23.

4 Pulmonary Ventilation
Pulmonary Ventilation
● Pulmonary ventilation is the flow of air into and out of the lungs.
● Air flows between the atmosphere and the pulmonary alveoli of the lungs.
● Pressure differences created by respiratory muscle contraction and
relaxation drive the flow of air.
● Alveolar surface tension, lung compliance, and airway resistance also
affect airflow and effort required for breathing.
Pressure Changes During Pulmonary Ventilation
● Air moves into the lungs when the air pressure inside the lungs is lower
than atmospheric pressure.
● Air moves out of the lungs when the air pressure inside the lungs is higher
than atmospheric pressure.
*Inhalation
● Inhalation (inspiration) is the process of breathing in.
● Before each inhalation, the air pressure inside the lungs is equal to
atmospheric pressure.
● For air to flow into the lungs, the pressure inside the pulmonary alveoli
must become lower than atmospheric pressure.
● Increasing the size of the lungs achieves this condition.
● Boyle’s law states that the pressure of a gas in a closed container is
inversely proportional to the volume of the container.
Muscles of Inhalation
● The main muscle of inhalation is the diaphragm.
● Contraction of the diaphragm causes it to flatten, increasing the vertical
diameter of the thoracic cavity.
● The diaphragm is innervated by fibers of the phrenic nerves, which
emerge from the spinal cord at cervical levels 3, 4, and 5.
● The external intercostals are also important muscles of inhalation.
● Contraction of the external intercostals elevates the ribs, increasing the
anteroposterior and lateral diameters of the chest cavity.
Intrapleural Pressure
● Intrapleural pressure is the pressure within the pleural cavity.
● The pleural cavity is the space between the parietal pleura and visceral
pleura.
● It functions as a vacuum due to its negative pressure (lower than
atmospheric pressure).
● The negative pressure attaches the visceral pleura to the chest wall.
Inhalation technique
● During inhalation, intrapleural pressure decreases due to the expansion of
the thoracic cavity.
● The volume of the pleural cavity increases.
● The parietal pleura is pulled outward, along with the lungs.
● The volume of the lungs increases, causing the alveolar pressure to drop.
Pressure Difference
● A pressure difference is established between the atmosphere and the
pulmonary alveoli.
● Air flows from a region of higher pressure (atmosphere) to a region of
lower pressure (alveoli).
● This causes inhalation to occur.
● Air continues to flow into the lungs as long as a pressure difference exists.
Accessory Muscles of Inhalation
● During deep, forceful inhalations, accessory muscles of inspiration
participate.
● Accessory muscles include sternocleidomastoid, scalene, and pectoralis
minor muscles.
● They elevate the sternum and ribs, further increasing the size of the
thoracic cavity.
● These muscles contract vigorously during exercise or forced breathing.
Active Inhalation
● Both normal quiet inhalation and inhalation during exercise involve
muscular contraction.
● The process of inhalation is considered active.
● Muscular contraction expands the thoracic cavity and increases lung
volume.
● This allows for efficient gas exchange and oxygenation of the blood.
**Exhalation
● Exhalation, also known as expiration, is the process of breathing out.
● It is a passive process during quiet breathing and an active process during
forceful breathing.
● Exhalation is aided by the elastic recoil of the chest wall and lungs, as well
as the forces of surface tension and elastic fibers.
Passive Exhalation
● Passive exhalation occurs during quiet breathing.
● No muscular contractions are involved.
● Exhalation results from the elastic recoil of the chest wall and lungs.
● Two forces contribute to elastic recoil: recoil of elastic fibers and inward
pull of surface tension.
Active Exhalation
● Active exhalation occurs during forceful breathing, such as playing a wind
instrument or exercise.
● Muscles of exhalation, including the abdominal and internal intercostals,
contract.
● This increases pressure in the abdominal region and thorax.
● Contraction of the abdominal muscles moves the diaphragm upward.
● Contraction of the internal intercostals pulls the ribs downward.
Pressure Changes
● During exhalation, the diaphragm and external intercostals relax.
● This decreases the thoracic cavity dimensions and lung volume.
● Alveolar pressure increases to about 762 mmHg.
● Air then flows from the pulmonary alveoli to the atmosphere.
Factors Affecting Pulmonary Ventilation
1. Air Pressure Differences
● Air pressure differences drive airflow during inhalation and exhalation.
2. Surface Tension of Alveolar Fluid
● Surface tension of the alveolar fluid affects pulmonary ventilation.
● A thin layer of alveolar fluid coats the luminal surface of the pulmonary
alveoli.
● Surface tension arises from the attraction of water molecules to each
other.
● Inwardly directed force due to surface tension causes the pulmonary
alveoli to assume the smallest possible diameter.
● During inhalation, surface tension must be overcome to expand the lungs.
● Surface tension also contributes to lung elastic recoil during exhalation.
3. Surfactant
● Surfactant is a mixture of phospholipids and lipoproteins present in
alveolar fluid.
● Surfactant reduces the surface tension of the alveolar fluid.
● Deficiency of surfactant in premature infants leads to respiratory distress
syndrome.
● Respiratory distress syndrome causes increased surface tension and
collapsing of pulmonary alveoli during exhalation.
4. Compliance
● Compliance refers to how much effort is required to stretch the lungs and
chest wall.
● High compliance means that the lungs and chest wall expand easily.
● Low compliance means that they resist expansion.
● Non-compliance: inability of the person to make good chest expansion
Factors Affecting Compliance
● Compliance in the lungs is related to two principal factors: 1. elasticity and
2. surface tension.
● Elastic fibers in lung tissue contribute to high compliance.
● Surfactant in alveolar fluid reduces surface tension and increases
compliance.
Conditions Affecting Compliance
● Decreased compliance can result from scarred lung tissue (e.g.,
tuberculosis), fluidfilled lung tissue (e.g., pulmonary edema), surfactant
deficiency, or lung expansion impairment (e.g., paralysis of intercostal
muscles).
● Increased compliance is seen in emphysema due to destruction of elastic
fibers in alveolar walls.
5. Airway Resistance
● Airway resistance affects the rate of airflow through the airways.
● Airflow is determined by the pressure difference and the resistance.
● Resistance is offered by the walls of the airways, especially the
bronchioles.
Changes in Airway Diameter
● During inhalation, the bronchioles enlarge as the lungs expand.
● Larger diameter airways have decreased resistance.
● During exhalation, the bronchioles narrow, increasing resistance.
Smooth Muscle Regulation
● Airway diameter is also regulated by the smooth muscle in the airway
walls.
● Sympathetic signals cause relaxation of bronchiolar smooth muscle,
resulting in decreased resistance.
● Parasympathetic signals cause contraction of bronchiolar smooth muscle,
resulting in increased resistance.
Conditions Affecting Airway Resistance
● Any condition that narrows or obstructs the airways increases
resistance.
● Asthma and chronic obstructive pulmonary disease (COPD) are
characterized by increased airway resistance due to airway obstruction or
collapse.
Breathing Patterns and Modified Breathing
Movements
1. Eupnea
● The term for the normal pattern of quiet breathing
● Consists of shallow, deep, or combined shallow and deep breathing
2. Costal Breathing
● Pattern of shallow (chest) breathing
● Upward and outward movement of the chest
● Caused by contraction of the external intercostal muscles
3. Diaphragmatic Breathing
● Pattern of deep (abdominal) breathing
● Outward movement of the abdomen
● Caused by contraction and descent of the diaphragm
Functions of Breathing
● Expressing emotions such as laughing, sighing, and sobbing
● Expelling foreign matter through sneezing and coughing
● Modified and controlled during talking and singing
Modified Breathing Movements
● Expressing emotion or clearing airways
● Reflexes that can also be initiated voluntarily
23.5 Lung Volumes and Capacities
Introduction
● During inhalation and exhalation, varying amounts of air move into and out
of the lungs
● These amounts depend on factors related to individuals' characteristics
and pulmonary disorders
Types of Air
● Two types: lung volumes and lung capacities
● Lung volumes can be measured directly using a spirometer
● Lung capacities are combinations of different lung volumes
Spirometer
● Apparatus used to measure lung volumes and capacities
● Also known as respirometer
● Records are called spirograms
Lung Volumes
● Tidal volume (VT): volume of one breath, averages around 500 mL
● Only about 70% of VT reaches the respiratory zone for gas exchange
● Remaining 30% stays in conducting airways as anatomic dead space
Inspiratory and Expiratory Reserve Volumes
● Inspiratory reserve volume (IRV): additional air that can be inhaled after a
normal breath
● Expiratory reserve volume (ERV): additional air that can be exhaled after a
normal breath
● IRV is about 3100 mL in males, 1900 mL in females
● ERV is about 1200 mL in males, 700 mL in females
Forced Expiratory Volume in 1 Second (FEV1)
● FEV1 measures the volume of air exhaled in 1 second with maximal effort
● COPD greatly reduces FEV1 due to increased airway resistance
Residual Volume
● Volume of air that remains in the lungs after maximal exhalation
● Cannot be measured by spirometry
● Around 1200 mL in males, 1100 mL in females
Minimal Volume
● Air remaining in the lungs after intrapleural pressure equalizes with
atmospheric pressure
● Cannot be measured but used as a tool to determine stillbirth
● Minimal volume can be detected if lung floats in water
Lung Capacities
● Combinations of specific lung volumes
● Inspiratory capacity (IC): sum of tidal volume and inspiratory reserve
volume
● Functional residual capacity (FRC): sum of residual volume and expiratory
reserve volume
● Vital capacity (VC): sum of inspiratory reserve volume, tidal volume, and
expiratory reserve volume
● Total lung capacity (TLC): sum of vital capacity and residual volume
Minute Ventilation and Alveolar Ventilation
● Minute ventilation (V): total volume of air inspired and expired each minute
● V = tidal volume × respiratory rate
● Normal minute ventilation is about 6000 mL/min
● Alveolar ventilation (V.A): volume of air per minute that reaches the
respiratory zone
● Typically around 4200 mL/min
23.6 Exchange of Oxygen and Carbon Dioxide
Introduction
● The exchange of oxygen and carbon dioxide between alveolar air and
pulmonary blood occurs via passive diffusion
● This diffusion is governed by two gas laws:
1. Dalton’s law
2. Henry’s law
1. Dalton’s Law
● According to Dalton’s law, each gas in a mixture of gases exerts its own
pressure as if no other gases were present
● The pressure of a specific gas in a mixture is called its partial pressure
(Px)
● The total pressure of the mixture is calculated by adding all of the partial
pressures
Atmospheric Air
● Atmospheric air is a mixture of gases
● The main gases in atmospheric air are nitrogen (N2), oxygen (O2), argon
(Ar), carbon dioxide (CO2)
● It also contains variable amounts of water vapor (H2O) and other gases
present in small quantities
2. Henry’s Law
● Henry’s law explains how the solubility of a gas relates to its diffusion
● The more soluble a gas is, the higher its diffusion rate
● Solubility of a gas is influenced by factors such as temperature and
pressure
● Atmospheric pressure is the sum of the pressures of all of these gases:
Calculation Method
● Multiply the percentage of gas in the mixture by the total pressure
● Example: Nitrogen 78.6% * Total Pressure
Components in Inhaled Air
● Nitrogen: 78.6%
● Oxygen: 20.9%
● Argon: 0.093%
● Carbon Dioxide: 0.04%
● Other Gases: 0.06%
● Variable Water Vapor (04%)
Partial Pressures in Inhaled Air
Variation of Water Vapor
● Water content varies from 0% in deserts to 4% over the ocean
● Approximately 0.3% on a cool, dry day
Gas Exchange in the Respiratory System
● Gas exchange occurs between the atmosphere, lungs, blood, and body
cells
● Each gas diffuses across a permeable membrane from an area of higher
partial pressure to an area of lower partial pressure
● The greater the difference in partial pressure, the faster the rate of
diffusion
Henry's Law
● The quantity of a gas dissolved in a liquid is proportional to its partial
pressure and solubility
● Higher partial pressure and solubility increase the gas's ability to stay in
solution
● CO2 has much higher solubility in blood plasma compared to O2
Nitrogen Solubility in Body Fluids
● As total air pressure increases, the partial pressures of all gases increase
● Scuba divers breathing compressed air with higher nitrogen partial
pressure may experience nitrogen narcosis
● Too rapid ascent can cause decompression sickness due to gas bubbles
forming in tissues
Symptoms of Decompression Sickness
● Joint pain, dizziness, shortness of breath, extreme fatigue, and paralysis
● Bubbles in nervous tissue can cause mild or severe effects
● Proper ascent and elimination of dissolved nitrogen are crucial
External Respiration
External Respiration: Overview
● External respiration is the diffusion of O2 from the lungs to the blood and
the diffusion of CO2 in the opposite direction.
● It occurs in the pulmonary alveoli of the lungs and the pulmonary
capillaries.
● The process involves converting deoxygenated blood into oxygenated
blood.
● Oxygenated blood is then transported back to the heart.
Oxygen Diffusion
● O2 diffuses from pulmonary alveolar air to the blood in pulmonary
capillaries.
● The partial pressure of O2 is higher in alveolar air (105 mmHg) and lower
in the blood.
● Diffusion continues until the partial pressures equalize (105 mmHg).
● Blood in the pulmonary veins has a slightly lower PO2 (100 mmHg) due to
mixing with nonexchanged blood.
Carbon Dioxide Diffusion
● CO2 diffuses from deoxygenated blood to pulmonary alveoli.
● The PCO2 is higher in deoxygenated blood (45 mmHg) and lower in
alveolar air (40 mmHg).
● Diffusion continues until the PCO2 of the blood decreases to 40 mmHg.
● Exhalation helps maintain alveolar PCO2 at 40 mmHg.
Impact of Exercise
● During vigorous exercise, cardiac output and blood flow increase.
● Transit time in pulmonary capillaries decreases.
● Still, the PO2 of blood in the pulmonary veins reaches 100 mmHg.
● Diseases affecting gas diffusion can disrupt equilibrium and lead to lower
PO2 and higher PCO2 in systemic arterial blood.
Internal Respiration
Introduction
● The left ventricle pumps oxygenated blood into the aorta and through the
systemic arteries to systemic capillaries.
● Exchange of O2 and CO2 between systemic capillaries and tissue cells is
called internal respiration or systemic gas exchange.
● Internal respiration occurs in tissues throughout the body, unlike external
respiration, which occurs only in the lungs.
Oxygen Diffusion
● PO2 of blood pumped into systemic capillaries is higher than the PO2 in
tissue cells.
● Oxygen diffuses out of the capillaries into tissue cells.
● Blood PO2 drops to 40 mmHg by the time the blood exits systemic
capillaries.
Carbon Dioxide Diffusion
● CO2 diffuses from tissue cells through interstitial fluid into systemic
capillaries.
● PCO2 of cells is higher than that of systemic capillary blood.
● CO2 diffuses into systemic capillaries until the PCO2 in the blood
increases to 45 mmHg.
Oxygenation and Deoxygenation
● Deoxygenated blood returns to the heart and is pumped to the lungs.
● External respiration occurs in the lungs.
● During exercise, more O2 diffuses from the blood into metabolically active
cells.
● Oxygen content of deoxygenated blood drops below 75%.
Factors Affecting Gas Exchange
1. Partial pressure difference of the gases affects gas diffusion.
2. Surface area available for gas exchange affects the rate of external
respiration.
3. Diffusion distance affects the speed of gas exchange.
4. Molecular weight and solubility of the gases play a role in diffusion.
23.7 Transport of Oxygen and Carbon Dioxide
Introduction
● Gas transport and exchange are vital processes in the body.
● They involve the movement of oxygen (O2) and carbon dioxide (CO2)
between the lungs and body tissues.
● Certain chemical reactions occur in the blood to facilitate these
processes.
Oxygen Transport
● Oxygen does not dissolve easily in water.
● Only about 1.5% of inhaled O2 is dissolved in blood plasma (mostly water).
● Approximately 98.5% of blood O2 is bound to hemoglobin in red blood
cells.
● Each 100 mL of oxygenated blood contains the equivalent of 20 mL of
gaseous O2.
● Based on these percentages, about 0.3 mL of O2 is dissolved in the
plasma and 19.7 mL is bound to hemoglobin.
Introduction
● The most important factor that determines how much O2 binds to
hemoglobin is the PO2
● Higher PO2 leads to more O2 binding with Hb
● Hemoglobin exists in two states: fully saturated (HbO2) and partially
saturated (mixture of Hb and Hb-O2)
● Percent saturation expresses the average saturation of hemoglobin with
oxygen
● Hemoglobin can bind a maximum of four O2 molecules
Oxygen-Hemoglobin Dissociation Curve
● Illustrates the relationship between percent saturation of hemoglobin and
PO2
● At high PO2, hemoglobin is almost 100% saturated
● At low PO2, hemoglobin is only partially saturated
● Greater PO2 leads to more O2 binding to hemoglobin
● In pulmonary capillaries (high PO2), a lot of O2 binds to hemoglobin
● In tissue capillaries (lower PO2), hemoglobin does not hold as much O2
and dissolved O2 is unloaded via diffusion into tissue cells
● Hemoglobin is still 75% saturated with O2 at a PO2 of 40 mmHg (average
PO2 of tissue cells at rest)
Hemoglobin Saturation Levels
● Between 60 and 100 mmHg, hemoglobin is 90% or more saturated with O2
● Blood picks up a nearly full load of O2 from the lungs even at low PO2
(e.g., 60 mmHg)
● People can perform well at high altitudes or with certain cardiac and
pulmonary diseases despite low PO2
● Hemoglobin saturation drops to 35% at a PO2 of 20 mmHg
● Large amounts of O2 are released from hemoglobin in response to small
decreases in PO2
● In active tissues, such as contracting muscles, PO2 may drop below 40
mmHg, leading to significant O2 release from hemoglobin
Acidity (pH)
● As acidity increases (pH decreases), the affinity of hemoglobin for O2
decreases
● Increasing acidity enhances the unloading of oxygen from hemoglobin
● The main acids produced by metabolically active tissues are lactic acid
and carbonic acid
● When pH decreases, the entire oxygen–hemoglobin dissociation curve
shifts to the right
● The Bohr effect: An increase in H+ in blood causes O2 to unload from
hemoglobin
Partial pressure of carbon dioxide
● CO2 can bind to hemoglobin, similar to H+
● As PCO2 rises, hemoglobin releases O2 more readily
● PCO2 and pH are related factors
● Low blood pH (acidity) results from high PCO2

As CO2 enters the blood, much of it is temporarily converted to carbonic acid

(H2CO3), a reaction catalyzed by an enzyme in red blood cells called carbonic
anhydrase (CA):
Carbonic Acid and pH
● Carbonic acid in red blood cells dissociates into hydrogen ions and
bicarbonate ions
● Increased H+ concentration leads to a decrease in pH
● Higher PCO2 levels create a more acidic environment
● Acidic environment helps release O2 from hemoglobin
Temperature
● Increased temperature leads to more O2 released from hemoglobin
● Metabolic reactions generate heat
● Heat released by contracting muscle fibers raises body temperature
● Higher demand for O2 and liberation of acids and heat promote O2
release
● Fever has a similar effect
● Remember: Hypothermia slows cellular metabolism, reduces O2 need,
and keeps more O2 bound to hemoglobin (shift to the left)
Introduction
● Under normal resting conditions, deoxygenated blood contains 53 mL of
gaseous CO2 per 100 mL of blood.
● Carbon dioxide is transported in the blood in three main forms:
1. Dissolved co2
2. Carbamino compounds
3. Bicarbonate ions
1. Dissolved CO2
● Approximately 7% of CO2 is dissolved in the blood plasma.
● In the lungs, dissolved CO2 diffuses into alveolar air and is exhaled.
2. Carbamino Compounds
● About 23% of CO2 combines with amino groups of amino acids and
proteins in blood.
● Carbamino compounds are formed.
● In blood, most of the CO2 bound to hemoglobin (inside red blood cells).
● CO2 binds to the terminal amino acids in the alpha and beta globin chains
of hemoglobin.

Hemoglobin that has bound CO2 is termed carbaminohemoglobin (Hb–CO2):

The formation of carbaminohemoglobin is greatly influenced by PCO2. For
example, in tissue capillaries PCO2 is relatively high, which promotes formation
of carbaminohemoglobin.

But in pulmonary capillaries, PCO2 is relatively low, and the CO2 readily splits
apart from globin and enters the alveoli by diffusion
3. Bicarbonate ions
• The greatest percentage of CO2—about 70%—is transported in blood
plasma as bicarbonate ions (HCO3−)
• As CO2 diffuses into systemic capillaries and enters red blood cells, it reacts
with water in the presence of the enzyme carbonic anhydrase (CA) to form
carbonic acid, which dissociates into H+ and HCO3
CO2 Transport in RBCs
• Thus, as blood picks up CO2, HCO3- accumulates inside RBCs.
• Some HCO3− moves out into the blood plasma, down its concentration gradient.
• In exchange, chloride ions (Cl−) move from blood plasma into the RBCs.
• This exchange of negative ions, which maintains the electrical balance between
blood plasma and RBC cytosol, is known as the chloride shift (see Figure 23.23b).
• The net effect of these reactions is that CO2 is removed from tissue cells and
transported in blood plasma as HCO3−.
• As blood passes through pulmonary capillaries in the lungs, all these reactions
reverse, and CO2 is exhaled.
Reversal of Reactions in Pulmonary Capillaries
● As blood passes through pulmonary capillaries in the lungs, the reactions
reverse
● CO2 is exhaled
23.8 Control of Breathing
Oxygen Consumption during Exercise
● At rest, about 200 mL of O2 is used each minute by body cells.
● During strenuous exercise, O2 use typically increases 15 to 20-fold in
normal healthy adults, and as much as 30-fold in elite endurance-trained
athletes.
Respiratory Center
● Breathing effort is matched to metabolic demand through several
mechanisms.
● Nerve impulses transmitted from the brain control the contraction and
relaxation of the breathing muscles in the thorax.
● The respiratory center, located in the brain stem, consists of the medullary
respiratory center and the pontine respiratory group.
Chemoreceptor Regulation of Breathing
● Chemoreceptors monitor levels of CO2, H+, and O2 in the blood
● Central chemoreceptors respond to changes in H+ concentration or PCO2
in cerebrospinal fluid
● Peripheral chemoreceptors respond to changes in PO2, H+, and PCO2 in
the blood
● Chemoreceptors participate in a negative feedback system to regulate
CO2, O2, and H+ levels
Proprioceptor Stimulation of Breathing
● Proprioceptors monitor movement of joints and muscles
● Proprioceptor input stimulates the medullary respiratory center
● Upper motor neurons from the primary motor cortex also stimulate the
medullary respiratory center
The Inflation Reflex
● Stretchsensitive receptors called baroreceptors are located in the walls of
bronchi and bronchioles
● Overinflation of the lungs activates the inflation reflex
● The inflation reflex inhibits the medullary respiratory center, leading to
exhalation
● As air leaves the lungs, the stretch receptors are no longer stimulated and
a new inhalation begins
Other Influences on Breathing
● Limbic system stimulation can increase the rate and depth of breathing
● Temperature changes affect breathing rate
● Pain can briefly affect breathing rate
● Stretching the anal sphincter muscle increases breathing rate
● Irritation of airways can cause cessation of breathing followed by
coughing or sneezing
● Blood pressure changes have a small effect on breathing