Vol. 22, No.
9 September 2000
CE Refereed Peer Review
FOCAL POINT
★The avian spleen differs from
the mammalian spleen in its
anatomy, function, and response
to disease; assessing the spleen
can aid in the diagnosis of
multisystemic illness.
KEY FACTS
■ The lack of lymph nodes in
most avian species increases the
relative importance of the spleen
in disease resistance.
■ In proportion to body weight, the
spleen is smaller in birds than in
mammals.
■ The avian spleen does not
function as a blood reservoir.
■ The spleen is primarily a
defensive organ and is frequently
involved in systemic disease.
■ The spleen is an excellent
cytologic sampling site for
detection of chlamydial
inclusions.
The Avian Spleen:
Anatomy, Physiology,
and Diagnostics
Timberlyne Animal Clinic, Chapel Hill, North Carolina
Lauren V. Powers, DVM
ABSTRACT: Literature reviewing the avian spleen, particularly that of psittacines, is scarce.
The anatomy, physiology, and response to disease of the avian spleen are significantly differ-
ent from those of the mammalian spleen. However, there are several methods to assess
splenic health, and results of such testing can aid in evaluating systemic disease as well as
specific diseases of the spleen. This article briefly discusses the anatomy and function of the
avian spleen and describes methods used to assess splenic health. A companion article will
discuss diseases of the avian spleen.
I
n birds, the thymus and bursa of Fabricius serve as primary lymphoid organs.
The bone marrow has been described as both a primary and a secondary
lymphoid organ.1,2 Secondary lymphoid tissue includes the spleen, all mu-
cosa-associated lymphoid tissue, and periarteriolar lymphoid aggregates. The
avian spleen is considered the largest secondary lymphoid organ. In most birds,
lymph nodes are absent or rudimentary (e.g., the small cervicothoracic nodes in
waterfowl). The lack of lymph nodes in most avian species increases the relative
importance of the spleen in disease resistance. To date, no reviews of avian
splenic histology and immune function of the psittacine spleen have been pub-
lished. Chicken and turkey spleens have been the most extensively studied.
There are published reviews of the splenic anatomy of ducks, brown-headed
cowbirds (Molothrus ater), pied flycatchers (Ficedula hypoleuca), and doves.3,4
ANATOMY
Location
The avian spleen lies dorsal to the right lobe of the liver between the proven-
triculus and ventriculus (Figure 1). At necropsy, the spleen can be isolated by ro-
tating the ventriculus to the right. The spleen is located at the angle between the
proventriculus, ventriculus, and liver. Accessory spleens are well recognized in
chickens and may hypertrophy after splenectomy.5 One or more small accessory
spleens may be cranial, adjacent, or caudal to the spleen. Accessory spleens have
not been described in psittacines.
Compendium September 2000
Figure 1—Anatomic arrangement of the avian spleen dorsal
to the proventriculus and between the proventriculus and
ventriculus in a 2-year-old cockatiel. (Photo courtesy of H. J.
Barnes, DVM, PhD, North Carolina State University,
Raleigh)
Size and Shape
The spleen is oval in psittacines and galliformes; bean
shaped in pigeons 6; elongated, tubular, or comma
shaped in passerines; and long and narrow in gulls. In
proportion to body weight, the spleen is smaller in
birds than in mammals.3 The diameter of a normal
chicken spleen is approximately one quarter the length
of the proventriculus. The size of the psittacine spleen
varies widely, ranging from 1 to 2 mm in budgerigars
and 3 to 4 mm lovebirds and cockatiels to 7 to 8 mm
in Amazon parrots. Because the psittacine spleen is
ovoid, these sizes should be used only as an estimate to
help determine whether splenomegaly is present. The
color varies from pink to red–brown in normal birds.7
Seasonal variation in spleen size associated with migra-
tion occurs in some passerines and columbiformes.3
MICROANATOMY
Among bird species, the gallinaceous spleen has been
the best studied. Other avian species, including exotic
birds, have been less well investigated. In many mam-
malian species, the spleen is surrounded by a muscular
capsule and partially divided into sections by trabecu-
lae. The avian spleen has a thin connective tissue cap-
sule that contains only rare smooth muscle fibers.3,4
True trabeculae do not occur. Unlike the mammalian
spleen, the avian spleen is considered to have little ca-
pacity for blood storage and is unlikely to contract to
release blood components into the circulation in re-
sponse to blood loss.2,3
The main blood supply to the chicken spleen comes
from the cranial and caudal splenic arteries, with smaller
TRABECULAE ■ ELLIPSOID ■ RED PULP ■ WHITE PULP
Small Animal/Exotics
Figure 2—Schematic representation of the histology of the
chicken spleen. (CA = central artery; CV = central vein; GC =
germinal center; PALS = periarterial lymphoid sheath; PC =
penicilliform capillary; PELS = periellipsoidal lymphoid
sheath; S = sinusoid; TA = trabecular artery; TC = terminal
capillary; TV = trabecular vein)
branches arising from the gastric and hepatic arteries.1 It
drains into the larger of the two hepatic portal veins en-
tering the liver.3 The vascular tree within the spleen be-
gins with a trabecular artery that gives rise to smaller
central arteries and arterioles; these arterioles still possess
a single muscular layer. Several penicilliform capillaries
branch from the central arterioles; these capillaries lack a
muscular layer and are surrounded by the ellipsoid (i.e.,
the Schweigger-Seidel sheath). Blood flows from the
penicilliform capillaries into the interstices of the red
pulp, and venous capillaries receive blood from the red
pulp. Venous capillaries fuse to become larger trabecular
veins that eventually leave the spleen.1
Red and white pulps constitute about 80% of the
chicken spleen. The spleen is roughly divided into
equal amounts of red and white pulp, although the his-
tologic division between the two is not as distinct as in
mammals.3,4 The red pulp occupies less of the spleen in
chickens than in mammals, accounting for approxi-
mately 40% to 45% of the chicken spleen compared
with 76% to 79% of the human spleen.3 The red pulp
is composed of blood-filled sinusoids and diffusely scat-
tered lymphoid and nonlymphoid cells (e.g., macro-
phages, granulocytes, plasma cells). The marginal zone
in the mammalian spleen consists of layers of reticulum
and lies between the red and white pulps, which is
where blood gains much of its access to pulp tissue. No
histologically distinct marginal zone exists in the chick-
en spleen.3,8
The white pulp is a denser lymphoid tissue and is
closely associated with the vascular tree (Figure 2). The
central arteries and arterioles are enclosed by the periar-
Small Animal/Exotics Compendium September 2000

terial lymphoid sheath. Lym- the differentiation of plasma

phoid tissue in this region is cells.11 Plasma cell proliferation
believed to be primarily com- appears to occur in the red pulp.
posed of T-dependent lym- There seems to be no correla-
phocytes (T cells). The cen- tion between the number of
tral arteries branch into the germinal centers and antibody-
penicillary arterioles, and the producing capacity.11 Although
muscular layer of these arteri- the spleen is an important
oles is replaced by the ellip- component of the lymphoid
soid. These vessels have tall en- system, splenectomy apparent-
dothelial cells, thick basement ly has no profound effect on
laminae, and an intimate asso- immunocompetence in chick-
ciation with reticular cells. The ens, possibly as a result of the
Figure 3—Histologic appearance of a normal psittacine
sheath is composed of two lay- spleen (adult macaw). The splenic capsule at the top of presence of accessory spleens
ers of cuboidal reticular cells. the photograph appears somewhat contracted (hema- and compensatory antibody
The ellipsoid is surrounded by toxylin & eosin stain; original magnification, ×18). (Pho- synthesis by the bone marrow.4
a layer of lymphoid cells called to courtesy of H. J. Barnes, DVM, PhD, North Carolina However, splenectomy in
the periellipsoidal lymphoid State University, Raleigh) young birds delays peak anti-
sheath, which is composed of body response.11
bursa-dependent lymphocytes
(B cells), plasma cells, and macrophages. Ellipsoidal cells, Granulopoiesis
the periellipsoidal lymphoid sheath, and surrounding When chickens hatch, the spleen is a granulocytopoi-
macrophages form a complex considered to be analogous etic organ but quickly transforms into a predominantly
to the marginal zone in mammalian spleens. Some lym- lymphocytic organ. The granulocytes vacate the spleen
phoid tissue surrounds venules and is bursa independent.9 at hatching and are replaced by lymphocytes.2,3 The
Germinal centers contain B cells and are often located ad- spleen is not known to be a site for embryonic lym-
jacent to central arteries within the periarterial lymphoid phopoiesis, and its major development occurs after
sheath.8 hatching.3

NONIMMUNE FUNCTIONS
IMMUNE FUNCTIONS
The spleen is a site of phagocytosis of senescent or
Antigen Processing
diseased erythrocytes by macrophages in the red pulp.12
The spleen is primarily a defensive organ and is fre-
Some erythropoiesis occurs in the chicken spleen dur-
quently involved in systemic disease. All antigen is ini-
ing early development but is believed to be confined to
tially localized in the ellipsoid, independent of the
the bone marrow in healthy mature birds.3 Extrame-
nature of the antigen. Antigenic activation of ellipsoid-
dullary hematopoiesis can occur in the psittacine spleen
associated cells triggers them to migrate to the periarte-
secondary to chronic blood loss.a
rial lymphoid sheath.2,10 Germinal centers are formed by
the agglutination of migrating ellipsoid-associated cells
KEY DIFFERENCES IN THE PSITTACINE SPLEEN
with B lymphocytes having complementary surface anti-
To date, there are no published reviews of the psittacine
body receptors. Ellipsoid-associated cells likely differen-
spleen and how it differs from the chicken model. Some
tiate into follicular dendritic cells.10,11 Dendritic cells
observed differences in the psittacine spleen include an
may prevent diffusion of virus into the host or converse-
indistinct red pulp without detectable sinusoids, scarcity
ly may facilitate viral transmission throughout the body.
of germinal centers in normal spleens, and probable dif-
Such antigens as lipopolysaccharide may also be ab-
ferences in the distribution and type of lymphoid tissue.
sorbed by red-pulp macrophages. Ellipsoid-associated
Most of the lymphoid cells are small, mature lymphocytes
cells may enter the general circulation, where they serve
that are not in obvious periellipsoidal or periarterial lym-
as antigen-presenting cells to other secondary lymphoid
phoid sheath arrangements (Figure 3). Blood flow and
tissue, via the red pulp.
function are presumed to be similar among avian species,
but the differences in structure suggest that there may
Memory and Plasma Cells also be differences in function.b Germinal centers have
The function of the germinal centers in chickens may a
Phalen D: Personal communication, Texas A&M University,
be the proliferation of memory B cells as opposed to College Station, 1999.

PERIELLIPSOIDAL AND PERIARTERIAL LYMPHOID SHEATHS ■ GERMINAL CENTERS

Compendium September 2000 Small Animal/Exotics

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CE CREDIT FROM MICHIGAN STATE UNIVERSITY Volume 21 Number 9 September 2000

Figure 4—Schematic representation of the radiographic ap-

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pearance of the avian spleen on the plain lateral view.

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been reported in psittacines affected with type 2 aden- at the NAPCC
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RADIOLOGY 508
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Radiography is useful in evaluating the coelomic or-
518
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Survey Radiographs
The spleen lies dorsal to the proventriculus on lateral
survey radiographs (Figures 4 and 5) and can be visual- The Most Widely Read
ized in approximately one third of normal large psit-
tacines using this view. The spleen cannot be visualized Journal in Its Field
in the ventrodorsal plane in normal psittacines. Spleno-
megaly is best detected using the lateral view. An enlarged ■ The technician’s right hand—the source
spleen might be seen through the hepatic tissue to the
right and slightly cranial to the ventriculus on the ven- they reach for first
trodorsal view, although splenomegaly is rarely docu- ■ Articles of interest to animal health
mented in this manner.14 Splenic neoplasia, notably in
professionals of all kinds: breeders,
budgerigars, may displace the ventriculus caudodorsally
on the lateral view.6 caretakers, trainers…
■ Medical and management articles of
Computed Tomography and
Magnetic Resonance Imaging interest to the dedicated pet owner
Computed tomography and magnetic resonance imag-
ing may be promising diagnostic tools to evaluate the
avian spleen. The spleen was identified among the SUBSCRIBE TODAY!
coelomic viscera in the golden eagle (Aquila chrysaetos) CALL 800-426-9119
using computed tomography (5-mm slices).15 Magnetic
resonance imaging can identify the spleen within the Veterinary Technician is published by
coelom. However, the spleen was difficult to identify in Veterinary Learning Systems
b
Barnes HJ: Personal communication, North Carolina State 275 Phillips Blvd.
University, Raleigh, 1999. Trenton, NJ 08618-1496
Price is in US dollars and is subject to change.

SPLENOMEGALY ■ SPLENIC NEOPLASIA

Small Animal/Exotics Compendium September 2000

all planes examined in a one of the splenic poles can

study of magnetic resonance help to differentiate the spleen
imaging techniques in pi- from a similarly colored,
geons; the authors suggested large mesenteric blood vessel
that this difficulty may be re- that is located in the same
lated to the shape of the area.19
spleen in pigeons, in which it
is a slender, oval structure.16 Biopsy Techniques
In psittacines, the spleen is Biopsy is indicated if
round and thus more easily splenomegaly is present or
identified using magnetic lesions are observed on the
resonance imaging.16 spleen during endoscopic ex-
amination. In most patients,
Ultrasonography Figure 5—Plain survey lateral radiograph of a Moluccan cock- a 5-Fr elliptic biopsy cup or
The spleen is difficult to atoo (Cacatua moluccensis). The spleen is visible dorsal to the fine-needle aspiration can be
visualize ultrasonographical- proventriculus. used to collect a sample of
ly in birds; a lateral approach splenic tissue.20 Alternatively,
is suggested. The spleen is a left lateral laparotomy inci-
slightly more echogenic than is the liver. The tissue has sion can be made; the spleen can then be accessed by
a fine, dense granularity and an even texture. An en- proceeding dorsally over the proventriculus, moving the
larged spleen is easier to visualize using a ventromedial intestines caudally, and exposing the organ.21 However,
approach. Homogenous enlargement is seen with infec- the laparoscopic approach is much more traumatic and
tion or trauma. Splenic neoplasia frequently has mixed indicated only when laparotomy is being performed for
echogenicity.17 Massive splenomegaly is uncommon in other purposes or when the spleen cannot be approached
psittacines but is easily identifiable by ultrasonography using standard endoscopic technique. If bleeding occurs
when present.a during biopsy, a small wedge of Gelfoam® (Pharmacia &
Ratites have no muscle on either side of the linea Upjohn, Kalamazoo, MI) or Surgicel™ (Ethicon, a
alba. This zone provides an adequate sonographic win- Johnson & Johnson company, Somerville, NJ) can be in-
dow for imaging the coelomic organs. The spleen is es- serted endoscopically and pressed against the biopsy site
pecially difficult to visualize ultrasonographically in until bleeding stops.
ratites if there is abundant gas in the gastrointestinal
tract or hyperinflation of the caudal coelomic air sacs.18 CYTOLOGY AND HISTOPATHOLOGY
Cytologic evaluation of the avian spleen can be per-
ENDOSCOPY formed on fine-needle aspiration or biopsy samples or at
Endoscopy is indicated for persistent unexplained necropsy. Before making impression smears, cut sections
splenomegaly detected radiographi- of the spleen should be blotted on
cally or persistent unexplained sys- filter paper or a paper towel to re-
temic disease. Psittacines should be duce blood contamination. In nor-
fasted for 3 to 6 hours and raptors mal birds, impression smears usually
for 12 to 20 hours before endoscop- reveal large amounts of peripheral
ic examination to ensure that the blood within the spleen. Many ma-
proventriculus is empty for the pro- ture lymphocytes can also be seen,
cedure. representing the abundant lymphoid
tissue present. Macrophages exhibit-
Approach ing erythrophagocytosis are com-
The spleen should be approached mon. Splenic stromal (reticular) cells
from the left abdominal air sac, which have a pale blue cytoplasm with an
can be entered from the left caudal indistinct cytoplasmic border. The
thoracic air sac or via a left caudal ab- nucleus is eccentric and round to
dominal air sac approach. The Figure 6—Endoscopic appearance of a nor- oval with a loose chromatin pat-
proventriculus is pushed ventrally mal pigeon spleen. (Photo courtesy of M. tern.22
with the tip of the endoscope to ex- Taylor, DVM, University of Guelph, On- The spleen is a dense, cellular or-
tario, Canada)
pose the spleen (Figure 6). Identifying gan and tends not to fix well unless

ECHOGENICITY ■ LAPAROSCOPIC APPROACH ■ IMPRESSION SMEARS

Compendium September 2000
sectioned before being placed in formalin. The spleen
should be sliced like a loaf of bread (i.e., the cut sec-
tions are not actually separated) in 2-mm sections to
best expose it to the fixative. Samples of splenic tissue
should be collected for microbiologic culture and virus
isolation before immersing in formalin. The spleen is
an excellent cytologic sampling site for detection of
chlamydial inclusions. Excessive amounts of iron accu-
mulation with erythrophagocytosis may be associated
with hemolytic anemias. The presence of hemosiderin
in the spleen is common in any chronic disease as well
as with iron storage disease.a Such etiologic agents as
bacteria and hemoparasites may be observed.23
ACKNOWLEDGMENT
The author extends deep appreciation to David Phalen,
DVM, PhD, Diplomate American Board of Veterinary
Practitioners (Avian Practice), Texas A&M University,
College Station, for reviewing an early draft of this
manuscript.
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3. John JL: The avian spleen: A neglected organ. Q Rev Biol
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Small Animal/Exotics
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16 Romagnano A, Shiroma JT, Heard DJ, et al: Magnetic reso-
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Iowa State University Press, 1995.
ABOUT THE AUTHOR
Dr. Powers is an exotics and general veterinary practition-
er at Timberlyne Animal Clinic, Chapel Hill, North Caroli-
na. She is a Diplomate of the American Board of Veteri-
nary Practitioners (Avian Practice).