Birds

Introduction and Evolution

Birds (Class Aves) are the second largest class of vertebrates with around 9,600
recognised species – 726 of these reside in Australia. The success of birds can be
significantly attributed to the evolution of endothermy and flight, both of which have
allowed them to adapt to a wide range of environments. Birds evolved from reptiles
over 150 million years ago with fossil evidence from the Jurassic period showing
clear intermediates between reptiles and modern birds – the most famous of these
is Archaeopteryx. Birds are evolutionarily highly derived having many adaptations
for flight. Those extant species which do not fly have lost the ability to do so
secondarily. Amongst extant species, birds are most closely related to the
crocodilians with whom they share several features including a muscular gizzard
and diapsid skull. Birds, crocodilians and dinosaurs all diverged from a common
ancestor during the Triassic period.

Some of you may find yourselves working as veterinarians in the commercial

poultry industry, or as specialist avian veterinarians, in which case a thorough
understanding of avian anatomy is essential. However, many of you who enter
mixed or small animal practice will be expected to treat avian patients as many
people now keep birds as pets, egg layers or as a small-scale commercial
breeding enterprise and so an understanding of avian anatomy is also important
for the general practice vet.

Integument

Skin
In birds, the skin is relatively thinner and more loosely attached to the underlying
muscles than it is in mammals. In feathered areas, the epidermis is only about ten
cells thick, with half of these being the dead, cornified layer. In the foot pads, the
skin is much thicker. The dermis is also relatively thin and is less well vascularised
and innervated than in mammals. The comb and wattles of the chicken, turkey and
many wild species are formed by a thickened and very vascular dermis with many
arterio-venous anastomoses. Similarly, the brood patch, which develops on the
breast of incubating birds, is formed by a thickening of the dermis, vascular
development within the dermis, and the loss of feathers, all of which serve to
increase heat transfer from the adult bird to the eggs.

Birds develop horny structures from several regions of the skin:

Beak (bill or rhamphotheca): derived from the skin, the beak is a hard, keratinised
structure that grows continually to replace losses caused by wear, and may require
clipping in cage birds. Poultry are often debeaked to prevent cannibalism (note:
there are ethical concerns over this procedure given the beak‟s profuse sensory
innervation). Newly hatched chicks have an egg tooth on the tip of the beak that
allows them to break the egg shell.

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Claws: are generally curved with a softer under surface. They may also require
clipping in cage birds. Some species also have claws on the digits of the wing
(emu and kiwi on 2nd digit; occasionally on 1st digit of fowl and goose).

Spurs: consist of a bony core covered with a keratinised sheath. They are well
developed in the male fowl and turkey; the bony core developing after 6 months of
age. Removal of the bud in chicks prevents the spur from developing.

Scales: consist of raised patches of highly keratinised epidermis separated by

folds of normally keratinised skin.

Uropygial (preen or oil) gland: lies dorsally on the tail and consists of two lobes
opening to a median papilla with a tuft of feathers (uropygial wick). Its secretion is
a holocrine lipoid material that keeps the feathers, beak and scales supple and
water repellant (particularly important in waterbirds which have very large
uropygial glands) – when birds preen they distribute the secretion over the
feathers. There is some suggestion that it may also be a source of vitamin D. It is
not present in all species.

Birds have no cutaneous glands apart from the uropygial, external ear (wax) and
vent (mucous) glands. However, the epidermis as a whole may release lipoid
sebaceous secretions. Birds have no sweat glands, so temperature regulation is
largely achieved by evaporative cooling from the respiratory tract.

Feathers
Feathers, like scales, are highly keratinised epidermal outgrowths. Pigments from
chromatophores within the epidermis are carried into the feather as it develops,
giving feathers their wide variety of colours. The iridescence characteristic of
feathers is created by the light refractive effects of the barbs and barbules (below).

Contour feathers are the most prominent feathers of the body and assist in
both shaping the bird aerodynamically and in providing protection to the skin.

Flight feathers are the main feathers involved in flight. They occur on the wings
(remiges) and tail (retrices). Wing feathers are divided into primaries, which
are found on the manus, and secondaries, which are located on the
antebrachium (ulna). Both contour and flight feathers have a similar structure
and are arranged in well-defined linear tracts called pterylae or feather tracts;
however, flight feathers are larger and stiffer than contour feathers.

In between the contour and flight feathers are the down feathers which provide
insulation. Some birds also have filoplumes which are long, slender feathers, with
barbs only at their tips, which originate from extensively innervated follicles and
serve a proprioceptive function; they also play a role in courtship and display
behaviours. Most birds possess short, stiff feathers known as bristles which are
found around the eyes and nostrils where they help keep out foreign materials;
insectivorous birds have particularly long bristles around their mouths which assist
in catching prey. Natal down feathers are present on chicks at hatching or sprout
shortly after.

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Structure
The main axis of the feather is formed by the quill, which has a hollow proximal
end, called the calamus, which is embedded in the follicle in the skin, and a distal
solid portion called the shaft or rachis. The dermis of the follicle projects slightly
into the calamus and forms a dermal papilla which is covered with a thin layer of
epidermis. This layer of epidermis forms an epidermal collar at the base of the
feather and, being continuous with the epidermis of the follicle, is the main
attachment of the feather.

The rachis bears two lines of stiff filaments called barbs which, in turn, give rise to
even finer filaments, the barbules. Each barbule carries tiny hooks which engage
with the ridges of adjacent barbuIes to form a unified surface - the vane. If the
vane is disrupted so that the hooks disengage, they are easily re-engaged during
preening. In birds that have lost the ability of flight, such as the emu, the barbules
lack hooks and so the feathers are fluffy rather than smooth and sleek.

In some more ancestral species there is a small opening at the junction of the
rachis and calamus near which a small afterfeather may arise – these feathers
provide further insulation.

Development
Feathers develop within dermal follicles and are folded within an epidermal
keratinised feather sheath. As the feather sheath ruptures through the epidermis,
the barbs unfurl to their normal position and the sheath is sloughed off.

Moulting
As feathers become worn, they are replaced by moulting. The fowl moults three
times in its first 6 months (2 complete and 1 partial moult) and then has a complete
moult each autumn. Other birds generally moult once a year after breeding
(occasionally twice a year), and usually only a few feathers are lost at a time and
then replaced so that flight is not disrupted. However, ducks and geese loose all
their flight feathers at once, making them temporarily flightless (and vulnerable).
Old feathers are ejected from the feather follicle by the emerging new feather.

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Figure 1. Feather Anatomy

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Skeleton

The avian skeleton is light but very strong, having a greater content of calcium
phosphate than mammalian bone. A unique characteristic of avian bone is the
invasion by diverticula of the air sacs – such bones are described as being
pneumatic and this enables bones to be proportionally large and strong, without
being correspondingly heavy. Diverticula extend through the pneumatic foramina
into the medullary cavities of the bones, consequently reducing the volume of bone
marrow. In most species the skull, vertebrae, pelvis, humerus, femur, sternum and
costal bones are pneumatic. In the chicken, only the humerus is pneumatic.

As in mammals, most bones are preformed as cartilage which later ossifies.

Another avian specialisation is the production by females of medullary bone on the
inner surface of the endosteum just prior to egg laying, which is thought to provide
a store of calcium for the formation of the egg shell. Its deposition and subsequent
mobilisation are under hormonal control.

Skull
The avian skull evolved from the diapsid reptilian skull but shows significant
modification:
- enlarged cranium
- jaws are toothless and covered by keratinised sheaths
- the skull is prokinetic: most of the upper jaw consists of the premaxilla and nasal
bone which forms a cartilaginous craniofacial hinge with the frontal bone allowing
the upper jaw to be raised as the lower jaw is depressed. This kinesis is especially
well developed in parrots.

Vertebrae
The number of cervical vertebrae is proportional to the length of the neck - there
are between 14 and 17 cervical vertebrae in the chicken, 8 in small birds and up to
25 in the swan. The neck is very flexible in contrast to the thoracic vertebrae which
are rigidly fused to each other, providing a strong fulcrum for the action of the
wings. Most cervical vertebrae (except the atlas) possess vestiges of ribs, but the
last two cervical vertebrae bear large, mobile ribs.

There are 5 thoracic vertebrae (and so 5 ribs) in the fowl and the first three, with
the addition of the last cervical vertebra, form a fused notarium. Thoracic vertebrae
are defined as having a complete rib consisting of both dorsal and ventral
components - the dorsal rib corresponds to the osseous portion, and the ventral rib
corresponds to the cartilaginous portion, of the mammalian rib. Most of the dorsal
ribs have an uncinate process that overlaps with the adjacent rib, providing an
increased area for muscle and ligament attachment, and additional support to the
thoracic wall. The fourth thoracic vertebra is freely mobile, articulating with the
notarium and the synsacrum.

The synsacrum is a fused section of 15-16 vertebrae from the thoracic, lumbar,
sacral and caudal regions. Distal to the synsacrum are 6 freely mobile caudal
vertebrae and a flattened pygostyle, comprised of 5-6 fused caudal vertebrae,
which supports the tail.

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The sternum has a large midventral keel in flying birds which provides attachment
for the flight muscles. This is absent in the flightless species (except penguin).

Forelimb
The pectoral girdle consists of a scapula, clavicle and coracoid. The scapula is
long and blade-like, and is firmly attached to the ribs. The clavicles unite to form
the furcula (wishbone) and the massive coracoids articulate with the sternum,
holding the wings out and preventing the thoracic cage from collapsing during
contraction of the pectoral muscles.

The humerus is large and is the point of insertion of the main flight muscles. The
head of the humerus articulates with the glenoid fossa formed by the scapula and
coracoid. The ulna is thicker and longer than the radius. During extension of the
elbow joint the radius slides proximally pulling the manus with it so that the whole
wing straightens.

The manus is much reduced with only three digits (II, III and IV) arising from a
fused carpometacarpus. The first digit (II) is associated with a series of feathers
known as the alula (function described later).

Hindlimb
The ilium, ischium and pubis are firmly united with each other and with the
vertebral column to form a sturdy pelvic girdle. However, the pubic bones and
ischia of each side do not fuse in the midline to form a pubic symphysis – this may
have evolved to allow for the passage of a large egg.

The acetabulum is large and dorsal to it is the anti-trochanter which provides

additional articulation for the femur and allows the bird to stand on one leg in spite
of weak adductor muscles.

The proximal row of tarsal bones is fused to the tibia to form a tibiotarsus, and the
distal row is fused to the metatarsus (itself a fusion of tarsals II, III, IV) to form the
tarsometatarsus. The fibula is reduced to a small spine which is fused to the
tibiotarsus. In the fowl, digits I to IV are present, with digit I directed backwards.

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Figure 2. Axial Skeleton of the Chicken

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Figure 3. Appendicular Skeleton of the Chicken

Muscles

Head, Neck and Trunk

The epaxial and hypaxial muscles of the neck are divided into a network of fine
bundles making the neck very mobile.

The muscles of the back (e.g. iliocostalis and longissimus dorsi) are much reduced
due to the fusion of the vertebrae. The main muscles of the tail are the levator and
depressor caudae. The four abdominal muscles (external oblique, internal oblique,
rectus abdominis and transversus abdominis) are similar to those in mammals but,
in the fowl, the rectus abdominis is reduced to a membrane.

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Wing
The main flight muscles are the pectoralis and supracoracoideus (these are what
you get when you ask for “chicken breasts”) which both have their origin on the
sternum. The pectoralis is responsible for the powerful downstroke of the wing and
inserts directly on the ventral surface of the humerus. The supracoracoideus,
which is responsible for the upstroke, sends its tendon through the triosseal canal
(formed by the scapula, coracoid and clavicle) to insert on the dorsal surface of the
humerus.

Figure 4. Wing Musculature of the Chicken

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Hind leg
Muscles of the hind leg are well developed in those species which are cursorial.
There is considerable difference of opinion on which (if any) of these avian
muscles are homologous with their mammalian counterparts.

Birds are able to sleep while perching because the digital flexor tendons pass over
the caudal surface of the intertarsal joint; flexion of this joint pulls on the tendon
and clamps the digits around the perch.

Birds have two types of muscle fibres, red and white, which give muscles their
characteristic dark (chicken leg or dark meat) or light (chicken breast or light meat)
appearance, respectively. Dark meat consists of a higher proportion of red fibres
which contain large amounts of myoglobin, have more mitochondria, lipid globules
and greater vascularity than the white fibres which predominate in light meat. Red
fibres use fat as an energy source which makes them capable of sustained effort,
while white fibres rely on glycogen which enables them to provide a rapid burst of
power.

Digestive System

Introduction
The digestive system of birds is very simple. The main modifications from the
general tetrapod plan, and which are associated with the evolution of flight, are the
loss of the teeth and heavy jaw bones and their associated muscles; and the
development of a bill, tongue and gizzard instead.

Birds eat a wide variety of foods and this has led to considerable interspecific
variation in the digestive system. However, as in mammals, the digestive tract has
a basic structural pattern.

Bill and Tongue

These are used for obtaining food and show many adaptive modifications. In a few
cases, there is mechanical treatment of the food by the bill, but mastication takes
place in the gizzard. The tongue in birds is considerably more variable than in
mammals. Consider the „furry‟ tongue of nectivores such as the rainbow lorikeet
as compared to the pestle-like tongue of the seed and nut eating parrots. It is
supported by the hyoid apparatus and often shows superficial keratinisation.

Oropharynx
There is no soft palate in birds and so there is no clear demarcation between
mouth and pharynx – hence the term oropharynx. The roof of the oropharynx is
formed by the hard palate. The choanal cleft in the hard palate opens to the nasal
cavity while the more caudal infundibular cleft leads to the auditory (eustachian)
canal. Numerous mechanical papillae are present on the hard palate which help to
move the bolus of food towards the oesophagus. Mucous secreting salivary
glands are well developed for lubrication.

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Figure 5. Head and Buccal Cavity

Oesophagus and Crop

The oesophagus is relatively long, wide and thin-walled. Proximally it is positioned
between the trachea and cervical muscles but very quickly deviates to the right
side of the neck – an important point to remember when crop feeding birds.
Longitudinal folds allow for significant distension. The crop is a clearly
differentiated, expansible portion of the oesophagus used for food storage,
especially in seed-eaters. It provides a regular supply of food to the stomach. Both
crop and oesophagus are lined with stratified squamous epithelium, with mucous
glands in the tunica.

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The pigeon has a 2-lobed crop which produces "crop-milk" for nestlings from fat-
laden epithelial cells which proliferate and then desquamate.

Stomach
The stomach in birds is probably the most variable aspect of the digestive tract
among the different families. It is divided into a glandular proventriculus and a
muscular ventriculus.

Proventriculus
The proventriculus is a thick-walled, spindle-shaped continuation of the
oesophagus with a mucosa which appears white on gross examination. The
lumen is lined with a mucous-secreting columnar epithelium. Also present are
raised papillae which contain the ducts of multilobular glands that contain
oxynticopeptic cells, which produce both HCl and pepsin (note these are two
distinct cell types in mammals).

Gizzard
The gizzard consists of two thick masses of muscle that give it an overall lens-
shaped appearance. The muscles are attached to a central aponeurosis, with the
fibres arranged in bundles. The outer longitudinal muscle layer is lost, so
Auerbach's plexus lies on the surface under the serosa. The lining of the gizzard
is thin but extremely durable, with a cuboidal epithelium and many tubular gizzard
glands in the lamina propria which produce koilin, a carbohydrate-protein complex
that solidifies on the luminal surface forming a hard cuticle. The koilin cuticle is
continually replenished from the glands below as it is subject to abrahasion at the
surface. The gizzard produces considerable crushing forces and, with the aid of
grit, masticates food in a similar way to that of the teeth in mammals. The food
passes repeatedly between the gizzard and the proventriculus.

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Figure 6. Gizzard and Koilin Lining

Small Intestine
The small intestine consists of the duodenum, jejunum and ileum. The narrow
duodenum passes caudally from the right side of the gizzard, is U-shaped and
encloses the pancreas which empties into the distal portion of the duodenum
along with the bile ducts. The jejunum is very thin-walled and the contents appear
greenish. At the junction between the ileum and the jejunum is the vitelline
diverticulum, a remnant of the yolk sac which provides nourishment to the chick
for several days after hatching.

All parts of the small intestine show a similar histological structure to that of
mammals except that there is generally poor development of the sub-mucosa.
Brunner's glands are absent but mucous is secreted by the many goblet cells. The
epithelium is thrown into villi with crypts of Lieberkuhn but no lacteals are present.
This is the major site for digestion of starch, sugars, proteins and fats. Large
quantities of diffuse lymphatic tissue and nodules occur in the lamina propria.

Large Intestine
The large intestine consists of two blind-ending caeca and the colon. The caeca
arise at the ileocolic junction and are the site of bacterial breakdown of cellulose.
Caeca are well developed in the chicken where the lymphatic tissue in the neck
forms a caecal tonsil. Passerines and pigeons have very short caeca and parrots
lack them altogether. The colon is a straight tube, lined with villi, leading to the
cloaca.

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Cloaca
The cloaca is a common chamber for the digestive and urogenital tracts. It is
divided into three sequential chambers by folds of tissue:

Coprodeum: a reservoir for faeces and urine

Urodeum: contains the openings of the ureters, oviduct in the female and deferent
ducts in the male

Proctodeum: contains a small opening in its dorsal wall which leads to the
cloacal bursa (bursa of Fabricius) which is the site of B lymphocyte maturation

The proctodeum ends in the external opening known as the vent which is lined
with a stratified squamous epithelium. On the ventral surface of the vent is the
phallus of the male.

Pancreas
The pancreas consists of three lobes and usually has three ducts which deliver
pancreatic secretions to the distal duodenum. Histologically and functionally, it is
similar to the mammalian pancreas (see also description under Endocrine
Organs).

Liver
The liver is divided into right and left lobes which are connected cranially, with the
left lobe having both dorsal and ventral parts. As there is no diaphragm, the
caudal surface of the heart lies against the liver. The right lobe supports the gall
bladder. Each lobe contains a bile duct which enters the duodenum but only the
duct of the right lobe is connected directly to the gall bladder. The gall bladder
stores and concentrates bile which neutralises acid in the chyme, emulsifies fats
and in the chicken also contains amylase. Several species including pigeons and
budgies lack a gall bladder.

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Figure 7. Digestive System of the Chicken

Respiratory System

Most organs of the avian respiratory system are homologous with those of
mammals but their function is often somewhat different. The avian system is also
considerably more efficient than that of mammals.

Nasal cavity
The position, size and surrounds of the external nares are very variable in birds.
The left and right nasal cavities are, like mammals, divided by a septum·which may
be perforated. Conchae arise from the lateral wall dividing the nasal cavity into
three compartments: rostral, middle and caudal. The infraorbital (maxillary) sinus is
a large triangular space in the upper jaw with dorsal exits to the nasal cavity.

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The nasal cavity is important in olfaction, filtering inspired air, and as a heat and
water exchanger.

The nasal gland (salt gland) opens at the rostral concha and, in marine birds, it
secretes a 5% NaCI solution.

Figure 7. Nasal Cavity of the Chicken

Larynx
The larynx is located on the floor of the oropharynx and consists of four cartilages,
which may become ossified: the paired dorsal arytenoids form the glottis and
articulate caudally with the procricoid and the cricoid. The main function of the
larynx is to prevent extraneous material from entering the trachea (note that birds
lack an epiglottis). It may also modulate sound, although there are no vocal
chords, and noise production occurs in the syrinx (see below).

Trachea
The trachea consists of a series of complete, overlapping cartilage rings. In most
birds the trachea starts medially, passes to the right, and then returns to enter the
thorax medially. In some long-necked species such as swans the trachea is longer
than the neck and forms a loop at the thoracic inlet.

The mucosa of the trachea, larynx and syrinx consists of pseudostratified

columnar epithelium with goblet cells and projecting ridges carrying cilia.

Syrinx
The syrinx is positioned at the caudal end of the trachea where it bifurcates into
primary bronchi. It shows considerable variation between different species of birds

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but consists of a series of cartilages and tympaniform membranes. The syrinx
produces sound by contraction of the surrounding muscles causing vibration of the
tympaniform membranes during exhalation.

Larynx

Syrinx

Figure 8. Cartilages and Muscles of the Larynx and Syrinx in the Chicken

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Lungs
The lungs are relatively small, flattened, rectangular, unlobed and dorsal, with
vertebral ribs buried in their dorso-medial surface. Avian lungs are much smaller in
volume (1/10) than comparable mammalian lungs but their weights are similar.

Primary Bronchi
The two primary bronchi arise from the syrinx and pass through the lung in an S-
shape, decreasing in diameter until they terminate caudally at the abdominal air
sac (see later). The primary bronchi each give rise to many secondary bronchi.

Secondary Bronchi
Four groups of secondary bronchi arise from the primary bronchi. They are named
according to the parts of the lung they supply:
Medioventral
Mediodorsal
Lateroventral
Laterodorsal

Secondary bronchi communicate with the air sacs.

Figure 9. Trachea, Lungs and Bronchi of the Chicken

Parabronchi and Air Capillaries

The secondary bronchi give off large numbers of parabronchi, which have a small
and uniform diameter (1 - 1.5 mm) and which anastomose with each other end-to-
end forming loops. Numerous extensions, called atria, develop from the
parabronchial lumen and extend into wall of the parabronchus. The atria are

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polygonal cavities which lead to funnel-shaped infundibula and the terminal air-
capillaries. Air capillaries (3 – 10 µm diameter) form an extensive network and are
the site of gas exchange, making them the functional equivalent, or homologue, of
the mammalian alveoli. Air capillaries and their associated blood vessels comprise
much of the parabronchial walls.

Figure 10. Transverse Section Through a Parabronchus of the Chicken

Air Sacs
Air sacs are blind-ending, thin-walled extensions of the bronchi which function in
respiration, but due to poorly vascularised walls do not contribute significantly to
gas exchange. The air sacs function as bellows, moving air through the lungs:
during inspiration, air is drawn into the lungs and air sacs; during expiration the air
sacs are compressed and air from caudal sacs passes through parabronchi while
that from cranial sacs is exhaled through the trachea. The air sacs are positioned
in close proximity to thoracic and abdominal viscera, and diverticula enter various
bones and extend between skeletal muscles.

In the chicken there are 8 air sacs: single cervical and clavicular sacs, and paired
cranial thoracic, caudal thoracic and abdominal sacs.

Cervical Sac
A small median sac positioned between the lungs and dorsal to the oesophagus. It
has a pair of tubular diverticula which extend cranially into, and alongside, the
vertebral column.

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Clavicular Sac
A large, complex median sac in the thoracic inlet. Diverticula extend along the
sternum, around the heart, oesophagus, trachea, syrinx, and bones and muscles
of the pectoral girdle to the humerus – this is clinically significant because fractures
to the humerus may thus introduce infection into other air sacs and the lungs.

Cranial Thoracic Sacs

Paired sacs in a dorso-lateral position within the thoracic cage. No diverticula.
Caudal Thoracic Sacs
Paired sacs caudal to the cranial thoracic sacs. No diverticula.

Abdominal Sacs
Paired sacs that pass caudally between the intestines. Pelvic diverticula extend
around the kidneys; femoral diverticula pass to the head and neck of the femur.

Figure 11. Lungs and Air Sacs of the Chicken

Pleural Cavity and Membranes

Pleural membranes are similar to those found in mammals but the pleural cavity is
largely obliterated by fibrous strands which run between the two pleura. These
strands will partially prevent collapse of the lung if the pleural cavity is penetrated.

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COMPARISON OF AVIAN AND MAMMALIAN RESPIRATORY SYSTEMS

Avian Mammalian
Trachea Length 2.7 1
Trachea Radius 1.29 1
Tracheal Dead 4.5 1
Space
Breathing Rate 1 3
Lung Weight 1 1
Lung Volume 1 10
Blood-Gas Barrier 0.3 1.4
(µm)
Area for Gas 18 1.8
Exchange (cm2/g
body weight)

Air Pathways
Airflow between the air sacs and the lungs appears to be unidirectional during both
inspiration and expiration. Functioning as a two-cycle pump, air is drawn into the
caudal sacs during inspiration, with a small amount of air also entering the lungs.
During the following expiration, air is moved through the parabronchi and into the
cranial air sacs where it is eventually expelled from the cranial air sacs into the
primary bronchi.

The control of this unidirectional air flow is uncertain, but passive aero-dynamic
factors, rather than muscular control, may be most important because:
it operates in dead, as well as living, lungs
there are no valves in the lung.

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Figure 12. Air Flow Through the Lungs and Air Sacs of the Chicken

Cardiovascular System

Introduction
The avian cardiovascular system, with a double circulation (pulmonary and
systemic), and a divided heart, appears very similar to the mammalian system.
The sinus venosus is much reduced in birds (and absent in mammals as it merges
with the right atrium during embryonic development). The conus arteriosus divides
to form the pulmonary trunk which exits from the right ventricle, and an aortic or
systemic trunk which originates from the left ventricle. In mammals and birds the
systemic arch is a single structure which loops to the right in birds, and to the left
in mammals, reflecting their evolution from different reptilian ancestors.

Given the higher metabolic rate of birds as compared to mammals, the avian
cardiovascular system has a higher efficiency and performance than that of
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mammals. The avian heart is much larger than that of comparably sized mammal:
a sparrow's heart forms 1.3% of its body weight while a mouse's heart forms only
0.5% of its body weight. The avian heart rate is also considerably faster than that
of mammals: 350-470 bpm in the chicken and 1000bpm in quail! Blood pressure in
birds is higher than that in mammals.

Heart
The heart lies in a median position in the thoracic cage surrounded by the liver
(not by the lungs as in mammals). It is enclosed in a tough, fibrous pericardial sac
which contains a serous fluid between the visceral and parietal layers.

The cranial vena cava is separated into right and left vessels with the left cranial
vena cava draining directly into the right atrium. The caudal vena cava and right
cranial vena cava drain into the primitive thin-walled sinus venosus which is not
fully incorporated into the right atrium and is separated from it by thin muscular
sinuatrial valves. The right atrium is significantly larger than the left. The thin walls
of the atria are strengthened by muscle bands which pass through the inter-atrial
septa and radiate into pectinate muscles.

Unlike the tri-cuspid valve of mammals, the right atrio-ventricular (AV) orifice is
closed by a muscular AV valve which almost encircles the orifice and operates as
a sphincter muscle without chordae tendinae attaching it to papillary muscles. A
similar structure is found in crocodiles and monotremes. In some species (duck
and kiwi), the valve is partially membranous. The right ventricle is thinner walled
than the left and does not extend to the apex of the heart. In cross-section it
appears crescent-shaped and partly encircles the more cylindrical left ventricle.
Blood leaves the right ventricle via the pulmonary trunk (guarded by three semi-
lunar valves). This divides into two pulmonary arteries which convey blood to the
lungs.

Oxygenated blood returns to the left atrium via pulmonary veins which form a
single trunk to enter the left atrium as a common pulmonary vein. The left AV
orifice is guarded by an AV valve consisting of three cusps (2 large, 1 small)
attached by chordae tendinae to papillary muscles on the walls of the ventricle.
(note in mammals this is a bi-cuspid, or mitral, valve). Contraction of the left
ventricle expels blood into the aortic arch past 3 semi-lunar valves.

~ 68 ~
a

Figure 13. Avian Heart. a) Longitudinal Section Through the Heart; b) Transverse
Section Through the Heart

~ 69 ~
Arterial System
The aorta is derived from the right fourth aortic (systemic) arch (left in mammals)
and thus the ascending aorta curves to the right. The first branches are the
coronary arteries which supply the heart itself, followed by the left and right
brachiocephalic arteries. Each of these then gives rise to a:

Subclavian artery, which branches into:

o Brachial artery (to wing), which branches into:
 Radial artery
 Ulnar artery, which branches into:
Digital artery and metacarpal artery

o Pectoral artery, which branches into:

 Cranial pectoral artery
 Caudal pectoral artery

Common carotid, which branches into:

o Vertebral artery (to base of skull in canal of transverse foramina of
the cervical vertebrae)
o Carotid artery (in ventral groove of cervical vertebrae); branches into:
 Internal carotid artery (to brain)
 External carotid artery (to neck, lower jaw, orbit and much of
the cranial integument)

As the dorsal aorta passes caudally, it gives off several small paired inter-costal and
lumbar arteries to the body wall.

The alimentary tract is supplied by the following arteries which are all branches of the
aorta:

Coeliac artery: branches to proventriculus, gizzard, liver, spleen, pancreas and

duodenum, ileum and caeca

Cranial mesenteric artery: supplies most of the small intestine including the
duodenum and caeca

Caudal mesenteric artery: to ileum and colon; anastamoses with the cranial
mesenteric artery at the caeca

The kidneys are supplied by cranial, middle and caudal renal arteries, the latter two
arising from the ischiatic artery, which is also a branch of the aorta.

The Gonads are supplied by the the testicular/ovarian arteries which arise from the
cranial renal artery.

The oviduct is supplied by four arteries: cranial, middle and caudal oviductal arteries
and a vaginal oviductal artery.

~ 70 ~
There is no common iliac artery in birds, and the internal and external iliac arteries
arise separately from the aorta.

External iliac artery: arises from the aorta before passing through the kidney
and into the leg where it then branches to form the circumflex artery, medial
femoral artery and pubic artery

Ischiatic artery: is the major artery of the leg; also branches to form the middle
and caudal renal arteries.

Internal iliac artery: arises at the end of the aorta and branches into the
pudendal artery and lateral caudal artery

Median Caudal artery: is the termination of the aorta

~ 71 ~
 Carotid

Brachial

Pectoral

Subclavian

Dorsal
Aorta Coeliac

Cranial
Renal Mesenteric

External
Iliac

Ischiatic

Internal
Iliac

Caudal
Median
Mesenteric
Caudal

Figure 14. Arterial System.

~ 72 ~
Venous System
The main venous drainage of the brain is via cerebral sinuses to the vertebral veins
and cervical sinuses, while blood from the rest of the head flows into the left and right
jugular veins; note that the right jugular vein is considerably larger than the left
jugular vein and provides a convenient site for venipuncture. Some species of small
birds lack a left jugular vein altogether! At the cranial end of the neck there is a large
anastomosis between the left and right jugular veins. At the base of the neck there is
another large anastomosis between the jugular veins and the vertebral sinus.

Cranial vena cavae: two cranial vena cavae are formed from the jugular veins
and the:
o brachial vein: from the wings; also a useful site for venipuncture
o pectoral vein: from the pectoral muscles
o internal thoracic vein: from the cranial body wall

Caudal vena cava: the caudal vena cava receives the left and right hepatic
veins from the liver

Hepatic portal veins: there are two hepatic portal veins, connected by a large
superficial vein on the dorsal surface of the liver:

o Left hepatic portal vein: drains the proventriculus and gizzard

o Right hepatic portal vein: drains most of the alimentary system
(proventriculus, gizzard, duodenum, small intestine, caeca) pancreas
and spleen. This is a functional homologue of the mammalian portal
vein.

Testicular and ovarian veins: drain into the caudal vena cava

Common iliac veins: large veins which enter the caudal vena cava after
draining:

o kidney (cranial and caudal renal vein)

o cranial part of the oviduct
o leg (external iliac veins)

~ 73 ~
Figure 15. Venous System.

Renal Portal System

Venous blood from most of the caudal part of the body (tail, legs, hindgut) enters the
renal portal system. A renal portal system is a fundamental feature of all vertebrates,
except mammals where it is lost early in embryonic life.

The renal portal veins, from the common iliac vein, form a venous ring around both
kidneys, joined cranially by an internal vertebral venous sinus and caudally by the
caudal mesenteric vein. The smaller portal branches become interlobular veins
forming a capillary network around the renal lobules. A muscular valve is situated
within the lumen of the common iliac vein which can direct portal flow away from the
kidney tissues and directly to the caudal vena cava when it is open, or into the renal
portal system when it is closed. Blood can also by-pass the kidney in the caudal renal
portal area by entering the caudal mesenteric vein, or in the cranial renal portal area
through the vertebral renal sinuses.
~ 74 ~
The large caudal mesenteric vein drains the hindgut and also forms a direct
connection between the renal and hepatic portal systems. This connection allows
blood to flow in either direction, switching between the liver and the kidney. The
significance of this is unknown.

Figure 16. Renal Arteries and Veins.

Collateral Channels
A general feature of the avian circulatory system is the number of well developed
collateral channels which can compensate for a blockage in a vessel. In an
experiment, 75% of chickens survived for over two weeks following the ligation of
both carotid arteries and both vertebral arteries, with no apparent brain damage (no

~ 75 ~
jokes re bird brains and how could you tell if a chicken had brain damage anyway...!).

Blood
Avian blood has similar functions to mammalian blood but has some differences in
composition. The bone marrow is the major site of erythrocyte and leukocyte
formation. The spleen is of minor importance. Blood accounts for about 10% of body
weight (7-8% in mammals).

Erythrocytes
Oval in shape and nucleated. Life span 28-45 days.

Thrombocytes
Similar to erythrocytes but smaller (about half the size) and variable in shape.
Densely staining with a rounded nucleus. Fragile and often breakdown during slide
preparation.

Leukocytes
Birds possess the same functional types of leukocytes as mammals, but with some
minor morphological differences.
Heterophils: second most common leukocyte after lymphocytes These are
equivalent in function to the mammalian neutrophil. Rounded, with a
bilobed nucleus and many acidophilic (pink) rod-shaped bodies

Eosinophils: similar to heterophils, with many strongly acidophilic (red)

spherical bodies

Basophils: similar in appearance to heterophils, with a large rounded

nucleus and large basophilic (blue) granules

Lymphocytes: have a large nucleus with varying amounts of cytoplasm –

very similar in appearance to mammalian lymphocytes. The most prevalent
leukocyte

Monocytes: large cells with relatively more cytoplasm than lymphocytes.

Similar appearance to those in mammals.

Proportions of Avian Leukocytes

Heterophils 13 - 26%
Eosinophils 1.5 - 3%
Basophils 2.5 - 4%
Lymphocytes 57 - 76%
Monocytes 6 - 11%

~ 76 ~
Lymphatic System

Introduction
Like mammals, the avian lymphatic system has two basic functions:
return of extravascular fluids to the blood via lymphatic vessels
reaction of lymphatic tissues to foreign antigens via the instigation of cell-
mediated or humoral immune responses

Lymphatic Vessels
Generally, pairs of lymphatic vessels closely follow the blood vessels, particularly the
veins (but also arteries in the coelom). Birds have relatively fewer lymphatic vessels
than mammals, and also have relatively fewer valves to prevent retrograde flow. The
thoracic ducts are the largest lymphatic vessels in the fowl but are usually under 1
mm diameter. All lymphatic vessels finally flow into the cranial vena cavae.

In the goose (and a few other species), there is a pair of contractile lymph hearts in
the pelvic region adjacent to the aorta. These are also present in the chick embryo
but disappear with maturity.

Lymph Nodes and Nodules

Lymph nodes are absent in chickens, but in aquatic species (including duck and
goose), there are two pairs:
cervicothoracic: near the thyroid gland at the junction of the jugular and
vertebral veins
lumbar: alongside the aorta near the kidneys

Solitary lymphatic nodules occur diffusely in the parenchymatous organs (pancreas,

liver, lungs, kidneys, etc.) of most birds. Aggregated lymphatic nodules, consisting of
diffuse masses, with germinal centres, are numerous in the walls of the digestive
tract. The most prominent are the caecal tonsils at the ileo-caecal junction, but
aggregations are also found around the choanal and infundibular openings, and the
ileum (Peyer's patches).

Thymus
The thymus consists of a string of three to eight pale pink, flattened lobes on either
side of the neck next to the jugular veins. Caudally, the lobes are closely associated
with the thyroid and parathyroid. In the fowl, the thymus reaches its maximum size at
sexual maturity and then decreases in size, but in wild birds, there may be a regrowth
after the breeding season. The thymus is responsible for T cell maturation.

Spleen
The spleen is a rounded, reddish body (2 cm diameter in the fowl) lying to the right of
the proventriculus. Histologically, it is similar to the mammalian spleen but the
distinction between red and white pulp is less obvious. The spleen produces
erythrocytes briefly in the embryo, but the white pulp produces lymphocytes
throughout life. Similarly to the mammalian spleen, it is important in antibody
production and the phagocytosis of effete erythrocytes (red pulp); however, unlike the
mammalian spleen, it is not a significant reservoir of blood.

~ 77 ~
Cloacal Bursa (Bursa of Fabricius)
The cloacal bursa is a dorso-medial diverticulum of the proctodeum which is unique
to birds. In the fowl, the maximum size (4 g) is reached at 10 weeks and at sexual
maturity it regresses (a small remnant may remain). The lymphatic tissue is divided
into a number of lobules with a dark cortex and pale medulla. The cloacal bursa is the
site of B cell development and antibody production.

Disease
Lymphoid leukosis is an important disease in poultry. Cells of the cloacal bursa are
target cells for attack by lymphoid leukosis viruses which results in the cells
becoming neoplastic. As the bird grows older and the bursa involutes, tumour cells
migrate from the bursa to other organs and at sexual maturity (16-24 weeks) the
tumour involvement is so extensive that the bird dies. Removal of the cloacal bursa
(either surgically or by hormone treatment) reduces losses to lymphoid leukosis, but
impairs the growth and quality of the birds.

Marek's disease also produces similar lymphoid tumours in visceral organs, but can
be distinguished from lymphoid leukosis by specific lesions in the autonomic and
peripheral nervous system.

Urinary System

Introduction
The avian kidney represents a transitional stage between reptilian and mammalian
kidneys, showing some features of both. An important factor is the excretion of
nitrogenous wastes as uric acid.

Kidney
The paired elongate kidneys lie in close contact with the dorsal body wall, extending
from the lungs to the synsacrum. Each kidney is divided into cranial, middle and
caudal divisions by grooves of the external iliac and ischial arteries. In some species,
the caudal divisions may be fused. The surface of the kidney is covered with rounded
projections, 1-2 mm in diameter, which are external expressions of the lobules, the
functional units of the kidney. All lobules drain to the ureter and there is no renal
pelvis.

Renal Lobule
The renal lobules form the basic units of the kidney. Histologically, they appear as
pear-shaped units of tissue, between the interlobular veins of the renal portal system
and the collecting tubules. They have a large cortical component and a small
medullary component.

The collecting tubules converge to form a conical bundle of tubules in the medullary
region of the lobule. The conical bundles of collecting tubules from several adjacent
lobules converge into a single cone-shaped assembly which drains into a secondary
branch of the ureter and represents the medullary region of a whole renal lobe. Clear
demarcation between medulla and cortex is not obvious due to the varying depths of
lobules and lobes.

~ 78 ~
Nephrons
The avian kidney has two types of nephron:

Cortical nephron: the most common of the two types of nephron, found in the
cortical region of the lobules. It is reptilian in form with a long proximal
convoluted tubule, a short intermediate segment and a distal convoluted
tubule

Medullary nephron: this second type of nephron is more mammalian in form

with proximal and distal convoluted tubules, as in the cortical nephron, but with
the intermediate segment expanded into a medullary loop, which is
functionally similar to the mammalian loop of Henle, and which often descends
into the medullary region of the lobule

Both types of nephron start with a renal corpuscle consisting of a glomerular capsule
(Bowman's) deeply indented by the glomerulus. Birds have a similar juxtaglomerular
apparatus to mammals. The intralobular arteries give rise to afferent arterioles which
form the glomeruli of the renal corpuscles.

Ureter
The ureter originates at the cranial division of the kidney and passes caudally on the
kidney's ventral surface, receiving many primary branches. Each primary branch
drains 5-6 secondary branches from the medullary components of the renal lobes.
The ureter opens into the urodeum of the cloaca. A bladder is absent in all birds.

Excretion
Birds have less concentrated urine than mammals but still produce urine which is
hypertonic to blood plasma (2 to 4 times more concentrated). The glomerular filtration
rate is lower in birds than in mammals, due to the smaller size and simplicity of the
glomeruli; however, this is compensated by:
a larger number of glomeruli
waste excretion largely dependent on tubular secretion (supplied by the
massive renal portal blood supply)
excretion of uric acid (as in reptiles) which is synthesised in the liver and is
excreted, partly by glomerular filtration, but mainly by tubular secretion. Uric
acid can form colloidal solutions which allows transport through the tubules
without precipitation. Urine in the ureter is viscous, probably resulting from the
secretion of mucous in the tubules which lubricates the ureter and assists in
the passage of excreta

The excretion of nitrogenous wastes as uric acid is probably less of an adaptation for
water conservation than a means of avoiding toxicity for the developing embryo in a
cleidoic (closed box) egg.

~ 79 ~
Figure 17. Renal Lobule.

Reproductive System
Male
The male reproductive system consists of paired testes, epididymides, deferent ducts
and a phallus. The testes are relatively large and are positioned adjacent to the
cranial kidneys. The epididymis does not have a distinct head, body and tail, as in
mammals, but rather appears as a slight bulge on the testis. Sperm pass from the
epididymis into the tightly coiled deferent duct which follows the course of the ureter
to terminate at a small papilla on the lateral wall of the urodeum. There are no
accessory sex glands and seminal fluid is produced by the testis, epididymis and
deferent duct.

The phallus is protrusible from the cloaca in some species (e.g. ducks and geese)
while in others it exists as a tubercle on the ventral wall of the vent. During
insemination, sperm passes from the papillae of the deferent ducts into a groove on
the phallus, the vent is everted and the phallus is pressed against the mucosa of the
female‟s cloaca.

~ 80 ~
Female
The female reproductive system consists of the left ovary, and the left oviduct – the
right ovary and oviduct regress during fetal development but the remnants of the right
oviduct may be found on the right side of the cloaca. The ovary contains follicles that
produce large, yolk-filled oocytes. No corpus luteum develops from the ruptured
follicle since there is no pregnancy to maintain as there is in mammals.

Fertilisation takes place in the oviduct and depending on the species, sperm may be
stored by the oviduct for several weeks. In the adult chicken, the oviduct is
approximately 60 cm long! The oviduct can be divided into 5 regions, each of which
has a dinstinct function:

infundibulum: applies the chalaziferous layer which is a thin layer of very

dense albumen immediately around the yolk. In the chicken, the transit time is
approximately 15 min

magnum: highly coiled. Adds the relatively thick layer of less dense albumen.
Transit time 3 hrs

isthmus: adds more albumen and the internal and external shell membranes.
Transit time 1 hr

uterus: also called the shell gland. Deposits the shell and cuticle. Transit time
20 hrs

vagina: short tube through which the egg passes when layed

~ 81 ~
Figure 18. Reproductive Tract of Female Chicken. From www.ag.ansc.purdue.edu

~ 82 ~
Nervous System

The avian spinal cord is the same length as the neural canal (no cauda equina)
and spinal nerves pass laterally to the intervertebral foramina. The spinal cord has
brachial and lumbosacral enlargements associated with nerve plexuses. The
brachial plexus is larger in flying birds, while in flightless species, the lumbosacral
plexus is larger. The lumbosacral enlargement contains a structure known as a
rhomboidal sinus which contains a glycogen-rich gelatinous body, the function of
which is unknown – not to be mistaken for a lesion at necropsy!

The meninges in all non-mammalian tetrapods consists of the dura mater and the
meninx - the latter splits to form the arachnoid and pia mater in mammals.
Cerebrospinal fluid (0.5 ml) can be obtained at the foramen magnum in the adult
fowl but copious haemorrhage is likely! The arrangement of grey and white matter
in the spinal cord is similar to that of mammals, but the ventral and lateral columns
of white matter are comparatively large.

Spinal Nerves
As with mammals the spinal nerves are numbered according to their associated
vertebra. The first spinal nerve emerges from the foramen magnum. The spinal
nerves form several important plexuses which are clinically important for the
diagnosis of the neural form of Marek's disease:
Brachial plexus: SN 13-16; supplies the main nerves of the wing
Lumbar plexus: SN 23-25/26; largely within the kidney; supplies the body
wall and cranial muscles of the thigh (femoral and obturator nerves)
Ischiatic (sacral) plexus: SN 25-30; within the kidney; supplies the hind limb
(ischiatic nerve)
Pudendal plexus: SN 30-32; supplies the oviduct, cloaca and tail.

Autonomic Nervous System

The parasympathetic system has two divisions:
Cranial: cranial nerves III, VII, IX and X
Sacral: spinal nerves 30-33; form the pudendal nerve which has a large
cloacal ganglion on the coprodeum

The sympathetic system consists of a chain of vertebral ganglia with a smaller

series of prevertebral ganglia associated with the aorta. The cranial end of the
chain consists of the very large cranial cervical ganglion associated with cranial
nerves IX and X.

Brain
The avian brain is relatively small and, in some species, is not much larger than
the eye. The avian and mammalian hind- and mid- brains show many homologies,
but the evolution of the forebrain has differed considerably.

Medulla oblongata: the ventral fissure is conspicuous but many features

which are prominent in mammals, such as the pons, are not

Cerebellum: is large and well developed. The cerebellum is particularly

~ 83 ~
 important for the regulation of posture and movement

Midbrain: the paired ventrolateral optic lobes (= mammalian rostral colliculi)

are particularly large in birds, reflecting the importance of sight

Diencephalon: the pineal gland lies dorsally between the cerebral

hemispheres and the cerebellum. It contains some cells which seem to be
rudimentary photoreceptors but most are secretory. It is, however, strongly
influenced by light, via both the eyes and the brain, and may control
reproductive function

Cerebral hemispheres: the rostral olfactory bulb is small, indicating the

limited importance of smell to birds (except in birds which scavenge). The
cerebral hemispheres, separated by a median fissure, are almost smooth,
lacking the sulci and gyri seen in mammals. They are covered by a very thin
layer of cortex quite undeveloped compared with that of mammals. The
enlargement of the avian cerebrum relative to that of reptiles has resulted
from the enormous development of the corpus striatum (basal ganglia)
which is involved in the integration of information

Cranial Nerves
The avian cranial nerves are similar in layout and function to those of mammals.
The caudal nerves IX, X and XII have numerous anastomoses and mixed fibres.

~ 84 ~
Figure 19. Brain of the Chicken.

~ 85 ~
Special Sense Organs
Eye:
Vision is very important in birds, and the weight of the eyes may equal, or exceed,
that of the brain. The eyes may be placed laterally, providing a wide field of vision
(300°) but little binocular vision (24°) (e.g. pigeon, fowl); or frontally, giving a
smaller field of vision (150°), but greater binocular vision (70°) (e.g. owl, hawk).

The shape of the eye varies from:

Flat: most diurnal birds with narrow heads (fowl)
Globular: smaller birds and diurnal birds of prey (sparrow, hawk)
Tubular: nocturnal birds of prey (owl)

As in mammals, the wall of the eyeball consists of:

Cornea and sclera forming the outer fibrous coat
Choroid, ciliary body and iris forming the middle vascular tunic
Retina forming the inner nervous layer

Adjacent to the cornea, the sclera is modified into a ring of small overlapping
bones, the scleral ossicles; elsewhere, the sclera is strengthened by a continuous
layer of hyaline cartilage.

The pupil is generally unresponsive to light.

The avian lens is softer than in mammals.
The retina is thick and, in contrast with mammals, is avascular (mammalian retina
is vascularised). At the level of the optic disc is an outgrowth of the retina called
the pecten; it appears as a black, pleated ridge which is rich in blood vessels and
so is thought to provide nutrition to the retina. The retina contains rods, cones,
bipolar cells and ganglion cells, with a particularly high concentration of cones
localised to one area for maximum optical resolution.

The nictitating membrane is usually transparent and may cover the eye during
flight. In diving species, it forms an additional refracting lens. Eye movements are
generally very limited since the eyeball largely fills the orbit. This is compensated
for by the mobility of the head and neck. Unlike mammals, the movement of each
eye is independent!

~ 86 ~
Figure 20. Eye of the Chicken.

Ear:
There is no pinna, but the external orifice is covered by small feathers called ear
coverts. The external ear is primarily the external acoustic meatus which is short
and straight. The middle ear is similar in structure to that of mammals and birds
posses a tympanic membrane. The inner ear is also similar to mammals except
that the cochlear duct is shorter and straighter.

~ 87 ~
 C
o

Figure 21. Ear of the Chicken.

Endocrine Organs
Avian endocrine organs are generally similar to those found in mammals:

Hypophysis (pituitary)
The avian hypophysis consists of an adenohypophysis and neurohypophysis as in
mammals.

Thyroid
The thyroids are paired, dark red bodies lying at the base of the neck, medial to
the jugular vein. As in all vertebrates, they consist of follicles containing colloid
(thyroglobulin) lined by epithelial cells.

They produce thyroxine (T4) and triiodothryronine (T3) which are important for:
General metabolism; response to temperature changes
Body growth; development of reproductive organs; egg-laying
Control of moulting

~ 88 ~
Parathyroid
The parathyroids consist of two pairs of small, yellowish bodies, caudal to the
thyroids. They may be fused together and associated with the thyroid and/or
carotid body. They produce parathyroid hormone (PTH) plays a key role in
regulating blood calcium levels, especially during egg-laying.

Ultimobranchial Glands
The ultimobranchial glands are small, rounded pink bodies lying just caudal to the
parathyroids. They produce calcitonin which also plays an important role in
regulating blood calcium levels.

Adrenal Glands
The paired adrenal glands lie cranial to the kidneys and dorsal to the gonads.
Cells of the medulla and cortex are intermingled. As in mammals, cells of the
medulla secrete adrenaline and noradrenaline, while cells in the cortex produce
cortisol and aldosterone.

Pancreatic Islets
Unlike the mammalian pancreas, there are two types of islets:
Dark islets: composed of alpha cells that secrete glucagon
Light islets: composed of beta cells that secrete insulin

~ 89 ~
Figure 21. Endocrine Organs of the Chicken.

~ 90 ~