AVIAN-The Avıan Spleen Anatomy, Physiology and Diagnostics
AVIAN-The Avıan Spleen Anatomy, Physiology and Diagnostics
AVIAN-The Avıan Spleen Anatomy, Physiology and Diagnostics
9 September 2000
I
n birds, the thymus and bursa of Fabricius serve as primary lymphoid organs.
■ The avian spleen does not The bone marrow has been described as both a primary and a secondary
function as a blood reservoir. lymphoid organ.1,2 Secondary lymphoid tissue includes the spleen, all mu-
cosa-associated lymphoid tissue, and periarteriolar lymphoid aggregates. The
■ The spleen is primarily a avian spleen is considered the largest secondary lymphoid organ. In most birds,
defensive organ and is frequently lymph nodes are absent or rudimentary (e.g., the small cervicothoracic nodes in
involved in systemic disease. 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
■ The spleen is an excellent splenic histology and immune function of the psittacine spleen have been pub-
cytologic sampling site for lished. Chicken and turkey spleens have been the most extensively studied.
detection of chlamydial There are published reviews of the splenic anatomy of ducks, brown-headed
inclusions. 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 Small Animal/Exotics
Size and Shape branches arising from the gastric and hepatic arteries.1 It
The spleen is oval in psittacines and galliformes; bean drains into the larger of the two hepatic portal veins en-
shaped in pigeons 6; elongated, tubular, or comma tering the liver.3 The vascular tree within the spleen be-
shaped in passerines; and long and narrow in gulls. In gins with a trabecular artery that gives rise to smaller
proportion to body weight, the spleen is smaller in central arteries and arterioles; these arterioles still possess
birds than in mammals.3 The diameter of a normal a single muscular layer. Several penicilliform capillaries
chicken spleen is approximately one quarter the length branch from the central arterioles; these capillaries lack a
of the proventriculus. The size of the psittacine spleen muscular layer and are surrounded by the ellipsoid (i.e.,
varies widely, ranging from 1 to 2 mm in budgerigars the Schweigger-Seidel sheath). Blood flows from the
and 3 to 4 mm lovebirds and cockatiels to 7 to 8 mm penicilliform capillaries into the interstices of the red
in Amazon parrots. Because the psittacine spleen is pulp, and venous capillaries receive blood from the red
ovoid, these sizes should be used only as an estimate to pulp. Venous capillaries fuse to become larger trabecular
help determine whether splenomegaly is present. The veins that eventually leave the spleen.1
color varies from pink to red–brown in normal birds.7 Red and white pulps constitute about 80% of the
Seasonal variation in spleen size associated with migra- chicken spleen. The spleen is roughly divided into
tion occurs in some passerines and columbiformes.3 equal amounts of red and white pulp, although the his-
tologic division between the two is not as distinct as in
MICROANATOMY mammals.3,4 The red pulp occupies less of the spleen in
Among bird species, the gallinaceous spleen has been chickens than in mammals, accounting for approxi-
the best studied. Other avian species, including exotic mately 40% to 45% of the chicken spleen compared
birds, have been less well investigated. In many mam- with 76% to 79% of the human spleen.3 The red pulp
malian species, the spleen is surrounded by a muscular is composed of blood-filled sinusoids and diffusely scat-
capsule and partially divided into sections by trabecu- tered lymphoid and nonlymphoid cells (e.g., macro-
lae. The avian spleen has a thin connective tissue cap- phages, granulocytes, plasma cells). The marginal zone
sule that contains only rare smooth muscle fibers.3,4 in the mammalian spleen consists of layers of reticulum
True trabeculae do not occur. Unlike the mammalian and lies between the red and white pulps, which is
spleen, the avian spleen is considered to have little ca- where blood gains much of its access to pulp tissue. No
pacity for blood storage and is unlikely to contract to histologically distinct marginal zone exists in the chick-
release blood components into the circulation in re- en spleen.3,8
sponse to blood loss.2,3 The white pulp is a denser lymphoid tissue and is
The main blood supply to the chicken spleen comes closely associated with the vascular tree (Figure 2). The
from the cranial and caudal splenic arteries, with smaller central arteries and arterioles are enclosed by the periar-
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.
Your Animal
Health Collection
Isn’t Complete
Without
CE CREDIT FROM MICHIGAN STATE UNIVERSITY Volume 21 Number 9 September 2000
504
HOTLINE HEROES
been reported in psittacines affected with type 2 aden- at the NAPCC
New
ovirus.13 506
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Radiography is useful in evaluating the coelomic or-
518
gans. Survey whole-body radiographs are indicated in Get a Handle
on HIRING
birds with persistent lethargy, weight loss, leukocytosis, 526
and elevated liver enzymes or plasma bile acids. The The Golden Years—
A Roundtable on
SENIOR CARE
spleen should be evaluated as part of a systemic radio- Check out TechMart.
See page 541.
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.
sectioned before being placed in formalin. The spleen tion of avian dendritic cells in the spleen using a monoclonal
should be sliced like a loaf of bread (i.e., the cut sec- antibody specific for chicken follicular dendritic cells. Anat
Rec 249(1):81–85, 1997.
tions are not actually separated) in 2-mm sections to 11. Payne LV, Powell PC: The lymphoid system, in Freeman
best expose it to the fixative. Samples of splenic tissue MB (ed): Physiology and Biochemistry of the Domestic Fowl,
should be collected for microbiologic culture and virus vol 5. London, Academic Press, 1984, pp 298–304.
isolation before immersing in formalin. The spleen is 12. Schmidt RE: Immune system, in Altman RB, Clubb SL,
an excellent cytologic sampling site for detection of Dorrestein GM, et al (eds): Avian Medicine and Surgery.
Philadelphia, WB Saunders Co, 1997, pp 645–652.
chlamydial inclusions. Excessive amounts of iron accu- 13. Gomez-Villamandos JC, Bautista MJ, Carrasco L, et al:
mulation with erythrophagocytosis may be associated Electron microscopic evidence for infection of splenic dendrit-
with hemolytic anemias. The presence of hemosiderin ic cells by adenovirus in psittacine birds. Res Virol 146(6):
in the spleen is common in any chronic disease as well 389–395, 1995.
14. Walsh MT: Radiology, in Harrison GJ, Harrison LR (eds):
as with iron storage disease.a Such etiologic agents as
Clinical Avian Medicine and Surgery. Philadelphia, WB
bacteria and hemoparasites may be observed.23 Saunders Co, 1986, pp 201–233.
15. Orosz SE, Toal RL: Tomographic anatomy of the golden ea-
ACKNOWLEDGMENT gle (Aquila chrysaetos). J Zoo Wildl Med 23(1):39–46, 1992.
The author extends deep appreciation to David Phalen, 16 Romagnano A, Shiroma JT, Heard DJ, et al: Magnetic reso-
nance imaging of the brain and coelomic cavity of the do-
DVM, PhD, Diplomate American Board of Veterinary
mestic pigeon (Columba livia domestica). Vet Radiol Ultra-
Practitioners (Avian Practice), Texas A&M University, sound 37(6):431–440, 1996.
College Station, for reviewing an early draft of this 17. Krautwald-Junghanns ME, Enders F: Ultrasonography, in
manuscript. Altman RB, Clubb SL, Dorrestein GM, et al (eds): Avian
Medicine and Surgery. Philadelphia, WB Saunders Co, 1997,
REFERENCES pp 200–209.
1. Jeurissen SH, Vervelde L, Janse EM: Structure and function 18. Williams J: Diagnostic imaging. Vet Clin North Am Food
of lymphoid tissues of the chicken. Poultry Sci Rev 5:183– Anim Pract 14(3):421–435, 1998.
207, 1994. 19. Harrison GJ: Endoscopy, in Harrison GJ, Harrison LR
2. Pope CR: Lymphoid system, in Riddell C (ed): Avian His- (eds): Clinical Avian Medicine and Surgery. Philadelphia, WB
topathology. Tallahassee, FL, American Association of Avian Saunders Co, 1986, p 242.
Pathologists, 1996, pp 18–44. 20. Taylor M: Biopsy techniques in avian medicine. Proc Annu
3. John JL: The avian spleen: A neglected organ. Q Rev Biol Conf Assoc Vet:275–280, 1995.
69(3):327–351, 1994. 21. McKibben JS, Harrison GJ: Clinical anatomy with emphasis
4. Nasu T, Shimizu K, Nakai M: Morphological study of the on the Amazon parrot, in Harrison GJ, Harrison LR (eds):
dove spleen. Poultry Sci 71:1527–1530, 1992. Clinical Avian Medicine and Surgery. Philadelphia, WB
5. Glick B, Sato K: Accessory spleens in the chicken. Poultry Saunders Co, 1986, pp 31–66.
Sci 43:1610–1611, 1964. 22. Campbell TW: Cytology, in Harrison GJ, Harrison LR
6. Krautwald-Junghanns ME: Avian radiology, in Rosskopf (eds): Clinical Avian Medicine and Surgery. Philadelphia, WB
WJ, Woerpel RW (eds): Diseases of Cage and Aviary Birds. Saunders Co, 1986, p 269.
Baltimore, William & Wilkins, 1996, pp 630–663. 23. Campbell TW: Avian Hematology and Cytology, ed 2. Ames,
7. Lowenstine LJ: Necropsy procedures, in Harrison GJ, Harri- Iowa State University Press, 1995.
son LR (eds): Clinical Avian Medicine and Surgery. Philadel-
phia, WB Saunders Co, 1986, p 307.
8. Oláh I, Glick B: Splenic white pulp and associated vascular ABOUT THE AUTHOR
channels in chicken spleen. Am J Anat 165:445–480, 1982. Dr. Powers is an exotics and general veterinary practition-
9. Eerola E, Veromaa T, Toivanen P: Special features in the
structural organization of the avian lymphoid system, in er at Timberlyne Animal Clinic, Chapel Hill, North Caroli-
Toivanen A, Toivanen P (eds): Avian Immunology: Basis and na. She is a Diplomate of the American Board of Veteri-
Practice, vol 1. Boca Raton, FL, CRC Press, 1987, pp 15–19. nary Practitioners (Avian Practice).
10. Gallego M, del Cacho E, Lopez-Bernad F, et al: Identifica-