activity of CD8
+
T cell clones correlates with their
levels of expression of adhesion molecules.
J. Exp. Med. 175, 895–905
22 Clyde, D.F. et al. (1973) Immunization of man
against sporozoite-induced falciparum malaria. Am.
J. Med. Sci. 266, 169–177
23 Krzych, U. et al. (1995) T lymphocytes from
volunteers immunized with irradiated Plasmodium
falciparum sporozoites recognize liver and blood
stage malaria antigens. J. Immunol. 155,
4072–4077
24 Hill, A.V. et al. (1991) Common West African HLA
antigens are associated with protection from severe
malaria. Nature 352, 595–600
25 Hill, A.V. et al. (1992) Molecular analysis of the
association of HLA-B53 and resistance to severe
malaria. Nature 360, 434–439
26 Connelly, M. et al. (1997) T-cell immunity to peptide
epitopes of liver-stage antigen 1 in an area of Papua
New Guinea in which malaria is holoendemic.
Infect. Immun. 65, 5082–5087
27 Bucci, K. et al. (2000) Influence of age and HLA
heterozygosity on IFN-γ responses to a naturally
occurring polymorphic epitope of Plasmodium
falciparum liver stage antigen-1. Clin. Exp.
Immunol. 122, 94–100
28 Luty, A.J.F. et al. (1999) Interferon-γ responses are
associated with resistance to reinfection with
Plasmodium falciparum in young African children.
J. Infect. Dis. 179, 980–988
29 Kurtis, J.D. et al. (1999) Interleukin-10 responses to
liver-stage antigen 1 predict human resistance to
Plasmodium falciparum. Infect. Immun. 67,
3424–3429
30 John, C.C. et al. (2000) Cytokine responses to
Plasmodium falciparum liver-stage antigen 1 vary
in rainy and dry seasons in highland Kenya. Infect.
Immun. 68, 5198–5204
31 Luty, A.J.F. et al. (1998) Parasite antigen-specific
interleukin-10 and antibody responses predict
accelerated parasite clearance in Plasmodium
falciparum malaria. Eur. Cytokine Netw. 9, 639–646
32 Domarle, O. et al. (1999) Factors influencing
resistance to reinfection with Plasmodium
falciparum. Am. J. Trop. Med. Hyg. 61, 926–931
33 Rousset, F. et al. (1992) Interleukin-10 is a potent
growth and differentiation factor for activated
human B lymphocytes. Proc. Natl. Acad. Sci. U. S. A.
89, 1890–1893
34 Jinquan, T. et al. (1993) Human IL-10 is a
chemoattractant for CD8
+
T lymphocytes and an
inhibitor of IL-8-induced CD4
+
T lymphocyte
migration. J. Immunol. 151, 4545–4551
35 Ockenhouse, C.F. et al. (1998) Phase I/IIa safety,
immunogenicity, and efficacy trial of NYVAC-Pf7, a
pox-vectored, multiantigen, multistage vaccine
candidate for Plasmodium falciparum malaria.
J. Infect. Dis. 177, 1664–1673
36 Heal, K.G. et al. (2000) Expression and
immunogenicity of a liver stage malaria epitope
presented as a foreign peptide on the surface of RNA-
free MS2 bacteriophage capsids. Vaccine 18, 251–258
37 Aidoo, M. et al. (1995) Identification of conserved
antigenic components for a cytotoxic T lymphocyte-
inducing vaccine against malaria. Lancet 345,
1003–1007
TRENDS in Parasitology Vol.17 No.5 May 2001
http://parasites.trends.com 1471-4922/01/$ – see front matter © 2001 Elsevier Science Ltd. All rights reserved. PII: S1471-4922(01)01895-5
223 Opinion
Leishmania parasites assemble an abundance of
distinctive glycoconjugates that are expressed on
their surface or secreted
1–3
(Fig. 1). Leishmania
produce a characteristic family of glycoconjugates
containing phosphoglycan (PG) that includes
membrane-bound lipophosphoglycan (LPG) and
proteophosphoglycan (PPG), as well as secreted PG,
PPG and acid phosphatase (sAP). In addition, the
surface of Leishmania contains an array of molecules
attached to glycosylphosphatidylinositol (GPI)
anchors (Fig. 1). These include free entities such as
glycosylinositolphospholipids (GIPLs), LPG itself and
proteins including the promastigote surface protease
(also known as leishmanolysin, PSP or gp63).
The abundance, location and uniqueness of these
Leishmania structures have led to the suggestion
that these glycoconjugates, particularly LPG, have
one or more important functions for the parasite
throughout its life cycle
2,4,5
. These include the procyclic
and metacyclic promastigotes carried by the sand fly
vector, and the amastigote stage, which resides
within the phagolysosome of macrophages (Fig. 2).
Most of the roles proposed for the glycoconjugates
have been deduced using purified molecules and
examining their effects on mouse macrophage
function and interactions with the sand fly vector.
From these in vitro experiments, there is substantial
evidence that LPG is required for survival during the
initial stage of establishment in the macrophage,
when the parasite is most vulnerable. For example,
LPG (or purified portions thereof ) has been shown to
bind macrophages and to inhibit macrophage signal
transduction and cytokine production
5,6
. However,
the distribution of PG units and GPI anchors across a
large number of different molecules (Fig. 1) raises
questions about the correct assignment of functions
for the individual glycoconjugates. Assays of LPG
function using purified LPG might mimic effects that
normally arise through other PGs or GPI-anchored
molecules. This problem is compounded by the fact
that LPG (or portions thereof ) is often tested outside
the biological milieu, at concentrations that might
exceed that attained during parasite invasion.
The use of parasite mutants provides an
opportunity to test the role of LPG in a biologically
relevant setting. Examples of these include the
Is lipophosphoglycan
a virulence factor?
A surprising diversity
between Leishmania
species
Salvatore J. Turco, Gerald F. Späth and
Stephen M. Beverley
Lipophosphoglycan is a prominent member of the phosphoglycan-containing
surface glycoconjugates of Leishmania. Genetic tests enable confirmation of its
role in parasite virulence and permit discrimination between the roles of
lipophosphoglycan and related glycoconjugates. When two different
lipophosphoglycan biosynthetic genes from Leishmania major were knocked
out, there was a clear loss of virulence in several steps of the infectious cycle
but,with Leishmania mexicana, no effect on virulence was found. This points to
an unexpected diversity in the reliance of Leishmania species on virulence
factors, a finding underscored by recent studies showing great diversity in the
host response to Leishmania species.