Respiratory Physiology

Introduction
Respiration is defined as the movement of oxygen from the outside environment to
the cells and removal of carbon dioxide in the opposite direction or the process by which
oxygen is taken in and carbon dioxide is given out or the physical and chemical processes
(such as breathing and diffusion) by which an organism supplies the oxygen to its cells and
tissues needed for metabolism and relieves them of the carbon dioxide formed in energy-
producing reactions.
As foetal lungs are non-functional, the first breath takes place only after birth.
Therefore, during intrauterine life the exchange of gases between foetal blood and mother’s
blood occurs through placenta. After the first breath, the respiratory process continues
throughout the life and the permanent stoppage of respiration occurs only at the time of death.
Types of respiration:
Respiration is classified into two types:
1. External: It is the type of respiration in which the exchange of respiratory gasses
(O2 & CO2) takes place between lungs and blood.
2. Internal: It is the type of respiration in which the exchange of gasses respiratory
(O2 & CO2) takes place between blood and body cells/ tissues.
Phases of respiration:
Respiration occurs in two phases:
1. Inspiration during which air enters the lungs from atmosphere.
2. Expiration during which air leaves the lungs.
During normal breathing, inspiration is an active process and expiration is a
passive process.
Functional Morphology of the Respiratory System
The respiratory system consists of the lungs and pleura and the air passages leading to
the lungs, including the nostrils, nasal cavities, pharynx, larynx, trachea, bronchi and
bronchioles.
Lungs and the pleura
 The lungs are the principal structures of the respiratory system. They are paired
structures and occupy all space in the thorax. When the thorax expands in volume, the
lungs also expand; this provides airflow into the lungs. They have an almost

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 friction‐free movement within the thorax because of the pleura (a smooth serous
membrane).
 The pleura consist of a single layer of cells fused to the surface of a connective tissue
layer. It envelops both lungs (visceral pleura).
 Pleura for the right and left lung meet near the midline and here it reflects upward
(dorsally), turns back on the inner thoracic wall and provides for its lining (costal/
parietal pleura).
 The space between the visceral and parietal layers of pleura is called intra-pleural
space. The intra-pleural space contains a thin film of serous fluid called intra-pleural
fluid, which is secreted by the visceral layer of the pleura.
Function:
1. It functions as the lubricant, to prevent friction between two layers of pleura
2. It is involved in creating the negative pressure called intra-pleural pressure
within intra-pleural space.
 The space between the respective visceral pleura layers as they ascend to the dorsal
wall is known as the mediastinal space. Within the mediastinal space are the venae
cavae, thoracic lymph duct, oesophagus, aorta and trachea. The mediastinal space is
closely associated with the intrapleural space (space between visceral and costal
pleura); thus, pressure changes in the intrapleural space are accompanied by similar
changes in the mediastinal space. Also, pressure changes within the mediastinal space
are accompanied by changes within the mediastinal structures, provided that their
walls are responsive to relatively low‐pressure distensibility.

Pleura and the pleural cavity

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Airway to the lungs
 Nostrils and Nasal Cavities
The nostrils (nares) are the paired external openings to the air passages. The
nostrils are the most pliable and dilatable in the horse and the most rigid in the pig.
The nostril dilatation is advantageous when more air is required, as in running and in
situations in which breathing is not done through the mouth. The horse is a runner and
open‐mouth breathing is not characteristic, therefore dilatable nostrils are
advantageous.

a b

e
c d

Nostrils of some domestic species

(a: horse, b: cow, c: sheep, d: pig, e:dog )

The nostrils provide the external openings for the paired nasal cavities. The nasal
cavities are separated from each other by the nasal septum and from the mouth by the hard
and soft palates. In addition, each nasal cavity contains mucosa‐covered turbinate bones
(conchae). The mucosa of the turbinate bones is well vascularized and serves to warm and
humidify inhaled air. Another function for the conchae/ turbinate bones involves cooling
blood through a counter-current heat exchange mechanism. Arteries that supply blood to the
brain divide into smaller arteries at the base of the brain. These are bathed in a pool of venous
blood coming from the walls of the nasal cavities where it has been cooled. This keeps brain
temperature 2-3 degrees cooler.
Pharynx
It is situated between mouth and nasal cavities from front and trachea and oesophagus
from behind and is a common passageway for air and food. The openings to the pharynx

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include two posterior nares, two eustachian tubes, a mouth (oral cavity), a glottis and an
oesophagus. The opening from the pharynx leading to the continuation of the respiratory
passageway is the glottis. Immediately caudal to the glottis is the larynx, organ of phonation/
sound production. In birds the larynx is called syrinx, located where the trachea bifurcates to
form bronchi.
Trachea
This is the primary passageway for air into the lungs. Cranially it is connected with
the larynx and caudally it bifurcates to form the left and right bronchi. The tracheal wall
contains cartilaginous rings to prevent collapse of the tracheal airway. Each tracheal ring is
incomplete (not joined dorsally), which permits variation in diameter for increased ventilation
requirements. The right and left bronchi and their subdivisions continue all the way to the
alveoli, the final and smallest subdivisions of the air passages.

Trachea

Right and left bronchi

 Subdivision of the trachea to the alveoli are:

 bronchi (Right and Left)
 bronchioles
 terminal bronchioles
 respiratory bronchioles
 alveolar duct
 alveolar sac

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 Pathway of the air via nasal cavities, pharynx and trachea

Path-way of the respiratory system from nose to alveoli

Respiratory unit
The portion of the lungs involved in gas exchange i.e. lung parenchyma, is formed by
respiratory unit that forms the terminal portion of respiratory tract. Respiratory unit is defined
as the structural and functional unit of lung. Exchange of gases occurs only in this part of the
respiratory tract.
The respiratory unit starts from the respiratory bronchioles and each respiratory
bronchiole divides into alveolar ducts. These alveolar ducts enter into an enlarged structure
called the alveolar sac. Space inside the alveolar sac is called antrum. Alveolar sac consists
of a cluster of alveoli. Few alveoli are present in the wall of alveolar duct also.

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 Respiratory unit
Pulmonary alveoli
They are the principal sites of gaseous diffusion between air and blood. The
separation of air and blood and thus the diffusion distance is minimal at the alveolar level.
Alveolar epithelium consists of two types cells called alveolar cells or pneumocytes, namely:
type I alveolar cells and type II alveolar cells.
Type I alveolar cells
Type I alveolar cells are the squamous epithelial cells and comprise about 95% of the
total number of alveolar cells. These cells form the site of gaseous exchange between the
alveolus and blood.
Type II alveolar cells
Type II alveolar cells are cuboidal in nature and comprise and comprise only about
5% of alveolar cells. These cells are also called granular pneumocytes. The function of these
cells is to secrete the alveolar fluid and surfactant.

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 Alveolar epithelium showing type I and type II pneumocytes or alveolar cells
Functions of the respiratory system:
 Respiratory functions:
 Pulmonary ventilation: means the inflow and outflow of air between the atmosphere
and the lung alveoli.
 Diffusion of oxygen and carbon dioxide between the alveoli and the blood.
 Transport of oxygen and carbon dioxide in the blood and body fluids to and from the
body’s tissue cells.
 Regulation of ventilation and other aspects of respiration.
 Non Respiratory functions:
 Olfaction: The olfactory receptors present on the mucous membrane of the nostrils
are responsible for olfaction.
 Vocalization: Larynx along with other structures forms the speech apparatus.
However, larynx alone plays major role in the process of vocalization. Therefore, it is
called sound box/ organ of phonation.
 In birds, the larynx is called syrinx.
 Prevention from dust particles: The filtration action of hairs present in the mucous
membrane of the nasal cavity prevents the entry of dust particles in to the lungs.
However, small dust particles that escape the nasal hairs and mucous membrane are
removed by phagocytosis of macrophages in the lung alveoli.
 Defence mechanism: The presence of various types of cells (leucocytes,
macrophages, mast cells, NK cells and dendritic cells) in the mucous membrane lining
the lung alveoli play an important role in immunological defence of body.

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  Maintenance of water balance
 Regulation of body temperature
 Regulation of acid-base balance
 Anticoagulant function: Mast cells, present in the lungs secrete an anticoagulant
called heparin. Heparin is a natural anticoagulant that prevents the intravascular
clotting.
 Secretion of Angiotensin converting enzyme (ACE)
 Synthesis of hormonal substances: Hormones like prostaglandins, acetylcholine and
serotonin are secreted from the alveolar/ lung tissues.
Respiratory cycles
A respiratory cycle consists of an inspiratory phase followed by an expiratory phase.
Inspiration involves an enlargement of the thorax and lungs, with an accompanying
inflow of air. The thorax enlarges by contraction of the diaphragm (the musculo-tendinous
separation between the thorax and abdomen) and by contraction of external intercostal
muscles (muscles located between the ribs). Under normal breathing conditions, inspiration
requires greater effort than expiration, and sometimes expiration might seem to be passive.
Expiration can become quite an active process, particularly during times of
accelerated breathing. The internal intercostal muscles contract to assist in expiration. Other
skeletal muscles can aid in either inspiration or expiration, such as the abdominal muscles.
The respiratory pattern or waveform, when recorded, varies little among
mammalian species. The inspiratory and expiratory phases of the cycles are generally smooth
and symmetrical. An exception to this general statement is the horse, in which there are two
phases during inspiration and two phases during expiration.
Complementary breathing cycles are characterized by a deep rapid inspiration
followed by expiration of longer duration. They occur normally in many species but
apparently not in the horse. This type of cycle has frequently been called a sigh. As it
naturally occurs, it is probably a compensatory mechanism for poor ventilation.
Types of breathing
There are two types of breathing: abdominal and costal.
Abdominal breathing is characterized by visible movements of the abdomen, in
which the abdomen protrudes during inspiration and recoils during expiration. Normally the
abdominal type of breathing predominates.
The other type is called costal breathing; it is characterized by pronounced rib
movements. During painful conditions of the abdomen such as peritonitis, in which

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movement of the viscera would aggravate the pain, costal breathing can predominate.
Similarly, during painful conditions of the thorax such as pleuritis, abdominal breathing
might be more apparent.
States of breathing
In addition to the different types of breathing, there are variations in breathing related
to the frequency of breathing cycles, depth of inspiration or both.
 Eupnea is the term used to describe normal quiet breathing, with no deviation in
frequency or depth.
 Dyspnea is difficult breathing, in which visible effort is required to breathe.
 Hyperpnea refers to breathing characterized by increased depth, frequency, or both,
and is noticeable after physical exertion.
 Polypnea is rapid shallow breathing, somewhat similar to panting. Polypnea is similar
to hyperpnea in regard to frequency, but is unlike hyperpnea in regard to depth.
 Apnea refers to a cessation of breathing. However, as used clinically, it generally
refers to a transient state of cessation of breathing.
 Tachypnea is excessive rapidity of breathing, and bradypnea is abnormal slowness
of breathing.
Respiratory frequency
Respiratory frequency refers to the number of respiratory cycles each minute. It is an
excellent indicator of health status because respiratory frequency increases during disease and
must be interpreted properly because it is subject to numerous variations.
In addition to variations observed among species, respiratory frequency can be
affected by other factors, such as body size, age, exercise, excitement, environmental
temperature, pregnancy, degree of filling of the digestive tract, and state of health. Pregnancy
and digestive tract filling increase frequency because they limit the excursion of the
diaphragm during inspiration. When expansion of the lungs is restricted, adequate ventilation
is maintained by increased frequency. For example, when cattle lie down, the large rumen
pushes against the diaphragm and restricts its movement, and respiratory frequency is seen to
increase.
Table: Respiratory frequency among different domestic species
S. No. Animal Species Resp. rate/ Min.
1 Cattle 25-30
2 Horse 8-16

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 3 Sheep 10-20
4 Goat 10-20
5 Pig 10-20
6 Dog 16-30
7 Cat 20-30
8 chicken 15-30
Lung sounds
It is obvious that considerable branching of the pulmonary airways occurs. Although
the branches may have smaller diameters than the parent branch, the combined cross‐
sectional area of the branches shows an increase over that of the parent. Consequently, the
velocity of airflow diminishes progressively from the trachea toward the bronchioles.
Listening for lung sounds with the aid of a stethoscope is termed auscultation. A
good‐quality stethoscope should be used in quiet surroundings.
The term breath sound applies to any sound that accompanies air movement through
the trachea-bronchial tree. Breath sounds vary randomly in intensity depending on whether
the sounds are produced over the larger airways or over the remaining lung parenchyma.
Adventitious sounds are extrinsic to the normal sound production mechanism of the
respiratory tract and are abnormal sounds superimposed on the breath sounds. Adventitious
sounds are further classified as crackles and wheezes. Diseases resulting in oedema or
exudates within the airways can result in crackles. Wheezes suggest airway narrowing (e.g.
bronchoconstriction, bronchial wall thickening, and external airway compression).
Pulmonary volumes and capacities
Lung volumes are either associated with the amount of air within them at any one
time or with the amount associated with a breath.
 Tidal volume is the amount of air breathed in or out during a respiratory cycle. It can
increase or decrease from normal, depending on ventilation requirements. Tidal
volume is probably used more frequently than other terms.
 Inspiratory reserve volume is the amount of air that can still be inspired after
inhaling the tidal volume.
 Expiratory reserve volume is the amount of air that can still be expired after
exhaling the tidal volume.
 Residual volume is the amount of air remaining in the lungs after the most forceful
expiration.

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 The combination of two or more volumes is called capacities and it is useful to
combine two or more of these volumes.
 Total lung capacity is the sum of all volumes.
 Vital capacity is the sum of all volumes over and above the residual volume; it is the
maximum amount of air that can be breathed in after the most forceful expiration.
 Inspiratory capacity is the sum of the tidal and inspiratory reserve volumes.
 Functional residual capacity is the sum of the expiratory reserve volume and the
residual volume.
Mechanics of Respiration
Respiration occurs in two phases namely inspiration and expiration. During
inspiration, thoracic cage enlarges and lungs expand so that air enters the lungs easily. During
expiration, the thoracic cage and lungs decrease in size and attain the preinspiratory position
so that air leaves the lungs easily.
During normal quiet breathing, inspiration is the active process and expiration is the
passive process.
Muscles of Respiration
Respiratory muscles are of two types: Inspiratory muscles and expiratory muscles.
However, respiratory muscles are generally classified into two types:
1. Primary or major respiratory muscles, which are responsible for change in size of
thoracic cage during normal quiet breathing
2. Accessory respiratory muscles that help primary respiratory muscles during forced
respiration.
1. Inspiratory Muscles: Muscles involved in inspiratory movements are known as
inspiratory muscles.
 Primary inspiratory muscles: include the diaphragm, which is supplied by phrenic
nerve and external intercostal muscles, supplied by intercostal nerves.
 Accessory inspiratory muscles: include Sterno-cleidomastoid, scalene, anterior serrati,
elevators of scapulae and pectorals are the accessory inspiratory muscles.
2. Expiratory Muscles: Muscles involved in expiratory movements are known as exspiratory
muscles.
 Primary expiratory muscles: include the internal intercostal muscles, which are
innervated by intercostal nerves.

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  Accessory expiratory muscles: include accessory expiratory muscles are the
abdominal muscles.
Movements of Thoracic Cage
Inspiration causes enlargement of thoracic cage. Thoracic cage enlarges because of
increase in all diameters viz. antero-posterior, transverse and vertical diameters.
 Antero-posterior and transverse diameters of thoracic cage are increased by the
elevation of ribs.
 Vertical diameter is increased by the descent of diaphragm.
 In general, change in the size of thoracic cavity occurs because of the movements of
four units of structures: thoracic lid, upper costal series, lower costal series and
diaphragm.
Movements of Lungs
During inspiration, due to the enlargement of thoracic cage, the negative pressure is
increased in the thoracic cavity. It causes expansion of the lungs.
During expiration, the thoracic cavity decreases in size to the preinspiratory
position. Pressure in the thoracic cage also comes back to the preinspiratory level. It
compresses the lung tissues so that, the air is expelled out of lungs.
Respiratory Pressures
Two types of pressures are exerted in the thoracic cavity and lungs during process of
respiration:
1. Intra-pleural pressure
2. Intra-alveolar pressure
1. Intrapleural Pressure
Intrapleural pressure is the pressure existing in pleural cavity, that is, in between the
visceral and parietal layers of pleura. It is exerted by the suction of the fluid that lines the
pleural cavity. It is also called intrathoracic pressure because it is exerted in the whole of
thoracic cavity.
2. Intra-Alveolar Pressure
Intra-alveolar pressure is the pressure existing in the alveoli of the lungs. It is also
known as intrapulmonary pressure. Normally, intraalveolar pressure is equal to the
atmospheric pressure, which is 760 mm Hg. It becomes negative during inspiration and
positive during expiration.

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 Recoiling Tendency of Lungs
Lungs are under constant threat to collapse even in resting conditions because of
certain factors.
There is a constant tendency for the lungs to collapse This recoil tendency is due to (i)
stretching of elastin and collagen fibers by lung inflation and (ii) surface tension of fluid
lining the alveoli. The stretching of elastic fibers is easy to visualize as a force that
contributes to recoil.
In spite of elastic property of lungs and surface tension in the alveoli of lungs, the
collapsing tendency of lungs is prevented by two factors: (1) Intrapleural pressure: It is the
pressure in the pleural cavity, which is always negative. Because of negativity, it keeps the
lungs expanded and prevents the collapsing tendency of lungs produced by the elastic tissues
and (ii) Surfactant: It is a substance secreted in alveolar epithelium. It reduces surface
tension and prevents the collapsing tendency produced by surface tension.
Surfactant
Surfactant is an agent that is responsible for lowering the surface tension of a fluid.
Surfactants are surface‐active substances for which water molecules have a lesser attraction.
Because of this property the surfactant molecules accumulate at the surface and a reduction in
surface tension occurs.
Surfactant is synthesized by the type II alveolar epithelial cells (secretory cells).
Pulmonary surfactant is a lipoprotein complex containing about 30% protein and 70% lipid.
Most of the lipid fraction is composed of the phospholipid dipalmitoyl lecithin. Proteins of
the surfactant are called specific surfactant proteins. There are four main surfactant proteins
called SPA, SPB, SPC and SPD. SPA and SPD are hydrophilic, while SPB and SPC are
hydrophobic. Surfactant proteins are vital components of surfactant and the surfactant
becomes inactive in the absence of proteins. Ions present in the surfactant are mostly calcium
ions.
Hence alveolar epithelium is not simply a passive membrane for the exchange of
oxygen and carbon dioxide, but an active metabolic unit. It is estimated that the lung may
account for as much as 8–10% of the basal oxygen consumption of the body. Surfactant is
formed relatively late in human fetal life and in some animal species. The time of its
formation is not known for most of the domestic species. In the premature human infant,
surfactant deficiency at birth leads to a respiratory distress syndrome characterized by
dyspnea, cyanosis, and expiratory grunts.

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 Lung compliance
Lung compliance is a measure of the distensibility of the lungs and is determined by
measurement of the lung volume change for each unit of pressure change. The standard units
for pulmonary compliance are milliliters per centimeter of water. If a compliance value in a
particular animal has decreased over a period of time, the tissues of the lung must be more
rigid and less distensible or certain abnormalities may have further reduced the expansibility
of the thorax.
Changes in surfactant (amount or composition) affect compliance values, and lack of
surfactant is associated with decreased compliance.
Pulmonary Ventilation
Ventilation is generally regarded as the process by which gas in closed places is
renewed or exchanged. When applied to the lungs, it is a process of exchanging the gas in the
airways and alveoli with gas from the environment. The main function of ventilation is to
refill oxygen and to remove carbon dioxide.
Terminology
 Pulmonary ventilation is defined as the volume of air moving in and out of
respiratory tract in a given unit of time during quiet breathing. It is also called minute
ventilation or respiratory minute volume (RMV).
 Alveolar ventilation is the amount of air utilized for gaseous exchange every minute.
 Total ventilation is the volume of gas moved in or out of the airways and alveoli over
a certain period of time.
 Minute ventilation is the total volume of gas moved in or out of the airways and
alveoli in 1 min.
 Normo-ventilation refers to normal ventilation in which a PaCo2 of about 40 mmHg
is maintained.
 Hyperventilation refers to alveolar ventilation increased beyond the metabolic needs
and a PaCo2 below 40 mmHg. Hyperventilation causes respiratory alkalosis.
 Hypoventilation is alveolar ventilation decreased below metabolic needs and a PaCo2
above 40 mmHg. Hypoventilation causes respiratory acidosis.
 Dead space ventilation: The tidal volume is used to ventilate not only the alveoli, but
also the airways leading to the alveoli. Because there is little or no diffusion of
oxygen and carbon dioxide through the membranes of most of the airways, they
compose part of what is called dead space ventilation.

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 Dead Space
Dead space is defined as the part of the respiratory tract, where gaseous exchange
does not take place. Air present in the dead space is called dead space air. Dead space is of
two types: (i) Anatomical dead space (ii) Physiological dead space.
(i) Anatomical Dead Space
Anatomical dead space extends from nose up to terminal bronchiole. It includes nose,
pharynx, trachea, bronchi and branches of bronchi up to terminal bronchioles. These
structures serve only as the passage for air movement. Gaseous exchange does not take place
in these structures.
(ii) Physiological Dead Space
Physiologic dead space is defined as the volume of gas inspired that does not takes
part in gas exchange in the airways and alveoli.

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