Highly Differential Expression of SN1,
a Bidirectional Glutamine
Transporter, in Astroglia and
Endothelium in the Developing
Rat Brain
JEAN-LUC BOULLAND, AMINA RAFIKI, LINE M. LEVY, JON STORM-MATHISEN,
AND FARRUKH A. CHAUDHRY*
Department of Anatomy, Institute of Basic Medical Sciences and Centre for Molecular Biology
and Neuroscience, University of Oslo, Oslo, Norway
KEY WORDS SN1; glutamine; glutamate; transporter; synaptogenesis; blood-brain
barrier
ABSTRACT The transmitters glutamate and GABA also subserve trophic action and
are required for normal development of the brain. They are formed from glutamine,
which may be synthesized in glia or extracted from the blood. In the adult, the glutamine
transporter SN1 is expressed in the astroglia. SN1 works in both directions, depending
on the concentration gradients of its substrates and cotransported ions, and is thought
to regulate extracellular glutamine and to supply the neurons with the transmitter
precursor. In this article, we have quantified the expression and studied the localization
of SN1 at different developmental stages. SN1 is expressed in astroglia throughout the
CNS from embryonic stages through adulthood. No indication of SN1 staining in neu-
ronal elements has been obtained at any stage. Quantitative immunoblotting of whole
brain extracts demonstrates increasing expression of SN1 from P0, reaching a peak at
P14, twice the adult level. A moderate and slower rise and fall of the expression levels
of SN1 occurs in the cerebellum. Strong transient SN1-like staining is also found in
Bergmann glia and vascular endothelium in the first postnatal weeks. Strong intracel-
lular staining in the same time period suggests a high rate of SN1 synthesis in the early
postnatal period. This coincides with the increasing levels of glutamate and GABA in the
CNS and with the time course of synaptogenesis. This study suggests that the expres-
sion of SN1 is highly regulated, correlating with the demand for glutamine during the
critical period of development. GLIA 41:260 –275, 2003. © 2003 Wiley-Liss, Inc.
INTRODUCTION
Neurotransmitters are endogenous regulators of
neurogenesis, neural migration, and synaptogenesis
(Spencer et al., 1998). Increased levels of the transmit-
ters and their specific receptors have been detected
especially during the critical period for synaptogenesis
in the rat brain (Kwong et al., 2000). A lack of activa-
tion or inhibition of these receptors leads to patholog-
ical development or cell death (Monti and Contestabile,
2000). Glutamate and GABA are the most abundant
excitatory and inhibitory neurotransmitters in the
brain, respectively. These transmitters are generated
from glutamine, which is generally believed to be syn-
thesized in the glia. Thus, impairment of glutamine
transport from glia to the neurons during ontogenesis
could be a limiting factor leading to reduced levels of
*Correspondence to: Farrukh A. Chaudhry, Department of Anatomy, Institute
of Basic Medical Sciences, University of Oslo, P.O. Box 1105 Blindern, N-0317
Oslo, Norway.
Received 1 July 2002; Accepted 17 October 2002
DOI 10.1002/glia.10188
GLIA 41:260 –275 (2003)
© 2003 Wiley-Liss, Inc.