INFECriON AND IMMUNITY, Mar. 1994, p. 1079-1088 Vol. 62, No. 3
0019-9567/94/$04.00+0
Copyright © 1994, American Society for Microbiology
Monoclonal Antibodies to Cryptococcus neoformans Capsular
Polysaccharide Modify the Course of Intravenous
Infection in Mice
SUSHOVITA MUKHERJEE,' SUNHEE LEE,2 JEAN MUKHERJEE,3 MATTHEW D. SCHARFF,3
AND ARTURO CASADEVALLl 4*
Departments of Microbiology and Immunology," Pathology, 2 and Cell Biology,3 and Division of Infectious
Diseases of the Department of Medicine,4 Albert Einstein College of Medicine, Bronx, New York 10461
Received 1 October 1993/Returned for modification 22 November 1993/Accepted 10 December 1993
Immunoglobulin Gl (IgGl) monoclonal antibodies (MAbs) to the capsular glucuronoxylomannan (GXM)
were studied for their ability to modify the course of intravenous Cryptococcus neoformans infection in mice. A/J
mice were given intraperitoneal injection of 1.0 mg of either a GXM-binding IgGl MAb (2H1 or 2DlOyl) or
the irrelevant isotype-matched control MAb 36-65 prior to intravenous infection. Parameters used to study
antibody efficacy were lung and brain tissue fungal burden, lung and brain weights, serum GXM levels, and
histopathological examination of lung, brain, heart, kidney, and spleen tissues. Mice given GXM-binding MAb
had significantly reduced lung tissue fungal burden as measured by CFU. In contrast to the reduction in lung
tissue burden, the reduction in brain tissue burden was small and did not achieve statistical significance.
Serum GXM levels were reduced in mice receiving GXM-binding MAb. Histopathological examination revealed
reduced numbers of granulomas and C. neofornans organisms in the lungs, brains, and kidneys of MAb
2H1-treated mice relative to control mice. The lungs and brains of mice receiving GXM-binding MAb weighed
significantly less than those of control animals, consistent with the reduced inflammation noted histologically.
Subendocardial inflammation and kidney cortical infarctions were present in control infected mice but not in
MAb 2H1-treated mice. Immunocytochemical staining for polysaccharide antigen revealed a marked reduction
in the amount of tissue polysaccharide in mice treated with MAb 2H1 relative to control mice. The results
support an useful role for passive antibody administration in C. neoformans infections.
Cryptococcus neoformans causes life-threatening meningo-
encephalitis in approximately 10% of patients with AIDS (77).
In the setting of AIDS, cryptococcal infections are usually
incurable because antifungal therapy does not eradicate the
infection (73) despite in vitro susceptibility of the organism to
antifungal drugs (6). This has renewed interest in the potential
of vaccines (9) and passive antibody therapy (15, 45, 58, 59, 67)
to prevent infection. Passive antibody is an attractive option
because it could enhance residual immunity in immunosup-
pressed patients (2). Precedent for the use of antibody in
therapy of human cryptococcosis exists: in 1925, Shapiro and
Neal used rabbit immune sera in an unsuccessful attempt to
treat a boy with meningitis (71); in 1959, Littman used human
gamma globulin and amphotericin B in a small number of
patients with encouraging results (48); and in 1963, Gordon
and Vedder used rabbit immune globulin and amphotericin B
to successfully treat two patients with cryptococcosis (23, 25).
Specific antibody can potentiate the effect of amphotericin B
against C. neoformans in mouse models (14, 24, 59a), suggest-
ing a rationale for combined therapy.
It is generally accepted that cellular immunity is the primary
line of defense against C. neoformans (1, 29, 31, 42, 45, 47, 54,
60) and that complement provides opsonins for phagocytosis
(1, 13, 43). In contrast, the importance of natural antibody
immunity has remained uncertain. Passive immunization with
polyclonal sera has produced evidence for (22, 28) and against
(49) a role for antibody in protection. Casting doubt on the
importance of antibody immunity are the observations that an
*
Corresponding author. Mailing address: Department of Medicine,
Albert Einstein College of Medicine, 1300 Morris Park Ave., Bronx,
NY 10461. Phone: (718) 430-4260. Fax: (718) 597-5814.
immunogenic vaccine failed to protect mice (26), B-cell defi-
cient mice are not at increased risk (55), and antibody is not an
absolute requirement for phagocytosis (1, 13) or killing of C.
neoformans (27, 52). However, the observation that antibodies
are potent opsonins (40, 41, 69) which are required for NK cell
(53, 61) and leukocyte (10, 11, 51) antifungal activity in vitro
suggests a role for antibody immunity. Other observations
supporting an important role for antibody are as follows: the
appearance of antibody in cerebrospinal fluid of rabbits is
temporally correlated with decreased colony counts in brain
tissue (36); mice succumb to infection when their serum
antibody titers decline (70); serum antibody is a good prognos-
tic indicator in humans (12); recovery from cryptococcal
meningitis is accompanied by the production of specific anti-
body in cerebrospinal fluid (44); and cryptococcosis in the
setting of hypogammaglobulinemia is occasionally reported
(30, 68). Three groups have shown that administration of
monoclonal antibody (MAb) to the capsular polysaccharide
can mediate protection in murine models of C. neoformans
infection (14-17, 57-59, 67). Antibody efficacy is dependent on
the quantity (15, 16), the isotype (58, 67), and the fine
specificity (58) of the MAb used, findings that could explain
the discordant results obtained in earlier experiments with
polyclonal sera.
In 1981, Graybill showed that the choice of animal model
was crucial for demonstrating efficacy with polyclonal sera (28).
The best animal model for identifying potential antibody
reagents against C. neoformans for human use is not known
and will not be known until human efficacy data are correlated
with data from animal studies. We have previously evaluated
the efficacy of some of our MAbs in intraperitoneal (i.p.) (57,
58) and intracerebral (i.c.) (59) murine models of C. neofor-
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