Biochimica et Biophysica Acta, 1145 (1993) 199-204 199
© 1993 Elsevier Science Publishers B.V. All rights reserved 0005-2736/93/$06.00
BBAMEM 75859
Effects of alkaline pH on the stimulation of glucose transport
in rat skeletal muscle
Jian-Ming Ren, Jang H. Youn, Eric A. Gulve, Erik J. Henriksen and John O. Holloszy
Department of Internal Medicine, Washington Uniuersity School of Medicine, St. Louis, MO (USA)"
(Received 7 October 1992)
Key words: Epitrochlearis muscle; Hypoxia; Insulin; Muscle contraction; 3-O-Methylglucose
Alkaline pH has been reported to cause release of Ca e+ from skeletal muscle sarcoplasmic reticulum (SR). Elevation of
sarcoplasmic Ca 2÷ concentration is thought to stimulate glucose transport in skeletal muscle. In this context, we examined the
effect of alkaline pH (extracellular pH of 8.6) on 3-O-methylglucose transport in skeletal muscle. Incubation of rat epitrochlearis
muscles at pH 8.6 for 45 min resulted in an approx. 3-fold increase in glucose transport activity, which was not affected by
reducing Ca z+ concentration in the incubation medium and essentially completely blocked by 25/zM dantrolene, an inhibitor of
SR Ca 2÷ release. In addition to stimulating glucose transport by itself, alkaline pH may partially inhibit the stimulation of sugar
transport by insulin hypoxia and contractions, as the combined effect of alkaline pH and the maximal effect of insulin,
contractions, or hypoxia on glucose transport are not different from the maximal effects of insulin, hypoxia, or contractions
alone. The maximal effects of insulin and contractions, and of insulin and hypoxia, on glucose transport are normally additive in
muscle. Alkaline pH completely prevented this additivity. In summary, our results show that alkaline pH stimulates glucose
transport activity in skeletal muscle and provide evidence suggesting that this effect is mediated by Ca 2÷. They further show that
alkaline pH blocks the additivity of the maximal effects of insulin and contractions or hypoxia suggesting that alkaline pH may
partially inhibit the stimulation of glucose transport by insulin, contraction and hypoxia.
Introduction
In skeletal muscle, glucose transport can be acti-
vated by insulin, contractile activity, and hypoxia. The
maximal effects on glucose transport activity of insulin
and contractions [1-4], and of insulin and hypoxia [5]
are additive, while the maximal effects of contractions
and hypoxia are not additive [5]. These findings have
been taken as evidence that contractions and hypoxia
stimulate glucose transport by the same pathway, and
that insulin and contractile activity/hypoxia stimulate
glucose transport by different pathways [1,2,5].
Evidence has accumulated suggesting that an in-
crease in cytoplasmic Ca 2+ concentration stimulates
glucose transport in skeletal muscle and that this is the
mechanism by which muscle contractions and hypoxia
induce an increase in glucose transport activity [5-14].
Alkaline pH triggers Ca 2+ release from skeletal muscle
sarcoplasmic reticulum [15], and stimulates glucose
transport in adipocytes and some cell lines [16-18].
Correspondence to: J.-M. Ren, Department of Internal Medicine,
Campus Box 8113, Washington University School of Medicine, 4566
Scott Avenue, St. Louis, MO 63110, USA.
However, we have not been able to find any informa-
tion regarding the effect of alkaline pH on glucose
transport in muscle. In this context, the present study
was undertaken to determine whether or not alkaline
pH stimulates glucose transport in skeletal muscle.
Our results show that alkaline pH does increase
glucose transport activity in skeletal muscle and sug-
gest that this effect is mediated by an increase in
cytoplasmic Ca 2+.
Methods
Animals and muscle preparation
Male Wistar rats weighing 100-120 g were obtained
from SASCO (Omaha, NE) and fed Purina Chow and
water. After an overnight fast, rats were anesthetized
with 5 mg/100 g body wt. of pentobarbital sodium
injected intraperitoneally, and the epitrochlearis mus-
cles [19] were dissected out.
Muscle incubations
The muscles were incubated in 2 ml of Krebs-
Henseleit bicarbonate buffer (KHB) [20] containing 8
mM glucose, 32 mM mannitol, 0.1% bovine serum
albumin and the additions indicated for each experi-