Expression of Mouse Sialic Acid on Gangliosides of a Human
Glioma Grown as a Xenograft in SCID Mice
Jeffrey A. Ecsedy, Kathryn A. Holthaus, *²Herbert C. Yohe, and Thomas N. Seyfried
Department of Biology, Boston College, Chestnut Hill, Massachusetts, *VA Medical and Regional Office Center, White River
Junction, Vermont, and ² Department of Pharmacology, Dartmouth Medical School, Hanover, New Hampshire, U.S.A.
Abstract: Ganglioside sialic acid content was examined
in the U87-MG human glioma grown as cultured cells and
as a xenograft in severe combined imunodeficiency
(SCID) mice. The cultured cells and the xenograft pos-
sessed N-glycolylneuraminic acid (NeuGc)-containing
gangliosides, despite the inability of human cells to syn-
thesize NeuGc. Human cells express only N-acetylneura-
minic acid (NeuAc)-containing gangliosides, whereas
mouse cells express both NeuAc- and NeuGc-containing
gangliosides. Small amounts of NeuGc ganglioside sialic
acid (2–3% of total ganglioside sialic acid) were detected
in the cultured cells, whereas large amounts (66% of total
ganglioside sialic acid) were detected in the xenograft.
The NeuGc in gangliosides of the cultured cells was
derived from gangliosides in the fetal bovine serum of the
culture medium, whereas that in the U87-MG xenograft
was derived from gangliosides of the SCID host. The
chromatographic distribution of U87-MG gangliosides
differed markedly between the in vitro and in vivo growth
environments. The neutral glycosphingolipids in the
U87-MG cells consisted largely of glucosylceramide,
galactosylceramide, and lactosylceramide, and their dis-
tribution also differed in the two growth environments.
Asialo-GM1 (Gg4Cer) was not present in the cultured
tumor cells but was expressed in the xenograft, suggest-
ing an origin from infiltrating cells (macrophages) from the
SCID host. The infiltration of mouse host cells and the
expression of mouse sialic acid on human tumor cell
glycoconjugates may alter the biochemical and immuno-
genic properties of xenografts. Key Words: Oligodendro-
glioma—Astrocytoma—N-Glycolylneuraminic acid—Brain
tumor—Asialo-GM1.
J. Neurochem. 73, 254 –259 (1999).
Glycosphingolipids (GSLs) are glycoconjugates
present on the outer leaflet of the cell membrane and
consist of a lipophilic ceramide conjugated to a hydro-
philic oligosaccharide chain. GSLs are classified into two
major groups: gangliosides and neutral glycosphingolip-
ids (NGSLs). The presence of sialic acid on gangliosides
distinguishes them from NGSLs. Sialic acids are nega-
tively charged derivatives of neuraminic acid, which
function in receptor mechanisms, cell– cell interactions,
and other cellular and molecular processes (Jeanloz and
Codington, 1976; Schauer, 1985, 1988; Hakomori, 1996;
Varki, 1997). Human gangliosides possess only the N-
acetylneuraminic acid (NeuAc) form of sialic acid,
whereas mouse gangliosides possess two forms of sialic
acid, NeuAc and N-glycolylneuraminic acid (NeuGc)
(Yu and Ledeen, 1970; Seyfried et al., 1978a, 1987;
Chou et al., 1998). NeuAc differs from NeuGc in having
an acetyl group instead of a glycolyl group attached to
the nitrogen on carbon number 5 (Schauer, 1985). The
absence of NeuGc-containing gangliosides in human tis-
sues arises from a partial deletion in the gene that en-
codes cytidine monophospho-N-acetylneuraminic acid
hydroxylase, which is necessary for synthesizing NeuGc
from the precursor NeuAc (Chou et al., 1998; Irie and
Suzuki, 1998).
The GSL composition of brain tumors differs mark-
edly from that of normal brain. This arises from GSL
abnormalities in proliferating tumor cells and from tu-
mor-infiltrating host cells (Seyfried et al., 1996, 1998).
The enrichment of GSLs on the cell surface and their
considerable structural diversity make them potential
targets for tumor immunotherapy and diagnosis (Irie and
Morton, 1986; Murray et al., 1994, 1996; Sung et al.,
1994; Wikstrand et al., 1994a,b; Suetake et al., 1995;
Uttenreuther Fischer et al., 1995; Nishinaka et al., 1996).
The utility of GSLs for tumor diagnosis and therapy will
depend to a large extent on their origin and cellular
location. In this regard, it is essential to understand the
influence of growth environment on tumor GSLs.
Fredman and co-workers (Mansson et al., 1986; Fred-
man, 1988) originally showed that NeuGc is the predom-
inant sialic acid present on gangliosides isolated from a
human glioma grown as a xenograft in nude mice. Ka-
Received October 1, 1998; revised manuscript received February 8,
1999; accepted March 3, 1999.
Address correspondence and reprint requests to Dr. T. N. Seyfried at
Department of Biology, Boston College, Chestnut Hill, MA 02467-
3811, U.S.A.
Abbreviations used: GalCer, galactosylceramide; GlcCer, glucosyl-
ceramide; GSL, glycosphingolipid; HPTLC, high-performance thin-
layer chromatography; LacCer, lactosylceramide; NeuAc, N-acetyl-
neuraminic acid; NeuGc, N-glycolylneuraminic acid; NGSL, neutral
glycosphingolipid; SCID, severe combined immunodeficiency.
254
Journal of Neurochemistry
Lippincott Williams & Wilkins, Inc., Philadelphia
© 1999 International Society for Neurochemistry