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Vol. 21, No.

10 October 1999 20TH ANNIVERSARY

CE Refereed Peer Review

The Avian
FOCAL POINT Respiratory System*
★ The respiratory system of birds
differs significantly from that of University of Tennessee
mammals; thus an understanding Susan E. Orosz, PhD, DVM
of the unique anatomy of the
avian respiratory system and
ABSTRACT: The avian respiratory system has important anatomic and physiologic differences
an awareness of the signs of
that are crucial for the veterinary clinician to understand. These principles are important in
respiratory illness in birds are
anesthesia, physical examination, diagnostic workup, and treatment. The respiratory system
imperative in effectively caring can be functionally and clinically divided into the upper and lower respiratory tracts. The prin-
for feathered patients. ciples of anatomy and physiology of the two components of the respiratory system as they re-
late to clinical medicine are reviewed.

KEY FACTS

B
ecause of the unique physiologic demands associated with flight, the
■ Birds are often presented for structure and function of the respiratory system in birds are significantly
treatment of an eye problem different from those in mammals. For example, birds have an extremely
when their condition is actually efficient mechanism for oxygen extraction, which enables them to maintain
caused by infraorbital sinusitis their high metabolic rates (necessary to cope with the physical stresses associated
secondary to nutritional with flight) and to fly at high altitudes (where oxygen is reduced).
deficiencies. This specialized system works extremely efficiently when a bird is healthy;
when a bird is ill, however, respiratory signs can be pronounced. A primary rea-
■ The syrinx is a common site son that birds are presented to clinics is for the treatment of obvious respiratory
for tracheal granulomas or distress. A basic understanding of the anatomy and physiology of the avian re-
entrapment of foreign bodies. spiratory system (Figure 1) is required to manage such a condition successfully.
The first step in assessing birds with respiratory signs is to determine whether
■ On radiographs, birds’ lungs the signs are localized to the upper or lower respiratory tract or whether both
have a spongy appearance and components are affected (see Components of the Respiratory Tract).
are best viewed from a lateral
projection. UPPER RESPIRATORY TRACT
Nares
■ Granulomatous reactions to Visual inspection of a bird’s head begins with an examination of the nares (i.e.,
infection are common in birds. nostrils), which are usually found medially within the area of the cere. The cere
is usually an unfeathered area on the dorsal surface of the upper bill or maxillary
rhamphotheca. The color of the cere sometimes indicates the sex of a bird; for
example, in the budgerigar (commonly known as a parakeet), the cere is blue in
males and pinkish-brown in females. Changes in the normal color pattern are
suggestive of a gonadal tumor.1
When examining the cere, the feathers above it should be examined. Matting
of the feathers with a serous exudate is highly suggestive of an upper respiratory
*Adapted from Orosz SE: The avian respiratory system. Perspectives May/June:47–55,
1994.
Small Animal/Exotics 20TH ANNIVERSARY Compendium October 1999

Figure 1—Relationship of the cranial and caudal respiratory structures in the macaw. The infraorbital sinus has extensive projec-
tions through the skull and is connected to the cervicocephalic air sac. The caudal respiratory tract includes the trachea, syrinx,
lungs, and pulmonary air sacs. Note that birds do not have a functional diaphragm. The inset shows the midsagittal section of the
head. The conchae project from the lateral wall in the nasal cavity to filter airborne particles and to warm and humidify air. Air
moves through the choanal slit into the glottis. The glottis is found just caudal to the base of the tongue and is not covered by an
epiglottis. (Illustration by Kevin Somerville, Jamestown, Rhode Island)

tract infection. Wet or matted feathers at the carpus chae (rostral, middle, and caudal). The middle concha
(i.e., “the wrist”) suggest a bird’s attempts to remove ex- is the largest, and its dorsal surface provides the open-
udate with its wing.1 The shape of the nares can also ing for drainage of the infraorbital sinus.2
alert practitioners to the presence of disease—chronic The infraorbital sinus, the only true paranasal sinus
respiratory tract infections can cause the nares to be- in birds, has extensive diverticula into the skull.2 The
come enlarged and misshapen or plugged and shrunk- rostral diverticulum extends into the maxillary beak,
en.1 the preorbital diverticulum is just rostral to the orbit
and extends ventrally and medially, the postorbital di-
Nasal Cavity verticulum is subdivided around the operculum or ex-
Air moves from the nares into the nasal cavity by ternal acoustic meatus of the ear canal, and the
passing over the operculum, which acts as a baffle to fil- mandibular diverticulum extends into the mandibular
ter out large particulate matter. The nasal cavity is often beak.1
divided by a septum. The lateral walls of the nasal cavi- The infraorbital sinus communicates caudally with the
ty are comprised of three highly vascularized nasal con- cervicocephalic air sac.1 This air sac, which varies in size

OPERCULUM ■ NASAL CONCHAE ■ CERVICOCEPHALIC AIR SAC


Compendium October 1999 20TH ANNIVERSARY Small Animal/Exotics

among species,2 is found just under the skin caudodor- be larger to compensate for the increased resistance to
sally on the skull in the caudal cervical region. It does airflow.3 This results in an increased tracheal dead space
not communicate with the pulmonary air sacs.2 before the oxygen-exchange surface of the lung can be
An understanding of the infraorbital sinus, its open- reached. To compensate, birds have a greater tidal vol-
ings, and its anatomic relationship with the skull is im- ume and a reduced respiratory rate.3 The increased
portant for examination of the upper respiratory tract, pressure differences created during inspiration require
irrigation and specimen collection, and surgical drain- that the trachea be of sufficient strength to prevent col-
age.1 The infraorbital sinus drains dorsally over the mid- lapse. Correspondingly, the tracheal rings (shaped like a
dle nasal concha. 2 It also series of signet rings) are complete and overlapping to
Components of the communicates with the cau- increase rigidity.2
Respiratory Tract dal nasal concha through a The anatomy of the avian glottis makes the trachea
dorsal opening.2 Although easy to intubate.3 Nevertheless, because the trachea is
Upper Respiratory Tract the sinus is lined with a cili- less compliant than in mammals, care should be taken
■ Nares ated epithelium, both of its to avoid overinflation when using a cuffed endotracheal
■ Nasal cavity openings are dorsal2; this ar- tube. Those who monitor anesthesia should ensure that
rangement does not allow the bird’s respiratory rate is slow and regular and that it
■ Infraorbital sinus
large amounts of inflam- is taking relatively deep breaths that mimic its natural
■ Cervicocephalic air sac matory products to be re- respiratory rate and tidal volume.3
moved.1 As a result, caseated
Lower Respiratory Tract pus can easily accumulate in Syrinx
■ Glottis the infraorbital sinus, dis- After entering the thoracic inlet, the trachea divides
■ Trachea tending it laterally so that into two primary bronchi. The syrinx (i.e., the voice
the area around the eye ap- box of birds) can be found in the vicinity of this bifur-
■ Primary bronchi
pears puffy. cation, although its exact location varies by species.2
■ Syrinx Poor nutrition can con- The syrinx has both internal and external membranes
■ Secondary bronchi tribute to infraorbital si- and associated syringeal muscles that are involved in
■ Parabronchi nusitis. Birds in a poor nu- the production of song.2
■ Lungs tritional plane are prone to Because the respiratory passageways narrow consider-
■ Pulmonary air sacs hypovitaminosis A, a condi- ably at this point, the syrinx is a common site for tra-
tion that results in squa- cheal granulomas or entrapment of foreign bodies (e.g.,
mous metaplasia of epithe- seeds). Blockage can cause acute, severe respiratory dysp-
lial surfaces, such as those in the respiratory tract. The nea, requiring emergency cannulation of a pulmonary
absence of ciliated epithelium makes a bird even more air sac. In birds with syringeal aspergillomas, acute
susceptible to infection. For this reason, birds are often signs include dyspnea, which may be preceded by a
presented for treatment of an eye problem when their voice change.1
condition is actually caused by infraorbital sinusitis sec-
ondary to nutritional deficiencies. Lungs and Air Sac System
The paired lungs are firmly attached to the ribs and
LOWER RESPIRATORY TRACT vertebral column dorsally, extending from the thoracic
Glottis, Larynx, and Trachea inlet caudally to the level of the adrenals and cranial di-
Unlike in mammals, the opening of the avian glottis vision of the kidneys.2 On radiographs, they have a
is not covered by an epiglottis; as a result, birds are more spongy appearance and are best viewed from a lateral
prone to aspiration pneumonia. The larynx is composed projection.1 The primary bronchi continue into the
of four partially ossified cartilages that are covered by lung, where they subsequently divide into secondary
mucosa (i.e., the laryngeal mound).2 The larynx of birds bronchi and the parabronchi. The parabronchi in birds
is not involved in the production of sound.2 are very different from those found in mammals in that
Because birds must use their beaks to preen as well as they freely anastomose with each other, have diverticula
to manipulate objects, their necks must be of sufficient or atria within their walls to allow air exchange, and
length to reach the tips of their body parts. Conse- maintain a constant mean diameter.2
quently, birds’ necks are considerably longer than those Secondary bronchi branch from the primary bronchi
of comparably sized mammals.2 This increased neck and are named for the region of the lung that they sup-
length has physiologic and biomechanical conse- ply.2 These secondary bronchi form a functional unit
quences. For example, the diameter of the trachea must that maintains the continuity of airflow. The medio-

INFRAORBITAL SINUS ■ HYPOVITAMINOSIS A ■ LARYNGEAL MOUND


Small Animal/Exotics 20TH ANNIVERSARY Compendium October 1999

Figure 2—Organization of the paleopulmonic and neopulmonic parabronchi and their role in respiration. On inspiration, air
moves into the pulmonary air sacs, particularly the abdominal and caudal thoracic sacs. Additionally, as a result of aerodynamic
valving, some air moves unidirectionally through the paleopulmonic secondary bronchi for exchange of gases in the adjacent air
capillaries. On expiration, air moves bidirectionally through the parabronchi in the neopulmonic system. It also travels back
through the neopulmonic system into the paleopulmonic secondary bronchi and parabronchi. (Illustration by Kevin Somerville,
Jamestown, Rhode Island)
Compendium October 1999 Small Animal/Exotics

Your comprehensive
Handle with Care guide to diagnostic
Because of birds’ bellowslike breathing cycle, it is
important to allow the sternum to move freely during ultrasonography
handling. Birds should be held firmly yet gently
Nautrup and Tobias
around the base of the head and kept upright as
much as possible. Many birds believed to have
died of “fright” during examination probably died of
suffocation because of being held tightly around the
chest wall. This understanding can be applied in
reverse to treat apneic birds: The sternum can be
compressed and lifted to assist ventilation.

dorsal and medioventral secondary bronchi are connect-


ed with the cranial pulmonary air sacs, which include
the cervical, clavicular, and cranial thoracic air sacs.
These air sacs and the arrangement of the secondary
bronchi and their parabronchi in this portion of the
lung form the paleopulmonic air sac system (Figure 2).3 New
In this system, there is a one-way flow of air through a
series of tubes into the cranial portion of the pulmo-
nary tissue.3 This gaseous flow is much more efficient

149
than the neopulmonic system of mammals,3 in which
both oxygenated and deoxygenated air passes in and out $
of the same tubes.
The neopulmonic system is found in the caudal por- Robert E. Cartee, Editor
tion of the respiratory system in most birds and in- 400 pages, hard cover
cludes most of the lateroventral secondary bronchi (i.e.,
branches of the distal bronchi), the caudal thoracic air 1597 illustrations
sacs, and the abdominal air sacs (into which the prima-
ry bronchi extend and empty) (Figure 2).2,3 The lat- ■ Sonographic diagnosis in dogs and cats,
eroventrally directed secondary bronchi and the contin- including ultrasound, M-mode, pulsed
uation of the primary bronchi into the abdominal air and color Doppler echography
sacs predispose the caudal portion of the respiratory
system to infection. This is because the primary ■ Echocardiography, abdominal and pelvic
bronchus feeds directly into the abdominal air sac and sonography, and fetal ultrasonography
the secondary bronchi are directed ventrally, making it
easier for foreign material to enter the caudal air sac ■ Case illustrations using conventional
system.1 The functional significance of these two pul- radiography, computed microfocal
monary systems can best be described by examining the tomography, specimen photography,
respiratory cycle.
and line drawings
RESPIRATORY CYCLE ■ Recognition of the disease process and
In birds, respiration does not rely on a change in the
volume of the thorax. Instead, pressure changes in the
courses of treatment
air sacs move the air through a nonexpanding lung.
These pressure changes result from volume changes in
the thoracoabdominal cavity and require muscular ex- CALL OR FAX TODAY TO ORDER
ertion during both inspiration and expiration.2,3 800-426-9119 • Fax: 800-556-3288
On inspiration, the ribs and sternum are drawn cra-
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Email: books.vls@medimedia.com

PALEO- AND NEOPULMONIC AIR SAC SYSTEM


Small Animal/Exotics 20TH ANNIVERSARY Compendium October 1999

change surfaces, the air capillaries (comparable to the


The Two-Breath Cycle
alveoli of mammals). The blood–gas barrier of birds is
First inspiration—Air flows into the trachea through anatomically similar to that of mammals except that it
the primary bronchi, and most of the air goes is much thinner in birds, thus enhancing efficient ex-
directly into the abdominal air sac and/or into the change. Blood flow is at a 90˚ angle from that of air-
flow, thereby acting physiologically as a cross-current
caudal thoracic air sac via the lateroventral
exchanger, which improves a bird’s ability to oxygenate
secondary bronchi. blood.3 ND E IU
First expiration—Air travels back through the MP

M’
20th

 CO

S
neopulmonary parabronchi and is then shunted via DEFENSE MECHANISMS 9 9 9
9 - 1

aerodynamic valving into the mediodorsal OF THE LOWER 1 9 7

ANNIVERSARY
secondary bronchi or the paleopulmonic system; RESPIRATORY TRACT
the air moves through the parabronchi of this
system and gas exchange occurs.
One of the first lines of
defense of the lower respi- A LookBack
ratory tract is the mucocili-
Second inspiration—Air is pulled from the ary escalator. 3 This ana- In the 20 years that have passed
mediodorsal secondary bronchi through the tomic and physiologic since the founding of the
escalator uses mucus secret- Association of Avian
medioventral secondary bronchi and into the cranial
Veterinarians, avian medicine
pulmonary air sacs. ed from the goblet cells to
has exploded. Research advances
trap foreign material, with
Second expiration—Air moves out of the cranial air have provided veterinary
both being swept away by
sacs, into the opening of the medioventral clinicians with new diagnostic
cilia for removal from the
secondary bronchi, and out of the trachea. tools and a range of treatment
respiratory tract. This de- modalities to improve the health
fense mechanism can be of avian patients. Anatomic
overwhelmed by infection. and physiologic studies of the
dominal wall is drawn ventrally.2,3 This action can be When this happens, the respiratory system of birds have
likened to that of a bellows (see Handle with Care): The mucociliary escalator can- been used to improve anesthesia
coracoid–sternal junction corresponds to the fixed end not move infectious materi- and provide treatment regimens
of the bellows and the distal end of the sternum to the al and products of inflam- for both upper and lower
more movable end. When the thoracoabdominal space mation out of the lung respiratory tract infections.
is expanded, the air rushes into the air sacs (particularly quickly enough to keep the Recent studies have correlated
the caudal sacs) as a result of the reduced pressure. On parabronchi clear.3 Nebu- the normal anatomy of the
expiration, the ribs and sternum move caudodorsally, lization reduces the in- respiratory system with
closing the “bellows” and reducing the thoracoabdomi- creased stickiness of the computed tomography images so
nal space. Consequently, the pressure in the air sacs is mucus associated with in- that abnormalities can be
increased, causing the air to be expelled.2 In those birds fection and helps the esca- diagnosed. Clinically relevant
with both neopulmonic and paleopulmonic systems, air lator work more efficiently.3 studies have improved
enters and exits the respiratory system in a two-breath In birds with vitamin A de- nebulization techniques, drug
cycle (see The Two-Breath Cycle).3 This two-breath cy- ficiency, cells undergo a therapy, and surgical approaches
cle allows a continuous flow of air through the para- change to a squamous type to the respiratory tree. New
bronchi to the gas-exchange surfaces, whereas the less ef- without mucus or cilia. arsenals will build on these
ficient mammalian breathing cycle allows gas exchange This results in a breakdown studies to provide even better
to occur only at the end of inspiration. of this fundamental system treatment options in
Avian respiratory function is more efficient in other to remove foreign material. years to come.
ways as well. In birds, the lungs do not expand or con- A second line of defense (Photograph courtesy of Phil
tract significantly—only the air sacs do.3 Thus the ex- involves the epithelial cells Snow, University of Tennessee
change surfaces can be smaller and more tightly packed themselves. These cells of College of Veterinary Medicine)
than are the alveoli of mammals. Because this results in the lung parenchyma act as
an increased surface area in birds compared with mam- fixed macrophages,3 engulf-
mals, birds are more efficient than are mammals in cap- ing foreign material and
turing oxygen and removing carbon dioxide. The gas- transferring it to interstitial
exchange surfaces of birds include the atria, which lie in macrophages for destruc-
the walls of the parabronchi and lead to the major ex- tion. Because macrophages

AIR CAPILLARIES ■ MUCOCILIARY ESCALATOR ■ GOBLET CELLS ■ NEBULIZATION


Small Animal/Exotics 20TH ANNIVERSARY Compendium October 1999

are involved early in defense, granulomatous reactions ing aware of the signs of respiratory illness in birds are
to infection are common in birds. imperative in effectively caring for these feathered pa-
When defense mechanisms of the lower respiratory tients.
tract are overwhelmed, birds often exhibit a number of
signs, including voice change (particularly when the sy- REFERENCES
rinx is involved), increased abdominal exertion (when 1. McKibben JS, Harrison GJ: Clinical anatomy (with empha-
the parabronchi are plugged with infection), tail bob sis on the Amazon parrot), in Harrison GJ, Harrison LR
(eds): Clinical Avian Medicine and Surgery. Philadelphia, WB
(associated with increased respiratory effort), and clicks Saunders Co, 1986, p 31.
and wheezes (caused by air moving through a lumen 2. King AS, McLelland J: Respiratory system, in: Birds, Their
that is reduced in size).1 If lower respiratory tract infec- Structure and Function. Philadelphia, Baillière Tindall, 1984,
tions obstruct the trachea, the abdominal or caudal p 110.
thoracic air sac can be cannulated for oxygenation of 3. Fedde MR: Respiration, in Sturkie PD (ed): Avian Phys-
iology, ed 4. New York, Springer-Verlag, 1986, p 191.
the respiratory system. Oxygen can be administered
into the clavicular air sac as well.
About the Author
CONCLUSION
Dr. Orosz is a professor of avian and exotic animal medi-
Although the respiratory system in birds operates
cine in the Department of Comparative Medicine, College
more efficiently than it does in mammals, birds appear
of Veterinary Medicine, University of Tennessee, Knox-
to have a greater predisposition to respiratory complica-
ville, Tennessee.
tions associated with infections. Understanding the
unique anatomy of the avian respiratory system and be-