© 2008 The Authors Doi: 10.1111/j.1742-7843.2008.00228.x
Journal compilation © 2008 Nordic Pharmacological Society . Basic & Clinical Pharmacology & Toxicology , 103, 36–42
Blackwell Publishing Ltd
Deficiency in Cytosolic Malic Enzyme Does Not Increase
Acetaminophen-Induced Hepato-Toxicity
Su Qian, Sheena Mumick, Peter Nizner, Michael R. Tota, Joseph Menetski, Marc L. Reitman and Douglas J. MacNeil
Department of Metabolic Disorders, Merck Research Laboratories, Rahway, NJ, USA
(Received June 8, 2007; Accepted November 14, 2007)
Abstract: Cytosolic malic enzyme (ME-1) is a nicotinamide adenine dinucleotide phosphate (NADP)-dependent enzyme
that generates NADPH. The activity of this enzyme, the reversible oxidative decarboxylation of malate to yield pyruvate,
links glycolytic pathway to citric acid cycle. The high level of ME-1 expression in liver, and its involvement in NADPH
production, suggests reduced ME-1 activity might compromise hepatic production of reduced glutathione (GSH) by the
NADPH-dependent enzyme glutathione reductase, and hence affect xenobiotic detoxification. The role of ME-1 in liver
detoxification was evaluated in Mod1 deficient mice (mod1
–/–
) by evaluating their sensitivity to acetaminophen-induced liver
injury. The results show that mod1
–/–
mice are not more sensitive to acetaminophen hepato-toxicity. Although GSH levels
were initially depleted more in the mod1
–/–
liver than in wild-type controls, the GSH levels recovered quickly. In conclusion,
our data indicate that ME-1 deficiency does not adversely affect GSH-dependent detoxification.
Cytosolic malic enzyme (ME-1) is a homotetrameric enzyme
catalysing reversible oxidative decarboxylation of malate
to yield pyruvate and carbon dioxide. The ME-1 reaction
is a step in the pyruvate–citrate shuttle by which acetyl-
CoA generated via pyruvate decarboxylation by pyruvate
dehydrogenase is exported out of the mitochondria and
made available for fatty acid synthesis in the cytosol. In the
process, ME-1 generates a significant fraction of the cytosolic
reducing agent NADPH, which is important for diverse
cellular functions, including de novo synthesis of long-chain
fatty acids [1,2]. In mammals, two additional isoforms of malic
enzyme reside in mitochondria. One malic enzyme isoform,
ME-2, uses nicotinamide adenine dinucleotide as the cofactor,
while the other isoform, ME-3, prefers nicotinamide adenine
dinucleotide but can also use nicotinamide adenine dinucleotide
phosphate (NADP). In lipogenic tissues, like liver and adipose
tissue, more than 90% of malic enzyme activity is present in
the cytoplasmic fraction [3].
Liver and adipose activity of ME-1 is induced by a high
carbohydrate/low-fat diet, and down-regulated by a high-
fat diet [2,4–6]. Due to its concerted induction with glucose-
6-phosphate dehydrogenase and 6-phospho-gluconate
dehydrogenase, two NADPH producing enzymes important
for fatty acid synthesis, ME-1 is also considered a lipogenic
enzyme [1–6]. Indeed, higher levels of liver ME-1 activity have
been associated with obese mouse and rat models [7,8]. High
ME-1 activity has also been observed in certain human
carcinoma cell lines [9,10], possibly reflecting altered energy
metabolism in cancer cells.
Cytosolic malic enzyme may play a significant role in
the liver’s detoxification of xenobiotics. Liver detoxification
of xenobiotics relies heavily on reduced glutathione (GSH)-
mediated peroxide reduction process. To maintain the liver’s
detoxification capacity, an adequate supply of NADPH is
crucial for GSH regeneration by glutathione reductase.
ME-1 is highly expressed in liver, and an increase in NADPH
consumption, due to xenobiotic detoxification, increases
the specific activity of ME-1 [11]. Thus, the inhibition or loss
of ME-1 may compromise the liver’s detoxification capacity,
leading to an increase in the sensitivity of liver to toxic agents.
To evaluate the effect of a reduction in ME-1 on the liver’s
detoxification capacity, we compared mod1
–/–
mice to wild-
type mice for their sensitivity to liver toxicity induced by
acetaminophen (APAP). A strain of spontaneous occurring
mod1
–/–
mice has been described [12]. The aberrant mod1
mRNA in the mutant mice carries a 522-bp duplication
corresponding to nucleotide 1027–1548 in the wild-type mod1
transcript [13]. The aberrant Mod1 protein accumulates to
only about 1–2% of the wild-type Mod1 level, presumably
due to reduced stability. No ME-1 activity was detected in
liver or kidney cytosol preparations from mod1
–/–
mice [12].
The mod1
–/–
mice do not exhibit overt abnormalities under
standard laboratory conditions.
Acetaminophen is a commonly used analgestic and
antipyretic drug, but this drug can cause life-threatening liver
damage when taken in excess. At therapeutic doses, APAP is
oxidized by cytochrome P-450 enzymes in the liver to produce
N-acetyl-p-benzoquinone imine (NAPQI), which is further
detoxified by conjugation with GSH [14]. After an overdose
of APAP, GSH is not regenerated fast enough and becomes
depleted. The unconjugated NAPQI triggers liver cell necrosis
by either covalent binding to cellular proteins and blocking
their activities, or increasing oxidative stress [15,16].
Author for correspondence: Su Qian, Department of Metabolic
Disorders, Merck Research Laboratories, P.O. Box 2000,
RY80M-213, Rahway, NJ 07065, USA (fax 732-594-6708, e-mail
su_qian@merck.com).