Endocrine Changes after Mating in Pregnant and Non-Pregnant Llamas and Alpacas

Plasma concentrations of oestradiol-17ß, progesterone, 15-keto–dihydro–PGF_2α and luteinizing hormone (LH) were monitored in llamas and alpacas after mating with an intact male. Concentrations of LH and PGF_2α metabolite were high immediately after copulation. Ovulation occurred in 92% of the animals. The first significant increases in progesterone were recorded on day 4 after mating. In non-pregnant animals the lifespan of the corpus luteum was estimated to be 8–9 days. Luteolysis occurred in association with the release of PGF_2α. In pregnant animals, a transient decrease in progesterone concentrations was observed between days 8 and 18 in both species. No significant changes in PGF_2α secretion were registered during this period. Oes– tradiol–17ß concentrations were high on the day of mating, declined to low values on day 4, and started to increase again on day 8. Peak values after luteolysis in non-pregnant animals were significantly higher than those registered in pregnant ones. Furthermore, concentrations of oestradiol-17ß were elevated for a longer period in non–pregnant than in pregnant animals. The results suggest that progesterone from the corpus luteum exerts a negative influence on follicular activity in pregnant animals by reducing oes– tradiol-17ß secretion.     

Relationship between the somatic cell count in milk and reproductive function in peripartum dairy cows

The aim of the present study was to examine the effect of the somatic cell count (SCC) in milk on reproductive performance, such as pregnancy status in the prepartum period and ovarian function in the postpartum period, in dairy cows. Blood samples were collected every week from one month prepartum to parturition in order to measure the concentrations of 13,14-dihydro-15-keto-PGF_2α (PGFM), estrone sulfate (E1S) and progesterone. Milk samples were collected three times per week in both the prepartum (for one month before the dry period) and postpartum periods (for 3 months immediately after parturition) to measure the SCC. Progesterone was also determined in the whole milk of postpartum cows to define the day of the first ovulation. In the prepartum period, the maximum SCC negatively correlated with the pregnancy period (r = –0.77), but not the calf birth weight. Positive and negative correlations were observed between the average SCC and PGFM or progesterone concentrations in plasma, respectively (r = 0.84 or –0.92, respectively), at 39 weeks of pregnancy. In the postpartum period, a correlation was observed between the day of the first ovulation and both the average and maximum SCC (r = –0.74 and –0.75, respectively), whereas days open was not related to the SCC. These results suggest that a high SCC in the prepartum period may advance parturition by increasing PGF_2α and decreasing progesterone and that the first ovulation in the postpartum period was affected by a high SCC.     

Uterine activity after ovariectomy in the camel (Camelus dromedarius): effect of exogenous administration of oestrogen and progesterone

During oestrous cycles of the camel, spontaneous uterine contractions were correlated significantly with plasma oestradiol-17β concentration. Ovariectomy in the camel resulted in a decreased plasma concentration of oestradiol-17β (<15 pg ml⁻¹) and progesterone (<0·1 ng ml⁻¹) and caused complete cessation of uterine activity. Daily administration of oestradiol benzoate (5 mg, intramuscularly) increased the plasma concentration of oestradiol-17β (>45 pg ml⁻¹) and increased the frequency and amplitude of uterine activity. Coadministration of progesterone (100 mg, intramuscularly) increased the plasma concentration of progesterone (>4 ng ml⁻¹) and increased the frequency but not amplitude of uterine activity. It is suggested that uterine activity in the camel is correlated with the circulating levels of oestradiol-17β and progesterone.     

Sequential Hormonal Changes in the Postpartum Dairy Cow

Peripheral plasma levels of 15-keto-13,14-dihydro-PGF_2α, progesterone, Cortisol, LH and prolactin were studied in 6 primiparous postpartum dairy cows. The cows were followed by hormone measurements and clinical examinations from parturition until pregnancy was established. Blood was collected 3 times per day. The cervix, uterus and ovaries were examined by rectal palpation at 6–10 days intervals. The cows were observed for signs of oestrus twice daily and were additionally teased with a bull to provoke standing heat.Four cows had a normal parturition and dropped their fetal membranes shortly afterwards. (NR group). The remaining 2 retained their fetal membranes for more than 24 h following parturition (RFM group). One out of 6 cows showed standing oestrus at the first ovulation, 4 animals were in oestrus at the second ovulation and all cows showed signs of oestrus at the third ovulation. Although the length of the first luteal phase varied from 9 to 22 days a corpus luteum was in all cases palpated. The secretion of progesterone during the first luteal phase was terminated by a PGF_2α release.A significant difference in 15-keto-13,14-dihydro-PGF_2α levels between the 2 groups was found on days 0–4 (2.39 vs 6.87 nmol/1 at Ρ < 0.06). Postpartum prostaglandin F_2α release as reflected by the level of 15-keto-13,14-dihydro-PGF_2α lasted shorter in the NR group than in the RFM group (15–17 vs 21 days). Significant positive correlations between 15-keto-13,14-dihydro-PGF_2α and Cortisol as well as between prolactin and Cortisol during the first 24 days postpartum were noted only in cows having normal parturition. The most pronounced daily prolactin variations occurred during the second luteal phase (NR group), when a significant difference between the times 8.00, 12.00 and 15.00 was recorded (14.7, 31.5 and 19.7 μg/l, respectively). Moreover, a partial negative correlation between log value of prolactin and arithmetical value of LH was found in these cows only during the first luteal phase after parturition.     

Effect on abortion of feeding Korean pine needles to pregnant Korean native cows

We studied the frequency and timing of abortion and the serum levels of 17β-estradiol and progesterone in Korean native cows fed pine needles during pregnancy. Fifteen pregnant cows were randomly assigned to groups of 5. The control group was fed a concentrate and rice straw, and the other 2 groups were fed, in addition, either 1.3 or 2.7 kg (dry weight) of Korean pine needles daily, starting at an average of 91 d of gestation and continuing until 245 d of gestation. The health status of the dams and the viability of the fetuses were ascertained by rectal palpation and ultrasound scanning during pregnancy. Simultaneously, blood samples were collected for analysis of serum 17β-estradiol and progesterone. Two abortions in mid-pregnancy (at 126 and 150 d of gestation) occurred in the group ingesting the higher daily amount of pine needles. Premature parturition occurred at 259 d of gestation in this group and at 241 and 252 d of gestation in the group ingesting the lower daily amount of pine needles. The serum 17β-estradiol concentration was significantly higher (P < 0.05) at 4 mo of gestation and the serum progesterone level significantly lower (P < 0.05) at 8 mo of gestation in the group ingesting more pine needles daily than in either of the other 2 groups. These results suggest that the ingestion of pine needles may play a role in abortion in Korean native cows by increasing the serum 17β-estradiol concentration and decreasing the serum progesterone concentration.     

Effect of β-carotene Supplementation on Italian Trotter Mare Peripartum

When the mare’s estrous cycle resumes in winter, the β-carotene content of hay is depleted. Sixty Italian trotter mares were randomly assigned to a Control or a Treated Group. Treated Group received 1g/d synthetic β-carotene for 15 days from parturition. Blood samples collected at parturition and on days 5, 10 and 15 after partum were analysed for β-carotene, vitamins A, progesterone, 17 β-estradiol, the energy parameters (glucose, cholesterol, NEFA), the protein profile (total protein, albumin, urea) and LDH. Some changes in these measures were attributable to treatment, which significantly affected β-carotene and 17 β-estradiol concentrations. A significant effect was also found on the resumption of estrous activity (χ² test=P<0.052).     

Plasma progesterone in reindeer in relation to ovariectomy and hysterectomy

Four semi-domesticated reindeer were ovariectomized and 3 both ovariectomized and hysterectomized, 30-70 days before expected parturition. Anaesthesia with etorphine/propyonylphenthiazine was followed by an increase in the plasma concentration of progesterone of peripheral blood. The concentration of progesterone returned to pre-anaesthetic values before ovariectomy. Ovariectomy did not result in any substantial change in the plasma progesterone concentration within an observation period of 35-120 min. Hysterectomy was followed by a marked decrease in peripheral plasma progesterone within 60 min. The progesterone concentration in ovarian venous blood was slightly higher than the jugular venous blood in 1 animal and 150 and 400 times higher in the 2 other animals examined. Thus, despite lack of marked changes of progesterone in jugular venous blood upon ovariectomy, it is concluded that the ovaries are a major production site of progesterone in pregnant reindeer. The progesterone in uterine venous blood was close to that in the jugular vein in 1 animal, and slightly higher in 2 others. This observation, and the maintenance of pregnancy in spite of ovariectomy in 2 out of 4 animals, point to the foeto-placental unit as an additional source of progesterone production.     

Preparation of protein-steroid conjugates and antisera against oestradiol-17

A conjugate of oestradiol-17β and bovine serum albumin (BSA) has been prepared by the mixed anhydride method. The conjugate was characterized by u.v.-analyses and immunoelectrophoresis, and the number of moles of oestradiol-17β per mole of BSA was found to be 25. The conjugate was used for immunization of two sheep and one rabbit in order to elict antisera against oestradiol-17β. Antibodies could be detected in all three animals after 5–16 weeks depending on the route of immunization. The antiserum from one sheep could be used in a radioimmunoassay system for oestradiol-17β in a dilution of 1:5000 and was found to cross-react with oestrone and oestradiol-17α to a lesser extent than another antiserum against oestradiol-17β.     

Estrone and Progesterone Plasma Levels of Normal Cows and Cows with Parturient Paresis

Blood plasma concentrations of estrone and progesterone, calcium and inorganic phosphorus were measured in 22 cows (Swedish Red and White Breed) from 4 weeks prepartum up to 6 days post partum. Ten cows with parturient paresis had Ca levels below 6 mg/100 ml. Data from plasma analyses in individual cows were grouped in the following periods: 28–22, 21–15, 14–8, 7–6, 5–4, 3, 2, 1 day(s) before parturition and 1, 2, 3–6 days after parturition. Statistical comparisons of the levels of the hormones and the ratio progesterone/estrone did not reveal any significant differences between the paretic and normal cows at any time period. The results do not support the theories that high systemic levels of estrogens or an imbalance between estrogen and progesterone predispose towards parturient paresis.     

Plasma fluctuation in estradiol-17β and bone resorption markers around parturition in dairy cows

Blood samples were obtained sequentially from 10 dairy cows around the time of parturition to assess plasma fluctuations in estradiol-17β (E₂) levels in association with those of several bone resorption markers. Plasma E₂ concentration increased sharply a few days prepartum and decreased quickly after parturition. In terms of bone resorption markers, the plasma level of tartrate-resistant acid phosphatase isoform 5b (TRAP5b) rose significantly, commencing 1 week prepartum, and was maintained at this level to a few days postpartum. The plasma concentration of carboxyterminal collagen cross-links of type-I collagen (CTx) increased significantly after parturition. These observations suggest that osteoclast-mediated bone resorption was activated after parturition when plasma E₂ concentrations decreased.     

Pregnancy and peripheral plasma progesterone levels in cows inseminated after synchronization of estrus with prostaglandin F2 alpha

Fifteen Holstein cows were treated with two doses of 25 mg of a prostaglandin F₂α (PGF₂α as dinoprost tromethamine) administered intramuscularly 11 days apart. The cows were then divided into three groups and inseminated either at 72, 80 or 72 and 96 hours after the second dose of PGF₂α. Thirteen cows ovulated after the second prostaglandin treatment. Eight cows were diagnosed pregnant by rectal palpation 42 days after insemination but only five calved. PGF₂α induced luteolysis in cows with active corpora lutea as evidenced by the dramatic decreases in the plasma progesterone concentrations after treatment. In contrast, following PGF₂α administration to cows in follicular or late luteal phase the concentrations of plasma progesterone either increased gradually or remained low for several days before increasing to maximal levels. The ovulatory rate after the two doses of PGF₂α11 days apart was high. However, the pregnancy rate after this treatment was influenced by other factors including abnormal ovarian function.     

The effect of a bacterial endotoxin or cloprostenol on the clinical status and hormonal levels in 80-100 days pregnant gilts

An experiment was conducted to examine the effect of a lipopoly-saccharide (LPS) of Salmonella typhimurium on the luteal function in 80 days pregnant gilts. Four animals were i.v. injected with 2 μg LPS/kg body weight and 3 animals were i.m. injected with 500 μg cloprostenol (CP). Gilts which maintained pregnancy after the initial injection were reinjected with GP around day 100. Clinical observations were made and plasma levels of 15-keto-13,14-dihydro-PGF₂α, progesterone, oestradiol-17β and oestrone sulphate were analysed by radioimmunoassay.The LPS induced a characteristic clinical endotoxemia. All LPS treated gilts maintained pregnancy until day 100 when 1 gilt aborted, 1 was emergency slaughtered and 2 reinjected. The comparative injections of CP induced abortion! within 48 h in 2 of 3 gilts at 80 days and in all reinjected animals at 100 days of pregnancy. Progesterone decreased immediately after both LPS and CP injections. In non-aborting gilts, the progesterone decrease had a transient character. The PGF_2α metabolite levels responded to LPS by a dramatic surge of approximately 4 h duration. All abortions were accompanied by a massive release of PGF_2α reaching peak levels during expulsion of the foetuses. Oestradiol-17ß and oestrone sulphate followed an ascendent pattern between days 80 and 100. Occasional transient decreases in oestradiol-17ß or increases in oestrone sulphate levels after LPS and CP injections were observed in several animals. Abortions were followed by a sharp decrease of both oestrogens. Post-abortum reproductive disorders occurred frequently. Endocrine changes associated with post-abortum ovarian activity were relevant to the clinical and morphological observations. The relationship between the stage of pregnancy in the pig and its endocrine response to abortifacient agents as well as some foetopathic effects of the endotoxin are discussed.     

Mammary secretion of oestrogens in the cow

Two experiments in vivo and one experiment in vitro were conduced to examine the mechanisms involved, which lead to mammary secretion of oestrogens and its importance for milk production and udder health in cows. In experiment 1 in six cows of the White-Black breed on day 268 of pregnancy catheters were inserted into uterine vein of pregnant horn, the abdominal aorta and the caudal superficial epigastric (milk) vein. Blood samples for estimation of oestrone, oestrone sulphate, oestradiol-17alpha and -17beta by RIA were obtained daily from day 7 pre-partum until day 1 post-partum. Only the concentration of oestradiol-17beta was statistically higher (P< or =0.01) in mammary venous plasma than in aortal and uterine plasma. In experiment 2 forty late-pregnant cows were divided into two groups according to their milk production in the previous lactation: group 1 (n=20) high-yielding cows (>6500kg milk per lactation), and group 2 (n=20) low-yielding cows (<3700kg milk per lactation). Blood samples for measurement of oestradiol-17beta by RIA were collected from milk and tail veins every fourth day during a period from day 20 prior to parturition to day 4 post-partum. The concentration of oestradiol-17beta was significantly higher (P< or =0.01) in the milk vein than in the peripheral plasma from day 12 pre-partum to parturition. In high-yielding cows the level of oestradiol-17beta in mammary venous blood was significantly higher (P< or =0.01) than in low-yielding cows. In six cows with pathological udder oedema ante-partum the concentration of oestradiol-17beta in milk vein was significantly higher (P< or =0.05) than in control cows. There were no statistically significant differences in the levels of oestradiol-17beta in cows with clinical mastitis (n=10) during 2 weeks after parturition and without it (P> or =0.05). In an in vitro experiment, homogenates of mammary tissue collected on day 7 pre-partum from two cows were incubated with 3H-androstendione. After incubation the samples were extracted and 3H-oestradiol-17beta was separated by HPLC. 3H-oestradiol-17beta was formed in a total yield of 37%. These results indicate that oestrone, oestrone sulphate and oestradiol-17alpha are not secreted by bovine mammary gland. Furthermore, the secretion of oestradiol-17beta starts about day 12 pre-partum and is associated with milk yield and udder oedema. Preliminary in vitro study suggests the synthesis of oestradiol-17beta by mammary tissue.     

Effect of Prolactin Suppression on the Ovarian Activity in the Lactating Sow

The effect of prolactin suppression during lactation was studied in 6 sows nursing 5–10 piglets. In order to inhibit prolactin secretion a dopamin agonist (bromocriptine, Parlodel® “Sandoz”) was given per os from about 4 weeks after parturition until weaning 9–17 days later. Plasma prolactin and plasma progesterone were determined by radioimmunoassays. In all animals plasma prolactin was suppressed during the bromocriptine treatment. Two sows resumed ovarian activity during treatment as indicated by elevated plasma progesterone, probably as a result of the low plasma prolactin levels. The results suggest that the elevated plasma prolactin levels during lactation is one of the factors responsible for lactational an oestrus in the sow.     

Prostaglandin Release at Dexamethasone Induced Parturitions in Cows

Three cows of the Swedish Red and White Breed (SRB) were used in the experiment. Cow no. 1 pregnant for 247 days, was given 20 mg dexamethasone twice with an interval of 48 hrs. Cows nos. 2 and 3, each pregnant for 254 days, received 20 mg of dexamethasone twice with an interval of 24 hrs. The cows delivered normal living calves 153, 138 and 137 hrs., respectively, after the second injection of dexamethasone. Blood samples were drawn from the jugular vein every third hour during the experimental period and the samples were analyzed for estrone, progesterone and 15-keto-13,14-dihydroprostaglandin F₂α. Following the dexamethasone injections there was a continuous increase in the blood plasma levels of estrone followed by a sharp decrease in conjunction with parturition. The blood plasma level of progesterone showed a slow but continuous decrease until about 24 hrs. before delivery when a marked drop occurred. The levels of the prostaglandin metabolite increased gradually until about 24 hrs. prior to delivery. This was followed by an abrupt rise, and high levels of the prostaglandin metabolite were recorded for up to four days following parturition. It is concluded that the estrone increase preceded that of the prostaglandin metabolite and that the final drop in the progesterone was synchronous with the final rise of the prostaglandin metabolite level.     

The Effect of Induced Standing Reflex,Cervical Stimulation and Insemination on Intraluminal Pressure Variations in the Isthmus of the Oviduct in Unrestrained Gilts

Four gilts were each equipped with 2 ultra-miniature pressure sensors, placed at 2 different points along the same isthmus of the oviduct, on the morning of the first day of standing oestrus (Day 1). Intraluminal pressure recordings were started the same afternoon. After an initial recording period, intraluminal pressure was recorded while the gilts showed standing oestrus and during cervical stimulation followed by insemination with either 100 ml saline or 100 ml boar semen. Monitoring of the pressure variations in the isthmus was continued for up to 6.5 h after the last insemination. Blood samples for monitoring oestradiol-17B, progesterone and 15-ketodihydroprostaglandin F_2α were collected before and after each manipulation of the gilt and every 30 min during the rest of the test period. None of the above manipulations had any consistent effect on the intraluminal pressure in the porcine isthmus, although, a clear 15-ketodihydroprostaglandin F_2α peak could be seen after insemination with boar semen.     

Effects of prostaglandin f(2alpha) on corpora lutea formation and function in mated bitches

Fifteen mated bitches were used to study the effects of prostaglandin F_2α on ovarian endocrine function during the early and midluteal phase. Five dogs were kept as controls, five were given 250 μg/kg prostaglandin F_2α twice daily between the first and fifth day of metestrus, and five were similarly treated with prostaglandin F_2α between 31 and 35 days of metestrus. Function of corpora lutea was monitored by measuring serum progesterone concentrations during the first 45 days of gestation.

Dogs treated with prostaglandin F_2α during the early luteal phase had progesterone concentrations similar to controls and pregnancies were undisturbed in both groups. A dramatic decrease in serum progesterone concentration and abortion resulted after prostaglandin F_2α administration at midpregnancy.

These results indicate that prostaglandin F_2α was not luteolytic during the early luteal phase and was therefore ineffective for preventing pregnancy at that time. However, at the dosage and frequency used in this study, prostaglandin F_2α was luteolytic and abortifacient at midgestation.

     

Treatment with Gonadotropin Releasing-Hormone in Prepubertal Gilts at Two Different Ages

To study the effect of GnRH in prepubertal gilts, seven crossbred gilts were treated with saline solution and 250 fig GnRH. In connection with saline and GnRH treatments blood was sampled every 15 min for 4 h, thereafter every 30 min for 2 h and every 60 min for 3 h, and finally every 3 h for 6 days. The ovaries were inspected by laparo-scopy just before and 6 days after GnRH treatment. The first GnRH treatment was undertaken when the gilts had a mean age of 141 days and mean body weight of 66 kg. One gilt was in prooestrus at this treatment. In the other 6 gilts the mean LH level was around 0.5 μg/l during a 4 h period after the saline injection. After the GnRH treatment a LH peak was seen with a mean duration of 4 h and a mean maximum level of 9.2 ± 2.07 μg/1. None of the gilts ovulated or showed oestrus within a week after GnRH treatment, which was confirmed by laparoscopy. The seventh gilt which was in prooestrus had high levels of oestradiol-17β (> 40 pmol/1) at GnRH treatment and no LH peak was seen during a 4 h period after treatment.Two gilts which had not shown oestrus at an age of 173 days and a mean body weight of 93 kg were treated a second time with 250 μg GnRH. The LH peak had a duration of 4 h and a mean maximum level of 5.3 ± 3.04 μg/l. Neither of these 2 gilts showed oestrus or ovulated within a week after GnRH injection. It was concluded that a single injection of GnRH results in a LH peak but is not enough to stimulate ovulation or oestrus in prepubertal gilts at a mean age either of 141 or 173 days.Key words: GnRH-treatment, prepubertal gilts, LH, oestradiol-17β     

Concentrations of Oestrone and Oestradiol-l 7β in the Peripheral Plasma of the Nanny Goat during the Oestrous Cycle, Gestation and Puerperium

The changes in the plasma levels of oestrone sulphate and oestradiol-17β during the oestrous cycle, gestation and puerperium in the goat are described. The oestrone sulphate concentrations remained fairly constant (250–350 pg/ml) throughout the oestrous cycle until day 20 when a sharp increase of the oestrone sulphate plasma levels occurred in pregnant goats which became significantly different at day 38 of gestation from nonpregnant values. Oestradiol-17β plasma levels were significantly lower at days 17–20 in pregnant than in nonpregnant does. Oestrone sulphate and oestradiol-17β concentrations rose until the 12th week of gestation and then declined to about 50% of the former ranges of concentrations before rising again to high values at weeks 17–20 of gestation. Increasing plasma levels of oestrone sulphate and oestradiol-17β were determined during the last ten days preceding parturition. The concentrations of oestrone sulphate returned to basal levels by the 2nd—4th day post partum whereas oestradiol-17β values reached base values 24 hours after parturition. Both oestrogen concentrations remained constant during the puerperium until day 51 post partum .     

Endocrine and Ovarian Changes in Response to the Ram Effect in Medroxyprogesterone Acetate-primed Corriedale Ewes During the Breeding and Nonbreeding Season

Two experiments were performed to determine the endocrine and ovarian changes in medroxyprogesterone acetate (MAP)-primed ewes after ram introduction. Experiment 1 was performed during the mid-breeding season with 71 ewes primed with an intravaginal MAP sponge for 12 days. While the control (C) ewes (n = 35) were in permanent contact with rams, the ram effect (RE) ewes (n = 36) were isolated for 34 days prior to contact with rams. At sponge withdrawal, all ewes were joined with eight sexually experienced marking Corriedale rams and estrus was recorded over the next 4 days. The ovaries were observed by laparoscopy 4–6 days after estrus. Four weeks later, pregnancy was determined by transrectal ultrasonography. In eight ewes from each group, ovaries were ultrasonographically scanned; FSH, LH, and estradiol-17β were measured every 12 hours until ovulation or 96 hours after estrus. The response to the rams was not affected by the fact that ewes had been kept or not in close contact with males before teasing. No differences were found in FSH, LH, estradiol-17β concentrations, growth of the ovulatory follicle, onset of estrus, ovulation rate, or pregnancy rate. Experiment 2 was performed with 14 ewes during the nonbreeding season. Ewes were isolated from rams for 1 month, and received a 6-day MAP priming. Ovaries were ultrasonographically scanned every 12 hours, and FSH, LH, estradiol-17β, and progesterone were measured. Ewes that ovulated and came into estrus had higher FSH and estradiol-17β levels before introduction of the rams than did ewes that had a silent ovulation. The endocrine pattern of the induced follicular phase of ewes that came into estrus was more similar to a normal follicular phase, than in ewes that had a silent ovulation. The follicle that finally ovulated tended to emerge earlier and in a more synchronized fashion in those ewes that did come into estrus. All ewes that ovulated had an LH surge and reached higher maximum FSH levels than ewes that did not ovulate, none of which had an LH surge. We conclude that (a) the effect of ram introduction in cyclic ewes treated with MAP may vary depending on the time of the breeding season at which teasing is performed; (b) patterns of FSH, and estradiol-17β concentrations, as indicators of activity of the reproductive axis, may be used to classify depth of anestrus; and (c) the endocrine pattern of the induced follicular phase, which is related to the depth of anestrus, may be reflected in the behavioral responses to MAP priming and the ram effect.