Introduction
There is increasing research interest in the role of inositol-
containing compounds in preimplantation embryo
development. Inositol is essential for continued cell
proliferation and growth of rabbit blastocysts (Fahy and
Kane, 1992) and it also stimulates hatching of hamster
blastocysts (Kane and Bavister, 1988). A major role of
inositol is as part of the phosphatidylinositol (PtdIns)
signal transduction system in which receptor-stimulated
activation of PtdIns-dependent phospholipase C (PLC) leads
to hydrolysis of phosphatidylinositol(4,5)bisphosphate
(PtdIns(4,5)P
2
) to produce the two second messengers:
inositol(1,4,5)trisphosphate (Ins(1,4,5)P
3
), which acts to
release Ca
2+
from internal Ca
2+
stores, and diacylglycerol,
which activates protein kinase C (for a review, see Berridge,
1992). Mouse (Kane et al., 1992), rabbit (Fahy and Kane,
1993) and cattle (Hynes et al., 2000) embryos, and mouse
embryonic stem (ES) cells (Duffy and Kane, 1996)
incorporate inositol into the phosphoinositides and inositol
phosphates of the PtdIns signal transduction system. In
addition, inhibitors of PLC inhibit mouse blastocyst
formation (Stachecki and Armant, 1996). Perhaps the most
important evidence for an essential role for phosphoinositide
compounds in preimplantation development is the demon-
stration that knocking out the PLC β3 gene prevents
development of mouse embryos to the blastocyst stage
(Wang et al., 1998).
Another role for inositol is in the formation of
glycosylphosphatidylinositol (GPI) lipids, which function as
anchors for the attachment of surface proteins to the plasma
membrane of various cell types (for a review, see Low, 1987).
The GPI anchors typically contain phosphatidylinositol
inserted in the cell membrane, connected to a carbohydrate
core structure consisting of glucosamine, mannose,
galactose and ethanolamine, to which the protein is
attached by a phosphodiester linkage (for reviews, see
Englund, 1993; McConville and Menon, 2000). Another
characteristic of the GPI anchors is that they may be cleaved
to release the protein by endogenous PtdIns-specific
phospholipases (Brewis et al., 1994).
Glycosylphosphatidylinositols are also involved in the
signal transduction pathways of numerous hormones and
growth factors such as insulin (Saltiel and Cuatrecasas,
1986), and growth factors and cytokines, including insulin-
like growth factor I (IGF-I), epidermal growth factor (EGF),
and transforming growth factor β1 (TGF-β1) as well as
interleukins 1 and 2 (reviewed by Jones and Varela-Nieto,
1998). Insulin and other growth factors activate PLC to
hydrolyse GPI, resulting in the release of the polar head
group, an inositol-containing oligosaccharide called an
Nature of glycosylphosphatidylinositols produced by mouse
embryonic stem cells
L. R. Quinlan and M. T. Kane*
Department of Physiology, National University of Ireland, Galway, University Road,
Galway, Ireland
Reproduction (2001) 122, 785–791
Research
Incorporation of [
3
H]inositol into mouse embryonic stem
cells of the CCE cell line leads to the labelling of the
three common phosphoinositides, phosphatidylinositol,
phosphatidylinositol 4-phosphate and phosphatidylinositol
4,5-bisphosphate, and a fourth unknown lipid (lipid X).
Incubation with [
3
H]glucosamine results in the labelling of
lipid X and at least one other lipid that co-migrates with
phosphatidylinositol (lipid Y), indicating that both of these
lipids are putative glycosylphosphatidylinositols. In this
study, the incorporation of other possible glycosylphos-
phatidylinositol precursors, ethanolamine, mannose and
galactose, into lipids X and Y was examined. Galactose was
incorporated into lipids X and Y, and ethanolamine and
mannose into lipid Y only. Inhibitors of glycosylphos-
phatidylinositol biosynthesis pathways, mannosamine and
2-fluoro-2-deoxyglucose, both significantly inhibited
ethanolamine incorporation into lipid Y. A high glucose
concentration (25 mmol l
–1
) abolished the action of both
inhibitors. Phospholipase C treatment of embryonic stem
cells that had been labelled in culture with
[
3
H]ethanolamine caused a large release of ethanolamine
label into the incubation medium and markedly decreased
the amount of ethanolamine-labelled lipid Y remaining in
the cell membranes. These effects were almost totally
abolished by incubation with mannosamine before
ethanolamine labelling. These studies strongly indicate
that lipid Y is a member of the protein anchor class of
glycosylphosphatidylinositol, whereas lipid X is a member
of the signal transduction inositol phosphoglycan class of
glycosylphosphatidylinositol.
© 2001 Society for Reproduction and Fertility
1470-1626/2001
*Correspondence
Email: michael.kane@nuigalway.ie