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Human Physiology An Integrated Approach 6/E PDF

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17 Mechanics of Breathing The Respiratory System Bones and Muscles of the Thorax Surround the Lungs Pleural Sacs Enclose the Lungs Airways Connect Lungs to the External Environment The Airways Warm, Humidify, and Filter Inspired Air Alveoli Are the Site of Gas Exchange Pulmonary Circulation Is High-Flow, Low-Pressure Gas Laws Air Is a Mixture of Gases Gases Move Down Pressure Gradients Boyle’s Law Describes Pressure-Volume Relationships Ventilation Lung Volumes Change During Ventilation During Ventilation, Air Flows Because of Pressure Gradients Inspiration Occurs When Alveolar Pressure Decreases Expiration Occurs When Alveolar Pressure Increases Intrapleural Pressure Changes During Ventilation Lung Compliance and Elastance May Change in Disease States Surfactant Decreases the Work of Breathing Airway Diameter Determines Airway Resistance Rate and Depth of Breathing Determine the Effi ciency of Breathing Gas Composition in the Alveoli Varies Little During Normal Breathing Ventilation and Alveolar Blood Flow Are Matched Auscultation and Spirometry Assess Pulmonary Function This being of mine, whatever it really is, consists of a little fl esh, a little breath, and the part which governs. — Marcus Aurelius Antoninus ( C . E . 121–180) B ackground Basics Ciliated and exchange epithelia Pressure, volume, fl ow, and resistance Pulmonary circulation Colored x-ray Surface tension of the lung Autonomic and somatic showing the motor neurons branching Velocity of fl ow airways. 600 Mechanics of Breathing I magine covering the playing surface of a racquetball court cavity to control their contact with the outside air. Internalization (about 75 m2) with thin plastic wrap, then crumpling up the creates a humid environment for the exchange of gases with the wrap and stuffi ng it into a 3-liter soft drink bottle. Impossible? blood and protects the delicate exchange surface from damage. Maybe so, if you use plastic wrap and a drink bottle. But the lungs Internalized lungs create another challenge, however: how to of a 70-kg man have a gas exchange surface the size of that plastic move air between the atmosphere and an exchange surface deep wrap, compressed into a volume that is less than that of the bottle. within the body. Air flow requires a muscular pump to create Th is tremendous surface area for gas exchange is needed to supply pressure gradients. More complex respiratory systems therefore the trillions of cells in the body with adequate amounts of oxygen. consist of two separate components: a muscle-driven pump and a A erobic metabolism in cells depends on a steady supply thin, moist exchange surface. In humans, the pump is the muscu- of oxygen and nutrients from the environment, coupled with loskeletal structure of the thorax. Th e lungs themselves consist of the removal of carbon dioxide. In very small aquatic animals, the exchange epithelium and associated blood vessels. simple diff usion across the body surface meets these needs. Dis- Th e four primary functions of the respiratory system are: tance limits diff usion rate, however, so most multicelled animals 1 E xchange of gases between the atmosphere and the blood. require specialized respiratory organs associated with a circula- Th e body brings in O for distribution to the tissues and tory system. Respiratory organs take a variety of forms, but all 2 eliminates CO waste produced by metabolism. possess a large surface area compressed into a small space. 2 2 H omeostatic regulation of body pH. The lungs can alter 17 Besides needing a large exchange surface, humans and body pH by selectively retaining or excreting CO . other terrestrial animals face an additional physiological chal- 2 3 P rotection from inhaled pathogens and irritating substances. lenge: dehydration. The exchange surface must be thin and Like all other epithelia that contact the external environ- moist to allow gases to pass from air into solution, and yet at ment, the respiratory epithelium is well supplied with de- the same time it must be protected from drying out as a result fense mechanisms to trap and destroy potentially harmful of exposure to air. Some terrestrial animals, such as the slug substances before they can enter the body. (a shell-less snail), meet the challenge of dehydration with be- 4 V ocalization. Air moving across the vocal cords creates havioral adaptations that restrict them to humid environments vibrations used for speech, singing, and other forms of and nighttime activities. communication. A more common solution is anatomical: an internalized respiratory epithelium. Human lungs are enclosed in the chest I n addition to serving these functions, the respiratory sys- tem is also a signifi cant source of water loss and heat loss from the body. These losses must be balanced using homeostatic compensations. RUNNING PROBLEM I n this chapter you will learn how the respiratory system Emphysema carries out these functions by exchanging air between the envi- ronment and the interior air spaces of the lungs. Th is exchange You could hear her whistling, wheezing breathing preceding is the b ulk fl ow of air, and it follows many of the same principles her down the hall. “Diagnosis: COPD,” reads Edna Wilson’s that govern the bulk fl ow of blood through the cardiovascular patient chart. COPD—chronic obstructive pulmonary disease—is the name given to diseases in which air system: exchange is impaired by narrowing of the lower airways. 1 F low takes place from regions of higher pressure to regions Most people with COPD have emphysema or chronic of lower pressure. bronchitis or a combination of the two. Individuals in whom 2 A muscular pump creates pressure gradients. chronic bronchitis predominates are sometimes called “blue 3 R esistance to air fl ow is infl uenced primarily by the diam- bloaters,” owing to the bluish tinge of their skin (from low eter of the tubes through which the air is fl owing. blood oxygen levels) and a tendency to be overweight. In contrast, patients with emphysema have been nicknamed A ir and blood are both fl uids. Th e primary diff erence be- “pink puff ers.” They tend to be thin, have normal (pink) skin tween air fl ow in the respiratory system and blood fl ow in the coloration, and often breathe out through pursed lips, which circulatory system is that air is a less viscous, compressible mix- helps open their airways. More than 12 million people in ture of gases while blood is a noncompressible liquid. the United States have COPD. Its most common cause is smoking, and most people can avoid the disease simply by not smoking. Unfortunately, Edna has been a heavy smoker for 35 of her 47 years. The Respiratory System T he word r espiration has several meanings in physiology ( Fig. 17.1 ). Cellular respiration refers to the intracellular reaction of oxygen with organic molecules to produce carbon 601 Mechanics of Breathing consists of structures involved in ventilation and gas exchange EXTERNAL RESPIRATION ( F ig. 17.2 ): The respiratory and circulatory systems coordinate to move oxygen 1 Th e c onducting system of passages, or airways , that lead and CO between the atmosphere and the cells. 2 from the external environment to the exchange surface of Exchange I: CO2 O2 the lungs. atmosphere 2 Th e alveoli (singular alveolus) {a lveus, a concave vessel}, to lung a series of interconnected sacs and their associated p ulmo- (ventilation) Airways nary capillaries. Th ese structures form the exchange sur- face, where oxygen moves from inhaled air to the blood, Alveoli of lungs and carbon dioxide moves from the blood to air that is CO2 O2 about to be exhaled. Exchange II: 3 Th e bones and muscles of the thorax (chest cavity) and ab- lung to blood CO2 O2 domen that assist in ventilation. Transport of Pulmonary Th e respiratory system can be divided into two parts. Th e gases in circulation the blood upper respiratory tract consists of the mouth, nasal cavity, pharynx, and larynx. Th e lower respiratory tract consists of the trachea, two primary bronchi { bronchos, windpipe; singular— bronchus}, their branches, and the lungs. Th e lower tract is also known as the thoracic portion of the respiratory system because Systemic it is enclosed in the thorax. circulation Bones and Muscles of the Thorax Exchange III: CO2 O2 Surround the Lungs blood to cells CO2 O2 Th e thorax is bounded by the bones of the spine and rib cage Cellular and their associated muscles. Together the bones and muscles Cells respiration are called the thoracic cage . Th e ribs and spine (the chest wall ) Nutrients form the sides and top of the cage. A dome-shaped sheet of skel- ATP etal muscle, the diaphragm , forms the fl oor ( Fig. 17.2 b). Two sets of intercostal muscles , internal and external, Fig. 17.1 connect the 12 pairs of ribs (F ig. 1 7.2a ). Additional muscles, the sternocleidomastoids and the s calenes , run from the head and neck to the sternum and fi rst two ribs. d ioxide, water, and energy in the form of ATP. External respi- F unctionally, the thorax is a sealed container fi lled with three ration, the topic of this chapter and the next, is the movement membranous bags, or sacs. One, the p ericardial sac, contains the of gases between the environment and the body’s cells. External heart. Th e other two bags, the p leural sacs, each surround a lung respiration can be subdivided into four integrated processes, il- {p leura, rib or side}. The esophagus and thoracic blood vessels lustrated in Figure 17.1 : and nerves pass between the pleural sacs ( Fig. 17.2 c). 1 Th e exchange of air between the atmosphere and the lungs. Pleural Sacs Enclose the Lungs Th is process is known as v entilation, or breathing. I nspi- ration (inhalation) is the movement of air into the lungs. Th e lungs ( Fig. 17.2 b, d) consist of light, spongy tissue whose Expiration (exhalation) is the movement of air out of the volume is occupied mostly by air-fi lled spaces. Th ese irregular lungs. Th e mechanisms by which ventilation takes place cone-shaped organs nearly fill the thoracic cavity, with their are collectively called the mechanics of breathing . bases resting on the curved diaphragm. Semi-rigid conducting 2 T he exchange of O and CO between the lungs and the 2 2 airways—the bronchi—connect the lungs to the main airway, blood. the trachea. 3 Th e transport of O and CO by the blood. 2 2 E ach lung is surrounded by a double-walled pleu- 4 Th e exchange of gases between blood and the cells. ral sac whose membranes line the inside of the thorax and External respiration requires coordination between the re- cover the outer surface of the lungs ( Fig. 17.3 ). Each pleu- spiratory and cardiovascular systems. Th e respiratory system ral membrane, or p leura, contains several layers of elastic 602 Mechanics of Breathing c onnective tissue and numerous capillaries. Th e opposing layers Concept Check Answers: End of Chapter of pleural membrane are held together by a thin fi lm of pleu- ral fl uid whose total volume is only about 25–30 mL in a 70-kg 1. What is the diff erence between cellular respiration and external man. Th e result is similar to an air-fi lled balloon (the lung) sur- respiration? rounded by a water-fi lled balloon (the pleural sac). Most illus- 2. Name the components of the upper respiratory tract and those of the trations exaggerate the volume of the pleural fl uid, but you can lower respiratory tract. appreciate its thinness if you imagine spreading 25 mL of water 3. Based on the total cross-sectional area of diff erent airways, where is evenly over the surface of a 3-liter soft drink bottle. the velocity of air fl ow highest and lowest? Pleural fluid serves several purposes. First, it creates a moist, slippery surface so that the opposing membranes can 4. Give two functions of pleural fl uid. slide across one another as the lungs move within the thorax. 5. N ame the components (including muscles) of the thoracic cage. List the Second, it holds the lungs tight against the thoracic wall. To vi- contents of the thorax. sualize this arrangement, think of two panes of glass stuck to- 6. Which air passages of the respiratory system are collapsible? gether by a thin fi lm of water. You can slide the panes back and forth across each other, but you cannot pull them apart because of the cohesiveness of the water. A similar fl uid bond between 17 the two pleural membranes makes the lungs “stick” to the tho- The Airways Warm, Humidify, racic cage and holds them stretched in a partially infl ated state, and Filter Inspired Air even at rest. During breathing, the upper airways and the bronchi do more than simply serve as passageways for air. Th ey play an important A irways Connect Lungs to the role in conditioning air before it reaches the alveoli. Condition- External Environment ing has three components: 1 W arming air to body temperature (37 (cid:2)C ), so that core Air enters the upper respiratory tract through the mouth and body temperature does not change and alveoli are not nose and passes into the p harynx, a common passageway damaged by cold air; for food, liquids, and air {p harynx, throat}. From the pharynx, 2 A dding water vapor until the air reaches 100% humidity, so air flows through the larynx into the trachea , or windpipe that the moist exchange epithelium does not dry out; and ( Fig. 1 7.2 b). The larynx contains the v ocal cords , connective 3 F iltering out foreign material, so that viruses, bacteria, and tissue bands that vibrate and tighten to create sound when air inorganic particles do not reach the alveoli. moves past them. T he trachea is a semiflexible tube held open by 15 to 20 I nhaled air is warmed by the body’s heat and moistened by C-shaped cartilage rings. It extends down into the thorax, where water evaporating from the mucosal lining of the airways. Un- it branches (division 1) into a pair of p rimary bronchi , one der normal circumstances, by the time air reaches the trachea, bronchus to each lung (F ig. 1 7.2b ). Within the lungs, the bronchi it has been conditioned to 100% humidity and 37 (cid:2)C . Breath- branch repeatedly (divisions 2–11) into progressively smaller ing through the mouth is not nearly as eff ective at warming and bronchi (F ig. 1 7.2e ). Like the trachea, the bronchi are semi-rigid moistening air as breathing through the nose. If you exercise tubes supported by cartilage. outdoors in very cold weather, you may be familiar with the Within the lungs, the smallest bronchi branch to become ache in your chest that results from breathing cold air through bronchioles, small collapsible passageways with walls of smooth your mouth. muscle. Th e bronchioles continue branching (divisions 12–23) Air is filtered both in the trachea and in the bronchi. until the respiratory bronchioles form a transition between the These airways are lined with ciliated epithelium whose cilia airways and the exchange epithelium of the lung. are bathed in a watery saline layer ( F ig. 17.5 ). The saline is T he diameter of the airways becomes progressively produced by epithelial cells when Cl- secreted into the lumen smaller from the trachea to the bronchioles, but as the individ- by apical anion channels draws Na+ into the lumen through the ual airways get narrower, their numbers increase geometrically paracellular pathway ( Fig. 17.5 c). Movement of solute from the ( F ig. 17.4 ). As a result, the total cross-sectional area in- ECF to the lumen creates an osmotic gradient, and water follows creases with each division of the airways. Total cross- the ions into the airways. Th e CFTR channel, whose malfunc- sectional area is lowest in the upper respiratory tract and tion causes cystic fi brosis, is one of the anion channels found on greatest in the bronchioles, analogous to the increase in cross- the apical surface of this epithelium. sectional area that occurs from the aorta to the capillaries in A sticky layer of mucus fl oats over the cilia to trap most the circulatory system. inhaled particles larger than 2 mm . Th e mucus layer is secreted 603 Fig. 17.2 ANATOMY SUMMARY The Lungs and Thoracic Cavity (a) Muscles of the thorax, neck, and abdomen (b) The respiratory system is divided create the force to move air during breathing. into upper and lower regions. Pharynx Nasal cavity Upper Vocal cords Sternocleido- respiratory mastoids Tongue system Scalenes Esophagus Larynx Trachea Lower respiratory system External Internal intercostals intercostals Diaphragm Abdominal muscles Left lung Right lung Muscles Muscles Diaphragm of inspiration of expiration Right bronchus Left bronchus (c) Sectional view of chest. Each lung is enclosed in (d) On external view, the right lung is divided two pleural membranes. The esophagus and aorta into three lobes, and the left lung is pass through the thorax between the pleural sacs. divided into two lobes. Apex Pleural Esophagus Aorta membranes Superior lobe Superior lobe Right Left lung lung Middle lobe Heart Inferior Inferior lobe lobe Right pleural Pericardial Left pleural cavity cavity cavity Base Cardiac notch Superior view 604 The Bronchi and Alveoli (e) Branching of airways creates (f) Structure of lung lobule. Each cluster of alveoli is about 80 million bronchioles. surrounded by elastic fibers and a network of capillaries. Larynx Bronchiole Branch of pulmonary artery Bronchial artery, Smooth muscle nerve and vein The trachea branches into Trachea two primary bronchi. Branch of Elastic Left primary pulmonary fibers Cartilage bronchus vein ring Capillary Lymphatic beds vessel The primary bronchus divides 22 more times, terminating in a cluster Secondary of alveoli. bronchus Alveoli Bronchiole Alveoli (g) Alveolar structure (h) Exchange surface of alveoli Capillary Elastic fibers Alveolar Nucleus of epithelium endothelial cell RBC Type I alveolar cell for gas exchange Capillary Endothelium Plasma Endothelial cell of capillary 0.1- 1.5 Type II alveolar μm cell (surfactant cell) synthesizes Alveolar surfactant. air space Alveolus Surfactant Fused Limited basement interstitial membranes fluid Alveolar Blue arrow represents gas exchange macrophage between alveolar air space and the plasma. ingests foreign material. 605 Mechanics of Breathing THE PLEURAL SAC Concept Check A nswers: End of Chapter The pleural sac forms a double membrane surrounding the lung, 7. Cigarette smoking paralyzes cilia in the airways and increases mucus similar to a fluid-filled balloon surrounding an air-filled balloon. production. Why would these eff ects cause smokers to develop a cough? Pleural membrane Air-filled Air space balloon of lung Alveoli Are the Site of Gas Exchange Fluid-filled balloon The pleural fluid has a much Th e alveoli, clustered at the ends of terminal bronchioles, make smaller volume than is up the bulk of lung tissue (F ig. 1 7.2f , g). Th eir primary function suggested by this illustration. is the exchange of gases between themselves and the blood. Fig. 17.3 E ach tiny alveolus is composed of a single layer of epithelium ( Fig. 17.2 g). Two types of epithelial cells are found in the alveoli. The smaller but thicker type II alveolar cells by goblet cells in the epithelium ( Fig. 17.5 b). Th e cilia beat with synthesize and secrete a chemical known as s urfactant. an upward motion that moves the mucus continuously toward Surfactant mixes with the thin fl uid lining of the alveoli to aid the pharynx, creating what is called the mucociliary escalator . lungs as they expand during breathing, as you will see later in Mucus contains immunoglobulins that can disable many patho- this chapter. Type II cells also help minimize the amount of fl uid gens. Once mucus reaches the pharynx, it can be spit out (e xpec- present in the alveoli by transporting solutes, followed by water, torated ) or swallowed. For swallowed mucus, stomach acid and out of the alveolar air space. enzymes destroy any remaining microorganisms. T he larger t ype I alveolar cells occupy about 95% of Secretion of the watery saline layer beneath the mucus is the alveolar surface area and are very thin so that gases can dif- essential for a functional mucociliary escalator. In the disease fuse rapidly through them (F ig. 1 7.2h ). In much of the exchange cystic fi brosis, for example, inadequate ion secretion decreases area, a layer of basement membrane fuses the alveolar epithe- fl uid movement in the airways. Without the saline layer, cilia be- lium to the capillary endothelium. In the remaining area only a come trapped in thick, sticky mucus. Mucus cannot be cleared, small amount of interstitial fl uid is present. and bacteria colonize the airways, resulting in recurrent lung The thin walls of the alveoli do not contain muscle be- infections. cause muscle fi bers would block rapid gas exchange. As a result, BRANCHING OF THE AIRWAYS Cross-sectional Name Division Diameter (mm) How many? area (cm2) Conducting system Trachea 0 15-22 1 2.5 Primary bronchi 1 10-15 2 Smaller 2 4 bronchi 3 4 1-10 5 6-11 1 x 104 2 x 104 100 Bronchioles 12-23 0.5-1 Exchange surface 8 x 107 5 x 103 Alveoli 24 0.3 3-6 x 108 >1 x 106 Fig. 17.4 606 Mechanics of Breathing AIRWAY EPITHELIUM (a) Epithelial cells lining the airways and submucosal (b) Cilia move the mucus layer toward the pharynx, removing trapped glands secrete saline and mucus. pathogens and particulate matter. Dust particle Ciliated Mucus layer traps epithelium inhaled particles. Watery saline layer allows cilia to push mucus toward pharynx. Cilia Goblet cell secretes mucus. Movement of mucus Nucleus of columnar 17 Mucus layer Submucosal epithelial cell Lumen of airway gland Basement membrane (c) One model of saline secretion by airway epithelial cells Sina lluinmee lanyer Na+ H2O Cl– 2 1 NKCC brings Cl– into epithelial cell from ECF. Anion Respiratory channel epithelial 2 Apical anion channels, cells including CFTR, allow Cl– to enter the lumen. 3 Na+ goes from ECF to lumen by the paracelllular pathway, drawn by the electrochemical gradient. K+ ATP 4 NaCl movement from ECF to 3 Na+ Na+ Na+2Cl–K+ K+ lumen creates a concentration H O gradient so water follows into ECF 2 1 the lumen. 4 Fig. 17.5 lung tissue itself cannot contract. However, connective tis- proximity of capillary blood to alveolar air is essential for the sue between the alveolar epithelial cells contains many elastin rapid exchange of gases. and collagen fi bers that create elastic recoil when lung tissue is stretched. Pulmonary Circulation Th e close association of the alveoli with an extensive net- Is High-Flow, Low-Pressure work of capillaries demonstrates the intimate link between the respiratory and cardiovascular systems. Blood vessels fi ll 80– Th e pulmonary circulation begins with the pulmonary trunk, 90% of the space between alveoli, forming an almost continuous which receives low-oxygen blood from the right ventricle. Th e “sheet” of blood in close contact with the air-fi lled alveoli. Th e pulmonary trunk divides into two pulmonary arteries, one to 607 Mechanics of Breathing CLINICAL FOCUS Concept Check Answers: End of Chapter 8. Is blood fl ow through the pulmonary trunk greater than, less than, or Congestive Heart Failure equal to blood fl ow through the aorta? When is a lung problem not a lung problem? The 9. A person has left ventricular failure but normal right ventricular answer: when it’s really a heart problem. Congestive heart function. As a result, blood pools in the pulmonary circulation, failure (CHF) is an excellent example of the interrelationships doubling pulmonary capillary hydrostatic pressure. What happens to among body systems and demonstrates how disruptions net fl uid fl ow across the walls of the pulmonary capillaries? in one system can have a domino eff ect in the others. The 10. Calculate the mean pressure in a person whose pulmonary arterial primary symptoms of heart failure are shortness of breath pressure is 25 8 mm Hg. ( dyspnea ), wheezing during breathing, and sometimes a > productive cough that may be pinkish from the presence of blood. Congestive heart failure arises when the right heart is a more eff ective pump than the left heart. When Gas Laws blood accumulates in the pulmonary circulation, increased volume increases pulmonary blood pressure and capillary Respiratory air fl ow is very similar in many respects to blood hydrostatic pressure. Capillary fi ltration exceeds the ability fl ow in the cardiovascular system because both air and blood of the lymph system to drain interstitial fl uid, resulting in pulmonary edema. Treatment of CHF includes increasing are fluids. Their primary difference is that blood is a non- urinary output, which brings yet another organ system into compressible liquid but air is a compressible mixture of gases. the picture. By current estimates, about 5 million Americans Figure 17.6 summarizes the laws that govern the behavior of suff er from CHF. To learn more about this condition, visit the gases in air and provide the basis for the exchange of air between American Heart Association web site ( www.americanheart. the external environment and the alveoli. org ) or MedlinePlus, published by the National Institutes of In this course, blood pressure and environmental air pres- Health ( www.nlm.nih.gov/medlineplus/heartfailure.html ). sure (a tmospheric pressure ) are both reported in millimeters of mercury (mm Hg). Respiratory physiologists sometimes report gas pressures in centimeters of water instead, where 1 mm Hg = 1.36 cm H O , or in kiloPascals (kPa), where 760 mm Hg = 2 each lung. Oxygenated blood from the lungs returns to the left 101.325 kPa. atrium via the pulmonary veins. A t sea level, normal atmospheric pressure is 760 mm Hg. A t any given moment, the pulmonary circulation contains However, in this course we follow the convention of designating about 0.5 liter of blood, or 10% of total blood volume. About 75 mL atmospheric pressure as 0 mm Hg. Because atmospheric pres- of this amount is found in the capillaries, where gas exchange sure varies with altitude and because very few people live ex- takes place, with the remainder in pulmonary arteries and veins. actly at sea level, this convention allows us to compare pressure Th e rate of blood fl ow through the lungs is much higher than the diff erences that occur during ventilation without correcting for rate in other tissues because the lungs receive the entire cardiac altitude. output of the right ventricle: 5 L min. Th is means that as much > blood fl ows through the lungs in one minute as fl ows through the entire rest of the body in the same amount of time! RUNNING PROBLEM Despite the high fl ow rate, pulmonary blood pressure is low. Edna has not been able to stop smoking, and her COPD Pulmonary arterial pressure averages 25 8 mm Hg, much lower > is a combination of emphysema and bronchitis. Patients than the average systemic pressure of 120 80 mm Hg. Th e right > with chronic bronchitis have excessive mucus production ventricle does not have to pump as forcefully to create blood fl ow and exhibit general infl ammation of the entire respiratory through the lungs because resistance of the pulmonary circulation tract. The mucus narrows the airways and makes breathing is low. Th is low resistance can be attributed to the shorter total diffi cult. length of pulmonary blood vessels and to the distensibility and large total cross-sectional area of pulmonary arterioles. Q1: What does narrowing of the airways do to airway resistance? N ormally, the net hydrostatic pressure fi ltering fl uid out of a pulmonary capillary into the interstitial space is low because of low mean blood pressure. Th e lymphatic system effi ciently re- moves fi ltered fl uid, and lung interstitial fl uid volume is usually minimal. As a result, the distance between the alveolar air space and the capillary endothelium is short, and gases diff use rapidly between them. 608 Fig. 17.6 ESSENTIALS Gas Laws This figure summarizes the rules that govern the behavior of gases in air. These rules provide the basis for the exchange of air between the external environment and the alveoli. (a) The ideal gas equation Where P is pressure, V is volume, n is the moles of gas, T is absolute PV = nRT temperature, and R is the universal gas constant, 8.3145 j/mol × K In the human body we can assume that the number of moles and temperature are constant. Removing the constants leaves the following equation: This relationship says that if the volume of gas V = 1/P increases, the pressure decreases, and vice versa. (b) Boyle’s Law Boyle’s law also expresses this inverse relationship between pressure and volume. For example, the container on the left is 1 L (V ) 1 P V = P V and has a pressure of 100 mm Hg (P ). 1 1 2 2 1 What happens to the pressure when the volume decreases to 0.5 L? 100 mm Hg × 1 L = P × 0.5 L 2 200 mm Hg = P 2 The pressure has increased ×2. The Ideal Gas law and Boyle’s law apply V = 1.0 L V = 0.5 L to all gases or mixtures of gases. 1 2 P = 100 mm Hg P = 200 mm Hg 1 2 (c) Dalton’s Law Dalton’s law says that the total pressure of a mixture of gases is the sum of the pressures of the individual gases. The pressure of an individual gas in a mixture is known as the partial pressure of the gas (P ). gas For example, at sea level, atmospheric pressure (P ) is 760 mm Hg, In humid air, water vapor “dilutes” the contribution atm and oxygen is 21% of the atmosphere. What is the partial pressure of of other gases to the total pressure. oxygen (P )? O2 Partial Pressures (P ) of Atmospheric Gases at 760 mm Hg To find the partial pressure of any one gas in a sample gas of dry air, multiply the atmospheric pressure (P ) by the gas’s atm Gas and its P in dry P in P in relative contribution (%) to P : gas gas gas atm percentage in air 25 ˚C air 25 ˚C air, 37 ˚C air, 100% humidity 100% humidity Partial pressure of a gas = Patm × % of gas in atmosphere Oxygen (O2) 21% 160 mm Hg 155 mm Hg 150 mm Hg Carbon dioxide PO2 = 760 mm Hg x 21% PO2 (CO2) 0.03% 0.25 mm Hg 0.24 mm Hg 0.235 mm Hg = 760 mm × 0.21 = 160 mm Hg Water vapor 0 mm Hg 24 mm Hg 47 mm Hg The partial pressure of oxygen (P ) in dry air at sea level O2 is 160 mm Hg. To calculate the partial pressure of a gas in humid air, you must first subtract the water vapor pressure from the total pressure. At 100% humidity and The pressure exerted by an individual gas is determined only by its 25° C, water vapor pressure (PH2O) is 24 mm Hg. relative abundance in the mixture and is independent of the molecular size or mass of the gas. P in humid air = (P – P ) × % of gas gas atm H2O P = (760 – 24) × 21% = 155 mm Hg O2 609

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Marcus Aurelius Antoninus . ration, the topic of this chapter and the next, is the movement slide across one another as the lungs move within the thorax. Second “Pump handle" motion increases anterior-posterior dimension of.
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