BRAIN
RESEARCH
ELSEVIER Brain Research 663 (1994) 329-334
Short communication
Anoxia induces an increase in intracellular sodium
in rat central neurons in vitro
Jonathan E. Friedman *, Gabriel G. Haddad
Department of Pediatrics, Section of Respiratory Medicine and Department of Cellular and Molecular Physiology, Yale University School of Medicine,
Fitkin Bldg. 517, 333 Cedar Str., New Haven, CT06520, USA
Accepted 16 August 1994
Abstract
Following our previous observations that anoxia induces a drop in extracellular Na ÷ in the brain slice and that removal of
extracelluar Na + prevents the anoxia-induced morphological changes in dissociated hippocampal neurons, we hypothesized that
intraceilular Na ÷ increases during anoxia in isolated neurons. Using the fluorophore Sodium Green in freshly dissociated rat
CA1 neurons, and SBFI in cultured cortical neurons, we found that 10 min of anoxia caused an increase in Na~- in both types of
cells, with a latency of about 2 min. In CA1 neurons, fluorescence increased by an average of 20.34% (n = 8). The mean baseline
Na~- level (determined using SBFI) was 25 + 2.5 raM, which increased to about an average of 52 + 3 mM after 3-4 rain. These
and our previous results strongly suggest that Na+-mediated events are involved in anoxia-induced nerve injury.
Keywords: Sodium Green; Cortex; Hippocampus; Volume
Oxygen deprivation has long been known to cause
irreversible damage to the mammalian brain. This
damage is believed to result primarily from a massive
influx of calcium into nerve cells, mediated in part by
the excitatory amino acid glutamate [6]. Calcium, by
virtue of its ability to activate, for example, enzymes
such as lipases, proteases and kinases has thus been
considered to cause cell damage. However, we, as well
as others, have shown in vitro that nerve damage from
anoxia still occurs in the absence of extraceliular Ca 2+,
and in the absence of any detectable rise in intra-
cellular Ca 2÷ [10,11]. Since (a) we have previously
shown that extracellular Na ÷ decreases in the brain
slice during anoxia [19], and (b) our previous data
indicate that extracellular Na ÷ is important in the
anoxia-induced depolarization [16], we entertained the
hypothesis that Na + enters into neurons during 0 2
deprivation.
In order to address this hypothesis, we have carried
out studies primarily using freshly dissociated (i.e. iso-
lated) CA1 hippocampal pyramidal neurons. This is an
optimal system for testing the direct effect of anoxia on
* Corresponding author. Fax: (1) (203) 785-6337.
0006-8993/94/$07.00 © 1994 Elsevier Science B.V. All rights reserved
SSDI 0006-8993(94)01000-5
cells, since potentially confounding factors such as
synaptic transmission and release of excitatory amino
acids are obviated. In addition, CA1 pyramidal neu-
rons are considered to be among the most sensitive
cells to anoxia [23,31]. Using this system, and the
Na÷-sensitive fluorophore Sodium Green (NaGrn), we
have monitored changes in Na~-. However, since Na-
Grn is not a ratiometric dye, we also employed another
Na+-sensitive dye, SBFI (sodium-binding benzofuran
isophthalate), in order to obtain quantitative results
[17,27].
Dissociated cell preparation and loading with Sodium
green. Cells were prepared as previously reported [11].
In brief, 21-30 day old Sprague-Dawley rats were
anesthetized by inhalation of methoxyfluorane (Pit-
mann-Moore), decapitated, the brain rapidly removed
and placed into ice-cold oxygenated dissociation buffer
(DB) (120 mM NaC1, 5 mM KC1, 1 mM CaC12, 1 mM
MgC12, 25 mM glucose and 20 mM HEPES (Sigma),
pH 7.4). The hippocampi were removed and cut into
400-500 /.~m slices, which were immediately placed
into a dissociation chamber containing 10 ml of DB +
10 mg trypsin (Sigma) for 40 min followed by the
addition of 4 mg Pronase E (Sigma) for another 20
min. The buffer was replaced with fresh DB, without