cells are not in direct contact with the external environment but are bathed by an internal body fluid. Insects and other animals with an open circulatory system have an internal pond composed of hemolymph, which bathes all body cells. In vertebrates and other animals with a closed circulatory system, the internal pond is interstitial fluid serviced by blood. Homeostatic mechanisms temper changes in an animal's body fluids, cushioning them from the potentially harmfiul impact of fluctuations in the extyernal environment (Fig. 40.1) nP880. 81 Controlling the internal environment 82 Cells require a balance between water uptake and loss: Water enter our body's by three ways: In our food In our drink Oxidative metabolism Metabolic water is produced by cellular respiration when electrons and hydrogen are added to oxygen. Lose water by: Urinating defecating And by evaporative loss due to sweating and breathing. Whether an organism inhabits land, fresh water, or salt water, or moves back and forth between these environments, one general problem occurs: The cells of the animal cannot survive a net water gain or loss. Water continuously enters and leaves an animal cell across the plasma membrane, however uptake and loss must balance. An animal cells swell and burst if there is a net uptake of water or shrivel and die if there is a net loss of water. Osmosis, is the movement of water across a selectively permeable membrane. It occurs whnever two solutions separated by the membrane differ in total solute concentration, or osmolarity (total solute concentration expressed as molarity, or moles of solute per liter of solution). Foe example, the osmolarity of human blood is about 300 mosm/L, while sea water commonly has an osmolarity of about 1000 mosm/L. Osmoconformers and osmoregulators There are two basic solutions to the problem of balancing water gain with water loss. One solution for a marine organism is to be isotonic with its saltwater environment. Such organisms, which do not aqctively adjust their internal osmolarity, are known as osmoconformers. Controlling the internal environment 83 By contrast, organisms whose body fluids are not isotonic with the outside environment, called osmoregulators, must either discharge excess water if they live in ahyptonic environement or continuously take in water to offset osmotic loss if they inhabit a hypertonic environment. The ability to osmoregulate enables animals to live, for example, in fresh water, where the osmolarity is too low to support cellular life, and on land, where water is usually in short supply. Most organisms, whether osmoconformers or osmoregulators, cannot tolerate substantial changes in external osmolarity. Such organisms are said to be stenohaline (stenos, "narrow": haline refers to salt). However, some organisms called euryhaline animals, do survive radical fluctuations of osmolarity in their surroundings. They either conform to the changes or regulate their internal osmolarity within a narrow range even as the external osmolarity changes. One example of euryhaline animal, a bony fish called tilapia can adjust to any salt concentrations between fresh water and twice that of seawater. All freshwater animals and many marine animals are osmoregulators. Humans and other terrestrial animals, also osmoregulators, must compensate for water loss. Maintanance water balance in different environments Marine animals. Animals Animals first evolved in the sea. Most marine invertebrates are osmoconformers, even these animals differ from seawater in their concentrations of specific salts. Among the vertebrates, the hagfishes, (jawless members of the class Agnatha) are isotonic with the surrounding seawater, but most marine vertebrates osmoregulate. Sharks and most cartilaginous fishes maintain internal salt concentrations lower than that of seawater. Their kidneys excrete some salts, and a salt-excreting organ called the rectal gland excretes sodium chloride out of the body through the anus. A large amounts of urea dissolved in the body fluids accounts for a shark's being slightly hypertonic to seawater. Sharks also produce and retain another Controlling the internal environment 84 organic compound, trimethylamine oxide (TMAO), which protects proteins from being damaged by the urea. Marine bony fishes constantly lose water by osmosis to their hypertonic surroundings (Fig. 40.2a) P882. They compensate by drinking large amounts of seawater, pumping out excess salts, and excreting urine in relatively small amounts. Freshwater animals Freshwater animals are constantly taking in water by osmosis because the osmolarity of their internal fluids is much higher than that of their surroundings. Amoeba and paramecium have contractile vacuoles that pump out excess water. Many freshwater animals, including fishes, bail out ( ) ينزحwater by excreting large amounts of very dilute urine and regaining lost salts in their food. (Fig.40.2b) P882. Controlling the internal environment 85 Life in temporary waters Dehydration kill most organisms, but some aquatic invertebrates living in temporary ponds and films of water around soil particles can lose almost all their body water and survive in a dormant state when their habitats dry up. This remarkable adaptation is called anhydrobiosis (life without water) or cryptobiosis (Hidden life). Example tardigrades, or water bears, tiny invetebrates less than 1mm long (fig. 40.3 P882). In their active, hydrated state, these organisms contain about 85% water by weight but can dehydrate to less than 2% water and survive in an inactive state, dry as dust, for a decade. Just add water, and within minutes the rehydrated tardigrades are moving about and feeding. Tardigrades survive drying out by containing a large amounts of sugars disaccharides called trehalose, which protect the cells by replacing the water associated with membranes and proteins. Many insects that survive freezing in the winter also utilize trehalose as a membrane protectant. Terrestrial animals Few terrestrial organisms are capable of anhydrobiosis. Human's die if they lose about 12% of their body water. Adaptations that have made it possible for organisms, which consist mostly of water, to survive on land is: 1. the waxy cuticle of plants on land. 2. Also insects have waxy exoskeletons. Controlling the internal environment 86 3. the shell of land snails. 4. and the multiple layers of dead, keratinized skin cells covering most terrestrial vertebrates. 5. behavioral adaptations as nervous, hormonal mechanisms which control thirst. 6. Desert animals are nocturnal, reduces dehydration. 7. The kidneys and other excretory organs of terrestrial organisms often exhibit adaptations that help conserve water. 8. Some mammals are so well adapted to minimizing water loss that they can survive in deserts without drinking. Example, Kangaroo rats lose so little water that they can recover 90% of the loss by using metabolic water . (Fig. 40.4 P883). Controlling the internal environment 87 Mosmoregulation depends on transport epithelia Specialized epithelia, called transport epithelia, regulate the solute movements. A transport epithelium is usually a single sheet of cells facing the external ebvironment or some channel that leads to the exterior through an opening on the body surface. The cells of the epithelium are joined by impermeable tight junctions (Fig. 7.34a), forming a continuous barrier at the tissue-environment boundary. The solute passing between the extracellular fluid and the environment must pass through the selectively permeable membranes of cells. Transport epithelia vary in their passive permeabilities to water and salts, and in number, type, and orientation of membrane proteins responsible for active transport. One of the most efficient transport epithilia is found in marine birds that spend months or years at sea and obtain water from the ocean (Fig. 40.5 P884). They have specialized salt-excreting glands whose transport epithelia are dedicated exclusively to osmoregulation-maintaining salt and water balance. Controlling the internal environment 88 Tubular systems function in osmoregulation and excretion in many invertebrates Protonephridia: The flame-Bulb system of flatworms Flatworms have simple tubular excretory systems called protonephridia. A protonephridium is a network of closed tubules lacking internal opening. (Fig. 40.6) P885. The flame-bulb systems of freshwater flatworms function mainly in osmoregulation, most metabolic wastes diffuse out from the body surface or are excreted into the gastrovascular cavity and eliminated through the mouth. Protonephridia are also found in rotifers, some annelids, and the larvae of mollusks. Controlling the internal environment 89 Metanephridia of earthworms Another type of tubular excretory system, the metanephridium, has internal openings that collect body fluids. Metanephridia are found in most annelids, including earthworms (Fig. 40.7) P885. Each segment of a worm has a pair of metanephridia, which are tubules immersed in coelomic fluid and enveloped by a network of capillaries. An earthworm's metanephridia have excretory and osmoregulatory functions. As the fluid moves along the tubule, the transport epithelium bordering the lumen pumps essential salts out of the tubule, and the salts are reabsorbed into the blood circulating through the capillaries. The urine that exists through the nephridiopore contains nitrogenous wastes and is hypotonic to the body fluid. Controlling the internal environment 90 Malpighian tubules in insects The excretory organs of insects and other terrestrial arthropods is called Malpighian tubules, remove nitrogenous wastes from the hemolymph and also function in osmoregulation (Fig. 40.8) P 886. The transport epithelium that lines a tubule pumps certain solutes, including salts and nitrogenous wastes, from the hemolymph into the lumen of the tubule. Controlling the internal environment 91 The kidneys of most vertebrates are compact organs with many excretory tubules The kidneys of vertebrates are compact organs containing numerous tubules that are not segmentally arranged. A dense network of capillaries intimately associated with the tubules is also part of the kidney. In vertebrates that osmoregulate, the tubules function in both excretion and osmoregulation. The kidneys, the blood vessels that serve them, and the structures that carry urine formed in the kidneys out of the body are the components of the vertebrate excretory system (Fig. 40.9) P887. The mammalian excretory system In humans, the kidneys are a pair of bean-shaped organs about 10 cm long (Fig. 40.9a). Blood enters each kidney via the renal artery and leaves each kidney via the renal vein. They receive about 20% of the blood pumped with each heartbeat. Urine exists the kidney through a duct called the ureter. The ureters of both kidneys drain into a common urinary bladder. During urination, urine leaves the body from the urinary bladder through a tube called the urethra. The nephron and associated structures The kidneys has two distinct regions, an outer renal cortex and an inner renal medulla. The nephron, which is the functional unit of the vertebrate kidney, consists of a single long tubule and a ball of capillaries called glomerulus. (See Fig. 40.9b, 40.9c, 40.9 d.) P887. Only mammals and birds have juxtamedullary nephrons; the nephrons of other vertebrates lack loops of Henle. The nephron and the collecting duct are lined by a transport epithelium that processes the filtrate to form the urine. From about 1100 to 2000 L of blood that flows through the human kidneys each day, the nephrons and collecting ducts process about 180 L of filtrate, but the kidneys excrete only about 1.5 L of urine. The rest of the filtrate, including about 99% of the water, is reabsorbed into the blood. Controlling the internal environment 92 Controlling the internal environment 93 The kidney's transport epithelia regulate the composition of blood Production of urine from a blood filtrate The transport epithelia of nephrons and collecting ducts regulate the composition of blood by a combination of three processes that transfer material between the tubules and the capillaries that serves them: filtration, secretion and reabsorption. Filtration of the blood: Blood pressure forces fluid from the capillaries of glomerulus across the epithelium of Bowman's capsule into the lumen of the nephron tubule. The porous capillaries, along with specialized cells of the capsule called podocytes, function as a filter, being permeable to water and small solutes but not to bloos cells or large molecules such as plasma proteins (Fig.40.10) P 889. Filtration is nonselective with regard to small molecules; any substance small enough to be forced through the capillary wall and between the podocytes by blood pressure enters the lumen of the nephron tubule. The filtrate contains solutes such as salts, glucose, and vitamins, nitrogenous wastes such as urea. Secretion as filtrate travels through the nephron tubule, it is joined by substances that are transported across the tubule epithelium from the surrounding interstitial fluid. The net effect of of renal secretion is the addition of plasma solutes to the filtrate within the tubule. The proximal and distal tubules are the most common sites of secretion. Reabsorption because filtration is nonselective, it is important that small molecules essential to the body be returned to the interstitial fluid and blood plasma. This selective transport of substances across the epithelium of the excretory tubule from the filtrate to the interstitial fluid is called reabsorption. The proximal and distal tubules and the loop of Henle all contribute to reabsportion as does the collecting duct. Controlling the internal environment 94 All sugars, vitamins and other organic nutrients present in the initial filtrate are reabsorbed. Most of the water of the filtrate is also reabsorbed in the kidneys of mammals and birds. Controlling the internal environment 95 Controlling the internal environment 96 The water-conserving ability of the mammalian kidney is a key terrestrial adaptation The osmolarity of human blood is about 300 mosm/L, but the kidney can excrete urine up to four times as concentrated-about 1200 mosm/L. The main two solutes responsible for this osmolarity gradient are NaCL, which is deposited in the renal medulla by the loop of Henle, and urea, which leaks across the epithelium of the collecting duct in the inner medulla. Fig. 40.12 P893. Controlling the internal environment Regulation of kidney function by feedback circuits Fig. 40.13 P895 97 Controlling the internal environment 98 Diverse adaptations of the vertebrate kidney have evolved in different habitats Mammals that excrete the most hypertonic urine, such as kangaroo rats and other mammals adapted to the desert, have exceptionally long loops of Henle. Long loops maintain steep osmotic gradients in the kindney, rsulting in urine becoming very concentrated. In contrast, beavers, which spend much of their time in fresh water and rarely face problems of dehydration, have nephrons with very short loops, resulting in dilute urine. Birds, like mammals, have kidneys with juxamedullary nephrons that specialized in conserving water. The kidneys of reptiles, having only cortical nephrons, produce urine that is, at best, isotonic to body fluids. Amphibians kidneys function much like those of fresh water fishes. When in fresh water, the skin of the frog accumulates certain salts from the water by active transport, and the kidneys excrete dilute urine., On land, where dehydration is the most pressing problem of osmoregulation, frogsconserve body fluid by reabsorbing water across the epithelium of the urinary bladder. Controlling the internal environment 99 Animal's nitrogenous wastes are correlated with its phylogeny and habitat Metabolism produces toxic by-products. The most troublesome is the nitrogencontaining waste from the metabolism of proteins and nucleic acids. The nitrogenous waste product is ammonia, a small and very toxic molecule. Excreting ammonia directly is bioenergetically efficient way to dispose of waste. Many organisms first convert ammonia to compounds such as urea or uric acids. Fig. 40.14 P897 Controlling the internal environment 100 Ammonia Most aquatic organisms excrete nitrogenous wastes as ammonia. Ammonia molecules are very soluble in water. In fishes, most of the ammonia is lost as ammonium ions (NH4+) across the epithelium of the gills.. In fresh water fishes, the epithelium of the gills takes up Na+ from the water in exchange for NH4+, which helps maintain concentration in the surrounding water. Urea Although it works in water, ammonia excretion is unsuitable for disposing of nitrogenous wastes on land. Mammals and most adult amphibians excrete urea. This substance can be tolerated in amuch more concentrated form because it is about 100,000 times less tocix than ammonia. Urea is produced in the liver by a metabolic cycle that combines ammonia and CO2. The circulatory system carries urea to the kidneys, where some of it is retained in kidneys, where it contributes to osmoregulation by helping maintain the osmolarity gradient that functions in water reabsorption. Amphibians that undergo metamorphosis generally switch from excreting ammonia to excreting urea during the transformation from an aquatic larva, the tadpole, to the terrestrial adult. This biochemical modification, however, is not always coupled with metamorphosis. Uric acid Land snails, insects, birds, and many reptiles excrete uric acid as the major nitrogenous waste. (Uric acid is less soluble in water than either ammonia and urea. In birds and reptiles, urine is eliminated in pastelike form along with faeces. Controlling the internal environment 101 Thermoregulation maintains body temperature within a range conductive to metabolism Thermoregulation is the maintenance of body temperature within a range that enables cells to function efficiently. For example, the rate of cellular respiration increases with temperature up to a certain point, and then declines when temperatures are high enough to begin denaturing enzymes. The properties of membranes also change with temperature. Heat transfer between organisms and their surroundings An organism exchanges heat with its external environment by four physical processes: conduction, convection, radiation, and evaporation. Conduction is the direct transfer of thermal motion (heat) between molecules of the environment and those of the body surface. Example: animal sits in a pool of cold water or on a hot rock. Heat will always be conducted from a body of higher temperature to one of lower temperature. Water is 50 to 100 times more effective than air in conducting heat. Convection is the transfer of heat by the movement of air or liquid past the surface of abody, as when a breeze contributes to heat loss from the surface of an animal with dry skin. Radiation is the emission of electromagnetic waves produced by all objects warmer than absolute zero, including an animal's body and the sun. example organism absorb heat from the sun. Evaporation is the loss of heat from the surface of a liquid that is losing some of its molecules as gas. Evaporation of water from an animal has a significant cooling effect on the animal's surface. Controlling the internal environment 102 Ectotherms derive body heat mainly from their surroundings and endotherms derive it mainly from metabolism An ectotherm warms it s body mainly by absorbing heat from its surroundings. Most invertebrates, fishes, amphibians, and reptiles are ectotherms. In contrast, an endotherm derives most or all of its body heat from its own metabolism. Mammals, birds some fishes, and numerous insects are endotherms. Many endothermsmaintain a consistent internal temperature even as the temperature of their surroundings fluctuates (Fig. 40.15) P899. Also, the term cold-blooded and warm-blooded are misleading. Many lizards, which are ectotherms, have higher body temperatures when active than mammals. So, the terms ectotherm and endotherm are not based on body temperature but rather on the main source of body heat. Controlling the internal environment 103 Thermoregulation involves physiological and behavioral adjustments Both ectothermic and endothermic organisms thermoregulate using some combination of up to four general categories of adaptations. 1. Adjusting the rate of heat exchange between the organism and its surroundings. Body insulation, such as hair, feathers, and fat located just beneath the skin, reduce an organism's heat loss. Other mechanisms that regulate heat exchange usually involve adaptations of the circulatory system. Example many endotherms and some ectotherms can alter the amount of blood flowing to their skin (vasodilation and vasoconstriction). Another type of adaptation that alters heat exchange is a special arrangement of arteries and veins called a countercurrent heat exchanger (fig. 40.16)P 900. 2. Cooling by evaporative heat loss. Terrestrial endotherms and ecotherms lose water in breathing and across their skin. If the humidity of the air is low enough, the water will evaporate and the organism will lose heat by evaporative cooling. Evaporation from the respiratory system can be increased by panting. Evaporative cooling via the skin can be increased by such means as sweating in mammals. 3. Behavioral responses. Many organisms can increase or decrease body heat loss by relocating. They will bask in the sun or on warm rocks in water; fine cool, damp areas or burrow in summer; or even migrate to a more suitable climate (Fig. 40.17.) P901. 4. Changing the rate of metabolic heat production. This is applied to endotherms, particularly mammals and birds. Many species of mammals and birds can double or triple their metabolic heat production when exposed to cold. Controlling the internal environment 104 Controlling the internal environment 105 Feedback Mechanisms in thermoregulation The regulation of body temperature in humans is an example of a complex homeostatic system facilitated by feedback mechanisms. Nerve cells that control thermoregulation, are concentrated in the hypothalamus. The hypothalamus serves as a thermostat, responding to changes in body temperature above and below a set point by activating mechanisms that promote heat loss or gain (Fig. 40.21) P905. Nerve cells that sense body temperatures are located in the skin, the hypothalamus itself, and some other parts of the nervous system. Some of these are warm receptors that signal the thermostat in the hypothalamus when the temperature of the skin or blood increases. Others are cold receptors that signal the thermostat when the temperature decreases. Responding to body temperature below the normal range, the thermostat inhibits heat-loss mechanisms and activates heat-saving ones such as the vasoconstriction of superficial vessels and the erection of fur, while stimulating heat-generating mechanisms (shivering and nonshivering thermogensesis). In response to elevated body temperature, the thermostat shuts down heat-saving mechanisms and promotes body cooling by vasodilation, sweating, or panting. Torpoer: Conseving energy during environmental extremes Torpor is an alternative physiological state in which metabolism decreases and the heart and respiratory system slow down. Many endotherms enter a state of torpor in which their body temperature declines. Hibernation is long-term torpor during which the body temperature is lowered as an adaptation to winter cold and food scaricity. Aestivation, or summer torpor, is characterized by slow metabolism and inactivity. It enables an animal to survive long periods of high temperatures and scarce water supplies. Many small mammals and birds exhibit a daily period of torpor that seems to be adapted to their feeding patterns. Example, most bats and shrews feed at night and go into torpor when they are inactive during daylight hours. Controlling the internal environment 106