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Association of oxytocin and arginine vasopressin with protein in bovine and rabbit neurohypophysial extracts has been studied by co-precipitation of the hormones with protein on addition of NaCl, by gel filtration and by dialysis. Although precipitates formed by addition of NaCl (2·5–20 g./100 ml.) to bovine neurohypophysial extracts at pH 3·1 contain both hormones, oxytocin and arginine vasopressin were partially separated at 5·0 g. NaCl/100 ml. when the proportion of oxytocin precipitated was approximately double that of arginine vasopressin. No precipitate formed on addition of NaCl (15 g./100 ml.) to bovine neurohypophysial extract at pH 5.8.

Experiments by gel filtration and dialysis showed that the binding of oxytocin and vasopressin to protein in neurohypophysial extracts is pH dependent and is maximal in the range pH 5·2–5·8. Dilution of a solution containing neurohypophysial hormones and protein results in dissociation of the complexes and this could account for differences observed in experiments with bovine and rabbit neurohypophysial extracts. It is suggested that the mode of binding between the hormones and protein is ionic association between the cationic free terminal NH2 of the cystine residue in the hormones and free carboxyl groups in protein.

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D. Schams, R. Koll and C. H. Li


The effect of insulin-like growth factor-I (IGF-I), epidermal growth factor (EGF), fibroblast growth factor (FGF) and nerve growth factor (NGF) on production of oxytocin and progesterone by cultured bovine granulosa and luteal cells was studied. Secretion of oxytocin was stimulated, in a dose-dependent manner, by IGF-I at 48 and 120 h of culture to levels much higher than those after stimulation with LH, FSH, EGF, FGF or NGF. A similar effect of IGF-I was observed for progesterone but, in contrast to oxytocin, secretion of progesterone was not increased by EGF, NGF or FGF. During primary culture, for 4 h, of dispersed bovine luteal cells obtained from corpora lutea between days 4 and 10 of the oestrous cycle, all the growth factors tested failed to stimulate secretion of oxytocin or progesterone. The data suggest the relevance of growth factors (especially IGF-I) for ovarian physiology and their possible importance for differentiation of follicles and luteinization.

J. Endocr. (1988) 116, 97–100

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E. L. M. Bolwerk and H. H. Swanson


Oxytocin is released during parturition and may also play a role in maternal behaviour. Oxytocin, injected in the cerebral ventricles, has been reported to accelerate the onset of maternal behaviour in oestrogen-pretreated virgin Sprague—Dawley rats within 2 h of injection. This study was an attempt to replicate and extend these findings in Wistar rats. In the first experiment, 16 virgin females were ovariectomized and a cannula was placed into the cerebral ventricle. Forty-eight hours after a single injection of 24 μg oestradiol benzoate (OB), 400 ng oxytocin or saline were injected into the ventricle. In the second experiment three groups were observed: an untreated control group plus two ovariectomized and cannulated groups treated with OB in a regimen designed to mimic pregnancy. After 10 days of OB administration they received an injection of either saline or oxytocin (400 ng) into the ventricle. Immediately after this injection they were exposed to the pups and observations started. In both experiments no rat became maternal in the first 1·5 h after the intracerebroventricular injection. Oxytocin therefore did not induce a rapid onset of maternal responsiveness in oestrogen-pretreated Wistar rats.

J. Endocr. (1984) 101, 353–357

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J Liebermann and D Schams


In the present investigation, the effect of recombinant (BST) and pituitary-derived (bGH) bovine somatotrophin on progesterone and oxytocin release was examined. Individual copora lutea (CL) were obtained from cows at different stages of the oestrous cycle (days 5–7, 8–12 and 15–18) and also from early pregnancy (days 60–120) and were implanted with an in vitro microdialysis system (MDS).

Perfusion with BST for 60 min (005, 0·5 and 5 μmol/l) induced a dose-dependent stimulation of progesterone release. Release of oxytocin from CL was significantly stimulated by BST at all dose levels. BST (0·5 μmol/l) stimulated progesterone release most during the early and mid-luteal phases and oxytocin release especially during the early luteal stage (days 5–7) of the oestrous cycle. CL from early pregnancy (days 60–120) treated with BST showed a significant response in progesterone and oxytocin release. bGH showed comparable effects.

Our results suggest that somatotrophin acts directly on the secretory function of bovine CL in the MDS, specifically during the early luteal stage (days 5–7) of the oestrous cycle and early pregnancy (days 60–120). Somatotrophin may therefore have physiologically relevant effects associated with the development and maintenance of luteal function.

Journal of Endocrinology (1994) 143, 243–250

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Lactating guinea-pigs were passively immunized with an antiserum to oxytocin of high titre, specificity and avidity. Single i.v. injections of 0·1–0·4 ml antiserum produced high titres which decayed slowly (half-life ≃7 days). Passively administered antiserum was effective in vivo; the clearance of exogenous oxytocin from plasma was greatly slowed in immunized animals. Passive immunization with 0·4 ml antiserum reduced milk transfer to the litter during suckling episodes of 10 min, and overall litter growth rates were significantly decreased. Non-immune serum was without effect. Plasma neurophysin levels showed the same large rises during suckling in immunized animals, indicating that neurohypophysial activation was unimpaired. Despite the presence of high titres of antiserum, some milk transfer still occurred at milk ejection. In-vitro experiments showed that more than 25% of oxytocin remained free 20 s after mixing with plasma taken from passively immunized animals. It is probable that the antiserum in the circulation was unable to bind all the oxytocin released from the posterior pituitary gland before it reached the mammary gland.

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A Kanda, K Takuwa-Kuroda, E Iwakoshi-Ukena, Y Furukawa, O Matsushima and H Minakata

We reported that the common octopus, Octopus vulgaris, in common with vertebrates, possesses two members of the oxytocin/vasopressin superfamily: octopressin (OP) and cephalotocin (CT). This was the first observation of its kind in invertebrates. As OP and CT have different biological activities, the presence of specific receptors has been proposed. We cloned the cDNA of an orphan receptor from Octopus brain and found it to encode a polypeptide of 397 amino acids that displays sequences characteristic of G-protein coupled receptors. The orphan receptor showed high homology to receptors of the oxytocin/vasopressin superfamily and seemed to conserve the agonist-binding pocket common to the oxytocin and vasopressin receptors. Xenopus oocytes that express the orphan receptor responded to the application of CT by an induction of membrane Cl(-) currents coupled to the inositol phosphate/Ca(2+) pathway. OP and the other members of the oxytocin/vasopressin superfamily did not activate this receptor. HPLC fractionation of the Octopus brain extract combined with an oocyte assay yielded a single substance that was identical to CT. On the basis of these results, we conclude that the cloned receptor is the CT receptor (CTR). Expression of CTR mRNA in Octopus was detected in the central and the peripheral nervous systems, the pancreas, the oviduct and the ovary. This receptor may mediate physiological functions of CT in Octopus such as neurotransmission, reproduction and metabolism.

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MF Walter, ML Forsling and DG Shirley

In order to determine the possible role of endogenous oxytocin in controlling electrolyte and water excretion in animals whose renal function is being assessed by invasive techniques, rats were anaesthetized and subjected to micropuncture surgery. Clearance measurements were made in the presence and absence of the potent oxytocin receptor antagonist d(CH(2))(5)[Tyr(Me)(2), Thr(4), Orn(8), Tyr(NH(2))(9)]-vasotocin. In rats infused with vehicle alone, glomerular filtration rate (GFR), sodium excretion and urine flow rate remained stable. In contrast, in antagonist-treated rats GFR was modestly reduced (P<0.05), and there were large falls in both absolute and fractional sodium excretion (P<0.01 in each case) and absolute and fractional water excretion (P<0.05 in each case), indicating effects on both filtered load and fractional tubular reabsorption. The antinatriuresis was not accompanied by a change in the fractional excretion of lithium, suggesting that proximal tubular function is unaffected by oxytocin receptor antagonism; nor was it accompanied by a change in the fractional excretion of potassium, suggesting that the tubular effect is located beyond the potassium secretory site, i.e. downstream of the cortical collecting tubule. We conclude that circulating plasma concentrations of oxytocin during anaesthesia and moderate surgery are sufficient to enhance GFR and reduce fractional tubular sodium and water reabsorption. This has important implications for the interpretation of invasive studies such as micropuncture.

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T Engstrom, P Bratholm, NJ Christensen and H Vilhardt

The objective of the present study was to further elucidate our previous observation that beta2-adrenoceptor activation induces oxytocin receptor (OTR) expression in rat myometrium. We wanted to investigate whether the mechanism behind this effect was under the influence of gonadal steroids. Ovariectomized non-pregnant rats were treated with estrogen, progesterone or a combination of both for 3 days. Some rats were concomitantly treated with isoproterenol. Estrogen treatment increased both OTR mRNA production and maximal binding of [3H]-oxytocin to isolated myometrial plasma membranes, but it did not affect contractility of isolated uterine strips challenged with oxytocin. When the estrogen regimen was combined with isoproterenol treatment, an augmented maximal contractile response (Emax) to oxytocin was observed although no further increase in OTR mRNA and binding was seen. Progesterone treatment did not in itself alter OTR mRNA, OTR binding or Emax. However, OTRs were induced at the level of gene expression when progesterone was supplemented with isoproterenol infusion. Finally, progesterone suppressed the effect of estrogen on OTR mRNA production and binding when the two compounds were administered together. However, when isoproterenol treatment was added this effect was abolished and Emax was enhanced more than that seen following treatment with estrogen alone. These data suggest that beta2-adrenoceptor activation represents an important regulator of OTR expression/function in estrogen- and progesterone-dominated rat myometrium.

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J. A. Russell, R. G. Gosden, E. M. Humphreys, R. Cutting, N. Fitzsimons, V. Johnston, S. Liddle, S. Scott and J. A. Stirland


Oxytocin secretion is inhibited by opioids, and oxytocin is important in parturition. The effects on parturition of morphine, a relatively selective μ-opioid receptor agonist, were studied in the rat. Morphine or vehicle with or without the opiate antagonist naloxone were administered immediately after the birth of the second pup and the subsequent course of parturition was recorded in a total of 80 rats. Both s.c. morphine (10 mg/kg) and intracerebroventricular (i.c.v.) morphine (18 μg through a previously implanted cannula) interrupted parturition, delaying the birth of the sixth pup after treatment to 187·3 ± 35·9 (s.e.m.) min and 195·4 ± 19·5 min respectively, compared with 46·4 ± 3·7 and 66·1 ± 17·5 min after vehicle alone. The dose of morphine given i.c.v. had no effect when given s.c. Naloxone given concurrently prevented the effects of morphine. Eventually the rate of parturition in the morphine-treated groups recovered. Perinatal pup mortality rate was not increased when morphine was given to the mothers, but it did inhibit the expression of normal intrapartum maternal behaviour.

Pup mortality was increased 48 h post partum by morphine given during parturition, and it reduced the proportion of rats with normal maternal behaviour 24 h post partum. Morphine did not affect spontaneous or oxytocin-stimulated contractile activity of the parturient uterus in vitro. The concentration of oxytocin in trunk blood plasma was decreased 40 min after i.c.v. morphine (24·3 ± 3·9 vs 39·3± 6·5 pmol/l in controls), as was vasopressin (7·2 ± 1·5 vs 19·7 ± 4·5 pmol/l in controls). Intravenous infusion of oxytocin (2–5 mU/min for 144·3 ± 8·2 min; total infused 448·5 ± 61·9 mU) after i.c.v. morphine re-started parturition; all pups were born to these rats (mean time to pup 6, 110·3 ± 12·7 min) before the i.v. vehicle-infused rats given i.c.v. morphine re-started (mean time to pup 6, 406·3±125·2 min).

It is concluded that morphine given during parturition acts centrally through opioid receptors to inhibit oxytocin secretion, and impairs the expression of maternal behaviour. Reversal of the effects of morphine on parturition by i.v. oxytocin demonstrates the important role of oxytocin in fetus ejection and expulsion.

Journal of Endocrinology (1989) 121, 521–536

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V. J. Ayad, E. L. Matthews, D. C. Wathes, T. J. Parkinson and M. L. Wild


The present study was designed to determine the localization of the endometrial oxytocin receptor during the ovine oestrous cycle, particularly on day 14, the time of initiation of luteal regression in the ewe. Samples were obtained from 29 ewes at different stages of the oestrous cycle (several during the luteal phase and on every day between day 14 (− 2) and day + 3 of the oestrous period).

Oxytocin receptors were localized autoradiographically in sections of uterine tissue, using the 125I-labelled oxytocin receptor antagonist [1-(β-mercapto-β,β-cyclopentamethylene propionic acid), 2-(ortho-methyl)-Tyr2,Thr4,Orn8,Tyr9-NH2]-vasotocin (125I-labelled OTA). There was some variation in the pattern of 125I-labelled OTA labelling between different uterine tissue samples from the same ewe and also between samples obtained from different ewes thought to be at the same stage of the oestrous cycle. A clear overall pattern did, however, emerge with 125I-labelled OTA-binding sites distributed between luminal epithelial cells, glandular epithelial cells and caruncular stromal cells to varying extents on different days of the cycle.

During the luteal phase (days 5–12) clear specific labelling of endometrial tissue was generally absent. On day 14 labelling was evident on the luminal epithelium, but only in nine tissue samples out of a total of 18 studied, indicating that the entire luminal surface did not contain oxytocin receptors at this time. Between the day before oestrus and day 3 of the oestrous cycle the luminal epithelium was consistently labelled. The most extensive labelling of the remaining endometrial tissue was observed on the day of oestrus, with 125I-labelled OTA-binding sites clearly present on the stromal cells within caruncles and on a large proportion of secretory epithelia. This contrasted with the day before and the day after oestrus when labelling of glandular tissue was confined to the superficial endometrium, and labelling of caruncular stromal cells, although sometimes evident, was never as intense as on day 0. On days 2 and 3 labelling varied between being similar to that found on day 1 and being confined to the luminal epithelium and very few superficial secretory glands.

The results of this study lead us to conclude that the oxytocin receptor shows a differential distribution between stromal cells, epithelial cells lining secretory glands and luminal epithelial cells during the oestrous cycle; that the steroidal regulation of the oxytocin receptor differs between endometrial cell types; and that control of the luminal epithelial oxytocin receptors is probably of particular importance to the regulation of prostaglandin F release at luteal regression and during the maternal recognition of pregnancy.

Journal of Endocrinology (1991) 130, 199–206