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SYSTEM
Your body’s cells continually use oxygen (O2) for the metabolic reac-
tions that release energy from nutrient molecules and produce ATP. At
the same time, these reactions release carbon dioxide (CO2). Because an
excessive amount of CO2 produces acidity that can be toxic to cells, ex-
cess CO2 must be eliminated quickly and efficiently. The cardio-
vascular and respiratory systems cooperate to supply O2 and
eliminate CO2. The respiratory system provides for gas ex-
change—intake of O2 and elimination of CO2—and the
cardiovascular system transports blood containing the gases
between the lungs and body cells. Failure of either system
disrupts homeostasis by causing rapid death of cells from
oxygen starvation and buildup of waste products. In addition
to functioning in gas exchange, the respiratory system also
participates in regulating blood pH, contains receptors for the
sense of smell, filters inspired air, produces sounds, and rids
the body of some water and heat in exhaled air. Like the diges-
tive and urinary systems that will be covered in subsequent chapters, in the respiratory sys-
tem there is an extensive area of contact between the external environment and capillary
blood vessels. This area of contact allows the body to constantly renew and replenish the inter-
nal fluid environment that surrounds and nourishes every body cell.
874
RESPIRATORY SYSTEM ANATOMY 875
Nose
Nasal cavity
Oral cavity
Pharynx
Larynx
Trachea
Right primary
Functions
bronchus
1. Provides for gas exchange—intake of O2
Lungs for delivery to body cells and elimination
of CO2 produced by body cells.
2. Helps regulate blood pH.
3. Contains receptors for the sense of smell, filters
inspired air, produces vocal sounds
(phonation), and excretes small amounts
of water and heat.
Larynx
Right common carotid artery Thyroid gland
Trachea
Subclavian artery
Right subclavian artery Phrenic nerve
Brachiocephalic artery Left common carotid artery
Superior vena cava
Arch of aorta
Rib (cut)
Liver Diaphragm
? Which structures are part of the conducting zone of the respiratory system?
876
Figure 23.2 Respiratory structures in the head and neck. (See Tortora, A Photographic Atlas of the Human Body,
Second Edition, Figures 11.2 and 11.3.)
As air passes through the nose, it is warmed, filtered, moistened, and olfaction occurs.
Bony framework:
Frontal bone
Nasal bones
Alar cartilage
Dense fibrous
connective and
adipose tissue
Sagittal Superior
plane Middle
Nasal meatuses Frontal sinus
Inferior Frontal bone
Olfactory epithelium
Sphenoid bone
Sphenoidal sinus
Superior
Internal naris Middle Nasal
Pharyngeal tonsil Inferior conchae
Lingual tonsil
Epiglottis
Mandible
Hyoid bone
LARYNGOPHARYNX
(hypopharynx) Ventricular fold (false vocal cord)
Vocal fold (true vocal cord)
Larynx
Esophagus
Nasopharynx
Thyroid cartilage
Oropharynx
Trachea Cricoid cartilage
Thyroid gland Laryngopharynx
877
878 CHAPTER 23 • THE RESPIRATORY SYSTEM
F I G U R E 23. 2 CO N T I N U E D
Periorbital fat
Frontal plane Ethmoidal cell
Eyeball
View
Superior nasal concha
Nasal septum:
Perpendicular
plate of ethmoid
? What is the path taken by air molecules into and through the nose?
Figure 23.3 Surface anatomy of the nose. The olfactory receptors lie in a region of the membrane lining
the superior nasal conchae and adjacent septum called the
The external nose has a cartilaginous framework
olfactory epithelium. Inferior to the olfactory epithelium, the
and a bony framework.
mucous membrane contains capillaries and pseudostratified
ciliated columnar epithelium with many goblet cells. As inhaled
air whirls around the conchae and meatuses, it is warmed by
blood in the capillaries. Mucus secreted by the goblet cells
moistens the air and traps dust particles. Drainage from the
nasolacrimal ducts also helps moisten the air, and is sometimes
assisted by secretions from the paranasal sinuses. The cilia move
1 the mucus and trapped dust particles toward the pharynx, at
3 which point they can be swallowed or spit out, thus removing
the particles from the respiratory tract.
2 4
䊉 CHECKPOINT
1. What functions do the respiratory and cardiovascular
systems have in common?
2. What structural and functional features are different in
the upper and lower respiratory systems? Which are the
same?
3. Compare the structure and functions of the external nose
and the internal nose.
Anterior view
Figure 23.4 Larynx. (See Tortora, A Photographic Atlas of the Human Body, Second Edition, Figures 11.5 and 11.6.)
The larynx is composed of nine pieces of cartilage.
Epiglottis
Hyoid bone
Thyrohyoid membrane
Epiglottis:
Leaf
Stem
Corniculate cartilage
Thyroid cartilage
(Adam’s apple)
Parathyroid
glands (4)
Tracheal cartilage
Epiglottis
Hyoid bone
Sagittal
Thyrohyoid membrane
plane
Thyrohyoid membrane
Cuneiform cartilage Fat body
Corniculate cartilage
Arytenoid cartilage Ventricular fold (false vocal cord)
Thyroid cartilage
Vocal fold (true vocal cord)
Cricoid cartilage Cricothyroid ligament
Cricotracheal ligament
Tracheal cartilage
Tongue
Thyroid cartilage
Epiglottis
Glottis:
Cricoid cartilage
Vocal folds
(true vocal cords)
Vocal ligament Rima glottidis
Ventricular folds
(false vocal cords)
Arytenoid cartilage
Cuneiform cartilage
Corniculate cartilage
Posterior
cricoarytenoid
Superior view of cartilages muscle View through a laryngoscope
and muscles
(a) Movement of vocal folds apart (abduction)
Lateral
cricoarytenoid
muscle
F I G U R E 23.5 CO N T I N U E D
Epiglottis
Rima
Larynx glottidis
Ventricular Cuneiform
folds (false cartilage
vocal chords) Corniculate
cartilage
Sound originates from the vibration of the vocal folds, but The layers of the tracheal wall, from deep to superficial,
other structures are necessary for converting the sound into are the (1) mucosa, (2) submucosa, (3) hyaline cartilage, and
recognizable speech. The pharynx, mouth, nasal cavity, and (4) adventitia (composed of areolar connective tissue). The
paranasal sinuses all act as resonating chambers that give the mucosa of the trachea consists of an epithelial layer of pseudo-
voice its human and individual quality. We produce the vowel stratified ciliated columnar epithelium and an underlying layer
sounds by constricting and relaxing the muscles in the wall of of lamina propria that contains elastic and reticular fibers.
the pharynx. Muscles of the face, tongue, and lips help us enun- Pseudostratified ciliated columnar epithelium consists of ciliated
ciate words. columnar cells and goblet cells that reach the luminal surface,
Whispering is accomplished by closing all but the posterior plus basal cells that do not (see Table 4.1E on page 117); it pro-
portion of the rima glottidis. Because the vocal folds do not vides the same protection against dust as the membrane lining
vibrate during whispering, there is no pitch to this form of the nasal cavity and larynx. The submucosa consists of areolar
speech. However, we can still produce intelligible speech while connective tissue that contains seromucous glands and their ducts.
whispering by changing the shape of the oral cavity as we The 16–20 incomplete, horizontal rings of hyaline cartilage
enunciate. As the size of the oral cavity changes, its resonance resemble the letter C, are stacked one above another, and are
qualities change, which imparts a vowel-like pitch to the air as it connected together by dense connective tissue. They may be felt
rushes toward the lips. through the skin inferior to the larynx. The open part of each
C-shaped cartilage ring faces posteriorly toward the esophagus
(Figure 23.6) and is spanned by a fibromuscular membrane.
• CLINICAL CONNECTION Laryngitis and Cancer
Within this membrane are transverse smooth muscle fibers,
of the Larynx
called the trachealis muscle, and elastic connective tissue that
Laryngitis is an inflammation of the larynx that is most often caused allow the diameter of the trachea to change subtly during inhala-
by a respiratory infection or irritants such as cigarette smoke. tion and exhalation, which is important in maintaining efficient
Inflammation of the vocal folds causes hoarseness or loss of voice by airflow. The solid C-shaped cartilage rings provide a semirigid
interfering with the contraction of the folds or by causing them to swell support so that the tracheal wall does not collapse inward (espe-
to the point where they cannot vibrate freely. Many long-term smokers cially during inhalation) and obstruct the air passageway. The
acquire a permanent hoarseness from the damage done by chronic in- adventitia of the trachea consists of areolar connective tissue that
flammation. Cancer of the larynx is found almost exclusively in individ- joins the trachea to surrounding tissues.
uals who smoke. The condition is characterized by hoarseness, pain on
swallowing, or pain radiating to an ear. Treatment consists of radiation
therapy and/or surgery. • • CLINICAL CONNECTION Tracheotomy and
Intubation
Trachea Several conditions may block airflow by obstructing the trachea. For ex-
The trachea (TRĀ-kē-a sturdy), or windpipe, is a tubular ample, the rings of cartilage that support the trachea may collapse due
passageway for air that is about 12 cm (5 in.) long and 2.5 cm to a crushing injury to the chest, inflammation of the mucous mem-
brane may cause it to swell so much that the airway closes, vomit or a
(1 in.) in diameter. It is located anterior to the esophagus
foreign object may be aspirated into it, or a cancerous tumor may pro-
(Figure 23.6) and extends from the larynx to the superior border
trude into the airway. Two methods are used to reestablish airflow past
of the fifth thoracic vertebra (T5), where it divides into right and
a tracheal obstruction. If the obstruction is superior to the level of the
left primary bronchi (see Figure 23.7).
RESPIRATORY SYSTEM ANATOMY 883
Figure 23.6 Location of the trachea in relation to the esophagus.
The trachea is anterior to the esophagus and extends from the larynx to the superior border of
the fifth thoracic vertebra.
Transverse
plane
Cartilage of
trachea
Esophagus
POSTERIOR
? What is the benefit of not having complete rings of tracheal cartilage between the trachea and the esophagus?
Figure 23.7 Branching of airways from the trachea: the bronchial tree. (See Tortora, A Photographic Atlas of the Human Body,
Second Edition, Figure 11.8.)
The bronchial tree begins at the trachea and ends at the terminal bronchioles.
BRANCHING OF
BRONCHIAL TREE
Larynx
Trachea
Trachea
Primary bronchi
Secondary bronchi
Tertiary bronchi
Visceral pleura
Bronchioles
Parietal pleura
Terminal bronchioles
Pleural cavity
Location of carina
Right primary
bronchus Left primary bronchus
Right bronchiole
Anterior view
? How many lobes and secondary bronchi are present in each lung?
2. Plates of cartilage gradually replace the incomplete rings of alveoli more quickly, lung ventilation improves. The parasympa-
cartilage in primary bronchi and finally disappear in the distal thetic division of the ANS and mediators of allergic reactions
bronchioles. such as histamine have the opposite effect, causing contraction
3. As the amount of cartilage decreases, the amount of smooth of bronchiolar smooth muscle, which results in constriction of
muscle increases. Smooth muscle encircles the lumen in spiral distal bronchioles.
bands. Because there is no supporting cartilage, however, muscle 䊉 CHECKPOINT
spasms can close off the airways. This is what happens during an 4. List the roles of each of the three anatomical regions of
asthma attack, which can be a life-threatening situation. the pharynx in respiration.
During exercise, activity in the sympathetic division of the 5. How does the larynx function in respiration and voice
production?
autonomic nervous system (ANS) increases and the adrenal
6. Describe the location, structure, and function of the
medulla releases the hormones epinephrine and norepinephrine;
trachea.
both of these events cause relaxation of smooth muscle in the
7. Describe the structure of the bronchial tree.
bronchioles, which dilates the airways. Because air reaches the
RESPIRATORY SYSTEM ANATOMY 885
Lungs • CLINICAL CONNECTION Pneumothorax and
The lungs ( lightweights, because they float) are paired Hemothorax
cone-shaped organs in the thoracic cavity. They are separated
In certain conditions, the pleural cavities may fill with air (pneumotho-
from each other by the heart and other structures in the
rax; pneumo- air or breath), blood (hemothorax), or pus. Air in the
mediastinum, which divides the thoracic cavity into two
pleural cavities, most commonly introduced in a surgical opening of the
anatomically distinct chambers. As a result, if trauma causes chest or as a result of a stab or gunshot wound, may cause the lungs to
one lung to collapse, the other may remain expanded. Each lung collapse. This collapse of a part of a lung, or rarely an entire lung, is
is enclosed and protected by a double-layered serous membrane called atelectasis (at-e-LEK-ta-sis; ateles- incomplete; -ectasis-
called the pleural membrane (PLOOR-al;
(P pleur- side). expansion). The goal of treatment is the evacuation of air (or blood)
The superficial layer, called the parietal pleura, lines the wall from the pleural space, which allows the lung to reinflate. A small pneu-
of the thoracic cavity; the deep layer, the visceral pleura, covers mothorax may resolve on its own, but it is often necessary to insert a
the lungs themselves (Figure 23.8). Between the visceral and chest tube to assist in evacuation. •
parietal pleurae is a small space, the pleural cavity, which
contains a small amount of lubricating fluid secreted by the
membranes. This pleural fluid reduces friction between the The lungs extend from the diaphragm to just slightly superior
membranes, allowing them to slide easily over one another dur- to the clavicles and lie against the ribs anteriorly and posteriorly
ing breathing. Pleural fluid also causes the two membranes to (Figure 23.9a). The broad inferior portion of the lung, the base,
adhere to one another just as a film of water causes two glass mi- is concave and fits over the convex area of the diaphragm. The
croscope slides to stick together, a phenomenon called surface narrow superior portion of the lung is the apex. The surface of
tension. Separate pleural cavities surround the left and right the lung lying against the ribs, the costal surface, matches the
lungs. Inflammation of the pleural membrane, called pleurisy or rounded curvature of the ribs. The mediastinal (medial) surface
pleuritis, may in its early stages cause pain due to friction be- of each lung contains a region, the hilum, through which
tween the parietal and visceral layers of the pleura. If the inflam- bronchi, pulmonary blood vessels, lymphatic vessels, and nerves
mation persists, excess fluid accumulates in the pleural space, a enter and exit (Figure 23.9e). These structures are held together
condition known as pleural effusion. by the pleura and connective tissue and constitute the root of the
Transverse
Sternum
plane
Left lung
Visceral pleura
Ascending aorta
Superior vena cava
Pulmonary arteries
Parietal pleura
Pulmonary vein
View
Right lung Esophagus
Body of T4
Spinal cord
LATERAL MEDIAL
POSTERIOR
Figure 23.9 Surface anatomy of the lungs. (See Tortora, First rib
A Photographic Atlas of the Human Body, Second Edition,
Figures 11.12 and 11.14.)
Apex of lung
The oblique fissure divides the left lung into
two lobes. The oblique and horizontal fissures Left lung
divide the right lung into three lobes.
Base of lung
Pleural cavity
Pleura
Apex
Superior lobe
Horizontal
fissure
Oblique fissure Oblique fissure
Cardiac notch
Inferior lobe
Middle lobe Inferior lobe
POSTERIOR POSTERIOR
Base
(b) Lateral view of right lung (c) Lateral view of left lung
Apex
Superior lobe
View (d)
Oblique fissure
(d) Medial view of right lung (e) Medial view of left lung
? Why are the right and left lungs slightly different in size and shape?
lung. Medially, the left lung also contains a concavity, the car- what shorter than the left lung because the diaphragm is higher
diac notch, in which the heart lies. Due to the space occupied by on the right side, accommodating the liver that lies inferior to it.
the heart, the left lung is about 10% smaller than the right lung. The lungs almost fill the thorax (Figure 23.9a). The apex of
Although the right lung is thicker and broader, it is also some- the lungs lies superior to the medial third of the clavicles and is
RESPIRATORY SYSTEM ANATOMY 887
the only area that can be palpated. The anterior, lateral, and Each lobe receives its own secondary (lobar) bronchus. Thus,
posterior surfaces of the lungs lie against the ribs. The base of the right primary bronchus gives rise to three secondary (lobar)
the lungs extends from the sixth costal cartilage anteriorly to the bronchi called the superior, middle, and inferior secondary
spinous process of the tenth thoracic vertebra posteriorly. The (lobar) bronchi, and the left primary bronchus gives rise to su-
pleura extends about 5 cm (2 in.) below the base from the sixth perior and inferior secondary (lobar) bronchi. Within the
costal cartilage anteriorly to the twelfth rib posteriorly. Thus, the lung, the secondary bronchi give rise to the tertiary (segmental)
lungs do not completely fill the pleural cavity in this area. bronchi, which are constant in both origin and distribution—
Removal of excessive fluid in the pleural cavity can be there are 10 tertiary bronchi in each lung. The segment of lung
accomplished without injuring lung tissue by inserting a needle tissue that each tertiary bronchus supplies is called a bron-
anteriorly through the seventh intercostal space, a procedure chopulmonary segment. Bronchial and pulmonary disorders
called thoracentesis (thor-a-sen-TE Ē¯-sis; -centesis puncture). (such as tumors or abscesses) that are localized in a bronchopul-
The needle is passed along the superior border of the lower rib monary segment may be surgically removed without seriously
to avoid damage to the intercostal nerves and blood vessels. disrupting the surrounding lung tissue.
Inferior to the seventh intercostal space there is danger of Each bronchopulmonary segment of the lungs has many
penetrating the diaphragm. small compartments called lobules; each lobule is wrapped
in elastic connective tissue and contains a lymphatic vessel, an
Lobes, Fissures, and Lobules arteriole, a venule, and a branch from a terminal bronchiole
One or two fissures divide each lung into lobes (Figure (Figure 23.10a). Terminal bronchioles subdivide into micro-
23.9b–e). Both lungs have an oblique fissure, which extends in- scopic branches called respiratory bronchioles (Figure 23.10b).
feriorly and anteriorly; the right lung also has a horizontal As the respiratory bronchioles penetrate more deeply into the
fissure. The oblique fissure in the left lung separates the supe- lungs, the epithelial lining changes from simple cuboidal to sim-
rior lobe from the inferior lobe. In the right lung, the superior ple squamous. Respiratory bronchioles in turn subdivide into
part of the oblique fissure separates the superior lobe from the several (2–11) alveolar ducts. The respiratory passages from the
inferior lobe; the inferior part of the oblique fissure separates the trachea to the alveolar ducts contain about 25 orders of branch-
inferior lobe from the middle lobe, which is bordered superiorly ing; branching from the trachea into primary bronchi is called
by the horizontal fissure. first-order branching, from primary bronchi into secondary
Terminal
bronchiole Terminal
bronchiole
Pulmonary
Pulmonary arteriole
venule
Lymphatic Blood
vessel vessel
Elastic
Respiratory
connective
bronchiole Respiratory
tissue
bronchiole
Alveolar
ducts Alveolar
ducts
Alveoli
Pulmonary
capillary
Alveolar Alveolar
Visceral sac sacs
pleura
Alveoli Visceral
pleura
LM about 30x
bronchi is called second-order branching, and so on down to the is surfactant (sur-FAK-tant), a complex mixture of phospho-
alveolar ducts. lipids and lipoproteins. Surfactant lowers the surface tension of
alveolar fluid, which reduces the tendency of alveoli to collapse
Alveoli (described later).
Around the circumference of the alveolar ducts are numerous Associated with the alveolar wall are alveolar macrophages
alveoli and alveolar sacs. An alveolus (al-VE EĒ-ō-lus) is a cup- (dust cells), phagocytes that remove fine dust particles and other
shaped outpouching lined by simple squamous epithelium and debris from the alveolar spaces. Also present are fibroblasts that
supported by a thin elastic basement membrane; an alveolar sac produce reticular and elastic fibers. Underlying the layer of type
consists of two or more alveoli that share a common opening I alveolar cells is an elastic basement membrane. On the outer
(Figure 23.10a, b). The walls of alveoli consist of two types surface of the alveoli, the lobule’s arteriole and venule disperse
of alveolar epithelial cells (Figure 23.11). The more numerous into a network of blood capillaries (see Figure 23.10a) that
type I alveolar cells are simple squamous epithelial cells that consist of a single layer of endothelial cells and basement
form a nearly continuous lining of the alveolar wall. Type II membrane.
alveolar cells, also called septal cells, are fewer in number and The exchange of O2 and CO2 between the air spaces in
are found between type I alveolar cells. The thin type I alveolar the lungs and the blood takes place by diffusion across the
cells are the main sites of gas exchange. Type II alveolar cells, alveolar and capillary walls, which together form the respiratory
rounded or cuboidal epithelial cells with free surfaces containing membrane. Extending from the alveolar air space to blood
microvilli, secrete alveolar fluid, which keeps the surface plasma, the respiratory membrane consists of four layers
between the cells and the air moist. Included in the alveolar fluid (Figure 23.11b):
Figure 23.11 Structural components of an alveolus. The respiratory membrane consists of a layer of type I and type II
alveolar cells, an epithelial basement membrane, a capillary basement membrane, and the capillary endothelium.
The exchange of respiratory gases occurs by diffusion across the respiratory membrane.
Monocyte
Reticular fiber
Elastic fiber
Type II alveolar
(septal) cell
Respiratory
membrane
Alveolus
(a) Section through an alveolus showing its cellular components (b) Details of respiratory membrane
RESPIRATORY SYSTEM ANATOMY 889
1. A layer of type I and type II alveolar cells and associated poxia (low O2 level). In all other body tissues, hypoxia causes
alveolar macrophages that constitutes the alveolar wall dilation of blood vessels to increase blood flow. In the lungs,
2. An epithelial basement membrane underlying the alveo- however, vasoconstriction in response to hypoxia diverts pul-
lar wall monary blood from poorly ventilated areas of the lungs to well-
ventilated regions. This phenomenon is known as ventilation–
3. A capillary basement membrane that is often fused to the
perfusion coupling because the perfusion (blood flow) to each
epithelial basement membrane
area of the lungs matches the extent of ventilation (airflow) to
4. The capillary endothelium alveoli in that area.
Despite having several layers, the respiratory membrane is Bronchial arteries, which branch from the aorta, deliver
very thin—only 0.5 m thick, about one-sixteenth the diameter oxygenated blood to the lungs. This blood mainly perfuses the
of a red blood cell—to allow rapid diffusion of gases. It has been muscular walls of the bronchi and bronchioles. Connections ex-
estimated that the lungs contain 300 million alveoli, providing ist between branches of the bronchial arteries and branches of
an immense surface area of 70 m2 (750 ft2 )—about the size of a the pulmonary arteries, however; most blood returns to the heart
racquetball court—for gas exchange. via pulmonary veins. Some blood, however, drains into
bronchial veins, branches of the azygos system, and returns to
Blood Supply to the Lungs the heart via the superior vena cava.
The lungs receive blood via two sets of arteries: pulmonary 䊉 CHECKPOINT
arteries and bronchial arteries. Deoxygenated blood passes 8. Where are the lungs located? Distinguish the parietal
through the pulmonary trunk, which divides into a left pul- pleura from the visceral pleura.
monary artery that enters the left lung and a right pulmonary 9. Define each of the following parts of a lung: base, apex,
artery that enters the right lung. (The pulmonary arteries are costal surface, medial surface, hilum, root, cardiac notch,
the only arteries in the body that carry deoxygenated blood.) lobe, and lobule.
Return of the oxygenated blood to the heart occurs by way of 10. What is a bronchopulmonary segment?
the four pulmonary veins, which drain into the left atrium 11. Describe the histology and function of the respiratory
(see Figure 21.29 on page 820). A unique feature of pulmonary membrane.
blood vessels is their constriction in response to localized hy-
Alveolar macrophage
Alveolus (dust cell)
Type II alveolar
(septal) cell
Alveolus
LM 1000x
Sternocleidomastoid
Scalenes
Sternum:
Exhalation
Internal
Inhalation
External intercostals
intercostals
Diaphragm
Diaphragm:
Exhalation
External Inhalation
oblique
Internal
oblique
Transversus
abdominis
Rectus
abdominis
(a) Muscles of inhalation and their actions (left); (b) Changes in size of thoracic cavity
muscles of exhalation and their actions (right) during inhalation and exhalation
? Right now, what is the main muscle that powers your breathing?
During quiet inhalations, the pressure between the two pleural ceral pleurae normally adhere tightly because of the subatmos-
layers in the pleural cavity, called intrapleural (intrathoracic) pheric pressure between them and because of the surface tension
pressure, is always subatmospheric (lower than atmospheric created by their moist adjoining surfaces. As the thoracic cavity
pressure). Just before inhalation, it is about 4 mmHg less than the expands, the parietal pleura lining the cavity is pulled outward in
atmospheric pressure, or about 756 mmHg at an atmospheric all directions, and the visceral pleura and lungs are pulled along
pressure of 760 mmHg (Figure 23.14). As the diaphragm and with it.
external intercostals contract and the overall size of the thoracic As the volume of the lungs increases in this way, the pressure
cavity increases, the volume of the pleural cavity also increases, inside the lungs, called the alveolar (intrapulmonic) pressure,
which causes intrapleural pressure to decrease to about drops from 760 to 758 mmHg. A pressure difference is thus es-
754 mmHg. During expansion of the thorax, the parietal and vis- tablished between the atmosphere and the alveoli. Because air
892 CHAPTER 23 • THE RESPIRATORY SYSTEM
Figure 23.14 Pressure changes in pulmonary ventilation. During inhalation, the diaphragm contracts, the chest expands,
the lungs are pulled outward, and alveolar pressure decreases. During exhalation, the diaphragm relaxes, the
lungs recoil inward, and alveolar pressure increases, forcing air out of the lungs.
Air moves into the lungs when alveolar pressure is less than atmospheric pressure, and out of the lungs
when alveolar pressure is greater than atmospheric pressure.
Alveolar Alveolar
pressure = pressure =
760 mmHg 758 mmHg
Intrapleural Intrapleural
pressure = pressure =
756 mmHg 754 mmHg
Alveolar
pressure =
762 mmHg
Intrapleural
pressure =
756 mmHg
? How does the intrapleural pressure change during a normal, quiet breath?
Atmospheric pressure
is about 760 mmHg
at sea level
Thoracic
cavity increases Thoracic cavity
in size and volume of During normal quiet decreases in size
lungs expands exhalation, diaphragm and and lungs recoil
external intercostals relax.
During forceful exhalation,
abdominal and internal
Alveolar pressure
intercostal muscles
decreases to 758 mmHg
contract.
depressed. These movements decrease the vertical, lateral, and surface tension of the alveolar fluid, compliance of the lungs,
anteroposterior diameters of the thoracic cavity, which decreases and airway resistance.
lung volume. In turn, the alveolar pressure increases to about
762 mmHg. Air then flows from the area of higher pressure
in the alveoli to the area of lower pressure in the atmosphere Surface Tension of Alveolar Fluid
(see Figure 23.14). As noted earlier, a thin layer of alveolar fluid coats the luminal
Exhalation becomes active only during forceful breathing, as surface of alveoli and exerts a force known as surface tension.
occurs while playing a wind instrument or during exercise. Surface tension arises at all air–water interfaces because the
During these times, muscles of exhalation—the abdominals polar water molecules are more strongly attracted to each other
and internal intercostals (see Figure 23.13a)—contract, which than they are to gas molecules in the air. When liquid surrounds
increases pressure in the abdominal region and thorax. a sphere of air, as in an alveolus or a soap bubble, surface
Contraction of the abdominal muscles moves the inferior ribs tension produces an inwardly directed force. Soap bubbles
downward and compresses the abdominal viscera, thereby “burst” because they collapse inward due to surface tension. In
forcing the diaphragm superiorly. Contraction of the internal the lungs, surface tension causes the alveoli to assume the small-
intercostals, which extend inferiorly and posteriorly between est possible diameter. During breathing, surface tension must be
adjacent ribs, pulls the ribs inferiorly. Although intrapleural overcome to expand the lungs during each inhalation. Surface
pressure is always less than alveolar pressure, it may briefly tension also accounts for two-thirds of lung elastic recoil, which
exceed atmospheric pressure during a forceful exhalation, such decreases the size of alveoli during exhalation.
as during a cough. The surfactant (a mixture of phospholipids and lipoproteins)
Figure 23.15b summarizes the events of exhalation. present in alveolar fluid reduces its surface tension below the
surface tension of pure water. A deficiency of surfactant in
premature infants causes respiratory distress syndrome, in
Other Factors Affecting Pulmonary Ventilation which the surface tension of alveolar fluid is greatly increased,
As you have just learned, air pressure differences drive airflow so that many alveoli collapse at the end of each exhalation. Great
during inhalation and exhalation. However, three other factors effort is then needed at the next inhalation to reopen the
affect the rate of airflow and the ease of pulmonary ventilation: collapsed alveoli.
894 CHAPTER 23 • THE RESPIRATORY SYSTEM
TA B L E 23 . 1
Modified Respiratory Movements
MOVEMENT DESCRIPTION
Coughing A long-drawn and deep inhalation followed by a complete closure of the rima glottidis, which results in a strong exhalation that
suddenly pushes the rima glottidis open and sends a blast of air through the upper respiratory passages. Stimulus for this reflex act
may be a foreign body lodged in the larynx, trachea, or epiglottis.
Sneezing Spasmodic contraction of muscles of exhalation that forcefully expels air through the nose and mouth. Stimulus may be an irritation
of the nasal mucosa.
Sighing A long-drawn and deep inhalation immediately followed by a shorter but forceful exhalation.
Yawning A deep inhalation through the widely opened mouth producing an exaggerated depression of the mandible. It may be stimulated by
drowsiness, or someone else’s yawning, but the precise cause is unknown.
Sobbing A series of convulsive inhalations followed by a single prolonged exhalation. The rima glottidis closes earlier than normal after each
inhalation so only a little air enters the lungs with each inhalation.
Crying An inhalation followed by many short convulsive exhalations, during which the rima glottidis remains open and the vocal folds vibrate;
accompanied by characteristic facial expressions and tears.
Laughing The same basic movements as crying, but the rhythm of the movements and the facial expressions usually differ from those of crying.
Laughing and crying are sometimes indistinguishable.
Hiccupping Spasmodic contraction of the diaphragm followed by a spasmodic closure of the rima glottidis, which produces a sharp sound on
inhalation. Stimulus is usually irritation of the sensory nerve endings of the gastrointestinal tract.
Valsalva (val-SAL-va) Forced exhalation against a closed rima glottidis as may occur during periods of straining while defecating.
maneuver
A lower-than-normal minute ventilation usually is a sign of spirogram. Inhalation is recorded as an upward deflection, and
pulmonary malfunction. The apparatus commonly used to exhalation is recorded as a downward deflection (Figure 23.16).
measure the volume of air exchanged during breathing and Tidal volume varies considerably from one person to another
the respiratory rate is a spirometer (spiro- breathe; meter and in the same person at different times. In a typical adult,
measuring device) or respirometer. The record is called a about 70% of the tidal volume (350 mL) actually reaches the
Figure 23.16 Spirogram of lung volumes and capacities. The average values for a healthy adult male and female are indicated, with
the values for a female in parentheses. Note that the spirogram is read from right (start of record) to left (end of record).
Lung capacities are combinations of various lung volumes.
6,000 mL
Inhalation
5,000 mL
INSPIRATORY INSPIRATORY VITAL TOTAL
RESERVE CAPACITY CAPACITY LUNG
VOLUME Exhalation 3,600 mL 4,800 mL CAPACITY
3,100 mL (2,400 mL) (3,100 mL) 6,000 mL
4,000 mL (1,900 mL) (4,200 mL)
3,000 mL
TIDAL
VOLUME 500 mL
EXPIRATORY
2,000 mL RESERVE
VOLUME End of Start of
1,200 mL record record
FUNCTIONAL
(700 mL) RESIDUAL
CAPACITY
1,000 mL RESIDUAL 2,400 mL
VOLUME (1,800 mL)
1,200 mL
(1,100 mL)
? If you breathe in as deeply as possible and then exhale as much air as you can, which lung capacity have you demonstrated?
896 CHAPTER 23 • THE RESPIRATORY SYSTEM
respiratory zone of the respiratory system—the respiratory bron- and inspiratory reserve volume (500 mL 3100 mL 3600 mL
chioles, alveolar ducts, alveolar sacs, and alveoli—and in males and 500 mL 1900 mL 2400 mL in females).
participates in external respiration. The other 30% (150 mL) Functional residual capacity is the sum of residual volume and
remains in the conducting airways of the nose, pharynx, larynx, expiratory reserve volume (1200 mL 1200 mL 2400 mL in
trachea, bronchi, bronchioles, and terminal bronchioles. males and 1100 mL 700 mL 1800 mL in females). Vital
Collectively, the conducting airways with air that does not capacity is the sum of inspiratory reserve volume, tidal volume,
undergo respiratory exchange are known as the anatomic (res- and expiratory reserve volume (4800 mL in males and 3100 mL
piratory) dead space. (An easy rule of thumb for determining in females). Finally, total lung capacity is the sum of vital
the volume of your anatomic dead space is that it is about capacity and residual volume (4800 mL 1200 mL 6000 mL
the same in milliliters as your ideal weight in pounds.) Not all in males and 3100 mL 1100 mL 4200 mL in females).
of the minute ventilation can be used in gas exchange because
䊉 CHECKPOINT
some of it remains in the anatomic dead space. The alveolar
16. What is a spirometer?
ventilation rate is the volume of air per minute that actually
17. What is the difference between a lung volume and a
reaches the respiratory zone. In the example just given, alveolar
lung capacity?
ventilation rate would be 350 mL/breath 12 breaths/min 18. How is minute ventilation calculated?
4200 mL/min. 19. Define alveolar ventilation rate and FEV1.0.
Several other lung volumes are defined relative to forceful
breathing. In general, these volumes are larger in males, taller
individuals, and younger adults, and smaller in females, shorter
individuals, and the elderly. Various disorders also may be
diagnosed by comparison of actual and predicted normal values EXCHANGE OF OXYGEN
for a patient’s gender, height, and age. The values given here are AND CARBON DIOXIDE
averages for young adults.
䊉 OBJECTIVES
By taking a very deep breath, you can inhale a good deal
• Explain Dalton’s law and Henry’s law.
more than 500 mL. This additional inhaled air, called the
• Describe the exchange of oxygen and carbon dioxide in
inspiratory reserve volume, is about 3100 mL in an average
external and internal respiration.
adult male and 1900 mL in an average adult female
(Figure 23.16). Even more air can be inhaled if inhalation The exchange of oxygen and carbon dioxide between alveolar
follows forced exhalation. If you inhale normally and then air and pulmonary blood occurs via passive diffusion, which is
exhale as forcibly as possible, you should be able to push out governed by the behavior of gases as described by two gas laws,
considerably more air in addition to the 500 mL of tidal volume. Dalton’s law and Henry’s law. Dalton’s law is important for
The extra 1200 mL in males and 700 mL in females is called the understanding how gases move down their pressure differences
expiratory reserve volume. The FEV1.0 is the forced expira- by diffusion, and Henry’s law helps explain how the solubility of
tory volume in 1 second, the volume of air that can be exhaled a gas relates to its diffusion.
from the lungs in 1 second with maximal effort following a
maximal inhalation. Typically, chronic obstructive pulmonary
disease (COPD) greatly reduces FEV1.0 because COPD increases Gas Laws: Dalton’s Law and
airway resistance.
Henry’s Law
Even after the expiratory reserve volume is exhaled, con- According to Dalton’s law, each gas in a mixture of gases exerts
siderable air remains in the lungs because the subatmospheric its own pressure as if no other gases were present. The pressure
intrapleural pressure keeps the alveoli slightly inflated, and some of a specific gas in a mixture is called its partial pressure (Px);
air also remains in the noncollapsible airways. This volume, the subscript is the chemical formula of the gas. The total pres-
which cannot be measured by spirometry, is called the residual sure of the mixture is calculated simply by adding all the partial
volume and amounts to about 1200 mL in males and 1100 mL pressures. Atmospheric air is a mixture of gases—nitrogen (N2),
in females. oxygen (O2), water vapor (H2O), and carbon dioxide (CO2), plus
If the thoracic cavity is opened, the intrapleural pressure rises other gases present in small quantities. Atmospheric pressure is
to equal the atmospheric pressure and forces out some of the the sum of the pressures of all these gases:
residual volume. The air remaining is called the minimal vol-
Atmospheric pressure (760 mmHg)
ume. Minimal volume provides a medical and legal tool for de-
PN2 PO2 PH2O PCO2 Pother gases
termining whether a baby is born dead (stillborn) or died after
birth. The presence of minimal volume can be demonstrated by We can determine the partial pressure exerted by each
placing a piece of lung in water and observing if it floats. Fetal component in the mixture by multiplying the percentage of the
lungs contain no air, so the lung of a stillborn baby will not float gas in the mixture by the total pressure of the mixture.
in water. Atmospheric air is 78.6% nitrogen, 20.9% oxygen, 0.04% car-
Lung capacities are combinations of specific lung volumes bon dioxide, and 0.06% other gases; a variable amount of water
(Figure 23.16). Inspiratory capacity is the sum of tidal volume vapor is also present, about 0.4% on a cool, dry day. Thus, the
EXCHANGE OF OXYGEN AND CARBON DIOXIDE 897
partial pressures of the gases in inhaled air are as follows: Because the partial pressure of nitrogen is higher in a mixture of
compressed air than in air at sea level pressure, a considerable
PN2 0.786 760 mmHg 597.4 mmHg
amount of nitrogen dissolves in plasma and interstitial fluid.
PO2 0.209 760 mmHg 158.8 mmHg
Excessive amounts of dissolved nitrogen may produce giddiness
PH2O 0.004 760 mmHg 3.0 mmHg
and other symptoms similar to alcohol intoxication. The condi-
PCO2 0.0004 760 mmHg 0.3 mmHg
tion is called nitrogen narcosis or “rapture of the deep.”
Pother gases 0.0006 760 mmHg 0.5 mmHg
If a diver comes to the surface slowly, the dissolved nitrogen
Total 760.0 mmHg
can be eliminated by exhaling it. However, if the ascent is too
These partial pressures determine the movement of O2 and rapid, nitrogen comes out of solution too quickly and forms gas
CO2 between the atmosphere and lungs, between the lungs and bubbles in the tissues, resulting in decompression sickness (the
blood, and between the blood and body cells. Each gas diffuses bends). The effects of decompression sickness typically result
across a permeable membrane from the area where its partial from bubbles in nervous tissue and can be mild or severe,
pressure is greater to the area where its partial pressure is less. depending on the number of bubbles formed. Symptoms include
The greater the difference in partial pressure, the faster the rate joint pain, especially in the arms and legs, dizziness, shortness
of diffusion. of breath, extreme fatigue, paralysis, and unconsciousness.
Compared with inhaled air, alveolar air has less O2 (13.6%
versus 20.9%) and more CO2 (5.2% versus 0.04%) for two
• CLINICAL CO N N EC TI O N Hyperbaric Oxygenation
reasons. First, gas exchange in the alveoli increases the CO2
content and decreases the O2 content of alveolar air. Second, A major clinical application of Henry’s law is hyperbaric oxygenation
when air is inhaled it becomes humidified as it passes along (hyper over; baros pressure), the use of pressure to cause more O2
the moist mucosal linings. As water vapor content of the air to dissolve in the blood. It is an effective technique in treating patients
increases, the relative percentage that is O2 decreases. In con- infected by anaerobic bacteria, such as those that cause tetanus and
trast, exhaled air contains more O2 than alveolar air (16% versus gangrene. (Anaerobic bacteria cannot live in the presence of free O2.) A
13.6%) and less CO2 (4.5% versus 5.2%) because some of the person undergoing hyperbaric oxygenation is placed in a hyperbaric
exhaled air was in the anatomic dead space and did not partici- chamber, which contains O2 at a pressure greater than one atmosphere
pate in gas exchange. Exhaled air is a mixture of alveolar air and (760 mmHg). As body tissues pick up the O2, the bacteria are killed.
inhaled air that was in the anatomic dead space. Hyperbaric chambers may also be used for treating certain heart disor-
Henry’s law states that the quantity of a gas that will dissolve ders, carbon monoxide poisoning, gas embolisms, crush injuries, cere-
in a liquid is proportional to the partial pressure of the gas and bral edema, certain hard-to-treat bone infections caused by anaerobic
bacteria, smoke inhalation, near-drowning, asphyxia, vascular insuffi-
its solubility. In body fluids, the ability of a gas to stay in
ciencies, and burns. •
solution is greater when its partial pressure is higher and when it
has a high solubility in water. The higher the partial pressure of a
gas over a liquid and the higher the solubility, the more gas will
stay in solution. In comparison to oxygen, much more CO2 is
External and Internal Respiration
dissolved in blood plasma because the solubility of CO2 is 24
times greater than that of O2. Even though the air we breathe External respiration or pulmonary gas exchange is the diffu-
contains mostly N2, this gas has no known effect on bodily func- sion of O2 from air in the alveoli of the lungs to blood in
tions, and at sea level pressure very little of it dissolves in blood pulmonary capillaries and the diffusion of CO2 in the opposite
plasma because its solubility is very low. direction (Figure 23.17a). External respiration in the lungs
An everyday experience gives a demonstration of Henry’s converts deoxygenated blood (depleted of some O2) coming
law. You have probably noticed that a soft drink makes a hissing from the right side of the heart into oxygenated blood (saturated
sound when the top of the container is removed, and bubbles rise with O2) that returns to the left side of the heart (see Figure 21.29
to the surface for some time afterward. The gas dissolved in on page 820). As blood flows through the pulmonary capillaries,
carbonated beverages is CO2. Because the soft drink is bottled it picks up O2 from alveolar air and unloads CO2 into alveolar
or canned under high pressure and capped, the CO2 remains air. Although this process is commonly called an “exchange” of
dissolved as long as the container is unopened. Once you gases, each gas diffuses independently from the area where its
remove the cap, the pressure decreases and the gas begins to partial pressure is higher to the area where its partial pressure
bubble out of solution. is lower.
Henry’s law explains two conditions resulting from changes As Figure 23.17a shows, O2 diffuses from alveolar air, where
in the solubility of nitrogen in body fluids. Even though the air its partial pressure is 105 mmHg, into the blood in pulmonary
we breathe contains about 79% nitrogen, this gas has no known capillaries, where PO2 is only 40 mmHg in a resting person. If
effect on bodily functions, and very little of it dissolves in blood you have been exercising, the PO2 will be even lower because
plasma because of its low solubility at sea level pressure. As the contracting muscle fibers are using more O2. Diffusion continues
total air pressure increases, the partial pressures of all its gases until the PO2 of pulmonary capillary blood increases to match the
increase. When a scuba diver breathes air under high pressure, PO2 of alveolar air, 105 mmHg. Because blood leaving pul-
the nitrogen in the mixture can have serious negative effects. monary capillaries near alveolar air spaces mixes with a small
898 CHAPTER 23 • THE RESPIRATORY SYSTEM
Figure 23.17 Changes in partial pressures of oxygen and carbon dioxide (in mmHg) during external and internal
respiration.
Gases diffuse from areas of higher partial pressure to areas of lower partial pressure.
Atmospheric air: CO2 exhaled
PO2 = 159 mm Hg
PCO = 0.3 mm Hg O2 inhaled
2
Alveolar air:
Alveoli PO2 = 105 mm Hg
PCO2 = 40 mm Hg
CO2 O
2
Pulmonary capillaries
Systemic capillaries
CO2 O2
? What causes oxygen to enter pulmonary capillaries from alveoli and to enter tissue cells from systemic capillaries?
volume of blood that has flowed through conducting portions of While O2 is diffusing from alveolar air into deoxygenated
the respiratory system, where gas exchange does not occur, the blood, CO2 is diffusing in the opposite direction. The PCO2 of
PO2 of blood in the pulmonary veins is slightly less than the PO2 deoxygenated blood is 45 mmHg in a resting person, and the
in pulmonary capillaries, about 100 mmHg. PCO2 of alveolar air is 40 mmHg. Because of this difference in
EXCHANGE OF OXYGEN AND CARBON DIOXIDE 899
PCO2, carbon dioxide diffuses from deoxygenated blood into the versus pulmonary blood increase during exercise. The larger
alveoli until the PCO2 of the blood decreases to 40 mmHg. partial pressure differences accelerate the rates of gas diffu-
Exhalation keeps alveolar PCO2 at 40 mmHg. Oxygenated blood sion. The partial pressures of O2 and CO2 in alveolar air also
returning to the left side of the heart in the pulmonary veins thus depend on the rate of airflow into and out of the lungs.
has a PCO2 of 40 mmHg. Certain drugs (such as morphine) slow ventilation, thereby
The number of capillaries near alveoli in the lungs is very decreasing the amount off O2 and CO2 that can be exchanged
large, and blood flows slowly enough through these capillaries between alveolar air and blood. With increasing altitude,
that it picks up a maximal amount of O2. During vigorous the total atmospheric pressure decreases, as does the
exercise, when cardiac output is increased, blood flows more partial pressure of O2—from 159 mmHg at sea level, to
rapidly through both the systemic and pulmonary circulations. As 110 mmHg at 10,000 ft, to 73 mmHg at 20,000 ft. Although
a result, blood’s transit time in the pulmonary capillaries O2 still is 20.9% of the total, the PO2 of inhaled air decreases
is shorter. Still, the PO2 of blood in the pulmonary veins normally with increasing altitude. Alveolar PO2 decreases correspond-
reaches 100 mmHg. In diseases that decrease the rate of gas ingly, and O2 diffuses into the blood more slowly. The com-
diffusion, however, the blood may not come into full equilibrium mon signs and symptoms of high altitude sickness—short-
with alveolar air, especially during exercise. When this happens, ness of breath, headache, fatigue, insomnia, nausea, and
the PO2 declines and PCO2 rises in systemic arterial blood. dizziness—are due to a lower level of oxygen in the blood.
The left ventricle pumps oxygenated blood into the aorta and • Surface area available for gas exchange. As you learned
through the systemic arteries to systemic capillaries. The earlier in the chapter, the surface area of the alveoli is huge
exchange of O2 and CO2 between systemic capillaries and tissue (about 70 m2 or 750 ft2). In addition, many capillaries
cells is called internal respiration or systemic gas exchange surround each alveolus, so many that as much as 900 mL of
(Figure 23.17b). As O2 leaves the bloodstream, oxygenated blood is able to participate in gas exchange at any instant.
blood is converted into deoxygenated blood. Unlike external Any pulmonary disorder that decreases the functional sur-
respiration, which occurs only in the lungs, internal respiration face area of the respiratory membranes decreases the rate of
occurs in tissues throughout the body. external respiration. In emphysema (page 913), for example,
The PO2 of blood pumped into systemic capillaries is higher alveolar walls disintegrate, so surface area is smaller than
(100 mmHg) than the PO2 in tissue cells (40 mmHg at rest) normal and pulmonary gas exchange is slowed.
because the cells constantly use O2 to produce ATP. Due to this
pressure difference, oxygen diffuses out of the capillaries into • Diffusion distance. The respiratory membrane is very thin,
tissue cells and blood PO2 drops to 40 mmHg by the time the so diffusion occurs quickly. Also, the capillaries are so
blood exits systemic capillaries. narrow that the red blood cells must pass through them in
While O2 diffuses from the systemic capillaries into tissue single file, which minimizes the diffusion distance from an
cells, CO2 diffuses in the opposite direction. Because tissue alveolar air space to hemoglobin inside red blood cells.
cells are constantly producing CO2, the PCO2 of cells (45 mmHg Buildup of interstitial fluid between alveoli, as occurs in
at rest) is higher than that of systemic capillary blood pulmonary edema (page 914), slows the rate of gas
(40 mmHg). As a result, CO2 diffuses from tissue cells through exchange because it increases diffusion distance.
interstitial fluid into systemic capillaries until the PCO2 in the • Molecular weight and solubility of the gases. Because O2
blood increases to 45 mmHg. The deoxygenated blood then has a lower molecular weight than CO2, it could be expected
returns to the heart and is pumped to the lungs for another cycle to diffuse across the respiratory membrane about 1.2 times
of external respiration. faster. However, the solubility of CO2 in the fluid portions of
In a person at rest, tissue cells, on average, need only 25% of the respiratory membrane is about 24 times greater than that
the available O2 in oxygenated blood; despite its name, of O2. Taking both of these factors into account, net outward
deoxygenated blood retains 75% of its O2 content. During exer- CO2 diffusion occurs 20 times more rapidly than net inward
cise, more O2 diffuses from the blood into metabolically active O2 diffusion. Consequently, when diffusion is slower than
cells, such as contracting skeletal muscle fibers. Active cells use normal, for example, in emphysema or pulmonary edema,
more O2 for ATP production, causing the O2 content of deoxy- O2 insufficiency (hypoxia) typically occurs before there is
genated blood to drop below 75%. significant retention of CO2 (hypercapnia).
The rate of pulmonary and systemic gas exchange depends 䊉 CHECKPOINT
on several factors. 20. Distinguish between Dalton’s law and Henry’s law and
give a practical application of each.
• Partial pressure difference of the gases. Alveolar PO2 must 21. How does the partial pressure of oxygen change as
be higher than blood PO2 for oxygen to diffuse from alveolar altitude changes?
air into the blood. The rate of diffusion is faster when the dif- 22. What are the diffusion paths of oxygen and carbon
ference between PO2 in alveolar air and pulmonary capillary dioxide during external and internal respiration?
blood is larger; diffusion is slower when the difference is 23. What factors affect the rate of diffusion of oxygen
smaller. The differences between PO2 and PCO2 in alveolar air and carbon dioxide?
900 CHAPTER 23 • THE RESPIRATORY SYSTEM
TRANSPORT OF OXYGEN certain chemical reactions occur that aid in gas transport and
gas exchange.
AND CARBON DIOXIDE
䊉 OBJECTIVE Oxygen Transport
• Describe how the blood transports oxygen and carbon
Oxygen does not dissolve easily in water, so only about 1.5% of
dioxide.
inhaled O2 is dissolved in blood plasma, which is mostly water.
As you have already learned, the blood transports gases between About 98.5% of blood O2 is bound to hemoglobin in red blood
the lungs and body tissues. When O2 and CO2 enter the blood, cells (Figure 23.18). Each 100 mL of oxygenated blood contains
Figure 23.18 Transport of oxygen (O2) and carbon dioxide (CO2) in the blood.
Most O2 is transported by hemoglobin as oxyhemoglobin (Hb–O2 ) within red blood cells; most CO 2 is transported
in blood plasma as bicarbonate ions (HCO3ⴚ).
Transport of CO2 Transport of O2
7% dissolved in plasma 1.5% dissolved in plasma
23% as Hb–CO2 98.5% as Hb–O2
–
70% as HCO3
Alveoli
CO2 O
2
7% 23%
1.5% 98.5%
70%
O2 Pulmonary
HCO3–
(dissolved) capillaries
Hb + O2
CO2+Hb Hb Red blood cell
Hb–O2
Hb–CO2
(a) External respiration: Plasma
pulmonary gas
exchange
CO2
(dissolved)
Hb–CO2 Hb–O2
(b) Internal respiration:
Hb systemic gas O2
exchange Systemic
O2
–
7% HCO3 (dissolved) capillaries
Hb
23%
70% 1.5%
Interstitial fluid
Systemic
CO2 O2 tissue cells
where the PO2 is lower, hemoglobin does not hold as much O2, 80
and the dissolved O2 is unloaded via diffusion into tissue cells
70
(see Figure 23.18b). Note that hemoglobin is still 75% saturated
with O2 at a PO2 of 40 mmHg, the average PO2 of tissue cells in 60
a person at rest. This is the basis for the earlier statement that 50
only 25% of the available O2 unloads from hemoglobin and is 40
used by tissue cells under resting conditions.
30
When the PO2 is between 60 and 100 mmHg, hemoglobin is
20
90% or more saturated with O2 (Figure 23.19). Thus, blood
picks up a nearly full load of O2 from the lungs even when the 10
PO2 of alveolar air is as low as 60 mmHg. The Hb–PO2 curve
0 10 20 30 40 50 60 70 80 90 100
explains why people can still perform well at high altitudes or
PO2 (mm Hg)
when they have certain cardiac and pulmonary diseases, even
though PO2 may drop as low as 60 mmHg. Note also in the curve ? What point on the curve represents blood in your
that at a considerably lower PO2 of 40 mmHg, hemoglobin is still pulmonary veins right now? In your pulmonary veins if you
75% saturated with O2. However, oxygen saturation of Hb drops were jogging?
902 CHAPTER 23 • THE RESPIRATORY SYSTEM
oxygen–hemoglobin dissociation curve shifts to the right; at hormones, such as thyroxine, human growth hormone, epi-
any given PO2, Hb is less saturated with O2, a change termed nephrine, norepinephrine, and testosterone, increase the forma-
the Bohr effect. The Bohr effect works both ways: An increase tion of BPG. The level of BPG also is higher in people living at
in H in blood causes O2 to unload from hemoglobin, and the higher altitudes.
binding of O2 to hemoglobin causes unloading of H from
hemoglobin. The
T explanation for the Bohr effect is that hemo-
globin can act as a buffer for hydrogen ions (H). But when H
ions bind to amino acids in hemoglobin, they alter its structure
slightly, decreasing its oxygen-carrying capacity. Thus, lowered Figure 23.20 Oxygen–hemoglobin dissociation curves
pH drives O2 off hemoglobin, making more O2 available for showing the relationship of (a) pH and (b) PCO2 to hemoglobin
tissue cells. By contrast, elevated pH increases the affinity of saturation at normal body temperature. As pH increases or PCO2
hemoglobin for O2 and shifts the oxygen–hemoglobin dissocia- decreases, O2 combines more tightly with hemoglobin, so that less
tion curve to the left. is available to tissues. The broken lines emphasize these
2. Partial pressure of carbon dioxide. CO2 also can bind to relationships.
hemoglobin, and the effect is similar to that of H (shifting As pH decreases or PCO2 increases, the affinity of
the curve to the right). As PCO2 rises, hemoglobin releases O2 hemoglobin for O2 declines, so less O2 combines
more readily (Figure 23.20b). PCO2 and pH are related factors with hemoglobin and more is available to tissues.
because low blood pH (acidity) results from high PCO2. As CO2
enters the blood, much of it is temporarily converted
d to carbonic 100 High blood pH
acid (H2CO3), a reaction catalyzed by an enzyme in red blood 90
(7.6)
70
Carbon monoxide (CO) is a colorless and odorless gas found in exhaust
60
fumes from automobiles, gas furnaces, and space heaters and in to-
50 Maternal
bacco smoke. It is a byproduct of the combustion of carbon-containing
materials such as coal, gas, and wood. CO binds to the heme group of 40
hemoglobin, just as O2 does, except that the binding of carbon monox- 30
ide to hemoglobin is over 200 times as strong as the binding of O2 to
20
hemoglobin. Thus, at a concentration as small as 0.1% (PCO 0.5
mmHg), CO will combine with half the available hemoglobin molecules 10
and reduce the oxygen-carrying capacity of the blood by 50%. Elevated
0 10 20 30 40 50 60 70 80 90 100
blood levels of CO cause carbon monoxide poisoning, which can cause
PO2 (mmHg)
the lips and oral mucosa to appear bright, cherry-red (the color of
hemoglobin with carbon monoxide bound to it). Without prompt treat-
ment, carbon monoxide poisoning is fatal. It is possible to rescue a vic-
tim of CO poisoning by administering pure oxygen, which speeds up the ? The PO2 of placental blood is about 40 mmHg. What are the
separation of carbon monoxide from hemoglobin. • O2 saturations of maternal and fetal hemoglobin at this PO2?
904 CHAPTER 23 • THE RESPIRATORY SYSTEM
3. Bicarbonate ions. The greatest percentage of CO2—about Thus, as blood picks up CO2, HCO3 accumulates inside RBCs.
70%—is transported in blood plasma as bicarbonate ions Some HCO3 moves out into the blood plasma, down its con-
(HCO3ⴚ). As CO2 diffuses into systemic capillaries and enters centration gradient. In exchange, chloride ions (Cl) move from
red blood cells, it reacts with water in the presence of the plasma into the RBCs. This exchange of negative ions, which
enzyme carbonic anhydrase (CA) to form carbonic acid, which maintains the electrical balance between blood plasma and RBC
dissociates into H and HCO3: cytosol, is known as the chloride shift (Figure 23.23b). The net
CA
effect of these reactions is that CO2 is removed from tissue cells
CO2 H2O H2CO3 H HCO3 and transported in blood plasma as HCO3. As blood passes
Carbon Water Carbonic Hydrogen Bicarbonate through pulmonary capillaries in the lungs, all these reactions re-
dioxide acid ion ion verse and CO2 is exhaled.
Figure 23.23 Summary of chemical reactions that occur during gas exchange. (a) As carbon dioxide (CO2) is ex-
haled, hemoglobin (Hb) inside red blood cells in pulmonary capillaries unloads CO2 and picks up O2 from
alveolar air. Binding of O2 to Hb ! H releases hydrogen ions (H). Bicarbonate ions (HCO3) pass into the
RBC and bind to released H, forming carbonic acid (H2CO3). The H2CO3 dissociates into water (H2O) and
CO2, and the CO2 diffuses from blood into alveolar air. To maintain electrical balance, a chloride ion (Cl)
exits the RBC for each HCO3 that enters (reverse chloride shift). (b) CO2 diffuses out of tissue cells that produce it and
enters red blood cells, where some of it binds to hemoglobin, forming carbaminohemoglobin (Hb–CO2). This reaction
causes O2 to dissociate from oxyhemoglobin (Hb–O2). Other molecules of CO2 combine with water to produce bicarbonate
ions (HCO3) and hydrogen ions (H). As Hb buffers H, the Hb releases O2 (Bohr effect). To maintain electrical balance,
a chloride ion (Cl) enters the RBC for each HCO3 that exits (chloride shift).
Hemoglobin inside red blood cells transports O2, CO2, and H⫹.
Reverse
chloride
Exhaled shift CO2 + Hb Hb–CO2
– –
Cl Cl
Carbonic anhydrase – +
CO2 CO2 CO2 CO2 + H2O H2CO3 HCO3 + H
–
HCO3
+
O2 O2 O2 O2 + Hb–H Hb–O2+ H
Interstitial Plasma
Inhaled fluid
Alveolus Pulmonary Red blood cell
capillary wall
Chloride
shift
Cl – Cl – CO2 + Hb Hb–CO2 + O2
Carbonic anhydrase – +
CO2 CO2 CO2 CO2 + H2O H2CO3 HCO3 + H
HCO3–
O2 O2 O2 O2 + Hb–H Hb–O2
Interstitial Plasma
fluid
Tissue cell Systemic Red blood cell
capillary wall
? Would you expect the concentration of HCO3 to be higher in blood plasma taken from a systemic artery or a systemic vein?
CONTROL OF RESPIRATION 905
The amount of CO2 that can be transported in the blood is in- At rest, about 200 mL of O2 are used each minute by body cells.
fluenced by the percent saturation of hemoglobin with oxygen. During strenuous exercise, however, O2 use typically increases
The lower the amount of oxyhemoglobin (Hb–O2), the higher 15- to 20-fold in normal healthy adults, and as much as 30-fold
the CO2 carrying capacity of the blood, a relationship known as in elite endurance-trained athletes. Several mechanisms help
the Haldane effect. Two characteristics of deoxyhemoglobin match respiratory effort to metabolic demand.
give rise to the Haldane effect: (1) Deoxyhemoglobin binds
to and thus transports more CO2 than does Hb–O2. Respiratory Center
(2) Deoxyhemoglobin also buffers more H than does
Hb–O2, thereby removing H from solution and promoting The size of the thorax is altered by the action of the respiratory
conversion of CO2 to HCO3 via the reaction catalyzed by muscles, which contract as a result of nerve impulses transmitted
carbonic anhydrase. to them from centers in the brain and relax in the absence of nerve
impulses. These nerve impulses are sent from clusters of neurons
Summary of Gas Exchange located bilaterally in the medulla oblongata and pons of the
and Transport in Lungs and Tissues brain stem. This widely dispersed group of neurons, collectively
called the respiratory center, can be divided into three areas on
Deoxygenated blood returning to the pulmonary capillaries in the basis of their functions: (1) the medullary rhythmicity area in
the lungs (Figure 23.23a on page 904) contains CO2 dissolved in the medulla oblongata; (2) the pneumotaxic area in the pons; and
blood plasma, CO2 combined with globin as carbaminohemoglo- (3) the apneustic area, also in the pons (Figure 23.24).
bin (Hb–CO2), and CO2 incorporated into HCO3 within RBCs.
The RBCs have also picked up H, some of which binds to and Medullary Rhythmicity Area
therefore is buffered by hemoglobin (Hb–H). As blood passes The function of the medullary rhythmicity area (rith-MIS-i-tē)
through the pulmonary capillaries, molecules of CO2 dissolved is to control the basic rhythm of respiration. There are inspi-
in blood plasma and CO2 that dissociates from the globin portion ratory and expiratory areas within the medullary rhythmicity
of hemoglobin diffuse into alveolar air and are exhaled. At the
same time, inhaled O2 is diffusing from alveolar air into RBCs
and is binding to hemoglobin to form oxyhemoglobin (Hb–O2).
Carbon dioxide also is released from HCO3 when H combines Figure 23.24 Locations of areas of the respiratory center.
with HCO3 inside RBCs. The H2CO3 formed from this reaction
The respiratory center is composed of neurons in
then splits into CO2, which is exhaled, and H2O. As the concen-
the medullary rhythmicity area in the medulla
tration of HCO3 declines inside RBCs in pulmonary capillaries,
oblongata plus the pneumotaxic and apneustic
HCO3 diffuses in from the blood plasma, in exchange for Cl.
areas in the pons.
In sum, oxygenated blood leaving the lungs has increased O2
content and decreased amounts of CO2 and H. In systemic cap-
illaries, as cells use O2 and produce CO2, the chemical reactions
reverse (Figure 23.23b).
䊉 CHECKPOINT
24. In a resting person, how many O2 molecules are Sagittal
plane
attached to each hemoglobin molecule, on average, in
blood in the pulmonary arteries? In blood in the
pulmonary veins?
25. What is the relationship between hemoglobin and PO2? RESPIRATORY
CENTER:
How do temperature, H, PCO2, and BPG influence the Midbrain
affinity of Hb for O2? Pneumotaxic area
26. Why can hemoglobin unload more oxygen as blood Apneustic area
flows through capillaries of metabolically active tissues, Pons
such as skeletal muscle during exercise, than is Medullary rhythmicity
unloaded at rest? area:
Inspiratory area Medulla
oblongata
area. Figure 23.25 shows the relationships of the inspi- the transition between inhalation and exhalation. One of these
ratory and expiratory areas during normal quiet breathing and sites is the pneumotaxic area (noo-mō-TAK-sik; pneumo- air
forceful breathing. or breath; -taxic arrangement) in the upper pons (see Figure
During quiet breathing, inhalation lasts for about 2 seconds 23.24), which transmits inhibitory impulses to the inspiratory
and exhalation lasts for about 3 seconds. Nerve impulses gener- area. The major effect of these nerve impulses is to help turn off
ated in the inspiratory area establish the basic rhythm of the inspiratory area before the lungs become too full of air. In
breathing. While the inspiratory area is active, it generates nerve other words, the impulses shorten the duration of inhalation.
impulses for about 2 seconds (Figure 23.25a). The impulses When the pneumotaxic area is more active, breathing rate is
propagate to the external intercostal muscles via intercostal more rapid.
nerves and to the diaphragm via the phrenic nerves. When
the nerve impulses reach the diaphragm and external inter- Apneustic Area
costal muscles, the muscles contract and inhalation occurs. Even Another part of the brain stem that coordinates the transition
when all incoming nerve connections to the inspiratory area between inhalation and exhalation is the apneustic area
are cut or blocked, neurons in this area still rhythmically (ap-NOO-stik) in the lower pons (see Figure 23.24). This area
discharge impulses that cause inhalation. At the end of 2 sec- sends stimulatory impulses to the inspiratory area that activate it
onds, the inspiratory area becomes inactive and nerve impulses and prolong inhalation. The result is a long, deep inhalation.
cease. With no impulses arriving, the diaphragm and external When the pneumotaxic area is active, it overrides signals from
intercostal muscles relax for about 3 seconds, allowing the apneustic area.
passive elastic recoil of the lungs and thoracic wall. Then, the
cycle repeats. Regulation of the Respiratory Center
The neurons of the expiratory area remain inactive during
quiet breathing. However, during forceful breathing nerve The basic rhythm of respiration set and coordinated by the
impulses from the inspiratory area activate the expiratory area inspiratory area can be modified in response to inputs from other
(Figure 23.25b). Impulses from the expiratory area cause brain regions, receptors in the peripheral nervous system, and
contraction of the internal intercostal and abdominal muscles, other factors.
which decreases the size of the thoracic cavity and causes
forceful exhalation.
Cortical Influences on Respiration
Because the cerebral cortex has connections with the respiratory
Pneumotaxic Area center, we can voluntarily alter our pattern of breathing. We can
Although the medullary rhythmicity area controls the basic even refuse to breathe at all for a short time. Voluntary control is
rhythm of respiration, other sites in the brain stem help coordinate protective because it enables us to prevent water or irritating
Figure 23.25 Roles of the medullary rhythmicity area in controlling (a) the basic rhythm of respiration and
(b) forceful breathing.
During normal, quiet breathing, the expiratory area is inactive; during forceful breathing, the inspiratory
area activates the expiratory area.
Activates
INSPIRATORY AREA
INSPIRATORY AREA EXPIRATORY AREA
ACTIVE INACTIVE ACTIVE
2 seconds 3 seconds
Internal intercostal
Diaphragm and external Diaphragm, and abdominal
Diaphragm and external intercostals relax, sternocleidomastoid, muscles contract
intercostals contract followed by elastic and scalene muscles
recoil of lungs contract
Normal quiet inhalation Normal quiet exhalation Forceful inhalation Forceful exhalation
? Which nerves convey impulses from the respiratory center to the diaphragm?
CONTROL OF RESPIRATION 907
gases from entering the lungs. The ability to not breathe, how- Figure 23.26 Locations of peripheral chemoreceptors.
ever, is limited by the buildup of CO2 and H in the body. When
PCO2 and H concentrations increase to a certain level, the Chemoreceptors are sensory neurons that respond
to changes in the levels of certain chemicals in
inspiratory area is strongly stimulated, nerve impulses are sent
the body.
along the phrenic and intercostal nerves to inspiratory muscles,
and breathing resumes, whether the person wants it to or not. It
is impossible for small children to kill themselves by voluntarily
holding their breath, even though many have tried in order to
get their way. If breath is held long enough to cause fainting,
breathing resumes when consciousness is lost. Nerve impulses
from the hypothalamus and limbic system also stimulate the
respiratory center, allowing emotional stimuli to alter respira-
tions as, for example, in laughing and crying.
100 mmHg but is still above 50 mmHg, the peripheral chemore- Figure 23.27 Regulation of breathing in response to changes
ceptors are stimulated. Severe deficiency of O2 depresses activity in blood PCO2, PO2, and pH (Hⴙ concentration) via negative
of the central chemoreceptors and inspiratory area, which then feedback control.
do not respond well to any inputs and send fewer impulses to the
An increase in arterial blood PCO2 stimulates the
muscles of inhalation. As the breathing rate decreases or breath-
inspiratory center.
ing ceases altogether, PO2 falls lower and lower, establishing a
positive feedback cycle with a possibly fatal result.
The chemoreceptors participate in a negative feedback system
that regulates the levels of CO2, O2, and H in the blood (Figure Some stimulus disrupts
homeostasis by
23.27). As a result of increased PCO2, decreased pH (increased
H), or decreased PO2, input from the central and peripheral
chemoreceptors causes the inspiratory area to become highly ac-
Increasing
tive, and the rate and depth of breathing increase. Rapid and
deep breathing, called hyperventilation, allows the inhalation of
Arterial blood PCO2
more O2 and exhalation of more CO2 until PCO2 and H are low- (or decreasing pH or PO2)
ered to normal.
If arterial PCO2 is lower than 40 mmHg—a condition called
hypocapnia or hypocarbia—the central and peripheral chemo-
receptors are not stimulated, and stimulatory impulses are not Receptors
sent to the inspiratory area. As a result, the area sets its own Central Peripheral
moderate pace until CO2 accumulates and the PCO2 rises to chemo- chemo-
40 mmHg. The inspiratory center is more strongly stimulated receptors receptors
in in aortic
when PCO2 is rising above normal than when PO2 is falling below medulla and
normal. As a result, people who hyperventilate voluntarily and carotid
bodies
cause hypocapnia can hold their breath for an unusually long
period. Swimmers were once encouraged to hyperventilate just
before diving in to compete. However, this practice is risky
because the O2 level may fall dangerously low and cause faint- Input Nerve
ing before the PCO2 rises high enough to stimulate inhalation. impulses
If you faint on land you may suffer bumps and bruises, but if Control center
you faint in the water you could drown. Inspiratory area in Return to homeostasis
medulla oblongata when response brings
arterial blood PCO2, pH,
• CLINICAL CONNECT ION Hypoxia and PO2 back to normal
TABLE 23.2
Voluntary hyperventilation controlled by the cerebral cortex and Voluntary hypoventilation controlled by the cerebral cortex.
anticipation of activity by stimulation of the limbic system.
Increase in arterial blood PCO2 above 40 mmHg (causes an increase in H) Decrease in arterial blood PCO2 below 40 mmHg (causes a decrease in H)
detected by peripheral and central chemoreceptors. detected by peripheral and central chemoreceptors.
Decrease in arterial blood PO2 from 105 mmHg to 50 mmHg. Decrease in arterial blood PO2 below 50 mmHg.
Increased activity of proprioceptors. Decreased activity of proprioceptors.
Increase in body temperature. Decrease in body temperature decreases the rate of respiration, and a
sudden cold stimulus causes apnea.
Prolonged pain. Severe pain causes apnea.
Decrease in blood pressure. Increase in blood pressure.
Stretching the anal sphincter. Irritation of pharynx or larynx by touch or chemicals causes brief apnea
followed by coughing or sneezing.
910 CHAPTER 23 • THE RESPIRATORY SYSTEM
EXERCISE AND THE decreases airflow into and out of the lungs. (2) Carbon monoxide in
RESPIRATORY SYSTEM smoke binds to hemoglobin and reduces its oxygen-carrying capabil-
ity. (3) Irritants in smoke cause increased mucus secretion by the mu-
䊉 OBJECTIVE cosa of the bronchial tree and swelling of the mucosal lining, both of
• Describe the effects of exercise on the respiratory system. which impede airflow into and out of the lungs. (4) Irritants in smoke
also inhibit the movement of cilia and destroy cilia in the lining of
The respiratory and cardiovascular systems make adjustments in
the respiratory system. Thus, excess mucus and foreign debris are
response to both the intensity and duration of exercise. The
not easily removed, which further adds to the difficulty in breathing.
effects of exercise on the heart are discussed in Chapter 20. Here (5) With time, smoking leads to destruction of elastic fibers in the
we focus on how exercise affects the respiratory system. lungs and is the prime cause of emphysema (described on page
Recall that the heart pumps the same amount of blood to the 913). These changes cause collapse of small bronchioles and trap-
lungs as to all the rest of the body. Thus, as cardiac output rises, ping of air in alveoli at the end of exhalation. The result is less effi-
the blood flow to the lungs, termed pulmonary perfusion, cient gas exchange. •
increases as well. In addition, the O2 diffusing capacity, a mea-
sure of the rate at which O2 can diffuse from alveolar air into the
䊉 CHECKPOINT
blood, may increase threefold during maximal exercise because
more pulmonary capillaries become maximally perfused. As a 30. How does exercise affect the inspiratory area?
result, there is a greater surface area available for diffusion of O2
into pulmonary blood capillaries.
When muscles contract during exercise, they consume large DEVELOPMENT OF THE
amounts of O2 and produce large amounts of CO2. During RESPIRATORY SYSTEM
vigorous exercise, O2 consumption and pulmonary ventilation
䊉
both increase dramatically. At the onset of exercise, an abrupt OBJECTIVE
increase in pulmonary ventilation is followed by a more gradual • Describe the development of the respiratory system.
increase. With moderate exercise, the increase is due mostly to The development of the mouth and pharynx are discussed in
an increase in the depth of ventilation rather than to increased Chapter 24. Here we consider the development of the other
breathing rate. When exercise is more strenuous, the frequency structures of the respiratory system that you learned about in
of breathing also increases. this chapter.
The abrupt increase in ventilation at the start of exercise is At about four weeks of development, the respiratory system
due to neural changes that send excitatory impulses to the begins as an outgrowth of the foregut (precursor of some diges-
inspiratory area in the medulla oblongata. These changes include tive organs) just anterior to the pharynx. This outgrowth is called
(1) anticipation of the activity, which stimulates the limbic the respiratory diverticulum or lung bud (Figure 23.28). The
system; (2) sensory impulses from proprioceptors in muscles, endoderm lining the respiratory diverticulum gives rise to the
tendons, and joints; and (3) motor impulses from the primary epithelium and glands of the trachea, bronchi, and alveoli.
motor cortex (precentral gyrus).
gyrus) The more gradual increase in Mesoderm surrounding the respiratory diverticulum gives
ventilation during moderate exercise is due to chemical and rise to the connective tissue, cartilage, and smooth muscle of
physical changes in the bloodstream, including (1) slightly these structures.
decreased PO2, due to increased O2 consumption; (2) slightly The epithelial lining of the larynx develops from the endo-
increased PCO2, due to increased CO2 production by contracting derm of the respiratory diverticulum; the cartilages and muscles
muscle fibers; and (3) increased temperature, due to liberation of originate from the fourth and sixth pharyngeal arches,
more heat as more O2 is utilized. During strenuous exercise, swellings on the surface of the embryo.
HCO3 buffers H released by lactic acid in a reaction that liber- As the respiratory diverticulum elongates, its distal end
ates CO2, which further increases PCO2. enlarges to form a globular tracheal bud, which gives rise to the
At the end of an exercise session, an abrupt decrease in pul- trachea. Soon after, the tracheal bud divides into bronchial
monary ventilation is followed by a more gradual decline to the buds, which branch repeatedly and develop with the bronchi. By
resting level. The initial decrease is due mainly to changes in 24 weeks, 17 orders of branches have formed and respiratory
neural factors when movement stops or slows; the more gradual bronchioles have developed.
phase reflects the slower return of blood chemistry levels and During weeks 6 to 16, all major elements of the lungs have
temperature to the resting state. formed, except for those involved in gaseous exchange (respira-
tory bronchioles, alveolar ducts, and alveoli). Since respiration
• CLINICAL CONNECT ION The Effect of Smoking is not possible at this stage, fetuses born during this time
on Respiratory Efficiency cannot survive.
During weeks 16 to 26, lung tissue becomes highly vascular
Smoking may cause a person to become easily “winded” during even and respiratory bronchioles, alveolar ducts, and some primitive
moderate exercise because several factors decrease respiratory effi- alveoli develop. Although it is possible for a fetus born near the
ciency in smokers: (1) Nicotine constricts terminal bronchioles, which
end of this period to survive if given intensive care, death
AGING AND THE RESPIRATORY SYSTEM 911
Figure 23.28 Development of the bronchial tubes and lungs. of alveolar fluid and thus reduce the tendency of alveoli to col-
lapse on exhalation. Although surfactant production begins by
The respiratory system develops from endoderm 20 weeks, it is present in only small quantities. Amounts
and mesoderm.
sufficient to permit survival of a premature (preterm) infant
are not produced until 26 to 28 weeks gestation. Infants born
Pharynx
Pharynx before 26 to 28 weeks are at high risk of respiratory distress
Respiratory syndrome (RDS), in which the alveoli collapse during exhala-
diverticulum Trachea
tion and must be reinflated during inhalation ((see page 894).
Tracheal bud At about 30 weeks, mature alveoli develop. However, it is es-
Esophagus timated that only about one-sixth of the full complement of alve-
Bronchial oli develop before birth; the remainder develop after birth during
buds
the first eight years.
Esophagus
As the lungs develop, they acquire their pleural sacs. The vis-
Fourth week ceral pleura and the parietal pleura develop from mesoderm.
The space between the pleural layers is the pleural cavity.
During development, breathing movements of the fetus cause
Left primary
bronchus the aspiration of fluid into the lungs. This fluid is a mixture of
amniotic fluid, mucus from bronchial glands, and surfactant. At
Trachea Left secondary
bronchi birth, the lungs are about half-filled with fluid. When breathing
Right primary begins at birth, most of the fluid is rapidly reabsorbed by blood
bronchus
and lymph capillaries and a small amount is expelled through
the nose and mouth during delivery.
䊉 CHECKPOINT
Right Right Left 31. What structures develop from the laryngotracheal bud?
secondary tertiary tertiary
bronchi bronchi bronchi
Muscular Increased rate and depth of breathing support increased activity of skeletal muscles
system during exercise.
Nervous Nose contains receptors for sense of smell (olfaction). Vibrations of air flowing across
system vocal folds produce sounds for speech.
Endocrine Angiotensin converting enzyme (ACE) in lungs catalyzes formation of the hormone
system angiotensin II from angiotensin I.
Cardiovascular During inhalations, respiratory pump aids return of venous blood to the heart.
system
Lymphatic Hairs in nose, cilia and mucus in trachea, bronchi, and smaller airways, and alveolar THE RESPIRATORY SYSTEM
system and macrophages contribute to nonspecific resistance to disease. Pharynx (throat)
immunity contains lymphatic tissue (tonsils). Respiratory pump (during inhalation)
promotes flow of lymph.
Reproductive Increased rate and depth of breathing support activity during sexual intercourse. Internal respiration provides oxygen
systems to developing fetus.
912
DISORDERS: HOMEOSTATIC IMBALANCES 913
Asthma tion during inhalation increases the size of the chest cage, resulting
Asthma (AZ-ma panting) is a disorder characterized by chronic in a “barrel chest.”
airway inflammation, airway hypersensitivity to a variety of stimuli, Emphysema is generally caused by a long-term irritation; cigarette
and airway obstruction. It is at least partially reversible, either sponta- smoke, air pollution, and occupational exposure to industrial dust are
neously or with treatment. Asthma affects 3–5% of the U.S. popula- the most common irritants. Some destruction of alveolar sacs may be
tion and is more common in children than in adults. Airway obstruc- caused by an enzyme imbalance. Treatment consists of cessation of
tion may be due to smooth muscle spasms in the walls of smaller smoking, removal of other environmental irritants, exercise training
bronchi and bronchioles, edema of the mucosa of the airways, under careful medical supervision, breathing exercises, use of bron-
increased mucus secretion, and/or damage to the epithelium of the chodilators, and oxygen therapy.
airway.
Chronic Bronchitis
Individuals with asthma typically react to concentrations of agents
too low to cause symptoms in people without asthma. Sometimes Chronic bronchitis is a disorder characterized by excessive secretion
the trigger is an allergen such as pollen, house dust mites, molds, or of bronchial mucus accompanied by a productive cough (sputum is
a particular food. Other common triggers of asthma attacks are emo- raised) that lasts for at least three months of the year for two succes-
tional upset, aspirin, sulfiting agents (used in wine and beer and to sive years. Cigarette smoking is the leading cause of chronic bronchi-
keep greens fresh in salad bars), exercise, and breathing cold air or tis. Inhaled irritants lead to chronic inflammation with an increase in
cigarette smoke. In the early phase (acute) response, smooth muscle the size and number of mucous glands and goblet cells in the airway
spasm is accompanied by excessive secretion of mucus that may clog epithelium. The thickened and excessive mucus produced narrows the
the bronchi and bronchioles and worsen the attack. The late phase airway and impairs ciliary function. Thus, inhaled pathogens become
(chronic) response is characterized by inflammation, fibrosis, edema, embedded in airway secretions and multiply rapidly. Besides a pro-
and necrosis (death) of bronchial epithelial cells. A host of mediator ductive cough, symptoms of chronic bronchitis are shortness of
chemicals, including leukotrienes, prostaglandins, thromboxane, platelet- breath, wheezing, cyanosis, and pulmonary hypertension. Treatment
activating factor, and histamine, take part. for chronic bronchitis is similar to that for emphysema.
Symptoms include difficult breathing, coughing, wheezing, chest
tightness, tachycardia, fatigue, moist skin, and anxiety. An acute attack
Lung Cancer
is treated by giving an inhaled beta2-adrenergic agonist (albuterol) to In the United States, lung cancer is the leading cause of cancer death
help relax smooth muscle in the bronchioles and open up the airways. in both males and females, accounting for 160,000 deaths annually. At
However, long-term therapy of asthma strives to suppress the underly- the time of diagnosis, lung cancer is usually well advanced, with dis-
ing inflammation. The anti-inflammatory drugs that are used most of- tant metastases present in about 55% of patients, and regional lymph
ten are inhaled corticosteroids (glucocorticoids), cromolyn sodium node involvement in an additional 25%. Most people with lung cancer
(Intal®), and leukotriene blockers (Accolate®). die within a year of the initial diagnosis; the overall survival rate is
only 10–15%. Cigarette smoke is the most common cause of lung
Chronic Obstructive Pulmonary Disease cancer. Roughly 85% of lung cancer cases are related to smoking, and
Chronic obstructive pulmonary disease (COPD) is a type of respiratory the disease is 10 to 30 times more common in smokers than non-
disorder characterized by chronic and recurrent obstruction of airflow, smokers. Exposure to secondhand smoke is also associated with lung
which increases airway resistance. COPD affects about 30 million cancer and heart disease. In the United States, secondhand smoke
Americans and is the fourth leading cause of death behind heart dis- causes an estimated 4000 deaths a year from lung cancer, and nearly
ease, cancer, and cerebrovascular disease. The principal types of COPD 40,000 deaths a year from heart disease. Other causes of lung cancer
are emphysema and chronic bronchitis. In most cases, COPD is pre- are ionizing radiation and inhaled irritants, such as asbestos and
ventable because its most common cause is cigarette smoking or radon gas. Emphysema is a common precursor to the development of
breathing secondhand smoke. Other causes include air pollution, lung cancer.
pulmonary infection, occupational exposure to dusts and gases, and The most common type of lung cancer, bronchogenic carcinoma,
genetic factors. Because men, on average, have more years of exposure starts in the epithelium of the bronchial tubes. Bronchogenic tumors
to cigarette smoke than women, they are twice as likely as women are named based on where they arise. For example, adenocarcinomas
to suffer from COPD; still, the incidence of COPD in women has develop in peripheral areas of the lungs from bronchial glands and
risen sixfold in the past 50 years, a reflection of increased smoking alveolar cells, squamous cell carcinomas develop from the epithelium
among women. of larger bronchial tubes, and small (oat) cell carcinomas develop
from epithelial cells in primary bronchi near the hilum of the lungs
Emphysema and tend to involve the mediastinum early on. Depending on the type
Emphysema (em-fi-SĒ-ma blown up or full of air) is a disorder of bronchogenic tumors, they may be aggressive, locally invasive, and
characterized by destruction of the walls of the alveoli, producing undergo widespread metastasis. The tumors begin as epithelial le-
abnormally large air spaces that remain filled with air during exhala- sions that grow to form masses that obstruct the bronchial tubes or
tion. With less surface area for gas exchange, O2 diffusion across the invade adjacent lung tissue. Bronchogenic carcinomas metastasize to
damaged respiratory membrane is reduced. Blood O2 level is some- lymph nodes, the brain, bones, liver, and other organs.
what lowered, and any mild exercise that raises the O2 requirements Symptoms of lung cancer are related to the location of the tumor.
of the cells leaves the patient breathless. As increasing numbers of These may include a chronic cough, spitting blood from the respira-
alveolar walls are damaged, lung elastic recoil decreases due to loss tory tract, wheezing, shortness of breath, chest pain, hoarseness, diffi-
of elastic fibers, and an increasing amount of air becomes trapped in culty swallowing, weight loss, anorexia, fatigue, bone pain, confusion,
the lungs at the end of exhalation. Over several years, added exer- problems with balance, headache, anemia, thrombocytopenia, and
914 CHAPTER 23 • THE RESPIRATORY SYSTEM
jaundice. For some people, there are relatively few or no dramatic sive nasal secretion, dry cough, and congestion. The uncomplicated
symptoms. common cold is not usually accompanied by a fever. Complications
Treatment consists of partial or complete surgical removal of a include sinusitis, asthma, bronchitis, ear infections, and laryngitis.
diseased lung (pulmonectomy), radiation therapy, and chemotherapy. Recent investigations suggest an association between emotional
stress and the common cold. The higher the stress level, the greater
Pneumonia the frequency and duration of colds.
Pneumonia is an acute infection or inflammation of the alveoli. It Influenza (flu) is also caused by a virus. Its symptoms include
is the most common infectious cause of death in the United chills, fever (usually higher than 101°F 39°C), headache, and muscu-
States, where an estimated 4 million cases occur annually. When cer- lar aches. Influenza can become life-threatening and may develop into
tain microbes enter the lungs of susceptible individuals, they release pneumonia. It is important to recognize that influenza is a respiratory
damaging toxins, stimulating inflammation and immune responses that disease, not a gastrointestinal (GI) disease. Many people mistakenly
have damaging side effects. The toxins and immune response damage report having “the flu” when they are suffering from a GI illness.
alveoli and bronchial mucous membranes; inflammation and edema
cause the alveoli to fill with fluid, interfering with ventilation and Pulmonary Edema
gas exchange. Pulmonary edema is an abnormal accumulation of fluid in the inter-
The most common cause of pneumonia is the pneumococcal stitial spaces and alveoli of the lungs. The edema may arise from
bacterium Streptococcus pneumoniae, but other microbes may also increased permeability of the pulmonary capillaries (pulmonary origin)
cause pneumonia. Those who are most susceptible to pneumonia are or increased pressure in the pulmonary capillaries (cardiac origin); the
the elderly, infants, immunocompromised individuals (AIDS or cancer latter cause may coincide with congestive heart failure. The most
patients, or those taking immunosuppressive drugs), cigarette smok- common symptom is dyspnea. Others include wheezing, tachypnea
ers, and individuals with an obstructive lung disease. Most cases of (rapid breathing rate), restlessness, a feeling of suffocation, cyanosis,
pneumonia are preceded by an upper respiratory infection that often pallor (paleness), diaphoresis (excessive perspiration), and pulmonary
is viral. Individuals then develop fever, chills, productive or dry cough, hypertension. Treatment consists of administering oxygen, drugs that
malaise, chest pain, and sometimes dyspnea (difficult breathing) and dilate the bronchioles and lower blood pressure, diuretics to rid the
hemoptysis (spitting blood). body of excess fluid, and drugs that correct acid–base imbalance;
Treatment may involve antibiotics, bronchodilators, oxygen therapy, suctioning of airways; and mechanical ventilation. One of the recent
increased fluid intake, and chest physiotherapy (percussion, vibration, culprits in the development of pulmonary edema was found to be
and postural drainage). “phen-fen” diet pills.
Asbestos-related Diseases ity in the mechanisms that control respiration or low levels of oxy-
Asbestos-related diseases are serious lung disorders that develop as gen in the blood. SIDS may also be linked to hypoxia while sleep-
a result of inhaling asbestos particles decades earlier. When asbestos ing in a prone position (on the stomach) and the rebreathing of ex-
particles are inhaled, they penetrate lung tissue. In response, white haled air trapped in a depression of a mattress. It is recommended
blood cells attempt to destroy them by phagocytosis. However, the that for the first six months infants be placed on their backs for
fibers usually destroy the white blood cells and scarring of lung tissue sleeping (“back to sleep”).
may follow. Asbestos-related diseases include asbestosis (widespread
Severe Acute Respiratory Syndrome
scarring of lung tissue), diffuse pleural thickening (thickening of the
pleurae), and mesothelioma (cancer of the pleurae or, less commonly, Severe acute respiratory syndrome (SARS) is an example of an emerg-
the peritoneum). ing infectious disease, that is, a disease that is new or changing.
Other examples of emerging infectious diseases are West Nile
Sudden Infant Death Syndrome encephalitis, mad cow disease, and AIDS. SARS first appeared in
Sudden infant death syndrome (SIDS) is the sudden, unexpected Southern China in late 2002 and has subsequently spread worldwide.
death of an apparently healthy infant during sleep. It rarely occurs It is a respiratory illness caused by a new variety of coronavirus.
before 2 weeks or after 6 months of age, with the peak incidence Symptoms of SARS include fever, malaise, muscle aches, nonproduc-
between the second and fourth months. SIDS is more common in tive (dry) cough, difficulty in breathing, chills, headache, and diarrhea.
premature infants, male babies, low-birth-weight babies, babies of About 10–20% of patients require mechanical ventilation and in some
drug users or smokers, babies who have stopped breathing and cases death may result. The disease is primarily spread through per-
have had to be resuscitated, babies with upper respiratory tract in- son-to-person contact. There is no effective treatment for SARS and
fections, and babies who have had a sibling die of SIDS. African the death rate is 5–10%, usually among the elderly and in persons
American and Native American babies are at higher risk. The exact with other medical problems.
cause of SIDS is unknown. However, it may be due to an abnormal-
MEDICAL TERMINOLOGY
Abdominal thrust maneuver First-aid procedure designed to clear the Bronchoscopy (bron-KOS-kō -pē) Visual examination of the bronchi
airways of obstructing objects. It is performed by applying a quick through a bronchoscope, an illuminated, flexible tubular instru-
upward thrust between the navel and costal margin that causes ment that is passed through the mouth (or nose), larynx, and
sudden elevation of the diaphragm and forceful, rapid expulsion of trachea into the bronchi. The examiner can view the interior of the
air in the lungs; this action forces air out the trachea to eject the trachea and bronchi to biopsy a tumor, clear an obstructing object
obstructing object. The abdominal thrust maneuver is also used to or secretions from an airway, take cultures or smears for micro-
expel water from the lungs of near-drowning victims before resusci- scopic examination, stop bleeding, or deliver drugs.
tation is begun. Previously called the Heimlich maneuver (HĪ M-lik Cheyne–Stokes respiration (CHĀN STŌ KS res-pi-RĀ-shun) A repeated
ma-NOO-ver). cycle of irregular breathing that begins with shallow breaths that
Asphyxia (as-FIK-sē-a; sphyxia pulse) Oxygen starvation due to low increase in depth and rapidity and then decrease and cease alto-
atmospheric oxygen or interference with ventilation, external respi- gether for 15 to 20 seconds. Cheyne–Stokes is normal in infants; it
ration, or internal respiration. is also often seen just before death from pulmonary, cerebral, car-
Aspiration (as-pi-RĀ-shun) Inhalation of a foreign substance such as diac, and kidney disease.
water, food, or a foreign body into the bronchial tree; also, the Dyspnea (DISP-nē-a; dys- painful, difficult) Painful or labored
drawing of a substance in or out by suction. breathing.
Avian influenza A respiratory disorder that has resulted in the deaths Epistaxis (ep-i-STAK-sis) Loss of blood from the nose due to trauma,
of hundreds of millions of birds worldwide. It is usually transmitted infection, allergy, malignant growths, or bleeding disorders. It can
from one bird to another bird through their droppings, saliva, and be arrested by cautery with silver nitrate, electrocautery, or firm
nasal secretions. Currently, avian influenza is difficult to transmit packing. Also called nosebleed.
from birds to humans; the few humans who have died from avian Hypoventilation (hypo- below) Slow and shallow breathing.
influenza have had close contact with infected birds. Also called
Mechanical ventilation The use of an automatically cycling device
bird flu.
(ventilator or respirator) to assist breathing. A plastic tube is
Black lung disease A condition in which the lungs appear black in- inserted into the nose or mouth and the tube is attached to a de-
stead of pink due to inhalation of coal dust over a period of many vice that forces air into the lungs. Exhalation occurs passively due
years. Most often it affects people who work in the coal industry. to the elastic recoil of the lungs.
Bronchiectasis (bron-kē-EK-ta-sis; -ektasis stretching) A chronic di- Rales (RĀLS) Sounds sometimes heard in the lungs that resemble
lation of the bronchi or bronchioles resulting from damage to the bubbling or rattling. Rales are to the lungs what murmurs are to
bronchial wall, for example, from respiratory infections. the heart. Different types are due to the presence of an abnormal
Bronchography (bron-KOG-ra-fē) An imaging technique used to type or amount of fluid or mucus within the bronchi or alveoli, or
visualize the bronchial tree using x-rays. After an opaque contrast to bronchoconstriction that causes turbulent airflow.
medium is inhaled through an intratracheal catheter, radiographs Respirator (RES-pi-rā-tor) An apparatus fitted to a mask over the nose
of the chest in various positions are taken, and the developed film, and mouth, or hooked directly to an endotracheal or tracheotomy
a bronchogram (BRON-kō -gram), provides a picture of the tube, that is used to assist or support ventilation or to provide
bronchial tree. nebulized medication to the air passages.
916 CHAPTER 23 • THE RESPIRATORY SYSTEM
Respiratory failure A condition in which the respiratory system either Sputum (SPŪ -tum to spit) Mucus and other fluids from the air
cannot supply sufficient O2 to maintain metabolism or cannot elim- passages that is expectorated (expelled by coughing).
inate enough CO2 to prevent respiratory acidosis (a lower-than- Strep throat Inflammation of the pharynx caused by the bacterium
normal pH in interstitial fluid). Streptococcus pyogenes. It may also involve the tonsils and mid-
Rhinitis (rı̄ -NĪ -tis; rhin- nose) Chronic or acute inflammation of the dle ear.
mucous membrane of the nose due to viruses, bacteria, or irritants. Tachypnea (tak-ip-NĒ-a; tachy- rapid; -pnea breath) Rapid
Excessive mucus production produces a runny nose, nasal conges- breathing rate.
tion, and postnasal drip.
Wheeze (HWEĒZ) A whistling, squeaking, or musical high-pitched
Sleep apnea (AP-nē-a; a- without; -pnea breath) A disorder in sound during breathing resulting from a partially obstructed airway.
which a person repeatedly stops breathing for 10 or more seconds
while sleeping. Most often, it occurs because loss of muscle tone
in pharyngeal muscles allows the airway to collapse.
Respiratory System Anatomy (p. 875) 12. The right lung has three lobes separated by two fissures; the left
lung has two lobes separated by one fissure and a depression, the
1. The respiratory system consists of the nose, pharynx, larynx, tra-
cardiac notch.
chea, bronchi, and lungs. They act with the cardiovascular system
13. Secondary bronchi give rise to branches called segmental bronchi,
to supply oxygen (O2) and remove carbon dioxide (CO2) from
which supply segments of lung tissue called bronchopulmonary
the blood.
segments.
2. The external portion of the nose is made of cartilage and skin and
14. Each bronchopulmonary segment consists of lobules, which
is lined with a mucous membrane. Openings to the exterior are the
contain lymphatics, arterioles, venules, terminal bronchioles, res-
external nares.
piratory bronchioles, alveolar ducts, alveolar sacs, and alveoli.
3. The internal portion of the nose communicates with the paranasal
15. Alveolar walls consist of type I alveolar cells, type II alveolar
sinuses and nasopharynx through the internal nares.
cells, and associated alveolar macrophages.
4. The nasal cavity is divided by a septum. The anterior portion of
16. Gas exchange occurs across the respiratory membranes.
the cavity is called the vestibule. The nose warms, moistens, and
filters air and functions in olfaction and speech. Pulmonary Ventilation (p. 890)
5. The pharynx (throat) is a muscular tube lined by a mucous mem-
brane. The anatomic regions are the nasopharynx, oropharynx, and 1. Pulmonary ventilation, or breathing, consists of inhalation and
laryngopharynx. exhalation.
6. The nasopharynx functions in respiration. The oropharynx and 2. The movement of air into and out of the lungs depends on pressure
laryngopharynx function both in digestion and in respiration. changes governed in part by Boyle’s law, which states that the
7. The larynx (voice box) is a passageway that connects the volume of a gas varies inversely with pressure, assuming that
pharynx with the trachea. It contains the thyroid cartilage (Adam’s temperature remains constant.
apple); the epiglottis, which prevents food from entering the 3. Inhalation occurs when alveolar pressure falls below atmospheric
larynx; the cricoid cartilage, which connects the larynx and pressure. Contraction of the diaphragm and external intercostals
trachea; and the paired arytenoid, corniculate, and cuneiform increases the size of the thorax, thereby decreasing the intrapleural
cartilages. pressure so that the lungs expand. Expansion of the lungs
8. The larynx contains vocal folds, which produce sound as they decreases alveolar pressure so that air moves down a pressure
vibrate. Taut folds produce high pitches, and relaxed ones produce gradient from the atmosphere into the lungs.
low pitches. 4. During forceful inhalation, accessory muscles of inhalation (ster-
9. The trachea (windpipe) extends from the larynx to the primary nocleidomastoids, scalenes, and pectoralis minors) are also used.
bronchi. It is composed of C-shaped rings of cartilage and 5. Exhalation occurs when alveolar pressure is higher than
smooth muscle and is lined with pseudostratified ciliated columnar atmospheric pressure. Relaxation of the diaphragm and external
epithelium. intercostals results in elastic recoil of the chest wall and lungs,
10. The bronchial tree consists of the trachea, primary bronchi, which increases intrapleural pressure; lung volume decreases
secondary bronchi, tertiary bronchi, bronchioles, and terminal and alveolar pressure increases, so air moves from the lungs to
bronchioles. Walls of bronchi contain rings of cartilage; walls of the atmosphere.
bronchioles contain increasingly smaller plates of cartilage and 6. Forceful exhalation involves contraction of the internal intercostal
increasing amounts of smooth muscle. and abdominal muscles.
11. Lungs are paired organs in the thoracic cavity enclosed by the 7. The surface tension exerted by alveolar fluid is decreased by the
pleural membrane. The parietal pleura is the superficial layer that presence of surfactant.
lines the thoracic cavity; the visceral pleura is the deep layer that 8. Compliance is the ease with which the lungs and thoracic wall can
covers the lungs. expand.
STUDY OUTLINE 917
9. The walls of the airways offer some resistance to breathing. 4. In each 100 mL of deoxygenated blood, 7% of CO2 is dissolved
10. Normal quiet breathing is termed eupnea; other patterns are costal in blood plasma, 23% combines with hemoglobin as carbamino-
breathing and diaphragmatic breathing. Modified respiratory move- hemoglobin (Hb–CO2), and 70% is converted to bicarbonate
ments, such as coughing, sneezing, sighing, yawning, sobbing, ions (HCO3).
crying, laughing, and hiccupping, are used to express emotions and 5. In an acidic environment, hemoglobin’s affinity for O2 is lower,
to clear the airways. (See Table 23.1 on page 895.) and O2 dissociates more readily from it (Bohr effect).
6. In the presence of O2, less CO2 binds to hemoglobin (Haldane
Lung Volumes and Capacities (p. 894) effect).
1. Lung volumes exchanged during breathing and the rate of respira-
Control of Respiration (p. 905)
tion are measured with a spirometer.
2. Lung volumes measured by spirometry include tidal volume, 1. The respiratory center consists of a medullary rhythmicity area in
minute ventilation, alveolar ventilation rate, inspiratory reserve the medulla oblongata and a pneumotaxic area and an apneustic
volume, expiratory reserve volume, and FEV1.0. Other lung volumes area in the pons.
include anatomic dead space, residual volume, and minimal 2. The inspiratory area sets the basic rhythm of respiration.
volume. 3. The pneumotaxic and apneustic areas coordinate the transition
3. Lung capacities, the sum of two or more lung volumes, include between inhalation and exhalation.
inspiratory, functional, residual, vital, and total lung capacities. 4. Respirations may be modified by a number of factors, including
cortical influences; the inflation reflex; chemical stimuli, such as
Exchange of Oxygen and Carbon Dioxide (p. 896) O2 and CO2 and H levels; proprioceptor input; blood pressure
changes; limbic system stimulation; temperature; pain; and irrita-
1. The partial pressure of a gas is the pressure exerted by that gas in
tion to the airways. (See Table 23.2 on page 909.)
a mixture of gases. It is symbolized by Px, where the subscript is
the chemical formula of the gas.
Exercise and the Respiratory System (p. 910)
2. According to Dalton’s law, each gas in a mixture of gases exerts its
own pressure as if all the other gases were not present. 1. The rate and depth of ventilation change in response to both the
3. Henry’s law states that the quantity of a gas that will dissolve in a intensity and duration of exercise.
liquid is proportional to the partial pressure of the gas and its 2. An increase in pulmonary perfusion and O2-diffusing capacity
solubility (given that the temperature remains constant). occurs during exercise.
4. In internal and external respiration, O2 and CO2 diffuse from areas 3. The abrupt increase in ventilation at the start of exercise is due to
of higher partial pressures to areas of lower partial pressures. neural changes that send excitatory impulses to the inspiratory area
5. External respiration or pulmonary gas exchange is the exchange of in the medulla oblongata. The more gradual increase in ventilation
gases between alveoli and pulmonary blood capillaries. It depends during moderate exercise is due to chemical and physical changes
on partial pressure differences, a large surface area for gas exchange, in the bloodstream.
a small diffusion distance across the respiratory membrane, and
the rate of airflow into and out of the lungs. Development of the Respiratory System (p. 910)
6. Internal respiration or systemic gas exchange is the exchange of
1. The respiratory system begins as an outgrowth of endoderm called
gases between systemic blood capillaries and tissue cells.
the respiratory diverticulum.
2. Smooth muscle, cartilage, and connective tissue of the bronchial
Transport of Oxygen and Carbon Dioxide (p. 900)
tubes and pleural sacs develop from mesoderm.
1. In each 100 mL of oxygenated blood, 1.5% of the O2 is dissolved
in blood plasma and 98.5% is bound to hemoglobin as oxyhemo- Aging and the Respiratory System (p. 911)
globin (Hb–O2).
1. Aging results in decreased vital capacity, decreased blood level of
2. The binding of O2 to hemoglobin is affected by PO2, acidity (pH),
O2, and diminished alveolar macrophage activity.
PCO2, temperature, and 2,3-bisphosphoglycerate (BPG).
2. Elderly people are more susceptible to pneumonia, emphysema,
3. Fetal hemoglobin differs from adult hemoglobin in structure and
bronchitis, and other pulmonary disorders.
has a higher affinity for O2.
918 CHAPTER 23 • THE RESPIRATORY SYSTEM
Fill in the blanks in the following statements. initial onset of exercise due to input to the inspiratory area from
1. Oxygen in blood is carried primarily in the form of _____; carbon proprioceptors. (3) When baroreceptors in the lungs are stimu-
dioxide is carried as _____, _____, and _____. lated, the expiratory area is activated. (4) Stimulation of the limbic
2. Write the equation for the chemical reaction that occurs for the system can result in excitation of the inspiratory area. (5) Sudden
transport of carbon dioxide as bicarbonate ions in blood: _____. severe pain causes brief apnea, while prolonged somatic pain
causes an increase in respiratory rate. (6) The respiratory rate in-
Indicate whether the following statements are true or false. creases during fever.
3. The three basic steps of respiration are pulmonary ventilation, ex- (a) 1, 2, 3, and 6 (b) 1, 4, and 5 (c) 1, 2, 4, 5, and 6
ternal respiration, and cellular respiration. (d) 2, 3, 4, 5, and 6 (e) 2, 4, 5, and 6
4. For inhalation to occur, air pressure in the alveoli must be less than 11. Place the steps for normal inhalation in order.
atmospheric pressure; for exhalation to occur, air pressure in the (a) decrease in intrapleural pressure to 754mmHg
alveoli must be greater than atmospheric pressure. (b) increase in the size of the thoracic cavity
Choose the one best answer to the following questions. (c) flow of air from higher to lower pressure
5. What structural changes occur from primary bronchi to terminal (d) outward pull of pleurae, resulting in lung expansion
bronchioles? (1) The mucous membrane changes from pseudo- (e) stimulation of primary breathing muscles by phrenic and inter-
stratified ciliated columnar epithelium to nonciliated simple costal nerves
cuboidal epithelium. (2) The number of goblet cells increases. (f) decrease in alveolar pressure to 758mmHg
(3) The amount of smooth muscle increases. (4) Incomplete rings (g) contraction of the diaphragm and external intercostals
of cartilage disappear. (5) The amount of branching decreases. (h) increase in the volume of the pleural cavity
(a) 1, 2, 3, 4, and 5 (b) 2, 3, and 4 (c) 1, 3, and 4 12. Match the following:
(d) 1, 3, 4, and 5 (e) 1, 2, 3, and 4 (a) functions as a passageway (1) nose
6. Which of the following would cause oxygen to dissociate more for air and food, provides a (2) pharynx
readily from hemoglobin? (1) low PO2, (2) an increase in H in resonating chamber for (3) larynx
blood, (3) hypercapnia, (4) hypothermia, (5) low levels of BPG speech sounds, and houses (4) epiglottis
(2,3-bisphosphoglycerate). the tonsils (5) trachea
(a) 1 and 2 (b) 2, 3, and 4 (c) 1, 2, 3, and 5 (b) site of external respiration (6) bronchi
(d) 1, 3, and 5 (e) 1, 2, and 3 (c) connects the laryngopharynx (7) carina
with the trachea; houses the (8) cricoid cartilage
7. Which of the following statements are correct? (1) Normal exhala- vocal cords (9) pleura
tion during quiet breathing is an active process involving intensive (d) serous membrane that (10) thyroid cartilage
muscle contraction. (2) Passive exhalation results from elastic re- surrounds the lungs (11) alveoli
coil of the chest wall and lungs. (3) Air flow during breathing is (e) functions in warming, (12) type I alveolar cells
due to a pressure gradient between the lungs and the atmospheric moistening, and filtering air; (13) type II alveolar cells
air. (4) During normal breathing, the pressure between the two receives olfactory stimuli;
pleural layers (intrapleural pressure) is always subatmospheric. is a resonating chamber
(5) Surface tension of alveolar fluid facilitates inhalation. for sound
(a) 1, 2, and 3 (b) 2, 3, and 4 (c) 3, 4, and 5 (f) simple squamous epithelial
(d) 1, 3, and 5 (e) 2, 3, and 5 cells that form a continuous
8. Which of the following factors affect the rate of external respira- lining of the alveolar wall;
tion? (1) partial pressure differences of the gases, (2) surface area sites of gas exchange
for gas exchange, (3) diffusion distance, (4) solubility and molecu- (g) forms anterior wall of
lar weight of the gases, (5) presence of bisphosphoglycerate the larynx
(BPG). (h) a tubular passageway for air
(a) 1, 2, and 3 (b) 2, 4, and 5 (c) 1, 2, 4, and 5 connecting the larynx to the
(d) 1, 2, 3, and 4 (e) 2, 3, 4, and 5 bronchi
(i) secrete alveolar fluid and
9. The most important factor in determining the percent oxygen satu-
surfactant
ration of hemoglobin is
(j) forms inferior wall of
(a) the partial pressure of oxygen.
larynx; landmark for
(b) acidity.
tracheotomy
(c) the partial pressure of carbon dioxide.
(k) prevents food or fluid from
(d) temperature.
entering the airways
(e) BPG.
(l) air passageways entering
10. Which of the following statements are true? (1) Peripheral and the lungs
central chemoreceptors are stimulated by an increase in PCO2 and (m) ridge covered by a sensitive
H and a decrease in O2. (2) Respiratory rate increases during the mucous membrane; irritation
triggers cough reflex
SELF-QUIZ QUESTIONS 919
13. Match the following: 15. Match the following:
(a) a deficiency of oxygen at (1) eupnea (a) prevents excessive (1) Bohr effect
the tissue level (2) apnea inflation of the lungs (2) Dalton’s law
(b) above-normal partial (3) hyperventilation (b) the lower the amount of (3) medullary
pressure of carbon dioxide (4) costal breathing oxyhemoglobin, the rhythmicity area
(c) normal quiet breathing (5) diaphragmatic higher the carbon dioxide (4) inspiratory area
(d) deep, abdominal breathing breathing carrying capacity of the (5) expiratory area
(e) the ease with which the (6) compliance blood (6) apneustic area
lungs and thoracic wall can (7) hypoxia (c) controls the basic rhythm (7) pneumotaxic area
be expanded (8) hypercapnia of respiration (8) Henry’s law
(f) hypoxia-induced (9) ventilation–- (d) active during normal (9) inflation
vasoconstriction to divert perfusion coupling inhalation; sends nerve (Hering–Breuer)
pulmonary blood from impulses to external reflex
poorly ventilated to well- intercostals and (10) Boyle’s law
ventilated regions of the diaphragm (11) Haldane effect
lungs (e) sends stimulatory
(g) absence of breathing impulses to the inspiratory
(h) rapid and deep breathing area that activate it and
(i) shallow, chest breathing prolong inhalation
14. Match the following: (f) as acidity increases, the
(a) total volume of air inhaled (1) tidal volume affinity of hemoglobin for
and exhaled each minute (2) residual volume oxygen decreases and
(b) tidal volume inspiratory (3) minute ventilation oxygen dissociates more
reserve volume expira- (4) expiratory reserve readily from hemoglobin;
tory reserve volume volume shifts oxygen-dissociation
(c) additional amount of air in- (5) inspiratory reserve curve to the right
haled beyond tidal volume volume (g) active during forceful
when taking a very deep (6) minimal volume exhalation
breath (7) inspiratory capacity (h) pressure of a gas in a
(d) residual volume expira- (8) vital capacity closed container is
tory reserve volume (9) functional residual inversely proportional
(e) amount of air remaining in volume to the volume of the
lungs after expiratory re- (10) total lung capacity container
serve volume is expelled (i) transmits inhibitory
(f) tidal volume inspiratory impulses to turn off the
reserve volume inspiratory area before
(g) vital capacity residual the lungs become too full
volume of air
(h) volume of air in one breath (j) the quantity of a gas that
(i) amount of air exhaled in dissolves in a liquid is
forced exhalation proportional to the partial
(j) provides a medical and le- pressure of the gas and its
gal tool for determining if a solubility
baby was born dead or died (k) relates to the partial
after birth pressure of a gas in a
mixture of gases whereby
each gas in a mixture
exerts its own pressure as
if all the other gases were
not present
920 CHAPTER 23 • THE RESPIRATORY SYSTEM
23.1 The conducting zone of the respiratory system includes the nose, 23.15 Normal atmospheric pressure at sea level is 760 mmHg.
pharynx, larynx, trachea, bronchi, and bronchioles (except the 23.16 Breathing in and then exhaling as much air as possible demon-
respiratory bronchioles). strates vital capacity.
23.2 The path of air is external nares : vestibule : nasal cavity : 23.17 A difference in PO2 promotes oxygen diffusion into pulmonary
internal nares. capillaries from alveoli and into tissue cells from systemic
23.3 The root of the nose attaches it to the frontal bone. capillaries.
23.4 During swallowing, the epiglottis closes over the rima glottidis, 23.18 The most important factor that determines how much O2 binds
the entrance to the trachea, to prevent aspiration of food and to hemoglobin is the PO2.
liquids into the lungs. 23.19 Both during exercise and at rest, hemoglobin in your pulmonary
23.5 The main function of the vocal folds is voice production. veins would be fully saturated with O2, a point that is at the up-
23.6 Because the tissues between the esophagus and trachea are soft, per right of the curve.
the esophagus can bulge and press against the trachea during 23.20 Because lactic acid (lactate) and CO2 are produced by active skele-
swallowing. tal muscles, blood pH decreases slightly and PCO2 increases when
23.7 The left lung has two lobes and two secondary bronchi; the right you are actively exercising. The result is lowered affinity of hemo-
lung has three of each. globin for O2, so more O2 is available to the working muscles.
23.8 The pleural membrane is a serous membrane. 23.21 O2 is more available to your tissue cells when you have a
23.9 Because two-thirds of the heart lies to the left of the midline, the fever because the affinity of hemoglobin for O2 decreases with
left lung contains a cardiac notch to accommodate the presence increasing temperature.
of the heart. The right lung is shorter than the left because the 23.22 At a PO2 of 40 mmHg, fetal Hb is 80% saturated with O2 and
diaphragm is higher on the right side to accommodate the liver. maternal Hb is about 75% saturated.
23.10 The wall of an alveolus is made up of type I alveolar cells, type 23.23 Blood in a systemic vein would have a higher concentration
II alveolar cells, and associated alveolar macrophages. of HCO3.
23.11 The respiratory membrane averages 0.5 m in thickness. 23.24 The medullary inspiratory area contains autorhythmic neurons
23.12 The pressure would increase fourfold, to 4 atm. that are active and then inactive in a repeating cycle.
23.13 If you are at rest while reading, your diaphragm is responsible 23.25 The phrenic nerves innervate the diaphragm.
for about 75% of each inhalation. 23.26 Peripheral chemoreceptors are responsive to changes in blood
23.14 At the start of inhalation, intrapleural pressure is about levels of oxygen, carbon dioxide, and H.
756 mmHg. With contraction of the diaphragm, it decreases to 23.27 Normal arterial PCO2 is 40 mmHg.
about 754 mmHg as the volume of the space between the two 23.28 The respiratory system begins to develop about 4 weeks after
pleural layers expands. With relaxation of the diaphragm, it fertilization.
increases back to 756 mmHg.