European Journal of Neuroscience, Vol. 11, pp. 1577–1586, 1999 © European Neuroscience Association
A novel P
0
glycoprotein transgene activates expression of
lacZ in myelin-forming Schwann cells
M. Laura Feltri,
1,2
Maurizio D’Antonio,
2
Angelo Quattrini,
1
Rita Numerato,
2
Marta Arona,
2
Stefano Previtali,
1,2
Shing-Yan Chiu,
3
Albee Messing
4
and Lawrence Wrabetz
1,2
1
Department of Neurology, San Raffaele Scientific Institute, via Olgettina 58, 20132 Milano, Italy
2
DIBIT, San Raffaele Scientific Institute, via Olgettina 58, 20132 Milano, Italy
3
Department of Physiology, Waisman Center, University of Wisconsin-Madison, Madison, WI 53706, USA
4
School of Medicine and Department of Pathobiological Sciences, Waisman Center, University of Wisconsin-Madison,
Madison, WI 53706, USA
Keywords: development, mice, peripheral nerve, transcription, Wallerian degeneration
Abstract
P
0
glycoprotein, the most abundant protein in peripheral nerve, is expressed specifically in the Schwann cell lineage. Upstream of
the rat P
0
gene 1.1 kb of DNA can activate expression of cDNAs specifically in Schwann cells in transgenic mice. However, the
expression of P
0
promoter-based transgenes has been inconsistent. As much as 9 kb of 5' flanking sequence fused to lacZ never
yielded detectable levels of β-galactosidase in multiple lines of mice. We describe transgenic mice that express lacZ in peripheral
nerve, using the complete mouse P
0
gene, including 6 kb of 5' flanking sequence, all exons and introns, and the natural
polyadenylation signal. This vector activated lacZ expression specifically in cultured Schwann cells, and myelin-forming Schwann
cells in four out of six transgenic lines. Transgene expression paralleled that of the endogenous P
0
gene, both during development
and after Wallerian degeneration. lacZ expression was lower than endogenous P
0
expression, and was not detected in neural crest
or Schwann cell precursors, where low levels of P
0
mRNA are present. However, when the same vector contained a small myc
tag instead of the 3.2-kb lacZ insert, the resulting transgenic mRNA was expressed at levels comparable to endogenous P
0
mRNA.
These data suggest that intragenic or 3' flanking sequences are necessary to generate the remarkable levels of endogenous P
0
gene expression.
Introduction
P
0
glycoprotein is an adhesion molecule that compacts the spiralling
Schwann cell (SC) plasma membrane to form myelin in the peripheral
nervous system. P
0
is indeed the most abundant protein of peripheral
myelin, where it constitutes over 50% of total protein (Greenfield
et al., 1973). P
0
is expressed by myelin-forming SC, cells that contact
a single axon and deposit a myelin sheath around it; but not by non-
myelin forming SC, cells that ensheath several axons without forming
myelin. Both types of cells derive from neural crest cells that
subsequently differentiate into SC precursors and embryonic SC
(Jessen et al., 1994; Dong et al., 1995). Basal levels of P
0
mRNA
are first seen in neural crest and remain in SC precursors and embryonic
SC (Bhattacharyya et al., 1991; Baron et al., 1994; Zhang et al., 1995;
Lee et al., 1997). The type of axon that a SC subsequently encounters
will determine its phenotype: large axons induce differentiation
into a myelin-forming SC, small axons – a non-myelin-forming SC.
Myelin-forming SC greatly upregulates the expression of P
0
glyco-
protein and other myelin-specific genes, so that after differentiation,
P
0
mRNA accounts for at least 8% of the mRNA synthesized (Stahl
et al., 1990).
SC-specific induction of P
0
expression is thought to be mediated
at the level of transcription (Lemke et al., 1988). In keeping with
this, the P
0
promoter successfully directed expression of foreign genes
Correspondence: M.L. Feltri, as above. E-mail: l.feltri@hsr.it
Received 4 August 1998, revised 8 December 1998, accepted 9 December 1998
to myelin-forming SC. In vitro, 1.1 kb of P
0
promoter drove SC-
specific expression of reporter genes (Lemke et al., 1988). Moreover,
in transgenic mice, 1.1 kb of P
0
promoter successfully activated
expression of toxins, hormones, oncogenes, transcription factors and
enzymes specifically in SC (Messing et al., 1992, 1994; Weinstein
et al., 1995; Nikitin et al., 1996). However, transgene expression
with the 1.1 kb promoter has been inconsistent. Thus, phenotypes
suggesting transgene expression have been obtained with potent
molecules (e.g. diphtheria toxin), although transgenic mRNA expres-
sion was low and was not measured as a direct proportion of
endogenous P
0
mRNA (Messing et al., 1992). Also, more consistent
expression was obtained when the transgene contained both exons
and natural introns from the heterologous gene, as for growth
hormone, SV40 T antigen, and connexin 32 (Messing et al., 1992,
1994; Bone et al., 1997). Instead, many attempts to express transgenes
containing P
0
promoter fragments fused to cDNAs or reporter genes,
even with heterologous introns (e.g. rabbit β-globin) included, have
had only limited success, sometimes requiring many independent
transgenic lines in order to obtain even low-level expression (Feltri
et al., unpublished observations; Maycox et al., 1997). For example,
P
0
promoter–lacZ transgenes have never been successfully expressed
in SC, even when 9 kb of 5' flanking sequence was included (Table 1).
Thus, we asked if the intragenic portion of the mouse P
0
gene
(Mpz) contributes to high-level P
0
mRNA expression in SC. As we
had no indication of where important regulatory elements may reside
outside of the promoter, we reconstructed the complete Mpz, including