Biology of the prolactin family in bovine placenta. II. Bovine prolactin-related proteins: Their expression, structure and proposed roles

The placenta produces various peptides and steroid hormones that regulate placental function and fetal growth. Prolactin‐related proteins are peptides that are produced by the placenta and belong to the growth hormone/prolactin family, and have structural similarity to prolactin and placental lactogen. Although several prolactin‐related protein genes have been detected in bovine placenta, their expression profiles and functions are not clear. The main difficulties in examining their biological function is the similarity between their genes and the lack of information about their proteins. Recently, molecular biology methods have been used to detect some new bovine prolactin‐related proteins, and elucidate their biological functions. This review focuses on the structures, expression profiles and conceivable functions of prolactin‐related proteins in bovine placenta. With respect to their expression profiles, bovine prolactin‐related proteins fall into four groups: (i) those expressed around the implantation period; (ii) those that reach peak expression in the middle of gestation; (iii) those that increase with the progress of gestation, reaching a peak in late gestation; and (iv) those that reach a plateau in early gestation and are maintained at that level throughout gestation. Data indicate that bovine prolactin‐related proteins have different biological roles in different periods of gestation. In situ monitoring suggests that bovine prolactin‐related protein‐I has a role in the attachment of trophoblast cells to endometrium during the early implantation period.     

The Possible Involvement of Salsolinol and Hypothalamic Prolactin in the Central Regulatory Processes in Ewes During Lactation

Salsolinol, a dopamine‐related compound and prolactin‐producing cells were found in the ovine hypothalamus. This study was designed to test the hypothesis that salsolinol, acting from the CNS level, is able to stimulate pituitary prolactin release as well as prolactin mRNA expression in the anterior pituitary cells (AP) and in the mediobasal hypothalamus (MBH) in lactating ewes. The intracerebroventricular infusions of salsolinol in two doses, total of 50 ng or 5 μg, were performed in a series of five 10‐min infusions at 20‐min intervals. All infusions were made from 12:30 to 15:00 and the pre‐infusion period was from 10:00 to 12.30 h. The prolactin concentration in plasma samples, collected every 10 min, was determined by radioimmunoassay; prolactin mRNA expression in AP and MBH tissues was determined by real‐time PCR. The obtained results showed that salsolinol infused at the higher dose significantly (p < 0.001) increased plasma prolactin concentration in lactating ewes, when compared with the concentration noted before the infusion and with that in lactating controls. In lactating ewes, the relative levels of prolactin mRNA expression in the AP and MBH were up to twofold and fivefold higher respectively than in non‐lactating ewes (p < 0.05). In our experimental design, salsolinol did not significantly affect the ongoing process of prolactin gene expression in these tissues. We conclude that in ewes, salsolinol may be involved, at least, in the process of stimulation of prolactin release during lactation and that hypothalamic prolactin plays an important role in the central mechanisms of adaptation to lactation.     

Arid-Zone adaptations of waders (Aves: Charadrii)

Waders (birds of the suborder Charadrii) have radiated into the arid zones of the world through the successful use of adaptations formerly evolved in response to a marine or brackish shoreline, habitat These preadaptations are traced through the rnajor lines of wader radiation, ending in such highly specialized taxa as the sandgrouse (family Pteroclidae), which refined existing adaptations or evolved further adaptations to suit them to a diet of seeds and the need to drink in.an environment poor .in surface water.     

Different origins of two corpora lutea recovered from a pregnant African elephant (Loxodonta africana)

Elephant ovaries contain multiple corpora lutea (CLs) throughout pregnancy. Two CLs (P‐1 and P‐2) collected from a pregnant African elephant were used to investigate their origin and physiological state in this study. The mRNA expressions of prolactin receptor, CYP11A and inhibin betaB subunit were higher in P‐2 than in P‐1, while LHCGR and inhibin betaA subunit mRNA were higher in P‐1 than in P‐2. Protein expression of cleaved caspase‐3 was detected in P‐1 but not in P‐2. These results suggest different origins for the two CLs in this one pregnant elephant, and we also demonstrated the production of bioactive prolactin by the elephant placenta.     

Physiological and hormonal aspects in female domestic pigeons (Columba livia) associated with breeding stage and experience

The present study examined the changes in serum biochemical values, hormone profiles and ovary prolactin receptor (PRLR) gene expression occurring in female domestic pigeons (Columba livia) under different breeding status and experience. The egg‐laying pigeons had lower calcium, total protein, albumin, prolactin levels and higher oestrogen levels than those of incubating birds (p < 0.05). First‐time breeders had higher (p < 0.05) progesterone levels and lower (p < 0.05) prolactin levels than that of experienced ones. The levels of oestrogen and follicle‐stimulating hormone (FSH) increased with age (p < 0.05). The very old birds showed a pronounced increase (p < 0.05) in PRL, FSH and progesterone and a little decrease in oestrogen. Serum prolactin level was not correlated with the ovary PRLR mRNA expression pattern among all the pigeons. Results showed that serum physiological profile of female pigeons was correlated with breeding status, whereas reproductive hormone levels were correlated with advancing breeding experience. It was concluded that female pigeons had a good ability of recovering from nutrient loss after each breeding attempts, and the degradation of reproductive performance might be attributed to changes in the endocrine system.     

Effect of different blood‐guided conditioning programmes on skeletal muscle ultrastructure and histochemistry of sport horses

The effects of three different blood‐guided conditioning programmes on ultrastructural and histochemical features of the gluteus medius muscle of 2‐year‐old sport horses were examined. Six non‐trained Haflinger horses performed three consecutive conditioning programmes of varying lactate‐guided intensities [velocities eliciting blood lactate concentrations of 1.5 (v_1.5), 2.5 (v_2.5) and 4 (v₄) mm respectively] and durations (25 and 45 min). Each conditioning programme lasted 6 weeks and was followed by a 5‐week resting period. Pre‐, post‐ and deconditioning muscle biopsies were analysed. Although training and detraining adaptations were similar in nature, they varied significantly in magnitude among the three different conditioning programmes. Overall, the adaptations consisted in significant increases in size of mitochondria and myofibrils, as well as a hypertrophy of myofibrillar ATPase type IIA muscle fibres and a reduction in number of type IIx low‐oxidative fibres. Together, these changes are compatible with a significant improvement in both muscle aerobic capacity and muscle strength. The use of v_1.5 and v_2.5 as the exercise intensities for 45 min elicited more significant adaptations in muscle, whereas conditioning horses at v₄ for 25 min evoked minimal changes. Most of these muscular adaptations returned towards the pre‐conditioning status after 5 weeks of inactivity. It is concluded that exercises of low or moderate intensities (in the range between v_1.5 and v_2.5) and long duration (45 min) are more effective for improving muscle features associated with stamina and power in sport horses than exercises of higher intensity (equivalent to v₄) and shorter duration (25 min).     

Efficacy of Ovsynch Protocol with Antiprolactin Treatment for Timed Artificial Insemination During Non‐breeding Seasons in Yaks (Poephagus grunniens L.)

An attempt was undertaken to investigate the efficacy of Ovsynch protocol for timed artificial insemination (TAI) with or without Norprolac (antiprolactin) treatment during non‐breeding season (winter months) in yaks (n = 25). During non‐breeding season, plasma prolactin profile has been reported high due to cold and nutritional stress. The Norprolac dose of i.m. administration was standardized for prolactin suppression. Three different doses viz. 2.5, 5.0 and 7.5 mg were attempted and the dose of 7.5 mg Norprolac i.m. per animal was found to be suitable for suppression of prolactin levels up to 30 h. Ovsynch treatment with Norprolac induced more number of oestrous symptoms per animal (4.8 vs 2.1), higher LH peak concentration (24.01 vs 16.16 ng/ml), longer duration of LH surge (6.8 vs 5.2 h) and higher conception rate (70 vs 30%) in Ovsynch plus Norprolac treated animals compared with animals treated with Ovsynch alone. Therefore, this study clearly indicates the opportunity for practical application of the Ovsynch plus Norprolac protocol for TAI in yaks during non‐breeding seasons.     

Immunohistochemical localization of steroidogenic enzymes and prolactin receptors in the corpus luteum and placenta of spotted seals (Phoca largha) during late pregnancy

To study luteal function in the late gestational period of Phocidae (seals), we analyzed the localization of steroidogenic enzymes (P450scc, 3betaHSD and P450arom) and prolactin receptors in the corpora lutea of pregnant spotted seals (Larga seal; Phoca largha) immunohistochemically. P450scc, 3betaHSD and prolactin receptors were present in all luteal cells of each corpus luteum, and most luteal cells were immunostained for P450arom. Although we analyzed only two specimens, P450scc, 3betaHSD and prolactin receptors were negatively immunostained in the placentae. P450arom was present in the syncytiotrophoblast of placentae. These findings suggest that 1) the corpus luteum of the spotted seal synthesizes pregnenolone, progesterone and estrogen during late gestational period, 2) the placenta of this species do not possess the capacity to synthesize progesterone, and 3) like other terrestrial carnivores, this species requires prolactin to maintain the corpus luteum during pregnancy. These characteristics support the recent classification of family Phocidae in the order Carnivora, and suggest a relationship between prolactin and reproductive failure during the post-implantation period in pinnipeds.     

L‐DOPA attenuates prolactin secretion in response to isolation stress in Holstein steers

To clarify endocrine responses to psychological stressors in cattle, the effects of isolation from familiar peers on plasma prolactin (PRL) and cortisol (CORT) concentrations, and the effect of 3,4‐dihydroxy‐L‐phenylalanine (L‐DOPA), a precursor of dopamine (DA), on stress‐induced PRL secretion were determined in Holstein steers. First, the potency of peripheral L‐DOPA administration on attenuation of central DA levels was confirmed. Cerebrospinal fluid (CSF) collected from a chronic cannula in the third ventricle and plasma were sampled 1 h before and 3 h after intravenous injection of L‐DOPA (100 mg/head). DA concentrations in CSF increased just after L‐DOPA injection with subsequent decrease in PRL secretion. Injection of L‐DOPA increased CORT secretion. Second, one experimental steer was isolated in its stall by removing its peers for 2 h with or without‐ pre‐injection of L‐DOPA. The concentration of PRL was elevated by isolation treatment, whereas the effect of isolation on CORT concentration could not be detected. The increase in PRL concentration after isolation was abolished by pre‐injection of L‐DOPA. These results suggest that PRL responds to isolation and that DA neurons in the central nervous system may regulate stress‐induced PRL secretion in steers.     

Plasma prolactin concentrations in broody turkeys: Lack of agreement between homologous chicken and turkey prolactin radioimmunoassays

1. Plasma concentrations of prolactin, measured using homologous chicken or turkey prolactin radioimmunoassays, increased 2‐ and 9‐fold, respectively in somatically mature turkey hens transferred from short (6 h light (L) :18 h dark (D)) to long (14 L:10 D) photoperiods.

2. A further increase in plasma prolactin was observed in laying hens 2 to 3 weeks before the onset of incubation. This increase was detected using the turkey prolactin assay but not the chicken prolactin assay.

3. The onset of incubation was not always associated with the cessation of egg production. Egg laying stopped in incubating hens only after plasma concentrations of prolactin were approaching their maximal values. These were 10‐fold higher than the prolactin concentrations at the onset of lay.

4. The removal of clutches of eggs from incubating hens resulted in a decrease in plasma concentrations of prolactin, which was associated with the cessation of incubation in two hens which had been incubating for 52 and 47 d. The decrease was only temporary in two hens which had been incubating for 8 and 32 d. The latter two hens continued to show incubation behaviour although they were not sitting on eggs.

5. In four turkey hens which did not become broody, egg production ceased after 12 weeks exposure to 14 L: 10 D. The concentration of plasma prolactin, as measured using the turkey prolactin assay, increased 2‐fold during the 2 weeks before these hens stopped laying.     

In vivo investigation of luteal function in dogs: Effects of cabergoline,a dopamine agonist,and prolactin on progesterone secretion during mid-pregnancy and -diestrus

The role of prolactin on luteal function in dogs was investigated in vivo. The function of prolactin in mid-luteal phase was compared in pregnant and nonpregnant dogs. A dopamine agonist, cabergoline, known for its prolactin secretion inhibitory effects, was injected subcutaneously at a dose of 5 μg/kg body weight in five pregnant and five nonpregnant Beagle bitches. Mean plasma prolactin and progesterone were dramatically suppressed for 4 to 5 days after injection in both groups when compared with control pregnant and non-pregnant animals, whereas no effect on luteinizing hormone (LH) secretion was observed. The decline in plasma progesterone occurred after that in prolactin, suggesting plasma progesterone was impaired by inhibition of prolactin secretion. These results confirm the luteotropic importance of prolactin in pregnant bitches, and also demonstrate its importance in luteal phase of the nonpregnant dog.Second, to demonstrate that the effects of cabergoline were mediated by prolactin inhibition and not by a direct action on the corpus luteum, concomitant administration on Day 30 of cabergoline and prolactin (375 μg IV twice daily on Days 30 and 31) or cabergoline and LH (750 μg IV twice daily on Days 30 and 31) was affected in two groups of five pregnant animals each. Results showed that only prolactin was able to reverse the negative effects of cabergoline on circulating progesterone. This confirms the indirect mode of action of the dopamine agonist, cabergoline on corpus luteum function.Third, further investigation on the precise luteotropic role of prolactin was made by IV injection of 375 μg pure canine prolactin twice daily in five pregnant bitches on Days 30 and 31, and in five pregnant bitches on Days 40 and 41. No direct stimulatory effect of prolactin on plasma progesterone secretion occurred. Nor was there a noticeable effect on plasma LH secretion. These results suggest that prolactin is unable to directly stimulate progesterone secretion by the corpus luteum of pregnancy.The results of this study suggest that prolactin is an essential luteotropin in the dog from mid-luteal phase in both pregnant and nonpregnant animals. However, it appears to act by sustaining corpus luteum lifespan and function rather than by direct stimulatory effects on progesterone secretion.     

Induced Lactation with a Dopamine Antagonist in Mares: Different Responses between Ovariectomized and Intact Mares

The aim of this study was to compare the effects of treatment with repeated injections of sulpiride (a dopamine D₂ antagonist) on prolactin secretion and induced lactation in ovariectomized and intact adult mares and to verify if this induction was possible at the beginning and at the end of the birth season. Two experiments were carried out in September [experiment (expt) 1], and in March (expt 2), in France (48°N). In expt 1, three groups of five mares were tested: intact‐control, intact‐treated and ovariectomized‐treated mares. In expt 2, mares previously subjected to artificial photoperiod were assigned in two groups: four intact‐control and five intact‐treated mares. The cyclicity of intact mares was previously synchronized with PGF_2α injections, then all the mares were in the follicular phase at the beginning of treatment. Sulpiride was intramuscularly injected (0.5 mg/kg of BW), twice a day. Mares were milked at 7:30, 11:45, 16:00 and 20:15 hours. Blood samples were collected every day during the treatment for progesterone, total oestrogen and prolactin assays. In the two experiments, only treated intact mares produced milk, with a large inter‐animal variability. Prolactin increase after sulpiride treatment was not so great in the ovariectomized‐treated mares as in the intact‐treated mares. The total correlations between prolactin, progesterone, oestrogen plasma concentrations and daily milk production were significant (0.57, 0.25, 0.17 respectively). This induction of lactation can be performed during the entire birth season in intact mares, but not in ovariectomized mares, indicating that steroids are necessary for this induction in mares treated by dopamine D2 antagonist.     

Feeding sorghum ergot (Claviceps africana) to sows before farrowing inhibits milk production

Objective To assess the impact of feeding different amounts of sorghum ergot to sows before farrowing. Design Fifty‐one pregnant sows from a continually farrowing piggery were sequentially inducted into the experiment each week in groups of four to seven, as they approached within 14 days of farrowing. Diets containing sorghum ergot sclerotia within the range of 0 (control) up to 1.5% w/w (1.5% ergot provided 7 mg alkaloids/kg, including 6 mg dihydroergosine/kg) were randomly allocated and individually fed to sows. Ergot concentrations were varied with each subsequent group until an acceptable level of tolerance was achieved. Diets with ergot were replaced with control diets after farrowing. Post‐farrowing milk production was assessed by direct palpation and observation of udders, and by piglet responses and growth. Blood samples were taken from sows on three days each week, for prolactin estimation. Results Three sows fed 1.5% ergot for 6 to 10 days preceding farrowing produced no milk, and 87% of their piglets died despite supplementary feeding of natural and artificial colostrums, milk replacer, and attempts to foster them onto normally lactating sows. Ergot inclusions of 0.6% to 1.2% caused lesser problems in milk release and neo‐natal piglet mortality. Of 23 sows fed either 0.3% or 0.6% ergot, lactation of only two first‐litter sows were affected. Ergot caused pronounced reductions in blood prolactin, and first‐litter sows had lower plasma prolactin than multiparous sows, increasing their susceptibility to ergot. Conclusion Sorghum ergot should not exceed 0.3% (1 mg alkaloid/kg) in diets of multiparous sows fed before farrowing, and should be limited to 0.1% for primiparous sows, or avoided completely.     

催乳素受体的研究进展

介绍催乳素受体的结构、功能,催乳素受体基因的定位、表达及调节,催乳素受体基因的多态性与生产性状和疾病的关系等。提示催乳素受体基因可作为生产性状及疾病的一个候选基因。     

The effect of xylazine on basal serum prolactin concentrations in male rats

In this study the effects of the major tranquilizer, xylazine, on serum prolactin level were studied in male rats. The results showed that, unlike all other tranquilizers, xylazine produced significant decrease (P less than 0.05) in serum prolactin levels. Xylazine at doses of 2.5, 5, and 10 mg/kg body weight (b.w.) induced significant decrease in serum prolactin levels 1 h after its intramuscular injection which remained significantly low for 4 h. The inhibition produced by 5 mg/kg and the 10 mg/kg b.w. doses was observed to be stronger in both magnitude and duration than that produced by the lower dose (2.5 mg/kg). This inhibitory effect of xylazine on prolactin levels may be attributable to its dopaminergic and/or alpha adrenergic antagonist activity. However, this point needs further investigation.     

Histological Assessment of Selected Blood Vessels of the Phocid Seals (Northern Elephant and Harbour Seals)

Phocid seals exhibit vascular adaptations that allow them to undertake prolonged deep dives. These vascular adaptations are either unique to phocids, or are modified vascular equivalents to those present in terrestrial mammals. One such adaptation, the aortic bulb, is a spherical enlargement of the ascending aorta specific to phocid seals. Its histological make‐up consists of a reinforced tunica media with circular and longitudinal layers of elastic fibres. This reinforcement enables multi‐axial deformation of the aortic bulb, thus complementing its function as a prominent elastic reservoir or ‘windkessel’. A second adaptation, the hepatic sinus, is an asymmetrical dilation of the abdominal portion of the caudal vena cava and accompanying hepatic veins. The hepatic sinus is comprised of a relatively thin tunica media, with a scant smooth muscle component. The bulk of the sinus wall is comprised of tunica adventitia. A third vascular adaptation distinctive to the phocids is the pericardial venous plexus, composed of convoluted veins circumnavigating the perimeter of the heart. Microscopically, these veins have a thick tunica media and also contain valves. Smaller arteries, venules and distinct capillary beds are observed interspersed in‐between these veins. It can be hypothesized, that in seals, certain vascular embryonic development may be arrested at an earlier embryonic stage, resulting in these unusual vascular formations. These modifications play a vital role in blood pressure regulation and distribution of oxygenated blood during prolonged deep diving. The purpose of this work was to elucidate the histological aspects of these unique vascular modifications and relate them to specific function.     

Growth and the plasma concentrations of growth hormone and prolactin in chicks: Effects of “environmental enrichment”, sex and strain

1. The effects of including novel objects in the environment ("environmental enrichment “) on body‐weight gain, relative body‐weight gain, gain : food ratio, plasma growth hormone (GH) and prolactin concentrations in male and female broiler and layer chicks was investigated.

2. Environmental enrichment improved body‐weight gain, relative body‐weight gain and gain: food ratio but had no effect on circulating GH or prolactin concentrations.

3. Weight gain and gain : food ratio were greater in the broilers than in the layer chicks, while plasma GH and prolactin (females only) concentrations were less.

4. There were no sex differences in weight gain and relative weight gain but gain : food ratio was significantly greater in females than in males. In both strains plasma GH concentrations were higher in males than in females and prolactin concentrations were higher in male than in female broilers.     

Effects of prostaglandin F2 alpha on prolactin secretion in the fowl

The plasma concentration of prolactin in immature cockerels was increased between 10 and 40 min after the intravenous administration of prostaglandin (PG) F2 alpha (200 micrograms/kg body weight). Lower doses had no effect on plasma prolactin concentration. The addition of PGF2 alpha (10(-9) to 10(-6) M) to incubation media had no effect on the basal release of pituitary prolactin but reduced the release of prolactin from pituitary-hypothalamus co-incubations. The addition of noradrenaline (10(-7) M), serotonin (10(-7) M), acetylcholine (10(-6) M) or histamine (10(-6) M) to the co-incubation increased the hypothalamus-induced prolactin release, although these effects were not observed in the presence of 10(-7) M PGF2 alpha. The in vitro release of pituitary prolactin was increased by adding chicken hypothalamic extract in the presence or absence of PGF2 alpha. These results suggest a dual effect on PGF2 alpha of prolactin secretion in the fowl; its stimulation in vivo may result from a peripheral action.     

Hypothalamic dopamine is required for salsolinol‐induced prolactin secretion in goats

The aim of the present study was to clarify the relationship between hypothalamic dopamine (DA) and salsolinol (SAL) for the secretion of prolactin (PRL) in goats. SAL or thyrotropin‐releasing hormone (TRH) was intravenously injected into female goats treated with or without the D₂ DA receptor antagonist haloperidol (Hal), which crosses the blood‐brain barrier, and the PRL‐releasing response to SAL was compared with that to TRH. PRL‐releasing responses to SAL, Hal, and Hal plus SAL were also examined after a pretreatment to augment central DA using carbidopa (Carbi) and L‐dopa. The PRL‐releasing response to Hal alone was greater than that to SAL or TRH alone. The PRL‐releasing response to Hal plus SAL was similar to that of Hal alone. In contrast, the PRL‐releasing response to Hal plus TRH was greater than that to TRH or Hal alone. The treatment with Carbi plus L‐dopa inhibited SAL‐ and Hal‐induced PRL secretion. The inhibition of the PRL‐releasing response to SAL disappeared when SAL was injected with Hal. These results indicate that the mechanisms underlying the SAL‐induced PRL response differ from those of TRH, and suggest that hypothalamic DA and its synthesis is associated in part with SAL‐induced PRL secretion in goats.