Neurotoxicology and Teratology, Vol. 15, pp. 107-115, 1993 0892-0362/93 $6.00 + .00
Printed in the U.S.A. All rights reserved. Copyright © 1993 Pergamon Press Ltd.
Fetal Nicotine Exposure Alters Ontogeny of
M -Receptors and Their Link to G-Proteins
E. A. ZAHALKA, I F. J. SEIDLER, J. YANAI AND T. A. SLOTKIN
Department of Pharmacology, Duke University Medical Center, Durham, NC 27710
The Melvin A. and Eleanor Ross Laboratory for Studies in Neural Birth Defects (JY), Department of
Anatomy and Embryology, Hebrew University, Hadassah Medical School, Jerusalem, Israel
Received 13 October 1992; Accepted 14 December 1992
ZAHALKA, E. A., F. J. SEIDLER, J. YANAI AND T. A. SLOTKIN. Fetal nicotine exposure alters ontogeny of
Mrreceptorsand theirlink to G-proteins. NEUROTOXICOL TERATOL 15(2) 107-115, 1993.-Prenatal nicotine exposure
has been shown to disrupt the development of cholinergic presynaptic tone and behaviors mediated through muscarinic
cholinergic receptors. The current study examines nicotine's effects on ontogeny of postsynaptic muscarinic Mrreceptors in
rat striatum and hippocampus after continuous maternal infusions of 2 mg/kg/day or 6 mg/kg/day from gestationai days 4
through 20. Although brain region weights were unaffected by nicotine exposure, significant alterations in receptor develop-
ment and receptor regulation by G-proteins were found. Postnatal development of striatal M,-receptor binding, as identified
with [3H]pirenzepine, was significantly impaired with either of the fetal nicotine regimens. Treatment with 2 mg/kg/day also
produced alterations in striatal receptor affinity state, characterized by enhanced ability of an agonist (oxotremorine-M) to
displace [3H]pirenzepine; raising the dose to 6 mg/kg/day masked the affinity shift by affecting G-protein regulatory mecha-
nisms, such that addition of the GTP analog, GppNHp, produced a larger decrease in agonist affinity. In the hippocampus,
no such effects on receptor binding, affinity state, or G-protein regulation were seen with either regimen. These data thus
indicate that fetal nicotine exposure, even at doses that do not cause overt signs of maternal/fetal/neonatal toxicity or
growth impairment, influences cholinergic receptor development and regulation of cell signaling mediated by G-proteins. The
selectivity of effects toward Mm-receptorsin the striatum, a region with a prenatal peak of neuronal mitosis, as compared to
hippocampus, where mitosis peaks postnatany, suggests that vulnerability to nicotine involves a critical phase of cell develop-
ment, rather than being targeted toward receptors of a given subtype.
Cholinergic receptors Effects of nicotine in developing brain G-proteins in developing brain
Development of cholinergic innervation Muscarinic receptor in developing brain Nicotine
Effects on cholinergic development Striatum Development of cholinergic innervation
Hippocampus
TOBACCO smoking during pregnancy leads to intrauterine
growth retardation, spontaneous abortion and low birth
weight (5,12,31,35). However, over and above its general peri-
natal toxicity, tobacco targets the fetal nervous system for
adverse psychological and behavioral effects (1,4,14,28,
44,58). Cigarette smoke contains a wide variety of potentially
injurious chemicals, including carbon monoxide (26) and
smoking causes fetal ischemia and hypoxia (8,29,40,49,53,55).
Nevertheless, nicotine itself is a neurobehavioral teratogen, as
demonstrated conclusively by animal models that use slow
infusions of nicotine that are devoid of other tobacco compo-
nents and do not cause hypoxia and ischemia (see reviews,
23,51). Because nicotine acts via highly selective and sensitive
neurotransmitter receptors that are present in fetal brain
(6,15,34,55), the developing nervous system is actually more
sensitive than are general indices of growth and development
(39), a relationship that runs counter to the standard principle
that systemic insults spare the developing brain relative to the
rest of the body (3,11).
In large measure, the alterations of central nervous system
development evoked by nicotine exposure reflect the fact that
acetylcholine plays a key role in the maturation of target cells
receiving cholinergic innervation, eliciting a switchover from
cell replication to differentiation and determining the subse-
quent architectural modeling of specific brain regions (17,
30,38). Fetal exposure to nicotine thus evokes the same de-
velopmental events, but because the drug administration is
occurring prior to the natural spike of cholinergic activity, the
processes occur prematurely and in a discoordinated fashion
(39,53,56). Accordingly, it is especially important to examine
the specific relationship of nicotine induced neuroteratologies
to development of cholinergic systems. Several studies have
Requests for reprints should be addressed to Dr. T. A. Slotkin, Box 3813, Department of Pharmacology, Duke University Medical Center,
Durham, NC 27710.
Supported by the SmokelessTobacco Research Council and USPHS DA-OS70.
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