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Physiological Reviews, Vol. 81, No. 2, April 2001, pp. 629-683
Copyright ©2001 by the American Physiological Society
Institut für Biochemie, Johannes Gutenberg Universität, Mainz, Germany
Gimpl, Gerald and
Falk Fahrenholz.
The Oxytocin Receptor System: Structure, Function, and
Regulation. Physiol. Rev. 81: 629-683, 2001.
The neurohypophysial peptide oxytocin (OT) and
OT-like hormones facilitate reproduction in all vertebrates at
several levels. The major site of OT gene expression is the
magnocellular neurons of the hypothalamic paraventricular and
supraoptic nuclei. In response to a variety of stimuli such as
suckling, parturition, or certain kinds of stress, the processed OT
peptide is released from the posterior pituitary into the systemic
circulation. Such stimuli also lead to an intranuclear release of OT.
Moreover, oxytocinergic neurons display widespread projections
throughout the central nervous system. However, OT is also synthesized
in peripheral tissues, e.g., uterus, placenta, amnion, corpus luteum, testis, and heart. The OT receptor is a typical class I G
protein-coupled receptor that is primarily coupled via
Gq proteins to phospholipase C-
. The high-affinity
receptor state requires both Mg2+ and cholesterol, which
probably function as allosteric modulators. The agonist-binding
region of the receptor has been characterized by mutagenesis and
molecular modeling and is different from the antagonist binding site.
The function and physiological regulation of the OT system is strongly
steroid dependent. However, this is, unexpectedly, only partially
reflected by the promoter sequences in the OT receptor gene. The
classical actions of OT are stimulation of uterine smooth muscle
contraction during labor and milk ejection during lactation. While the
essential role of OT for the milk let-down reflex has been
confirmed in OT-deficient mice, OT's role in parturition is
obviously more complex. Before the onset of labor, uterine sensitivity
to OT markedly increases concomitant with a strong upregulation of OT
receptors in the myometrium and, to a lesser extent, in the decidua
where OT stimulates the release of PGF2
. Experiments
with transgenic mice suggest that OT acts as a luteotrophic hormone
opposing the luteolytic action of PGF2. Thus, to
initiate labor, it might be essential to generate sufficient
PGF2 to overcome the luteotrophic action of OT in late
gestation. OT also plays an important role in many other
reproduction-related functions, such as control of the estrous cycle length, follicle luteinization in the ovary, and ovarian steroidogenesis. In the male, OT is a potent stimulator of spontaneous erections in rats and is involved in ejaculation. OT receptors have
also been identified in other tissues, including the kidney, heart,
thymus, pancreas, and adipocytes. For example, in the rat, OT is a
cardiovascular hormone acting in concert with atrial natriuretic peptide to induce natriuresis and kaliuresis. The central actions of OT
range from the modulation of the neuroendocrine reflexes to the
establishment of complex social and bonding behaviors related to the
reproduction and care of the offspring. OT exerts potent antistress
effects that may facilitate pair bonds. Overall, the regulation by
gonadal and adrenal steroids is one of the most remarkable features of
the OT system and is, unfortunately, the least understood. One has to
conclude that the physiological regulation of the OT system will remain
puzzling as long as the molecular mechanisms of genomic and nongenomic
actions of steroids have not been clarified.
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