Biochemical Pharmacology, Vol. 35, No. 24. pp. 4537-4542, 1986. 0006-2952/86 $3.00 + 0.00
Printed in Great Britain. © Pergamon Journals Ltd.
PROTECTION AGAINST PARAQUAT-INDUCED INJURY BY
EXOGENOUS GSH IN PULMONARY ALVEOLAR TYPE II
CELLS
TORY M. HAGEN,* LOU ANN BROWNt and DEAN P. JONES*~
Departments of *Biochemistry and +Pediatrics, Emory University School of Medicine, Atlanta, GA
30322, U.S.A.
(Received 27 March 1986; accepted 9 June 1986)
Abstract--Exogenous GSH provided rat alveolar type II cells with significant protection against injury
induced by paraquat. This protection was also observed in cells treated with acivicin to inhibit GSH
degradation and buthionine sulfoximine to inhibit GSH synthesis. Exogenous GSH was transported into
cells by a Na+-dependent system. Addition of inhibitors of this transport system, 7-glutamyl-glutamate
and probenecid, prevented the protection against injury afforded by GSH. Thus, the results indicate
that alveolar type II cells can supplement endogenous synthesis of GSH with uptake of exogenous GSH
to protect against paraquat-induced injury.
Oxidative injury in the lung occurs from inhaled air
pollutants, oxygen therapy, antitumor compounds,
and other redox cycling agents such as paraquat
[1-3]. Considerable effort has been made to under-
stand and minimize oxidative injury in lung.
However, because of the large number of different
cell types, understanding the critical susceptibility of
the individual cell types has been difficult. Over the
past several years, preparations of some of these cell
types have become available. We have focused on
the alveolar type II cells because they function in the
production and release of surfactant into the alveolar
space and also are progenitors of type I cells which
form most of the epithelial surface of the alveolus
[4, 5]. While type II cells are susceptible to oxidative
injury, they are more resistant than are type I cells,
but the reasons for this apparent resistance are as
yet unclear [6].
Glutathione (GSH) constitutes the major source
of low molecular weight thiol in mammalian tissue
and functions in detoxification of electrophilic com-
pounds and protection against oxidative injury.
Numerous reports show that GSH depletion occurs
prior to cell injury for several types of toxic sub-
stances [7-9]. Much of the effort has focused on
hepatic injury, and there is now substantial under-
standing of the toxicological consequences involving
thiol oxidation, such as inhibition of Ca 2+ ATPases
[10], disruption of cytoskeletal elements [11], and
alteration of metabolic regulation [12].
Molddus and co-workers [13] found that isolated
perfused lung can synthesize GSH from the precursor
amino acids and that the lung can also use exogenous
GSH for this purpose. However, inhibitor studies
indicated that the utilization of exogenous GSH was
largely due to extracellular breakdown and sub-
sequent resynthesis.
We have found recently that renal epithelial cells
and small intestinal epithelial cells contain a sodium-
Author to whom all correspondence should be sent.
dependent uptake system for intact GSH [14, 15]. In
the small intestinal cells, exogenous GSH provides
effective protection against oxidative injury due to
either t-butylhydroperoxide or menadione [15].
In the present study, we investigated whether
intact GSH can provide protection in type II cells
against the strong oxidant, paraquat, and, if so, the
mechanism by which GSH may offer this protection.
The results show that intact exogenous GSH was
taken up by isolated type II cells and that it provided
substantial protection against paraquat-induced
injury.
MATERIALS AND METHODS
GSH, Percoll, paraquat, (1,1-dimethyl-4,4'-bypy-
ridinium dichloride), HEPES (N-2-hydroxyethyl-
piperazine-N'-2-ethanesulfonic acid), probenecid, ),-
L-glutamyl-L-glutamate and heparin were purchased
from Sigma. [Glycine-2-3H]GSH (specific activity
l.lCi/mmole) was purchased from New England
Nuclear. Penicillin G potassium was obtained from
the Emory University Hospital Pharmacy. Oph-
thalmic acid was purchased from Bachem Inc., Torr-
ence, CA. Acivicin [L-(olS,5s)-cr-amino-3-chloro-4,5-
dihydro-5-isoxazoleacetic acid; AT-125] was a gift
from Dr. J. P. McGovren (Upjohn Co., Kalamazoo,
MI). Buthionine-SR-sulfoximine was purchased
from the Chemical Dynamics Corp.
Alveolar type II cells were obtained from male
Sprague-Dawley rats (King outbred albino), weigh-
ing approximately 200-250 g, that were kept in air-
filtered chambers to reduce the risk of lung infec-
tions. Type II cells were isolated by trypsinization
and purified by discontinuous albumin density gradi-
ent centrifugation as previously described [16]. When
examined by polychrome staining [17], 80-85% of
the cells contained lamellar bodies, indicating that
the preparation is largely type II cells. Primary cul-
tures established with this preparation yield approxi-
mately 95% type II cells which produce surfactant
4537