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S E C T I O N 8.1 The Task of Respiration E X P E C TAT I O N S Describe gas exchange in simple organisms. Explain why some organisms do not need specialized respiratory systems. List the respiratory adaptations that enable organisms to live on land. Describe the main types of animal respiratory systems. Figure 8.1 Single-celled organisms, like the amoeba shown here, do not require a specialized respiratory system. Like most organisms on earth, the amoeba is aerobic — that is, it requires oxygen to survive. Whether an aerobic organism is unicellular, like the amoeba, or multicellular, each cell in the organism must have a supply of oxygen. Oxygen is necessary to carry out cellular respiration, the process that releases the energy needed to drive all cell functions. Each cell must also rid itself of the carbon dioxide that is the waste product of cellular respiration. The basic function of the respiratory system is to make sure that oxygen can enter each cell in the organism, and that carbon dioxide can leave each cell. This process is known as gas exchange. REWIND To review the concept of cellular respiration, turn to Chapter 3, Section 3.3. Gas Exchange at Its Simplest Most single-celled aerobic organisms do not have a distinct respiratory system. Such organisms, including protists, algae, fungi, and some bacteria, rely instead on diffusion to meet their gas exchange requirements. These organisms need moist environments, and are found either in aquatic environments or in other moist places such as within the body of a host organism. Oxygen, which is dissolved in the water around the cell, diffuses through the outer membrane of each cell and thereby becomes available for cellular respiration. At the same time, carbon dioxide diffuses out. For these organisms, the cell membrane itself provides a moist surface area big enough to accommodate the organism’s gas exchange requirements. Math Different organisms have different kinds of respiratory systems to accomplish the task of gas exchange. Every respiratory system, however, shares two requirements. First, the surface area available for gas exchange, or respiratory surface, must be big enough for the exchange of oxygen and carbon dioxide to occur at a rate that will meet the organism’s metabolic needs. Second, respiration must take place in a moist environment, so that the oxygen and carbon dioxide are dissolved. The respiratory systems you will examine in this chapter all demonstrate variations of these two basic features. 250 MHR • Internal Systems and Regulation LINK A number of factors affect the rate of diffusion of dissolved gases. These include the surface area available for diffusion, the concentration gradient, the size of the diffusing molecules, and the length of the diffusion path. In one set of experiments, a group of researchers found that, under a particular set of conditions, a molecule of oxygen dissolved in water diffused 1 µm in 0.0001 s, but took 100 s to diffuse 1 mm. Write an equation showing the relationship between the diffusion time and the length of the diffusion path. Now estimate the distance from your nose to your writing hand when your arm is fully extended. How long would it take oxygen to diffuse that distance, assuming the same conditions as in the experiment above? Colonial algae, such as volvox, and simple animal organisms such as hydra (see Figure 8.2A and B) also lack specialized tissues for gas exchange. Although they are multi-celled organisms, almost all of the cells of these organisms are in direct contact with the surrounding aquatic medium. As a result, each individual cell can obtain sufficient oxygen through diffusion. In contrast to volvox or hydra, the worm-like planarian (see Figure 8.2C) has all three germ layers, or tissue types, found in higher animals. This tiny creature can still manage without a respiratory system, however, because of its thin, flat shape. A planarian’s body is at most only a few cells deep, so oxygen has a short distance to travel in order to diffuse into every living cell. At the same time, the width of the body provides a large surface area for gas exchange. B C Figure 8.2 Volvox (A), hydra (B) and planarian (C). These simple organisms can also depend on diffusion alone to meet their gas exchange requirements. A Unicellular Terrestrial Organisms Some unicellular terrestrial organisms also lack a distinct respiratory system. Bacteria and fungi living within the soil, as well as terrestrial protists, depend on the air in the tiny spaces in the soil for their source of oxygen. The organism’s requirement for a moist respiratory surface is satisfied by water derived from the soil solution. As long as there is water in the soil, the air spaces will remain humid enough for the cell membrane to stay moist. Many bacteria and fungi live on the surface of the soil, often in association with organic debris. These organisms depend on simple diffusion of gases to and from the surrounding air. As in their aquatic or soil-borne counterparts, this exchange takes place directly across the cell membrane. In general, it is the availability of moisture (either as soil moisture or as water vapour in the surrounding air) that restricts the habitat of such organisms. Figure 8.3 This environment supports many unicellular terrestrial organisms. The Breath of Life • MHR 251 The Specialized Respiratory System The larger an organism is, the more oxygen it requires to supply its cells, and the greater the distance over which that oxygen must travel in order to reach all of the cells. This raises two problems. First, diffusion of oxygen is only effective over a distance of a few cells, so an organism with a body more than a few cells thick must use another mechanism to bring oxygen to all its cells. Second, as body parts become differentiated for uses such as locomotion or reproduction, the body surface that can be dedicated to gas exchange is reduced. Different kinds of adaptations have made it possible for larger, more complex organisms to meet their need for oxygen. All of the higher animals have evolved a specialized respiratory system. An animal’s respiratory system includes a respiratory surface, any passageways that connect this surface to the external environment, and any muscular structures that are used to help bring the respiratory medium into contact with the respiratory surface. The following pages examine several types of specialized respiratory structures found in animals. PAUSE mouth carbon dioxide diffuses out oxygen diffuses in network of blood vessels and capillaries lying beneath the skin gut Figure 8.4 The earthworm makes use of its moist skin surface to exchange gases between its circulatory system and its environment. The exchange of gases across the skin of the animal is called skin respiration. As in volvox and planaria, the gas exchange occurs across the organism’s entire body surface. The crucial difference is that the earthworm uses a circulatory system to carry oxygen to the cells that could not rely on diffusion to meet their needs. RECORD Multicellular animals have evolved ways of storing nutrients and, to a lesser extent, water in their bodies. Your own body can survive for several days without water, and for weeks without food, but only for a few minutes without oxygen. What factors might prevent your body from storing oxygen? List your ideas and explain briefly. Hint: what does the term “oxidization” mean? Skin Respiration One adaptation is demonstrated by animals in the phylum Annelida, which includes the segmented worms such as earthworms and leeches. Segmented worms can survive only in water or in damp earth, because their skin must remain moist. To meet their oxygen requirements, earthworms make use of the moist surface and their simple circulatory mechanism. The skin is lined with many tiny capillary vessels, and these capillaries make contact with the skin surface all over the body of the worm. At each of these points of contact, respiratory gases diffuse into and out of the circulatory vessels (Figure 8.4). The oxygen is then carried through the circulatory vessels to other parts of the organism. 252 MHR • Internal Systems and Regulation Figure 8.5 The aquatic tube worm cannot exchange gases through its protective casing. Gills In contrast to the earthworm, whose whole body surface is available for respiration, most of the body of a tube worm is covered with a protective casing. Many other aquatic animals also have protective coats. These coats provide defence against predators or help maintain the animals’ internal environments. In such organisms the relative area over which diffusion of respiratory Figure 8.6 The fish draws water through its mouth and over its gills. As water passes over the gills, oxygen diffuses into the adjacent blood vessels. At the same time, carbon dioxide diffuses out. gills water and oxygen gill water and carbon dioxide artery to gills artery from gills oxygenated blood deoxygenated blood water flow gill filament gases can occur is considerably reduced. Two adaptations help these organisms accomplish the task of respiration. First, structural changes have increased the surface area of the body part involved in gas exchange. Second, a mechanism has evolved that enables the animal to ventilate this surface — that is, to move the oxygen-containing aquatic medium over the respiratory surface. This allows the respiratory surface to be constantly exposed to a fresh supply of oxygen. Many aquatic organisms, including molluscs, crayfish, mud puppies, tadpoles, and fish, have developed gills, which are feathery tissue structures that invariably consist of numerous delicate branches. When the gills are exposed to water, gases are exchanged across the thin gill membranes. The oxygen diffuses into the organism’s vascular system for circulation around the body. Gills ensure that a considerable surface area is available for gas exchange in a very limited space. Gilled animals have evolved various means of ventilation. In some animals, such as the tube worm, the gill is moved through the water. In others, such as fish and clams, water is drawn or pumped over the gill. In most gilled animals, the water flows only one way over the gills. This reduces the amount of energy the animal must expend to move the water over the respiratory surface. Figure 8.6 shows the respiratory organ of a typical fish. Water moves in through the mouth and out behind the animal’s head, passing over the gills on the way. Adjacent to the gills are a network of tiny capillaries that absorb oxygen and release carbon dioxide from the animal’s circulatory system. The ventilation mechanisms of some aquatic animals are linked with other functions. In a number of species, including many molluscs, feeding and ventilation are accomplished together. Food particles are captured as the gill moves through the water, and are filtered out by the organism at the same time that gas exchange takes place. Other animals use the currents produced by respiratory action for locomotion. A squid, for example, can move rapidly forward or backward by forcing water out of its gill siphon. Moving out of the Water Given how much equipment we need in order to breathe under water, it may be hard to imagine that, from an evolutionary perspective, respiration in air is actually the bigger challenge. In the water, the need for a respiratory surface that stays moist is rarely a problem. On land, however, moisture from a respiratory surface exposed to the air will evaporate, so the animal would have to release more water constantly to keep the surface moist. A terrestrial animal that made a large moist surface area available for gas exchange would therefore risk excessive water loss. Before living organisms could Figure 8.7 People must take air with them in order to breathe under water, along with special equipment to dispense it. The Breath of Life • MHR 253 begin to colonize terrestrial environments, adaptations were needed that could solve this critical problem. In much the same way that scuba divers take some of their usual respiratory medium (air) along on the dive, all terrestrial animals have evolved some way to carry a moist respiratory environment wherever they go. In the previous paragraphs, you learned that most aquatic animals have developed some way to ventilate their respiratory surface. The same is true of most terrestrial animals. The act of ventilating a respiratory surface with air is called breathing. The breathing of terrestrial organisms relies on a basic law of physics: air will move from a region of higher pressure to a region of lower pressure until equilibrium is reached (Figure 8.8). Different terrestrial animals have evolved different ways to make use of this principle in their respiratory systems. both containers same size, same pressure The Tracheal Respiratory System In insects, the problem of how to maintain a moist respiratory surface without losing too much water is resolved by an internal respiratory system that consists of a series of external pores called spiracles. The spiracles lead to an internal network of tubes called tracheae (singular trachea). All of the spiracles are controlled by valves. Although the specific configuration of spiracles and tracheae varies considerably from one species to the next, the basic elements of the tracheal system are common to most insects and many other nonvertebrate terrestrial animals. If you can, observe a living grasshopper. Note that the abdomen contracts and relaxes rhythmically. These are the breathing movements of the animal. As described in Figure 8.9, the animal’s breathing causes an orderly movement of air through the tracheae, from the front of the animal to the back. trachaeole O2 A At equilibrium, air pressure is equal in both vessels. left container larger, lower pressure; air moves in cell O2 air + O2 spiracle (pore through skin) A As the abdomen expands, air pressure drops within the tracheae. At the same time, the anterior four pairs of spiracles open, while the posterior six pairs of spiracles remain closed. Air therefore flows in through the anterior spiracles. trachaeole B If the volume of one vessel increases, the air pressure inside it decreases. Air flows from the vessel with the higher pressure into the vessel with the lower pressure. CO2 CO2 cell same pressure in both (but lower than above) air + CO2 spiracle (pore through skin) B When the abdomen contracts, the reverse situation occurs. The anterior four pairs close and the posterior six pairs open. Air pressure is now higher inside the animal than in the surrounding environment, so air flows out through the posterior spiracles. C A new equilibrium is reached with air pressure once again equal in each vessel. Figure 8.8 Air will move from a region of higher pressure to a region of lower pressure. 254 MHR • Internal Systems and Regulation Figure 8.9 The tracheal respiratory system of a grasshopper In insects, the respiratory and circulatory systems are separate from one another. The tracheal system of an insect does not require a circulatory system to transport the respiratory gases. Instead, the many branches of the tracheal tubes ensure that the respiratory surface is in close enough contact with all of the living cells to allow gas exchange to occur by diffusion across the moist tracheal walls. Because most of the tracheal system remains inside the animal, the risk of excessive water loss is reduced. lining of the mouth. This explains why a frog cannot live for any extended period away from either an aquatic or a very humid environment. Because it uses its skin as a respiratory surface, it has no satisfactory means of controlling water loss through the skin. The Lung The third general type of respiratory system, and the one that is characteristic of air-breathing vertebrates, is the internal lung arrangement. The lung is an internal respiratory surface connected to the air by means of internal passageways. Lung systems vary from species to species in terms of both structure and efficiency. However, all lung systems consist of three basic elements: one or two lungs that have a moist respiratory surface; some means of forcibly bringing air in contact with the lung surface; and a circulatory system to carry the gases between the lungs and the other cells of the body. Figure 8.10 outlines the various general lung types found in a number of animal species. The frog uses a simple lung system ventilated by the movement of muscles in the nostrils, mouth, and abdomen. To breathe in, the frog first closes its mouth and opens its nostrils or nares, then lowers the floor of the mouth, drawing air into the mouth cavity. The nostrils are then closed, and the floor of the mouth raised. As a result the air is forced into the lungs where gas exchange takes place. In the frog, as in other amphibians, the lung system is used in conjunction with both skin respiration and the exchange of gases across membranes in the SECTION Figure 8.10 Examples of the types of lungs found in animals such as amphibians, spiders, and reptiles. Spiders’ lungs are often referred to as “book lungs” because their many folds are arranged like the pages of a book. Snakes and other reptiles that flourish on land (and even in desert environments) do not use the skin as a site for the exchange of gases. They confine this exchange to the internal lung or lungs. All these organisms possess muscle and skeletal arrangements that enable them to force air out of the lungs and draw air back over the lung surface to ventilate the respiratory tract. REVIEW 1. Describe how gas exchange takes place in an amoeba. 2. K/U Describe two ways in which the problem of getting respiratory gases to individual cells has been overcome in larger, more complex organisms. 3. Imagine an argument between a tube worm and a grasshopper about the best way to respire. Working with a partner, try to create an imaginary dialogue between the two animals (assume that the two animals can speak the same language). Where are the greatest misunderstandings likely to arise? How would you suggest the two settle their argument? K/U C lungs 4. 5. Populations of grasshoppers and locusts occasionally skyrocket, creating plagues that can cause considerable damage to farm crops. How could knowledge of the way grasshoppers breathe be used to control their numbers? MC K/U Can a frog respire under water? Why or why not? 6. Account for the fact that you see many earthworms on the ground after a rainstorm. 7. I A new species of insect is discovered. This insect has 12 pairs of spiracles along its abdomen. Design an experiment to test whether the spiracles work in concert, in sequence, or some type of intermediate mechanism. K/U The Breath of Life • MHR 255