© 2007 The Authors 241
Journal compilation © 2007 Blackwell Publishing Ltd
Parasite Immunology , 2007, 29, 241–249 DOI: 10.1111/j.1365-3024.2007.00939.x
Blackwell Publishing Ltd ORIGINAL ARTICLE Immune responses in chronic schistosomiasis
Early responses associated with chronic pathology in
murine schistosomiasis
C. B. CÊTRE-SOSSAH,
1
M. A. MONTESANO,
2
G. L. FREEMAN, Jr.,
3
M. T. WILLARD,
4
D. G. COLLEY
5
& W. E. SECOR
Division of Parasitic Diseases, Centers for Disease Control and Prevention, Public Health Service, U.S. Department of Health and Human
Services, Atlanta, GA, USA
SUMMARY
Inbred male CBA/J mice infected with Schistosoma mansoni
develop either hypersplenomegaly syndrome (HSS) or moderate
splenomegaly syndrome (MSS) by 20 weeks of infection.
Pathologically and immunologically, MSS and HSS closely
parallel the intestinal and hepatosplenic clinical forms of
schistosomiasis in humans, respectively. By 6 weeks after infec-
tion, mice that eventually will become MSS develop T cell-
stimulatory, cross-reactive idiotypes (CRI) while HSS mice
never produce CRI. Because presence of CRI is useful to predict
degree of chronic pathology, we used this measure to investigate
what other early immunological events occurred in animals
destined to develop severe morbidity. At 8 weeks of infection,
there was a strong inverse correlation between CRI and
splenomegaly, egg counts, and liver hydroxyproline. Similarly,
phorbol myristate acetate (PMA)- and ionomycin-stimulated
intracellular cytokine expression of IL-4, IL-5, and GM-CSF
in splenic CD4
+
T cells was inversely correlated with serum
CRI and directly correlated with spleen size. In contrast,
spleen cell intracellular TNF-α and peritoneal cell production
of nitric oxide demonstrated positive correlations with CRI
and inverse correlations with measures of morbidity. Surpris-
ingly, IL-10 and IFN-γ were not correlated with CRI levels.
These studies link chronic pathology to certain immunological
responses during the acute phase of schistosomiasis.
Keywords cytokines, idiotypes, mice, schistosomiasis,
splenomegaly
INTRODUCTION
Most persons with chronic Schistosoma mansoni infections
may suffer a variety of subtle morbidities, but do not develop
severe hepatosplenic disease (1). Patients with this ‘intestinal’
form of schistosomiasis experience fatigue and only occa-
sional gastrointestinal discomfort and anaemia despite the
continued presence of adult schistosome worms in the mesenteric
vasculature. In contrast, a relatively small proportion of infected
individuals progress to develop the severe hepatosplenic form
of the disease, which is characterized by hepatosplenomegaly,
periportal fibrosis, portal hypertension, ascites, esophageal
varices, collateral circulation, haematemesis, and, if untreated,
death (2). The factors that determine whether an individual
develops the intestinal or hepatosplenic form of chronic
schistosomiasis are poorly understood, but many studies have
implicated a role for immunoregulatory mechanisms in this
process (3–6). Investigation of this hypothesis has been limited
by the absence of an animal model that predictably develops
such a spectrum of disease.
Over the last several years, we have studied inbred male CBA/
J mice with chronic S. mansoni infections that develop one of
two distinct pathophysiological syndromes by 20 weeks of
infection: hypersplenomegaly syndrome (HSS) or moderate
splenomegaly syndrome (MSS) (7,8). MSS and HSS remark-
ably parallel the intestinal and hepatosplenic clinical forms,
respectively, observed in human schistosomiasis. HSS affects
approximately 20% of these mice and presents with massive
splenomegaly, ascites, thymic atrophy, severe anaemia, and
cachexia. The remaining majority of mice develop only moder-
ate splenomegaly. Histopathological features of HSS include
Correspondence: W. Evan Secor, Division of Parasitic Diseases,
Centers for Disease Control and Prevention, 4770 Buford
Highway, NE, MS-F13, Atlanta, GA 30341-3724, USA
(e-mail: was4@cdc.gov).
1
Current Address: Virology Unit, CIRAD-EMVT, Campus
International de Baillarguet, TA 30/G, 34398 Montpellier Cedex 5,
France.
2
Current Address: Division of Laboratory Science, National Center
for Environmental Health, Centers for Disease Control and
Prevention, 4770 Buford Highway, NE, Atlanta, GA 30341, USA.
3
Deceased.
4
Current Address: Winship Cancer Institute, Emory University
School of Medicine, Atlanta, GA 30322, USA.
5
Current Address: Center for Tropical and Emerging Global
Diseases, 145 Coverdell Center, University of Georgia, Athens,
GA 30602, USA.
Received: 30 November 2006
Accepted for publication: 1 December 2006