Professional Documents
Culture Documents
Cestosis y Cestodos PDF
Cestosis y Cestodos PDF
Cestosis y Cestodos PDF
00
CHLAMYDIAL INFECTIONS IN
SMALL RUMINANTS
Jerome C. Nietfeld, DVM, PhD
dense than elementary bodies. Elementary bodies attach to cells and are
endocytosed into cytoplasmic, membrane-bound vacuoles where they
inhibit fusion of the vacuoles with lysosomes, thus evading host cell
defenses. la, 37, 46 After 6 to 8 hours, the elementary bodies transform into
reticulate bodies, which divide multiple times by binary fission for
approximately 24 hours and then begin to transform into elementary
bodies. The result is an intracytoplasmic inclusion filled with elementary
and reticulate bodies as well as forms that are intermediate between the
two. Forty eight to 72 hours after infection, progeny are released by lysis
of the host cell or by fusion of the inclusion with the host cell mem-
brane.lO, 37, 46
All chlamydiae share lipopolysaccharide antigens that are serologi-
cally and structurally related to lipopolysaccharide antigens of rough
mutants of gram-negative bacteria of the family Enterobacteriaceae. 46 The
chlamydial lipopolysaccharides also possess a chlamydia-specific anti-
gen that is the basis for the complement fixation test used for serologic
diagnosis of chlamydial infections. I, 46
Until 1999, the order Chlamydiales contained one family, the Chlamyd-
iaceae, with one genus, Chlamydia, that contained four species, trachomatis,
pneumoniae, psittaci, and pecorum. IO C. trachomatis consisted of strains that
primarily infect humans, but also included strains isolated from mice,
hamsters, and swine. C. pneumoniae also consisted of strains that primar-
ily infected humans, but included isolates from koalas and a horse. C.
pecorum strains originated from ruminants, pigs, and koalas, and were
not known to infect humans. C. psittaci was the most diverse species
and included strains with reservoirs in a wide variety of birds and
mammals, and which were capable of being transmitted to and causing
disease in humans. Because of increasing evidence of great genetic
diversity, the genus Chlamydia was reorganized with the addition of one
new genus and five new species. lO Strains of C. trachoma tis retained the
genus name Chlamydia and the other three species were assigned to the
new genus Chlamydophila (Chla my do' philo a).l0 Human biovars of
0 0 0
Transmission
Placentas and fetal fluids from affected animals are massively in-
fected with C. abortus. Uterine secretions of ewes contain chlamydiae
from approximately 1 day before abortion to 2 to 3 weeks after abortion,1,
29, 35 and secretions from goats can contain chlamydiae from as early as
9 days before to 2 weeks after abortion. 32 During typical spring weather
when lambing usually occurs, elementary bodies can remain infectious
in the environment for several days, and if the temperature is freezing
or near freezing, for months. 1 Most animals become infected by ingestion
of organisms in contaminated feed or water, licking animals contami-
nated with placental fluids or tissues, or inhalation of aerosols created
in the contaminated environments. 19, 55 If infected 5 to 6 weeks before
parturition, sheep and goats can develop clinical disease during the
current pregnancy; however, if infected later in gestation, many animals
apparently develop latent infections and are normal until the next birth-
ing season when they experience reproductive failure due to C.
abortus. 54,55 Lambs infected in the perinatal period with C. abortus are
clinically normal until their first pregnancy, when some of them
abort. 54, 55
Initial attempts at venereal transmission by mixing C. abortus with
semen and by inoculation of C. abortus into the vagina after natural
breeding resulted in typical signs of enzootic abortion in only a minority
of cases. 4,55 It was concluded that venereal transmission was possible
but probably not common because of the management techniques used
in raising sheep; however, more recent experiments in sheep indicate
that venereal transmission may be more important than previously
thought. Intravaginal infection with C. abortus 6 weeks before breeding
or at 60 days of gestation can result in chlamydiaI placentitis and
birth of small, weak lambs or abortion. 28 After experimental and natural
abortion due to C. abortus, affected ewes conceive and lamb normally,
but they shed C. abortus from their reproductive tracts for at least 2.5 to
3 years. 27, 29, 31 Organisms can be detected for 3 to 4 days before and
after ovulation but not at any other time, including after subsequent
lambings. 29, 31 In addition, some virgin ewe lambs born into an infected
flock shed C. abortus from their reproductive tract. 25 Because rams can
become infected and shed C. abortus in their semen after breeding an
infected ewe,26 it is possible that clinically normal ewes with persistent
reproductive tract infection are a source of transmission within and
between flocks.
Chlamydophila abortus has not been found to be transmitted in milk
or colostrum from sheep,18, 47, 48,50 but it is said to be transmitted in goat's
milk. 23 C. abortus is occasionally isolated from the feces of ruminants,
raising the possibility of intestinal infection and fecal-oral transmis-
CHLAMYDIAL INFECTIONS IN SMALL RUMINANTS 305
sion. 34, 45 Most fecal isolates are C. pecorum,12, 34 however, which makes
the importance of intestinal infection by C. abortus unknown.
Pathogenesis
that suppresses multiplication and induces latency. During the last tri-
mester of pregnancy, C. abortus is reactivated and multiplies in the
placenta, resulting in restimulation of the immune system. After abor-
tion, persistent infection of the reproductive tract causes persistent stim-
ulation of the immune system, resistance to clinical disease, and mainte-
nance of high antibody titers, but not elimination of infection.
In vitro studies may provide clues as to the mechanisms involved
in establishing latent infections. Under a variety of conditions, C. tracho-
matis, C. psittaci, and C. abortus can establish inapparent, persistent
infections in cell culture. 6, 9, 33 Persistently infected cell cultures have been
maintained for more than 1 year, and when the conditions used to
induce persistence were removed, there was resumption of chlamydial
growth that resulted in inclusion formation and celllysis. 6 Maintenance
of C. abortus in persistently infected McCoy cells for 6 months did not
reduce virulence for pregnant sheep.33 Interferon gamma (IFN-')') from
activated T cells restricts chlamydial growth in macrophages, fibroblasts,
and epithelial cells, and low levels of INF-')' added to cell cultures
infected with chlamydiae, including C. abortus, cause persistent infec-
tion. 6,9 Removal of the INF-')' results in resumption of chlamydial multi-
plication. Chlamydial infections induce INF-')' production, and depletion
of INF-')' in mice results in increased in vivo multiplication of C. tracho-
matis and C. abortus. 9
If the immune system of sheep can suppress chlamydial multiplica-
tion, something occurs during pregnancy that allows reactivation and
multiplication of C. abortus. 17 The factors that allow reactivation are
currently unknown. 9, 17 It is known that pregnancy results in immuno-
suppression of the dam and that sera from pregnant humans, dogs, and
mice depresses several lymphocyte functions. 9 Hormonal changes during
pregnancy also could directly stimulate or inhibit growth of C. abortus.
The finding that sheep persistently infected with C. abortus excrete the
organism from their reproductive tracts only during estrus29,31 seems to
indicate that hormones influence in vivo multiplication of C. abortus.
Diagnosis
into PCR methodology and verified as reliable, PCR will likely become
the method of choice for fast and accurate diagnosis.
The complement fixation (CF) test is the most widely used serologic
test, but it has some limitations. 34 The CF test detects antibodies to
antigens common to all chlamydiae and cannot differentiate between
titers to C. abortus, C. pecorum, or C. psittaci, which occasionally infects
ruminants. Antibody titers to C. abortus usually remain low or negative
until the termination of pregnancy when there is a marked rise in titer
that peaks in 14 to 21 days, at which time the CF titer of most animals
that have aborted is at least 1 : 80.40 The test is best used with paired
samples taken at the time of abortion and 2 to 3 weeks later, and when
used for diagnosis on a flock rather than on an individual basis. 34
The CF test cannot reliably identify infected rams and lambs. 34 Other
techniques such as immunofluorescence, microimmunofluorescence, and
ELISA have been evaluated for serologic diagnosis. These tests are
generally a little more sensitive and specific than the CF test, but they
still have some of the same problems. 34 A study that compared five
methods of detecting antibodies against C. abortus concluded that none
of the tests were entirely satisfactory.20 All tests could identify infected
flocks, but they were not capable of reliably identifying infected sheep
on an individual basis. Another problem is that in the United States,
standardized reagents for serologic tests are not available commercially
or through government laboratories. Recently, a commercial ELISA kit
(PANCLABORT; YabA Ltd, Bush Loan, Penicuik, Scotland) said to be
specific for antibodies to C. abortus has been marketed in Europe, but it
is not available in the United States.
tive to individual animal treatment. 23,40 Oral tylosin (200 mg/head/ day)
has also been used successfully.40 Oxytetracycline and tylosin inhibit
growth of chlamydiae and help limit additional placental damage, but
they do not completely eliminate the infection or reduce the severity of
existing placental damage, so additional abortions and stillbirths are
possible.
After abortion, affected animals persistently shed C. abortus from
their reproductive tract during estrus and are sources of infection for
other flock members, subsequent new additions, and breeding male
animals. 29,31 If not culled, it is best to segregate previously affected
animals and to use separate rams for the infected and clean groups, or
place the ram with the infected animals after the clean group has been
bred. 1 After breeding, it is best to keep clean and infected animals
separated through gestation and parturition. 1
Vaccination with killed, whole-cell bacterins before breeding sig-
nificantly reduces the incidence of abortion, but it does not prevent all
abortions, eliminate the disease, or reduce shedding at lambing. 1,34 Thus,
the cycle of infection is maintained and vaccination of infected herds
needs to be continued indefinitely. A temperature-sensitive, avirulent C.
abortus vaccine that gives strong, long-lasting protection and significantly
reduces shedding is available in the United Kingdom and a few Euro-
pean countries, but not the United States. 1, 23 The vaccine is licensed for
sheep but not goats and, because it is live, precautions should be taken
to prevent human exposure. 1, 23
Zoonotic Potential
20% or less DNA homology with the three species of the genus Chla-
mydia, and they were renamed Chlamydia pecorum in 1992 and reclassified
as Chlamydophila pecorum in 1999.11,12 The C. pecorum strains are a diverse
group, but they share at least two common antigens that are recognized
by species-specific monoclonal antibodies. 34 Strains of C. pecorum are
widely distributed in ruminants and have been isolated from a wide
variety of tissues and associated with a wide variety of diseases. Most
ruminant isolates of C. pecorum, however, are from the intestinal tract of
healthy animals, are noninvasive, and are associated with clinically
inapparent intestinal infections or localized infections of mucosal sur-
faces. 34 A few strains have been shown to be invasive and to cause
systemic infections after oral inoculation. 45 Strains of C. pecorum are
recognized as important causes of keratoconjunctivitis in sheep and
goats and polyarthritis in sheep. They also have been reported to cause
pneumonia in sheep and goats, but their role as primary respiratory
pathogens is far from certain.
Keratoconju nctivitis
Clinical Signs
Initially, there are hyperemia of the vessels of the conjunctiva and
sclera, serous lachrymation, and blepharospasm. 14, 15,52 Many cases spon-
taneously regress at this point. 15, 42 If the disease progresses, hyperplastic
lymphoid follicles develop in the conjunctiva, and inflammation spreads
from the sclera onto the cornea, resulting in neovascularization of the
comea. 14, 15, 52 The exudate becomes purulent and small corneal erosions
may develop, but corneal ulceration is uncommon. 14, 15 Usually the dis-
ease is more severe in adult sheep than in lambs. 15 In the United States,
concurrent polyarthritis can occur in 10% to 85% of lambs with chlamyd-
ial keratoconjunctivitis,14 but arthritis is not a problem in the United
Kingdom or Europe. 49
Diagnosis
Specific diagnosis can be made by identification of chlamydiae by
fluorescent antibody staining of exfoliated cells in conjunctival scrapings,
or by isolation of C. pecorum in cell culture or embryonated eggs. 15, 24, 42
CHLAMYDIAL INFECTIONS IN SMALL RUMINANTS 311
Treatment
Polyarthritis
Diagnosis
Treatment
If treated early, chlamydiaI arthritis responds well to intramuscular
oxytetracycline (250 mg for 2 days), tylosin (200 mg for 2 days), and
penicillin G (300,000 units once).45 Long-lasting oxytetracycline should
also produce a good response. Oral medication with 150 to 200 mg
tetracycline/head/ day helps reduce the incidence and severity of arthri-
tis. As the disease becomes chronic and joint damage becomes more
severe, the response to treatment decreases.
Zoonotic Potential
Chlamydophila pecorum has not been reported to cause illness in
humans.
CONCLUSIONS
References
1. Aitken ID: Chlamydial abortion. In Martin WB, Aitken ID (eds): Diseases of Sheep, ed
3. Oxford, Blackwell Science, 2000, pp 81-86
2. Amin JD, Wilsmore AJ: Detection of Chlamydia psittaci (ovis) antigen in tissue sections
and McCoy cells using streptavidin-biotin and the IMAGEN™ staining method. British
Veterinary Journal 150:555-560, 1994
3. Amin JD, Wilsmore AJ: Studies on the early phase of the pathogenesis of ovine
enzootic abortion in the non-pregnant ewe. British Veterinary JoumaI151:141-155, 1995
4. Appleyard WT, Aitken ID, Anderson IE: Attempted venereal transmission of Chlamydia
psittaci in sheep. Vet Rec 116:535-538, 1985
5. Appleyard WT, Aitken ID, Anderson I: Outbreak of chlamydial abortion in goats. Vet
Rec 113:63, 1983
6. Beatty WL, Morrison RP, Byrne GI: Persistent chlamydiae: From cell culture to a
paradigm for chlamydial pathogenesis. Microbiol Rev 58:686-699, 1994
7. Bowen RA, Spears P, Storz J, et al: Mechanisms of infertility in genital tract infections
due to Chlamydia psittaci through contaminated semen. J Infect Dis 138:95-98, 1978
8. Buxtom D, Barlow RM, Finlayson J, et al: Observations on the pathogenesis of Chla-
mydia psittaci infection of pregnant sheep. J Comp Pathol102:221-237, 1990
CHLAMYDIAL INFECTIONS IN SMALL RUMINANTS 313
9. Entrican G, Brown J, Graham S: Cytokines and the protective host immune response
to Chlamydia psittaci. Comp Immun Microbiol Infect Dis 21:15-26, 1998
10. Everett KDE, Bush RM, Andersen AA: Emended description of the order Chlamydiales,
proposal of Parachlamydiaceae fam. nov. and Simkaniaceae fam. nov., each containing
one monotypic genus, revised taxonomy of the family Chlamydiaceae, including a new
genus and five new species, and standards for the identification of organisms. Int J
Syst Bacteriol 49:415-440, 1999
11. Fukushi H, Hirai K: Chlamydia pecorum-the fourth species of genus Chlamydia. Micro-
bioI Immunol 37:515-522, 1993
12. Fukushi H, Hirai K: Proposal of Chlamydia pecorum sp. nov. for Chlamydia strains
derived from ruminants. Int J Syst BacterioI42:306-308, 1992
13. Gunson DE, Acland HM, Gillette DM, et al: Abortion and stillbirth associated with
Chlamydia psittaci var ovis in dairy goats with high titers to Toxoplasma gondii. J Am Vet
Med Assoc 183:1447-1450, 1983
14. Hopkins JB, Stephenson EH, Storz J, et al: Conjunctivitis associated with chlamydial
polyarthritis in lambs. J Am Vet Med Assoc 163:1157-1160, 1973
15. Hosie BD: Ocular diseases. In Martin WB, Aitken ID (eds): Diseases of Sheep, ed 3.
Oxford, Blackwell Science, 2000, pp 301-305
16. Johnson LW: Llama reproduction. Vet Clin North Am Food Anim Pract 5:159-182,1989
17. Jones GE: Chlamydia psittaci: Prevailing problems in pathogenesis. British Veterinary
Journal 151:115-118, 1995
18. Jones GE, Anderson IE: Chlamydia psittaci excretion in ovine milk tested. Vet Rec
124:562, 1989
19. Jones GE, Anderson IE: Chlamydia psittaci: Is tonsillar tissue the portal of entry in ovine
enzootic abortion? Res Vet Sci 44:260-261, 1988
20. Jones GE, Low JC, Machell J, et al: Comparison of five tests for the detection of
antibodies against chlamydial (enzootic) abortion of ewes. Vet Rec 141:164-168, 1997
21. Jorgensen DM: Gestational psittacosis in a Montana sheep rancher. Emerg Infect Dis
3:191-194, 1997
22. Kirkbride CA: Diagnoses in 1,784 ovine abortions and stillbirths. J Vet Diagn Invest
5:398-402, 1993
23. Matthews J: Abortion. In Diseases of the Goat. Oxford, Blackwell Science, 1999, pp
22-36
24. Matthews J: Eye disease. In Diseases of the Goat. Oxford, Blackwell Science, 1999,
pp 280-284
25. Morgan KL, Wills JM, Howard P, et al: Isolation of Chlamydia psittaci from the genital
tract of lambs: A possible link with enzootic abortion of ewes. Vet Rec 123:399-400,1988
26. Papp JR, Shewen PE: Chlamydia psittaci infection in sheep: A paradigm for human
reproductive tract infection. J Reprod Immunol 34:185-202, 1997
27. Papp JR, Shewen PE: Localization of chronic Chlamydia psittaci infection in the repro-
ductive tract of sheep. J Infect Dis 174:1296-1302, 1996
28. Papp JR, Shewen PE: Pregnancy failure following vaginal infection of sheep with
Chlamydia psittaci prior to breeding. Infect Immun 64:1116-1125, 1996
29. Papp JR, Shewen PE, Gartley CJ: Abortion and subsequent excretion of chlamydiae
from the reproductive tract of sheep during estrus. Infect Immun 62:3786-3792, 1994
30. Papp JR, Shewen PE, Gartley CJ: Chlamydia psittaci infection and associated infertility
in sheep. Can J Vet Res 57:185-189, 1993
31. Papp JR, Shewen PE, Thorn CE, et al: Immunocytologic detection of Chlamydia psittaci
from cervical and vaginal samples of chronically infected ewes. Can J Vet Res 62:72-
74, 1998
32. Rodolakis A, Boullet C, Souriau A: Chlamydia psittaci experimental abortion in goats.
Am J Vet Res 45:2086-2089, 1984
33. Rodolakis A, Bernard F, Souriau A, et al: Relationship between virulence of Chlamydia
psittaci strains and establishment of persistent infection of McCoy cells. Vet Microbiol
19:65-73, 1989
34. Rodolakis A, Salinas J, Papp J: Recent advances in ovine chlamydial abortion. Vet Res
29:275-288, 1998
35. Sanderson TP, Andersen AA: Evaluation of an enzyme immunoassay for detection of
314 NIETFELD
Chlamydia psittaci in vaginal secretions, placentas, and fetal tissues from aborting ewes.
J Vet Diagn Invest 1:309-315, 1989
36. Sanderson TP, Andersen AA: Evaluation of a commercial solid-phase enzyme immuno-
assay for the detection of ovine Chlamydia psittaci. J Vet Diagn Invest 4:192-193, 1992
37. Schachter J: Chlamydia. In Gorbach SL, Bartlett JG, Blacklow NR (eds): Infectious
Diseases. Philadelphia, WB Saunders, 1992, pp 1633-1641
38. Sharma KN, Mehrotra KP, Mehrotra PN: Chlamydial infections in sheep: Pneumonitis
and abortions. Indian J Comp Microbiol Immunol Infect Dis 4:145-148, 1983
39. Shupe JL, Storz J: Pathologic study of psittacosis-lymphogranuloma polyarthritis of
lambs. Am J Vet Res 25:943-951, 1964
40. Smith MC, Sherman DM: Chlamydiosis. In Goat Medicine. Philadelphia, Lea & Febiger,
1994, pp 421-422
41. Smith MC, Sherman DM: Musculoskeletal system. In Goat Medicine. Philadelphia,
Lea & Febiger, 1994, pp 63-121
42. Smith MC, Sherman DM: Ocular system. In Goat Medicine. Philadelphia, Lea &
Febiger, 1994, pp 179-192
43. Souriau A, Rokolakis A: Rapid detection of Chlamydia psittaci in vaginal swabs of
aborted ewes and goats by enzyme linked immunosorbent assay (ELISA). Vet Micro-
bioI 11:251-259, 1986
44. Storz J: Chlamydial abortions. In Kirkbride CA (ed): Laboratory Diagnosis of Livestock
Abortions, ed 3. Ames, lA, Iowa State University Press, 1990, pp 37-48
45. Storz J, Kaltenboeck B: Diversity of chlamydia-induced diseases. In Woldehiwet Z,
Ristic M (eds): Rickettsial and Chlamydial Diseases of Domestic Animals. New York,
Pergamon Press, 1993, pp 363-393
46. Storz J, Kaltenboeck B: The chlamydiales. In Woldehiwet Z, Ristic M (eds): Rickettsial
and Chlamydial Diseases of Domestic Animals. New York, Pergamon Press, 1993,
pp 27-64
47. Thomas R, Davison HC, Wilsmore AJ: Use of the IDEIA ELISA to detect Chlamydia
psittaci (avis) in material from aborted fetal membranes and milk from ewes affected
by ovine enzootic abortion. British Veterinary Journal 146:364-367, 1990
48. Venables C, Dawson M, Baskerville M: Chlamydia in ovine milk. Vet Rec 125:137, 1989
49. Watkins GH: Arthritis. In Martin WB, Aitken ID (eds): Diseases of Sheep, ed 3. Oxford,
Blackwell Science, 2000, pp 249-254
50. Wilsmore AJ: Chlamydia in ovine milk. Vet Rec 124:618-619, 1989
51. Wilsmore AJ, Davidson I: "Clearview" rapid test compared with other methods to
diagnose chlamydial infection. Vet Rec 128:503-504, 1991
52. Wilsmore AJ, Dagnall JR, Woodland RM: Experimental conjunctival infection of lambs
with a strain of Chlamydial psittaci isolated from the eyes of a sheep naturally affected
with keratoconjunctivitis. Vet Rec 127:229-231, 1990
53. Wilsmore AJ, Dawson M, Arthur MJ, et al: The use of a delayed hypersensitivity test
and long-acting oxytetracycline in a flock affected with ovine enzootic abortion. British
Veterinary Journal 142:557-561, 1986
54. Wilsmore AJ, Izzard KA, Wilsmore BC, et al: Breeding performance of sheep infected
with Chlamydia psittaci (avis) during their preceding pregnancy. Vet Rec 126:40-41, 1990
55. Wilsmore AJ, Parsons V, Dawson M: Experiments to demonstrate routes of transmis-
sion of ovine enzootic abortion. British Veterinary Jouma1140:380-391, 1984
56. Zeman DH, Kirkbride CA: Histopathology of food animal abortions. In Kirkbride CA
(ed): Laboratory Diagnosis of Livestock Abortions, ed 3. Ames, lA, Iowa State Univer-
sity Press, 1990, pp 191-201
Address reprint requests to
Jerome C. Nietfeld, DVM, PhD
Department of Diagnostic Medicine/Pathobiology
Mosier Hall
College of Veterinary Medicine
Kansas State University
Manhattan, KS 66506
e-mail: nietfeld@Vet.ksu.edu