Introduction
The eutherian mammalian zona pellucida (ZP) is composed
of three sulphated glycoproteins: ZPA (or ZP2), ZPB (or ZP1)
and ZPC (or ZP3), each of which is encoded by a gene that is
highly conserved among species (Harris et al., 1994).
Although the protein components of the zona pellucida
glycoproteins are highly homologous among species,
interspecific differences in the sugar components of the N-
and O-linked oligosaccharides are evident, and these may
relate to species specificity of sperm-zona pellucida binding
where this occurs. The importance of the oligosaccharides for
sperm binding is well established and supported by the
finding that addition of various saccharides and lectins to the
in vitro fertilization environment can prevent sperm-zona
pellucida binding and fertilization (Shalgi et al., 1986;
Cornwall et al., 1991; Mori et al., 1993; Tulsiani et al., 1997).
In mice, the O-linked oligosaccharides of ZPC facilitate
primary sperm binding (Florman and Wassarman, 1985),
either through a terminal α-galactose (α-Gal) (Bleil and
Wassarman, 1988) or N-acetylglucosamine (GlcNAc) (Miller
et al., 1992). In pigs, an N-linked oligosaccharide attached to
Asn220 of ZPB in a ZPB–ZPC heterocomplex appears to be
the primary binding site for spermatozoa (Kudo et al., 1998;
Yurewicz et al., 1998) while, in humans, sperm–zona
pellucida binding may be mediated through a selectin-like
ligand on the zona pellucida which recognizes fucose (Fuc)-
rich oligosaccharides (Patankar et al., 1993; Clark et al., 1996;
Oehninger et al., 1998). Removal of terminal sialic acid
residues (desialylation) from both the human zona pellucida
and sperm surface results in an increase in sperm binding
(Lassalle and Testart, 1994; Banjeree and Chowdhury, 1997;
Ozgur et al., 1998).
Lectin histochemistry has been used to identify the
saccharides and their distribution within eutherian zonae
pellucidae (Nicolson et al., 1975; Skutelsky et al., 1994).
Observational studies on the zonae pellucidae of rats (Shalgi
et al., 1991), humans (Bar-Shira Maymon et al., 1994) and river
buffalo (Parillo et al., 1998) have demonstrated interspecific
variation in the intensity of lectin-binding patterns and the
distribution of various saccharide components.
The zonae pellucidae of marsupials appear to be thinner
and more readily solubilized by proteases than those of
eutherians (Bedford, 1991, 1996; Bedford and Breed, 1994).
The three genes encoding the zona pellucida proteins in
Journal of Reproduction and Fertility (2000) 119, 111–120
Interspecific variation of zona pellucida glycoconjugates in several
species of marsupial
J. A. Chapman
1
, O. W. Wiebkin
2
and W. G. Breed
1
*
Departments of
1
Anatomical Sciences and
2
Medicine, University of Adelaide, South Australia 5005, Australia
The zona pellucida glycoconjugate content of several marsupial species was
investigated using differential lectin histochemistry. Ovaries from fat-tailed dunnarts, a
southern brown bandicoot, grey short-tailed opossums, brushtail possums, ringtail
possums, koalas and eastern grey kangaroos were fixed, embedded in paraffin wax,
sectioned and stained with ten fluorescein isothiocyanate-conjugated lectins. Sections
were also incubated with either neuraminidase or saponified, respectively, before
incubation with the lectins to identify saccharide residues masked by sialic acids or O-
acetyl groups on sialic acids. The zonae pellucidae surrounding the oocytes of the
marsupials demonstrated interspecific variation in glycoconjugate content, with
mannose-containing glycoconjugates exhibiting the greatest variation. Some of the
zona pellucida glycoconjugates of all species, except those of the opossums, were
masked by sialic acid with an increase in fluorescence with lectins from Arachis
hypogea (PNA), and Glycine max (SBA), after desialylation. The disaccharide β-
galactose(1-4)N-acetyl-D-glucosamine appeared to be conformationally masked by O-
acetyl groups of sialic acids in the zonae pellucidae of all species, with an increase in
fluorescence with the lectin from Erythrina cristagalli (ECA), after saponification. Similar
intensity and localization of β-(1-4)-N-acetyl-D-glucosamine, as shown by staining of
the lectin from Triticum vulgaris (WGA), to the inner and outer regions of the zona
pellucida, were found to those reported in eutherian species. WGA fluorescence became
uniform throughout the zonae pellucidae after saponification, indicating differential O-
acetylation of sialic acids on the internal compartment of the zonae pellucidae.
© 2000 Journals of Reproduction and Fertility Ltd
0022–4251/2000
*Correspondence.
Revised manuscript received 19 November 1999.