AVIAN RESPIRATION – MORPHOLOGY, PHYSIOLOGY AND

FUNCTIONING

Sneha kumari
Department of biosciences ,chandigarh university(UIBT)
mohali, NH-05, PUNJAB-140413
snehaa2503@gmail.com

Submitted by : Sneha Kumari Submitted to: Dr. Shivam Jasrotia

UID= 23MSZ10046
Sub= Comparative anatomy of vertebrates

AVIAN RESPIRATION – MORPHOLOGY, PHYSIOLOGY AND

FUNCTIONING
Abstract
The avian respiratory system contains some fundamental differences to the
mammalian system. Abstract The avian respiratory apparatus is separated into a gas
exchanger (the lung) and ventilators (the air sacs). Synchronized bellows-like
movements of the cranial and caudal air sacs ventilate the lung continuously and
unidirectionally in a caudocranial direction. With the lungs practically rigid, after their
insertion into the ribs and the vertebrae and on attaching to the membranous horizontal
septum, surface tension is not a constraining factor to the intensity that the gas
exchange tissue can subdivide. Delicate, transparent, capacious and avascular, the air
sacs are not directly involved in gas exchange. The airway system comprises of a
three-tiered system of passageways, namely a primary bronchus, the secondary
bronchi and the tertiary bronchi (parabronchi). The crosscurrent system is formed by
the perpendicular arrangement between the mass (convective) air flow in the
parabronchial lumen and the centripetal (inward) flow of the venous blood in the
exchange tissue; the countercurrent system consists of the centrifugal (outward) flow
of air from the parabronchial lumen into the air capillaries and the centripetal (inward)
flow of blood in the blood capillaries, and; the multicapillary serial arterialization
system is formed by the blood capillaries and the air capillaries where venous blood is
oxygenated in succession at the infinite number of points where the respiratory
units contact exchange tissue. Together with the aforementioned systems, features like
large capillary blood volume, extensive respiratory surface area and thin blood.gas
barrier accord high pulmonary diffusing capacity of O2 that supports the high
metabolic capacities and energetic lifestyles of birds.

Keywords
Birds ,Lung ,Air sacs , Respiration ,Development ,Flight , Oxygen

INTRODUCTION
The avian lung exhibits a special architecture and upon our understanding of this
architecture will depend our conception of its physiology. Locy and Larsell (1916).
The avian respiratory apparatus, the lung-air sac system, has been continuously
investigated for about four and half centuries (Coitier 1573). Notwithstanding this
unremitting inquiry, regarding some aspects of its biology, few organ systems have
remained as stubbornly intractable (Brown et al. 1995; Maina et al. 2009). Among the
most contro versial organs-systems, Farner (1970) ranked the avian respiratory
system very high. Incontrovertibly, the avian respiratory system is structurally the
most complex and functionally the most efficient gas exchanger that has evolved
among the extant air-breathing vertebrates (King 1966; Duncker 1971; Scheid 1979;
McLelland 1989; Maina 2005, 2006): the respiratory system of birds is more efficient
than that of mammals, transferring more oxygen with each breath (Scheid 1979; Fedde
1980). Structurally and functionally, the avian respiratory apparatus is sepa rated int o
a gas exchanging part (the lung) and a ventilator one (the air sacs). In active
vertebrates, locomotion exacts the greatest demands on the respiratory system (Banzett
et al. 1992).

Avian Nasal Cavity and Oropharynx

The nostrils of the bird, which lead into the nasal cavity, may have a flap of horn to
protect them, known as the Operculum. The Oral Cavity and the Nasal Cavity of the
bird are interconnecting via a slit in the hard palate called the choana. Birds lack a soft
palate.

©Nottingham 2008 Saggital Section of an Avian skull showing the conchae.

There are rostral, middle and caudal conchae arising from the lateral wall, filling part
of the nasal cavity. The rostral conchae in the vestibular region are lined with
stratified squamous epithelium. The middle conchae in the respiratory region are
lined with respiratory epithelium. The caudal conchae in the olfactory region are
lined with olfactory epithelium. The infraorbital sinus is a triangular cavity under the
skin, rostroventral to the eye. Some marine birds have a salt gland (nasal gland) which
excretes sodium.The larynx is on the floor of the oropharynx supported by cricoid and
paired arytenoid cartilages which are different in structure to those in mammals.
There is no epiglottis and there are no vocal folds - birds vocalise using a syrinx.The
avian trachea is composed of tightly stacked cartilages which are shaped similarly to
signet rings. They are complete with no dorsal space as in the mammalian trachea and
overlap considerably. The trachea can be palpated on the right side of the neck; it runs
alongside the oesophagus. The trachea is lined with respiratory epithelium. The
trachea bifurcates into two main bronchi as in mammals. The syrinx is formed by this
terminal part of the trachea.

Avian Lungs
Avian lungs are relatively compact, with a bird's lungs being approximately 50% as
large as the lung of a mammal of a similar size. The lungs are unlobed and do not have
the capacity to expand due to the close arrangement between the finite gas exchange
structures, i.e.the air capillaries and blood capillaries and scanty connective tissue. The
lungs are positioned in the craniodorsal region of the body, and are deeply indented by
both the thoracic vertebrae and ribs. Birds do not have a pleural cavity as the lungs do
not expand, thus the membranes are not necessary. One primary bronchus from
the trachea enters each lung, narrowing as it travels through, and communicates with
the abdominal air sac. This bronchus gives off branches as it travels through the lung,
known as secondary bronchi. Each of these gives off a further 400-500 parabronchi.
The parabronchichi give rise to atria. The atria form infundibulae from which the air
capillaries emerge. It is in the walls of the latter structures that gaseous
exchange takes place.
Air Sacs
Birds lack a diaphragm, and their thoracic and abdominal cavities are continuous. The
bird has a number of thin walled, easily distensible air sacs which can extend to
approximately 10x the volume of the lungs. Theye are present within body cavities,
and extend into some specific bones to take the place of the bone marrow. This has the
added function of reducing the weight of the bone, as they are essentially filled with
air. The air sacs create unidirectional flow of air to maximise oxygen extraction and
reduce heat production during flight.The chicken has 8 air sacs:Cervical - extends
within the cervical and thoracic vertebrae.Clavicular - lies within the thoracic inlet,
surrounding the heart, and within the humerus in the forelimb.Cranial Thoracic (x2) -
these are ventral to the lungs. Caudal Thoracic (x2) - located between the body wall
and the thoracic air sacs. abdominal (x2) - these are the largest air sacs and fill the
caudodorsal region of the abdomen, in contact with small and large
intestines, kidneys and reproductive organs. In addition, these airsacs utilise space
within the acetabulum and synsacrum The air sacs originate as blister-like structures at
different times and places on the cranial, the ventral and the caudal edges of the avian
lung (Maina 2003a; Figs. 19, 20). Except for the abdominal air sacs which enter the
postpulmonary septum to lie in the coelomic cavity, all the other air sacs enter the
septum, separating it into horizontal and oblique septa (King 1966; King and Molony
1971; Duncker 1978). In the chick embryo, six pairs of primordial air sacs initially
develop (Romanoff 1960). At maturity, however, the total number of air sacs is
smaller because some of the air sacs merge. The primordia of the abdominal air sacs
appear between days 5 and 7 of incubation while those of the cervical air sacs appear
between days 6 and 8 (Maina 2003b). From day 15, the abdominal air sacs are the
largest air sacs. At hatching (day 21), except for the clavicular air sacs, all the other air
sacs, namely the cervical, the craniothoracic, the caudothoracic and the abdominal air
sacs are paired. The sites where the air sacs connect to the lung are called ostia (Figs.
19, 20). In different species of birds, different bones are pneumatized to various
extents by different air sacs at different times of development (e.g., Hogg 1984;
Farjado et al. 2007; Dumont 2010). The mechanism by which it happens is, however,
unclear and needs to be investigated. The cervical, clavicular and cranial thoracic air
sacs form one functional group - cranial and the caudal thoracic and abdominal air
sacs forming another, caudal functional group. The air sacs have a vital role
in ventilation, but do not have the capacity for gaseous exchange.

Avian Ventilation
Ventilation in birds is strikingly different to that of mammals in that air flows through
the lungs in the same direction during both inspiration and expiration. In addition, both
the intaking of air and the expelling of air are active processes,
requiring muscle contraction.

During inspiration, the ribs are drawn forwards and the sternum lowered,the caudal
air sacs receiving fresh air. Simultaneously, the Cranial air sacs receive air which was
inhaled at the previous inhalation which is drawn from the lungs, this air has lost much
of its oxygen content.During expiration, the sternum is drawn caudal and dorsal, the
air sacs are compressed, air from the caudal air sac passes through the lungs, while the
air in the cranial air sac leaves via the trachea. Thus, oxygenated air passes through the
lungs on both inspiration and expiration.

Avian gaseous exchange

Avian Gas exchange takes place not in alveoli, as in mammals, but within air
capillaries which are extensions of the parabronchial lumen. They are an
interconnecting network of loops, and closely intertwine with blood capillaries. The
air capillaries and blood capillaries are arranged so that flow is crosscurrent. This
makes the gaseous exchange, which occurs from one to the other, extremely
efficient.Function of the avian respiratory system The directions of mass air flow in
the parabronchial lumen relative to the centripetal (inward) flow of the venous
(deoxygenated) blood are essentially perpendicular (Figs. 56–58): the arrangement
forms a crosscurrent system. A countercurrent system is formed by the centripetal
(inward) flow of venous (deoxygenated) blood in the blood capillaries and outward
(centrifugal) diffusion of O2 in the air capillaries (Figs. 59–61). The role of the
countercurrent system to the gas exchange process in the avian lung is uncertain.
Piiper and Scheid (1973) dismissed the countercurrent system as a ‘countercurrent-
like mechanism’ and Scheid (1979) termed it ‘an auxiliary mechanism superimposed
on, and independent of, the basic crosscurrent arrangement between the capillary
blood flow and the bulk parabronchial gas flow’. It is, however, because convective
flow of blood occurs in the blood capillaries and diffusion of air occurs in the air
capillaries. The partial pressure of O2 in the blood capillaries equilibrates with that
in air that flows through the parabronchial lumen and then moves centrifugally
(peripherally) into the air capillaries. At the entrance of a paleopulmonic
parabronchus, i.e., the part of the parabronchus that connects to the mediodorsal
secondary bronchus, deoxygenated blood is exposed to air at a high partial pressure of
O2 (PO2) while at the opposite end, i.e., the part that connects to the medioventral
secondary bronchus, the oxygenated blood is exposed to air with a low PO2. The
arrangement of the blood capillaries and the air capillaries in the exchange tissue of a
parabronchus forms the multicapillary serial arterialization system (Scheid 1979;
Maina 2005) (Figs. 58–61). The quantity of O2 in the blood which returns to the heart
(arterial blood) through the pulmonary vein derives from accretion of very small
quantities of O2 exchanged at an infinite number of points in the exchange tissue
where the air capillaries and the blood capillaries contact. In the lung of the Domestic
fowl (total lung volume 27 cm3 ) (Maina 2005), the parabronchi span a length of 32
meters, if they are connected end-to-end (Maina unpublished observations). Because
of the close entwining between the air and the blood capillaries and the dense
packing of the parabronchi in the avian lung, the multicapillary serial arterialization
system increases the volume of the gas exchange tissue in which air and blood are
exposed to each other across a blood-gas barrier. They increase the quantity of
respiratory gases (O2 and CO2) that is exchanged
Conclusion

Together, properties like unidirectional and continuous ventilation of the gas exchange
tissue, gas exchange designs like the crosscurrent and the multicapillary serial
arterialization systems and highly refined pulmonary mor phometric parameters such
as a thin blood-gas barrier and a large respiratory surface area and capillary blood
volume accord high gas exchange efficiency in the avian lung, permitting active flight
even under extreme conditions of high altitude (Maina 2000)
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