ELSEVIER
PII S0361-9230(96)00042-2
BrainResearchBulletin, Vol. 40, No. 3, pp. 167-174, 1996
Copyright© 1996Elsevier ScienceInc.
Printedin the USA.All rightsreserved
0361-9230/96$15.00 + .00
Expression of the fll and fl2(AMOG) Subunits of the
Na, K-ATPase in Neural Tissues: Cellular and
Developmental Distribution Patterns
EMILIA LECUONA,* SONIA LUQUJN, t JULIO AVILA,* LUIS M. GARCiA-SEGURA t
AND PABLO MARTJN-VASALLO .1
*Laboratorio de Biologfa del Desarrollo, Departamento de Bioqufmica y Biologfa Molecular, Universidad de La
Laguna, 38206 La Laguna, Tenerife, Spain and tlnstituto Cajal, C.S.I.C. Dr. Arce 37, 28002 Madrid, Spain
[Received 23 May 1995; Accepted 23 January 1996]
ABSTRACT: We have used isoform-specific antisera against the
Na,K-ATPase/]1 (SpETbl) and/~2(AMOG) (SpETb2) subunit iso-
forms in order to establish their specific cellular and subcellular
localization in several developmental stages of the rat central
nervous system. Immunocytochemical preparations revealed/~1
isoform protein in most neural cells, being predominantly lo-
cated in the soma of neurons and astrocytes, with no apprecia-
ble developmental variations. In the newborn rat,/~2(AMOG) im-
munoreactivity was present in cellular processes of astroglia
and in the somas of neurons and decreasing in intensity with
maturation until adulthood, where no/~2 isoform was detected
in neurons. The differential location of these isoforms, both de-
velopmentally and at the cellular level suggest a complex reg-
ulation of their genes expression and mechanisms of subcellular
distribution, as well as functional differences.
KEY WORDS: Na,K-ATPase/~ subunit isoforms, AMOG, Adhe-
sion molecule on glia.
INTRODUCTION
The Na,K-ATPase is the plasma membrane-bound enzyme that,
utilizing ATP as an energy source, is responsible for maintaining
the K ÷ and Na ÷ concentrations, osmotic balance, and membrane
electrical potentials characteristic of most animal cells. Na,K-
ATPase is present in high concentrations in the central nervous
system (CNS), maintaining the cation gradients necessary for
neural impulse conduction in neurons and potassium buffering
and neurotransmitter uptake in glial cells. The enzyme is com-
posed of two subunits, (called a and/5). The a, ( 112 kDa), or
catalytic subunit, contains the binding sites for ATP and the ions
implicated in the pumping process and serves as the receptor for
cardiac glycosides. The/5 (45 kDa) subunit is a highly glyco-
sylated protein and has no known role in ion transport, although
the expression of both c~ and/5 subunits is required for Na,K-
ATPase activity [ 20,15 ]. Two different isoforms of the/5 subunit
(/~1 and f12) have been described in mammals [14] and a/53
isoform in Xenopus laevis [10] and Bufo marinus [11]. The/52
isoform is an adhesion molecule on glial cells (AMOG) and is
specifically involved in neuron-astrocyte adhesion [7], with no
To whom requests for reprints should be addressed.
obvious adhesion function in other cell types. No adhesion func-
tion has been reported for the/~ 1 isoform. A role of the/5 subunit
as the receptor for the newly synthesized a-subunit has been
proposed [12], and interdependency of a and/5 subunit seems
to be necessary for their transport out of the endoplasmic retic-
ulum [6].
While a and /5-subunit mRNA expression [14,23], immu-
noblotting [4], 3H-ouabain binding [3], and a subunit immu-
nolocalization in neural tissue [15,4] and cultured neural cells
[ 3 ] have been broadly described, very little is known about the
specific cellular and subcellular localization of each/5 isoform in
neural cells and their developmental patterns "in vivo". In order
to further understand the expression, localization, and functional
implications of these proteins, isoform-specific antibodies against
the human Na,K-ATPase /51 (SpETbl) and /52(AMOG)
(SpETb2) isoforms [8,9] have been used to probe inununoblots
of brain microsomes and to perform immunohistochemistry on
preparations of central nervous system from newborn, 6-day-, 10-
day-, 15-day-, and 90-day-old (adult) rats.
EXPERIMENTAL PROCEDURES
Antibodies and Reagents
Antibodies specific for truncated protein Na,K-ATPase /51
(SpETbl) and/52(AMOG) (SpETb2) subunits [8,9] have been
described. Both sera (S) were raised against the ectodomain of
the human Na,K-ATPase /51 and/52(AMOG) subunits which
had been generated as truncated proteins by pET expression vec-
tors in E. coli [ 8,9,19 ].
Western Blots
Microsomal fractions were prepared from whole brains of
newborn and adult rats. Protein concentration was determined
according to Bradford [ 1 ]. Na,K-ATPase activity was determined
as described by B. Forbush, III [5]. One Na,K-ATPase unit =
/~mol Pi h ~ per mg of protein. Protein samples, loaded so that
each lane contained between 0.15 and 0.30 Na, K-ATPase units,
were subjected to electrophoresis in 10% polyacrylamide-SDS
167