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This research paper addresses the hypothesis that oxytocin (OT) could be released during suckling and during milking with and without the presence of a calf and that this release could be regulated by maternal behaviour. Plasma concentration patterns of OT and cortisol (CORT) were measured in six Tunisian dromedary camels during 2 suckling episodes, 2 manual milking episodes with calves beside the mother and 2 machine milking episodes without calves present. Various patterns of OT release were observed between each camel including specific two peak release patterns. Higher plasma OT concentrations were found during the suckling and hand-milking episodes with simultaneous suckling of calves, than during the machine milking episodes without calves. Exclusive mechanical milking episodes also evoked significant mean OT release, although greatly reduced compared to suckling and hand milking. The low basal levels and classical CORT release patterns suggested non-stressful management practices were used and there were very limited differences in udder stimulation between managements. The OT release induced by exclusive suckling and suckling together with hand-milking gives a reference point for what a good milk ejection stimulation is in camels. The important and specific reduction of OT release during machine milking without the calf present could be a physiological consequence of the maternal behaviour (selectivity for the own young) and to a lesser extent explained by a lower stimulation by machine milking.

The neuropeptide oxytocin (OT) promotes maternal care and social affiliation in adults but its importance in infant attachment still remains unknown. True animal models of infant attachment are extremely rare, and the sheep (in complement to non-human primates) is one of the few that provides the opportunity to investigate its neuroendocrinological basis. In the lamb, access to the udder has strong rewarding properties for the establishment of a preferential relationship with the mother. Therefore, the present study explored the possible involvement of OT through its release during close social contacts with the mother. The first experiment revealed that lambs having free access to the udder from birth onward developed, by 12 h of age, a clear preference for their mothers over another maternal ewe. Delaying access to the udder for six, four or even only 2 h starting at birth, by covering the ewe's udder, resulted in the lack of such a preference without affecting general activity. These effects persisted in most cases at 24 h but by 72 h of age a bond with the mother was clearly expressed. Experiment two showed that social interactions with the mother were followed by a release of OT in the plasma when lambs had the possibility to suckle. Non-nutritive interactions were without effects. Preliminary data on two subjects suggested that OT might also increase in the cerebrospinal fluid after suckling. Finally, in the third experiment, oral administration of a non-peptide OT receptor antagonist (L-368-899, Merck) over the first 4 h after birth led to decreased exploration of the mother's body compared to lambs receiving saline, and impaired the expression of a preference for the mother at 24 h. The effects were no longer observed at 48 h. Our findings demonstrate that both delayed access to the mother's udder and OT receptor antagonist alter the onset of mother preference in newborn lambs. This suggests that central OT facilitates the development of filial attachment through its release during suckling., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Nowak, Lévy, Chaillou, Cornilleau, Cognié, Marnet, Williams and Keller.)

Friendly interactions between humans and animals such as gentling or petting have been shown to have positive behavioural and physiological consequences in many species. In primates, rodents and dogs, oxytocin has been associated with tactile contact and anti-stress effects that may influence bonding and responses to stress situations. However the activation of the oxytocinergic system in other human-animal interactions such as with herbivores, had not yet been studied. Sixteen female lambs were reared by artificial feeding reinforced with 3× 30 s daily stroking sessions. At 6 weeks of age, the test consisted in measuring first plasma oxytocin and cortisol responses in lambs during a first 6-min phase in the home pen where the familiar caregiver gently stroked the lamb, and then physiological and behavioural responses in a test pen during a 20-min - phase of social isolation followed by a 20-min - phase of reunion with its familiar caregiver. The lambs expressed behavioural agitation during the whole period of isolation. A strong affiliative response towards the human and a sustained reduction of the agitation behaviour were observed during reunion. Lambs' behaviours when isolated and when in contact with the human were correlated suggesting a response to social separation from the familiar caregiver more than to social isolation from congeners. No significant changes in cortisol levels were observed during the test. Oxytocin levels did not vary during human contact, but increased when the familiar caregiver left the lamb alone in the test pen. In conclusion, lambs displayed affiliative responses towards their caregiver, and the lack of cortisol response during isolation while oxytocin was released suggest an anti-stress effect of oxytocin., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

This study evaluated the effect of exclusive machine milking on oxytocin (OT) and cortisol (CORT) release, and on milk yield and residual milk in Gir (group Gir), Holstein (group Hol) and crossbred animals (group GirHol). Six animals from each group were submitted to experimental milkings. As expected, milk yield was significantly higher in the Hol group than in the groups GirHol and Gir, and group GirHol produced more milk than the Gir group. In contrast, all groups exhibited significant but similar levels of OT, although OT increased more rapidly during milking for the groups Hol and GirHol than for the Gir group. In addition, CORT levels measured during and after milking were significantly influenced by the group. The Gir group showed higher levels of CORT than the groups GirHol and Hol. The lower performances of the Gir breed can not be explained by a less efficient milk ejection reflex because all cows studied released enough OT and had an effective milk ejection.

Oxytocin plays a major role in lactation mainly by its action on milk ejection via the contraction of myoepithelial cells. The effect of oxytocin on milk production and the presence of oxytocin receptors on different epithelial cells suggest that this hormone may play a role in mammary epithelial cells. To determine precisely the various roles of oxytocin, we studied localization of oxytocin receptors in lactating rabbit and rat mammary tissue and the influence of oxytocin on secretory processes in lactating rabbit mammary epithelial cells. Immunolocalization of oxytocin receptors on mammary epithelial cells by immunofluorescence and in mammary tissue by immunogold in addition to in situ hybridization showed that lactating rat and rabbit mammary epithelial cells expressed oxytocin receptors. Moreover, oxytocin bound specifically to epithelial cells. To determine whether oxytocin had an effect on lactating rabbit mammary epithelial cells, isolated mammary fragments were incubated in the presence or absence of 10(-6) i.u. ml(-1) of oxytocin. After 1 min of incubation with oxytocin, the morphology of epithelial cells and the localization of caseins and proteins associated with the secretory traffic suggested a striking acceleration of the transport leading to exocytosis, whereas the contraction of myoepithelial cells was only detectable after 7 min. Addition of 10(-8) g ml(-1) of atosiban before the addition of oxytocin prevented the oxytocin effect on secretory processes and on myoepithelial cell contraction. Addition of 10(-6) i.u. ml(-1) of vasopressin to the incubation medium did not mimic the stimulating effect of oxytocin on secretory traffic. These results show that lactating rabbit and rat mammary epithelial cells express oxytocin receptors and that oxytocin binds to these receptors. They strongly suggest that oxytocin has a dual effect on lactating mammary tissue: an acceleration of the intracellular transfer of caseins in mammary epithelial cells followed by the contraction of myoepithelial cells.

The aims of the present study were (1) to investigate the influence of insulin-like growth factor-I (IGF-I) on follicular size, on the secretion of oxytocin (OT), progesterone (P), estradiol (E), IGF binding protein-3 (IGFBP-3), inhibin A, inhibin B and cAMP and on the expression of proliferation-associated peptide PCNA, ERK-related mitogen activated protein kinase (MAPK/ERK1, 2) and protein kinase A (PKA) in cultured porcine ovarian follicles; (2) to examine the effects of OT on IGF-I and on these functions; and (3) to determine whether the effects of IGF-I can be mediated by OT. To define the involvement of OT in mediating IGF-I action, we compared responses of porcine ovarian follicles to IGF-I and OT and examined whether blockade of endogenous OT by specific antiserum can affect IGF-I action. It was observed that IGF-I (1, 10 or 100 ng/ml) was able to prevent a decrease in the size of ovarian follicles during culture and caused an increase in the diameter of some follicles. It also stimulated the secretion of OT, P, IGFBP-3, inhibin A and cAMP, decreased the secretion of E and inhibin B (RIA/EIA/ELISA), and induced the expression of PCNA, PKA, MAPK/ERK1, but not MAPK/ERK2 (Western blotting). Like IGF-1, OT (100 ng/ml) prevented decrease in follicular size and increased the diameter of some follicles. It also stimulated the secretion of P and IGF-I, but not E. Antiserum against OT (1%), when given alone, did not affect the reduction of follicular size but slightly increased the percentage of follicles increasing their diameter during culture. The antiserum also inhibited secretion of OT and cAMP but not the secretion of P, E, IGFBP-3 or the expression of PKA, MAPK/ERK1 or 2. When given together with IGF-I, the antiserum prevented the stimulatory action of IGF-I on the proportion of enlarged follicles and on OT, IGFBP-3 and MAPK/ERK1. It augmented the effect of IGF-I on P, but not the effect on E, cAMP, PKA or MAPK/ERK2. These observations demonstrate the involvement of IGF-I and OT in the control of ovarian follicular size and follicular cell proliferation, progestagen, estrogen, IGFBP-3, inhibin A and B secretion and in cAMP/PKA- and MAPK/ERK1-dependent intracellular mechanisms. Furthermore, the reciprocal stimulation of IGF-I and OT and the similarity of some their effects, together with the prevention or augmentation of some IGF-I effects after OT blockade, suggest that IGF-I action can be mediated by OT.

In postparturient goats, olfactory recognition of the young allows the establishment of a selective bond between the mother and her kids. Once this bond is formed, the mother rejects alien young that attempt to suckle. We tested whether the development of the maternal selective bond in goats modulates prolactin (PRL) and oxytocin (OT) release in response to suckling. On day 37 of lactation, serial blood samples were taken during nursing of the mother's own or alien kid(s) in 10 intact/selective goats and in 10 goats rendered anosmic/nonselective through prepartum peripheral ZnSO(4) irrigation. Spontaneous nursing behavior was also studied weekly from day 7 to 30 of lactation, at which time milk production was measured. Maternal selectivity had no effect on PRL release, in contrast to OT release, which was significantly affected by this factor. Intact mothers released OT only when nursing their own kids, but not with aliens, while anosmic/nonselective dams showed an increase in OT levels regardless of the identity of the kids. In addition to these effects on maternal selectivity, the amplitude of the response of both hormones was lower in anosmic mothers than in intact mothers. Finally, nursing behavior and milk production were not significantly affected by anosmia. We conclude that maternal selective behavior in goats, which relies on the individual olfactory signature of the kid, modulates the OT, but not the PRL, response to suckling. In addition, perception of the smell of the young appears to have a general facilitatory effect, independent of the kid's identity, on the release of both hormones., (Copyright 2002 Elsevier Science (USA))

This study evaluated the effect of different milking managements on oxytocin, prolactin and growth hormone release in Gir x Holstein cows. Six cows were suckled by their calves, before and after milking (SM group); another six cows were submitted to exclusive milking (M group). High OT levels were observed during suckling of the SM group, however both groups had similar levels of OT during milking. The SM group presented PRL levels significantly higher than the M group, however only during suckling. For GH levels, the SM group showed higher levels than the M group, however this difference was significant only during the first suckling, cleaning of the teats and at the beginning of machine milking. Despite these results, SM cows produced more milk (milking plus suckling) than M cows, however milk obtained by milking was similar for both groups.

The objective was to compare in the ewe the effects of easy and difficult procedures for artificial insemination (AI) (as related to rapid or poor accessibility of the cervix, respectively) on plasma cortisol (CORT) and oxytocin (OT), and uterine motility. All AI were simulated using a catheter empty of semen to study genital and environmental stimuli only. In experiment 1, 40 ewes were sampled after Al, and whether it was an easy or difficult procedure was reported for each animal. While CORT concentrations rose to a similar amount in all ewes, whatever the Al procedure, a significant OT response occurred after a difficult procedure only (n = 18) (17.4 +/- 1.7 versus 12.7 +/- 0.7 pg x mL(-1) before Al, p < 0.05). In experiment 2, uterine activity was monitored in 4 ewes using an implantable telemetric transmitter equipped with an intrauterine pressure catheter. An increased uterine activity occurred during 2 +/- 1 min after an easy Al (n = 5), whereas the evoked activity lasted for 15 +/- 4 min after a difficult Al (p < 0.001, n = 7). A similar long-lasting response occurred after OT administration (100 mIU, i.v.). We concluded that the increase in uterine motility after a difficult Al resulted from a reflex release of OT, and not to a "stress" effect.

This review describes the effects of milking (routine and management) on milk yield and milk quality on dairy ruminants and examines the physiological effects of milking on the synthesis and secretion of milk. During milking, differences in the composition of milk as a result of milk ejection reflex are observed: the cisternal milk, immediately available, contains little fat, then milk ejection provokes active transport of high-fat content alveolar milk, into the cisternal compartment. Milking frequency has the capacity to affect milk production too. So, an increase in milking frequency augments milk yield whereas a decrease in milking frequency decreases milk production, with effects on milk composition. The milk ejection reflex is mediated by oxytocin, which induces myoepithelial cell contraction. Nevertheless, other actions of oxytocin may exist, such as a direct effect on proliferation and differentiation of myoepithelial cells and on secretory processes in the mammary epithelial cells.

Milk yield and plasma oxytocin (OT), prolactin (PRL), and cortisol (CORT) during suckling and machine milking were measured in multiparous ewes subjected to a mixed management system of 3 sucklings and two daily milkings. Peak hormones were significantly increased and were similar during suckling and milking for PRL (181 vs. 163.3 ng x mL(-1)) and CORT (12.5 vs. 11.5 ng x mL(-1)). During the period of exclusive suckling, OT was always significantly released (90.3 pg x mL(-1)); however, during the period of mixed management, OT concentrations only increased during suckling compared to milking (91.7 vs. 13.1 pg x mL(-1)). The mean volume of milk obtained during suckling (632 mL) was significantly higher than during milking (255 mL). Thus, during a mixed management system, oxytocin and prolactin releases are not under similar central regulation. A mixed system, without OT release during milking, does not contribute to accelerate the conditioning of ewes for machine milking.

The aim of this work was to investigate whether luteal oxytocin released after non-luteolytic prostaglandin F2 alpha (PGF2 alpha) challenge could explain the intramammary pressure (IMP) rises previously described in pregnant ewes after the normal life span of a corpus luteum. Blood oxytocin levels and IMP after challenge were measured in cycling and pregnant lactating ewes until the response ceased (approximately 55 d post oestrus). Oxytocin release was not significantly different for cycling and pregnant ewes, and fell to its lowest levels on days 16-18 post oestrus in both groups, i.e. with or without luteolysis. However, although IMP response fell with luteolysis in cycling ewes, the response persisted until day 55 in pregnant ewes. This lack of correlation between oxytocin release and IMP response raised interesting questions including: is there an unknown oxytocic compound that can be released after PGF2 alpha stimulation of the corpora lutea?

The pure tetrameric form of Acetylcholinesterase (EC-3.1.1.7) from the electric eel electrophorus electricus has been covalently coupled to oxytocin. This conjugate has been used as tracer in a heterologous competitive immunoassay. Microtiter plates coated with a mouse monoclonal anti-rabbit immunoglobulin antibody were used to separate bound and free moieties of the tracer. Acetylcholinesterase activity bound to the solid phase was measured by a colorimetric assay. The minimum detectable concentration was 0.075 pg/well (ie 1.5 pg/ml) and precision was less than 8% at concentration above 0.15 pg/well. An extraction step improved sensitivity up to 10 times with good recoveries. To assess the validity of this assay, basal levels of oxytocin were measured during the oestrous cycle of a cow.

This experiment was conducted in 59 Lacaune breed ewes in order to compare milk production and milk distribution between alveolar and cisternal storage after superovulation. After a corpora lutea (CL)-free control period, the ewes were superovulated by different treatments (experimental period) and 5 classes were differentiated according to the number of corpora lutea observed (0, 1, 2, 3 to 6 and > 6 CL respectively (group A (n = 20), B (n = 7), C (n = 14), D (n = 7), E (n = 11)). Our results showed a positive correlation between the number of corpora lutea and the oxytocin and progesterone levels in plasma, total milk production and cisternal volume, and a negative correlation with alveolar volume. The milk production at the evening milking for groups A and E (388.6 and 384.8 ml respectively during the control period) respectively reached 321.7 ml (-17.2%) and 413.4 ml (+7.4%) during the experimental period; ie, a 24.6% difference between these 2 extreme groups. These results could likely be explained by oxytocin levels reaching those obtained during milking and by the effect of milk transfer from the alveolar to the cisternal lumen. Additionally, progesterone could act directly on milk synthesis at the level of the secretory cells.

The oxytocic effect of a prostaglandin F2 alpha analogue, fenprostalene, was assessed in 4 ovariectomized ewes fitted with electrodes in both uterine horns and in the cervix. In the absence of estradiol priming, significant motility changes were not elicited by fenprostalene. Conversely, when ewes were primed with 17-beta-estradiol, fenprostalene markedly increased the electrical activity in the uterus and cervix. After a single subcutaneous fenprostalene administration (5 micrograms/kg), activity values remained about twice that of the control values during 8.52 +/- 3.31 hours. When the same dosage was administered IM, similar post-injection activity values were obtained, but only during 5.88 +/- 0.72 hours. Oxytocic effects of fenprostalene were far longer than those elicited by a single IM administration of 50 micrograms of prostaglandin F2 alpha (tham salt)/kg (0.91 +/- 0.32 hours) or by a single IM administration of 1 microgram of cloprostenol/kg (1.88 +/- 0.81 hours). Using the dose-effect relationship curve obtained from the same ewes by continuous IV infusions of oxytocin (OXT), the postinjection activity values reached after a single subcutaneous administration of fenprostalene were equivalent to those of an IV infusion of OXT at an average dose of 4.09 ImU of OXT/kg/hr for 6 to 13 hours, according to the values of the particular ewe concerned. These long-lasting oxytocic properties, in addition to its luteolytic capabilities, would make fenprostalene a suitable drug for promoting effective evacuation of the uterus when required in daily veterinary practice.