Effect of rumen degradable bolus containing melatonin or progestagen pessary plus pregnant mare serum gonadotropin on estrus response and lambing rate in ewes

Two practical regimens designed to induce estrus and ovulation in ewes in late anestrus were compared. Forty ewes were given a soluble glass rumen bolus containing 150 mg melatonin on July 9 and were joined with two vasectomized rams on July 23 and with three fertile rams on August 6. A second group of 40 ewes was treated with an intravaginal progestagen pessary (60mg medroxy-progesterone acetate) on July 23. Following pessary removal after 12 d, ewes were given 750 IU of pregnant mare serum gonadotropin (PMSG). Five fertile rams were joined with these ewes 48 h after progestagen removal. Melatonin concentrations were determined in single blood samples collected in early afternoon of July 21. Mating dates, lambing dates and litter sizes were recorded. Date of mating was significantly later in ewes treated with melatonin compared with those treated with progestagen plus PMSG (P<0.0001). All ewes given melatonin were mated within 4 wk, and those on progestagen plus PMSG treatment within one day of fertile ram introduction. Thirty-four ewes (85%) allocated to melatonin treatment and 36 (90%) allocated to progestagen plus PMSG treatment lambed (P>0.05). Mean (+/-SEM) lambing date was later in melatonin-treated ewes (January 17+/-1.2 d) compared to those given progestagen plus PMSG (December 30+/-0.6 d; P<0.0001). Mean litter size was lower in melatonin-treated ewes (1.5+/-0.1) compared with those given progestagen plus PMSG (2.0+/-0.1; P<0.001). Plasma melatonin concentrations indicated that 9 of 40 ewes treated with melatonin had circulating melatonin concentrations of less than 16 pg/ml. It is concluded that under conditions that existed in this experiment, treatment with progestagen plus PMSG in late anestrus resulted in more synchronous mating and lambing and a higher litter size than that following administration of a soluble glass rumen-degradable bolus containing melatonin.     

Influence of classification levels of ram sexual activity on spring breeding ewes

Ram lambs (7–8 months old) and mature rams (19–20 months old) were used to evaluate the effect of classification levels of male sexual performance on reproductive performance of ewes during spring breeding. In Exp. 1, sexually active ram lambs with high (1.8±0.3; n=5) and low (0.9±0.2; n=5) sexual performance scores (HP and LP; mean±S.E.M.) were used in single sire breeding pens. Ewes (n=305) were stratified by age and assigned to 10 pens for 34 days starting in late March. For Exps. 2 and 3, two replicates were conducted for 2 years with sexually active mature rams in a single sire mating scheme. For Exp. 2, HP rams (n=5) averaged 3.6±0.2 ejaculations and LP rams (n=7) 1.8±0.2 ejaculations for sexual performance scores based on nine, 30 min serving capacity tests (SCT). Polypay ewes (n=152 to 153 per year) were stratified by age and assigned to pens each year for 34–38 days starting in late March for years 1 and 2. For Exp. 3, HP rams (n=6) averaged 3.7±0.1 ejaculations and LP rams (n=10) 2.3±0.1 ejaculations for sexual performance scores based on 18, 30 min SCT. Polypay ewes (n=229 in year 3 and n=244 in year 4) were stratified by age and assigned to pens each year for 34 days starting in late March. In Exp. 1, lambing rates for ewes bred to HP versus LP ram lambs did not differ (65.8 versus 53.0; P=0.20). Prolificacy tended (P=0.06) to be increased by 0.1 lambs in ewes bred by LP ram lambs. Total number of lambs born per ewe present at lambing, and lambing distribution were not altered by HP and LP ram lambs. In Exp. 2, lambing rates for fall-lambing ewes bred to mature HP or LP rams did not differ (58.1 versus 60.1; P=0.78). In Exp. 3, lambing rates for fall-lambing ewes bred to mature HP or LP rams did not differ (74.3 versus 69.0; P=0.35). There was no difference (P>0.10) between years for Exp. 2 or Exp. 3, and mature HP and LP rams did not affect the other reproductive variables monitored. Analyses of the combined data for Exps. 2 and 3 indicated only a year difference (P<0.001) in lambing rates and total lambs born. Present studies indicate that different sexual performance classifications for ram lambs and mature rams did not alter lambing rates or distribution of lambing of Polypays bred in late March to April. These results indicate that HP and LP, sexually active, Polypay rams and ram lambs with average to high quality semen can provide a source of rams for spring breeding Polypays in ambient conditions and that there was no advantage to using HP over sexually active LP ram lambs or rams.     

Short term, repeated exposure to rams during the transition into the breeding season improves the synchrony of mating in the breeding season

The ram effect is widely used in Mediterranean breeds of sheep but its use in temperate genotypes is restricted by breed seasonality. However, ewes from these highly seasonal genotypes are sensitive to stimulation by rams close to the onset of the natural breeding season. In this study we developed a pre-mating protocol of repeated, short-term exposure to rams (fence-line contact or vasectomised rams) beginning during late anoestrus and continuing into the breeding season. We hypothesised that this pre-mating protocol would synchronise the distribution of mating of North of England Mule ewes during the breeding season above that observed in ewes isolated from rams prior to mating. Ram-exposed ewes were given contact with rams (Experiment 1: fence-line; FR, n = 94 and Experiment 2: vasectomised rams; VR; n = 103) for 24 h on Days 0 (10 September), 17 and 34 of the experiment. Control ewes (Experiment 1; FC, n = 98 and Experiment 2; VC; n = 106) remained isolated from rams prior to mating. In Experiment 2, a subset of VR (n = 35) and VC ewes (n = 35) were blood sampled twice weekly to monitor their pre-mating progesterone profiles. At mating, harnessed entire rams were introduced, 17 or 16 days after the last ram exposure (Experiments 1 and 2) and raddle marks were recorded daily. The median time from ram introduction to mating was reduced in ewes given both fence-line and vasectomised ram contact (P < 0.001), leading to a more compact distribution of mating and lambing (At least P < 0.01). In the blood sampled VR ewes, there was a progressive decline in the number of days from ram exposure to the onset of dioestrus (at least P < 0.05). This observation indicates that the cycles in VR ewes became increasingly synchronised over the pre-mating period, a pattern not evident in VC ewes. In conclusion, repeated, short-term exposure of ewes to rams during the transition into the breeding season is an effective method of synchronising the distribution of mating during the breeding season.     

Importance of behavioural stimuli in ram-induced ovulation in seasonally anovular Corriedale ewes

Three groups of about 65 seasonally anovular Corriedale ewes were penned and exposed to rams on November 4, 1989. A fourth group (n=64) was maintained in a 3-ha paddock and also exposed to rams on the same date. The sexual behaviour exhibited by rams in contact with penned ewes was recorded during nine simultaneous observation sessions lasting 2 h each.

Five days after ram introduction, incidence of ovulation and ovulation rate were higher (P < 0.01) in adult than in 1.5-year-old ewes. Among penned adult ewes, a greater (P < 0.05) incidence of ovulation was recorded in ewes maintained in contact with rams and oestrous ewes (95.8%) than in ewes exposed to either rams with or without recent experience with oestrous ewes (68.7 and 57.1%, respectively). The fourth group, exposed to rams in field conditions, exhibited a lower (P < 0.05) incidence of ovulation (31.9%) than any other group. No differences among treatments were detected in 1.5-year-old ewes. The incidence of ovulation was associated (r=0.99, P<0.05) with the sexual behaviour displayed by the rams.

It was concluded that the sexual behaviour of the rams plays an important role in ram-induced ovulation and that the continuous presence of oestrous ewes is necessary for the maximum efficacy of the technique in Corriedale ewes.     

Effects of lactation status, progestogen and ram exposure on response to cloprostenol in ewes during the anestrous season

Progestogen pretreatment and introduction of rams were used to prepare 432 Rasa aragonesa ewes for synchronization of estrus with prostaglandin (PG) during anestrus. The experiment was a 3 x 3 x 2 factorial with lactation status, ram/progestogen treatment and treatment with pregnant mare's serum gonadotropin (PMSG; 250 IU) at injection of PG as main effects. At ram introduction (Day 0), ewes were dry (Group 1), weaned (Group 2) or suckling a lamb (Group 3). They received either norgestomet implants for 12 d (Days -12 to Day 0 (Group A), ram introduction for 12 d (Days 0 to 12) (Group B) or both (Group C). Half the ewes received PMSG with PG (100 mug Cloprostenol) on Day 12. Pregnancy rate was higher at first service in dry (Group 1; 42%) than in recently-weaned ewes (Group 3; 24% and Group 2; 31%; P<0.01). Occurrence of estrus and conception and pregnancy rates to first service were higher (P<0.01) in ewes previously exposed to rams (Groups B and C) than in ewes treated only with progestogen (Group A). There were no effects of PMSG, no interactions among the three variables and no differences in prolificacy (1.12 at first service).     

A new strategy for superior reproductive performance of ewes bred out-of-season utilizing progestagen supplement prior to withdrawal of intravaginal pessaries

Two experiments were conducted to examine the effect of progestagen supplement 24h prior to intravaginal pessary withdrawal on reproductive performance of seasonal anestrous ewes. Ewes in each experiment were allocated to treatment and control and all were induced to estrus using either intravaginal MAP (Exp. 1; n=24) or CIDR-G (Exp. 2; n=28) pessaries for 12 days. Half of the ewes in each experiment were supplemented 24h before withdrawal of pessaries with either 10mg oral MAP tablets (Exp. 1) or 25mg i.m. progesterone (P(4)) administration (Exp. 2; P(4)-supplement-treated group). Fertile rams were allowed with the ewes at sponge removal (Day 0, 0h) and estrus was monitored at 6-h intervals for 3 days. Blood samples were collected for measurements of P(4) (Exp. 1 and Exp. 2) and LH (Exp. 2). In both experiments, the percent of ewes in estrus was greater (P<0.05) and intervals to estrus were longer (P<0.05) in progestagen-supplement-treated than control ewes. In Exp. 2, the occurrence and magnitude of LH surges were greater (P<0.01) and intervals to onset of LH surge were longer (P<0.01) in P(4)-supplement-treated than control ewes. In Exp. 2, P(4) supplement elevated P(4) levels from 1.8+/-0.1ng/mL on Day -1 to 4.2+/-0.3 on Day 0 (P<0.001). Following pessaries removal, P(4) concentrations fell to basal values on Day 1 in both groups and remained low until Day 5. Then, P(4) concentrations increased and remained elevated through Day 19 in all (100%) progestagen-supplement-treated in Exp. 1 (12/12) and Exp. 2 (14/14) and in only 5/12 (41.7%) and 6/14 (42.9%) control ewes, respectively. These ewes were confirmed pregnant by ultrasonography and lambed on Day 149.2+/-0.2 following Day 0. In conclusion, progestagen supplement 24h prior to removal of pessary can be used successfully to improve reproductive performance of ewes bred out-of-season.     

Embryo production and endocrine response in ewes superovulated with PMSG, with or without monoclonal anti-PMSG administered at different times

Mature nonlactating Altamurana ewes (n = 168) were synchronized in the seasonal anestrus period with FGA-impregnated intravaginal pessaries for 12 d. In Experiment 1, 48 ewes were divided into a 3 x 4 factorial design for anti-PMSG monoclonal antibody (AP) bioassay test. Concomitant injections of PMSG (1000, 1500, 2000 IU) and AP (0, 1, 2, 3 microl/IU PMSG) were given, and ovarian response was evaluated by laparoscopy. In Experiment 2, 120 ewes were divided into 8 experimental groups (n = 15 per group). The ewes treated with 1000 or 1500 IU PMSG at -24 h from sponge removal were given AP intravenously at 50 h after pessary withdrawal, 12 or 24 h after the onset of estrus, while the controls did not receive AP. Blood samples were collected from ewes (n = 6) treated with 1500 IU PMSG with or without anti-PMSG. Ovarian response and embryo production were evaluated on Day 7 after sponge removal upon laparotomy. It was found that 1 microl AP was effective in neutralizing 1 IU PMSG. No significant differences in serum concentrations of progesterone were observed among the groups of superovulated ewes. Estradiol-17 beta levels were reduced following AP treatment 12 h after the onset of estrus. At a lower dosage of superovulatory treatment (1000 IU PMSG), AP injected at 12 or 24 h after the onset of estrus significantly lowered large follicles (P < 0.01) and increased the rate of ovulation (P < 0.05). Moreover, embryo production showed a more than two-fold increase (P < 0.01) of viable embryos following AP injection at 12 or 24 h after the onset of estrus (3.2 to 3.3 vs 1.3, with vs without anti-PMSG). It is concluded that superovulatory treatment with 1000 IU PMSG plus AP administered at a fixed time after the onset of estrus may improve ovarian response and the yield of viable embryos in ewes.     

Ram-induced ovulation to improve artificial insemination efficiency with frozen semen in sheep.

Ram effect, defined as shortening of seasonal anestrus in ewes by exposure to the ram, is now well recognized but the underlying mechanisms are still unclear. Little information also exists whether the ram is able to influence the estrus cycle and ovulation. Three experiments were conducted to investigate endocrine response, time of ovulation and pregnancy rate of ewes in proestrus, exposed to the ram (treated) or an adult ewe (control). In the first experiment, ewes (n = 20) were treated with fluorgestone acetate pessaries for 12 days and were given eCG and cloprostenol one day before withdrawal of pessaries. On the day after removal of the pessaries ewes in the treated group (n = 10) were exposed to the ram and those in the control group (n = 10) were exposed to an adult ewe. Blood samples were taken for LH assay every 20 min from 2 h before to 24 h after ram exposure. In the second experiment, ewes (n = 120) were induced into proestrus and on the day after removal of the pessaries were exposed to either a ram (n = 60) or a ewe (n = 60) as described above and were laparoscoped 50, 60 or 70 h after pessary withdrawal (n = 20 at each time interval). In the third experiment ewes (n = 90) were induced and exposed to the ram (n = 45) or an adult ewe (n = 45) and inseminated via a laparoscope whit frozen-thawed semen at 50 or 60 h after pessary removal, respectively. Exposure to the ram was followed in 2 h by a marked rise in LH, equivalent to a preovulatory surge in duration and amplitude. It was also followed by concentrated ovulation within 25 to 30 h and by an increased pregnancy rate in exposed ewes (73.3 vs. 53.3%).     

Effect of ram exposure at the end of progestagen treatment on estrus synchronisation and fertility during the breeding season in ewes

Two experiments were conducted to examine the effects of ram exposure during the breeding season, in combination with progestagen treatment on estrus synchronization, fertility the LH surge and ovulation in ewes. Experiment 1 was subdivided into experiments 1a and 1b. In all experiments cross-bred ewes were treated with an intravaginal sponge for 12-14 days and three days before sponge withdrawal ewes were divided into control (no further treatment; n=191, 103 and 50 for experiments 1a, 1b and 2, respectively) or ram exposed (three mature rams per 50 ewes were introduced; +Ram; n=187, 99 and 49 for experiments 1a, 1b and 2, respectively). At sponge withdrawal ewes in Experiments 1a and 2 received 500 IU eCG and rams were removed from all the +Ram groups. In Experiments 1a and 1b, raddled, entire rams were introduced to ewes 48 h after sponge withdrawal. The timing of mating was recorded and ewes were maintained until lambing. In Experiment 2, estrus behavior was determined every 4 h and the time of the LH surge and ovulation were determined from a subset of 10 ewes per group. In Experiment 1a, less +Ram ewes were bred by 48 h after ram introduction (control 98% versus +Ram 89%, P<0.001) and in Experiments 1a and 1b 14% fewer (P<0.05) of the ewes bred in the first 3 h after ram introduction lambed to that service. In Experiment 1a, ram exposed ewes had a lower litter size than control ewes (1.93+/-0.06 versus 1.70+/-0.06 lambs per ewe; P<0.05). In Experiment 2, rams advanced (P<0.05) estrus, the LH surge and ovulation by 2-6 h compared with control ewes. We speculate that exposure of ewes to rams increased LH secretion and that this in turn increased follicle development and the production of oestradiol that led to a more rapid onset of estrus, the LH surge and ovulation compared to control ewes. Unexpectedly, ewes that were bred had lower fertility in the +Ram groups than control groups.     

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.     

Induction of estrus and superovulation in seasonally anestrous ewes

The induction of estrus in 17 previously cycling nulliparous ewes, 9 to 10 months of age, was attempted with Medroxyprogesterone acetate (MAP) pessaries during the early anestrous period (March-April). Ewes were verified to be anestrous by the lack of estrous behavior in the presence of a vasectomized ram and by a radioimmunoassay for serum progesterone in two samples taken 7 days apart showing less than 1 ng/ml serum progesterone. Superovulation was attempted with injections of either FSH or FSH + LH. MAP vaginal pessaries remained in place for a period of 12 days and FSH was administered to all ewes (IM) at 12 hr intervals over a 3 day period; 5 mg was injected twice on day 11 after pessary insertion, followed by 4 and 3 mg injections twice daily on each succeeding day, for a total of 24 mg per ewe. Nine ewes were given 25 mg LH (IV) within 8 hrs after the onset of behavioral estrus in addition to FSH. Ewes were hand-mated to several rams at 12 hr intervals throughout the estrus period. Ovulation and fertilization rates were recorded for each ewe following midline laparotomy and embryo collection. All ewes were in estrus between 36 and 48 hrs after removal of the MAP pessaries. In ewes injected with FSH only, 8 of 8 ovulated with a mean ovulation rate of 6.0 +/- 4.4 and a fertilization rate of 70%. Nine of 9 ewes receiving both FSH + LH ovulated with a mean ovulation rate of 13.9 +/- 13.1 and a fertilization rate of 72%. Statistical analysis by Students t-test resulted in differences in number of ova recovered (P<.05) between FSH only and FSH + LH treated ewes and a trend towards increased ovulation rate in FSH + LH treated ewes. These results show that early seasonally anestrous ewes can be successfully induced and synchronized for estrus with MAP pessaries and the number of ova recovered is increased with the inclusion of LH in the superovulation regime.     

Patterns of estrous cycles, estrous behavior, and circulating prolactin in spring and summer in ewes selected for autumn lambing and exposed to ambient or long-day photoperiods

Estrous behavior in response to ambient and long-day photoperiods was evaluated in ewes developed by 10 years of selection for ability to lamb in autumn. Following October lambing, 67 ewes were moved indoors and exposed to long-day (16L:8D) or ambient photoperiods from February 2 until July 6. Two vasectomized rams with marking harnesses were housed with each group. Estrous behavior was monitored twice weekly. Ewes from the selection line were unresponsive to long days, with no effects on estrous behavior, frequency of ovulation, or circulating prolactin. Adult ewes were anestrus for only 34±3 d, but 2- and 3-years-old ewes were anestrus for 72±7 and 57±10 d, respectively. Frequencies of ovulation based on circulating progesterone concentrations in March, May, and June were 97%, 95% and 52%, respectively, indicating that many ewes that did not exhibit estrus still ovulated. Prolactin concentrations increased from 10 ng/ml in February to 27 ng/ml in March and 173 ng/ml in June but were not affected by light treatment. Ten ewes that failed to exhibit estrus behavior for at most 24 d during the main study were then monitored for 74 additional long days. Nine of 10 ewes did not exhibit estrus for periods similar to 1 or 2 estrus cycles during this period, but eight ewes re-initiated cycles by the end of the study on September 18. Selection for ability to lamb in autumn thus resulted in ewes with an abbreviated seasonal anestrus and reduced sensitivity to long days.     

A note on ram preference by oestrous ewes: Influence of rams’ age and sexual performance

Ewes in oestrus, besides being receptive to rams, develop an active search for them. We performed two experiments to determine if oestrous ewes prefer: (1) high or low sexual-performance rams, and (2) 2-year-old or aged (>8-years-old) rams. In the first experiment, the preference of ewes for rams with high or low sexual performances was compared in 16 tests. In the second experiment, 17 tests were performed comparing ewes’ preference for aged rams (>8-years-old) or young rams (2-years-old). In each test, two rams were tied at equal distances from the point of entrance of the ewe, with a maximum mobility for each ram of 2 m. The first approach to a ram, the quantity of approaches toward each ram (less than one body length), and the total time that the ewe remained near each ram were registered during 5 min. Oestrous ewes did not display any preference according to sexual performance of rams, but spent significantly more time (P = 0.02), and did it in more tests (P = 0.025), with young than with aged rams. Ewes also tended to approach first (P = 0.07), and more times in more tests (P = 0.06), to young than to aged rams. We conclude that ram's sexual performance seems not to influence oestrous ewes preference, and that ewes’ prefer young (2-years-old) rather than aged rams (>8-years-old).     

Further development of a transcervical technique for artificial insemination in sheep using previously frozen semen

A transcervical technique (the Guelph System for transcervical AI) was used to inseminate 2060 ewes on 65 farms (average 31 ewes, range 5 to 107) in Ontario, Canada, from October 1990 to September 1992, using previously frozen semen. Estrus was synchronized using progestagen pessaries and PMSG with median inseminations done at 54 h from pessary removal. Maiden ewes were not included. Only ewes in which the cervix could be penetrated were inseminated with 150 million spermatozoa per insemination. A total of 1809 were penetrated and inseminated (penetration rate 87.8%). Success of penetration increased from 76.3% in the first 500 ewes to 97.9% in the last 500 (P=0.01). Cervical penetration was more successful in ewes in the accelerated lambing program (92.3%, average 3.1 mo since the previous lambing) than those in the annual lambing program (82.4%, average 7.0 mo since the previous lambing; P=0.06). The lambing rate for ewes bred during the combined traditional breeding seasons (Fall of 1990, 1991, 1992) was 50.7% compared to 24.4% for ewes bred at other periods (P=0.00001). The average time required for handling and insemination decreased from 8.62 min in the first 500 ewes to 3.62 min in the last 500 ewes. The Guelph System for Transcervical AI was found to be successful for cervical penetration in most ewes. Penetration success was affected by period since the last lambing and by inseminator experience. The lambing rate was higher for ewes bred during the traditional Fall breeding seasons than during other times of the year.     

Oestradiol and the preovulatory surges of luteinising hormone and follicle stimulating hormone in ewes during the breeding season and transition into anoestrus

This study was designed to see if giving exogenous oestradiol, during the follicular phase of the oestrous cycle of intact ewes, during the breeding season or transition into anoestrus, would alter the occurrence, timing or magnitude of the preovulatory surge of secretion of luteinising hormone (LH) or follicle stimulating hormone (FSH). During the breeding season and the time of transition, separate groups of ewes were infused (intravenously) with either saline (30 ml h⁻¹; n = 6) or oestradiol in saline (n = 6) for 30 h. Infusion started 12 h after removal of progestin-containing intravaginal sponges that had been in place for 12 days. The initial dose of oestradiol was 0.02 μg h⁻¹; this was doubled every 4 h for 20 h, followed by every 5 h up to 30 h, to reach a maximum of 1.5 μg h⁻¹. Following progestin removal during the breeding season, peak serum concentrations of oestradiol in control ewes were 10.31 ± 1.04 pg ml⁻¹, at 49.60 ± 3.40 h after progestin removal. There was no obvious peak during transition, but at a time after progestin removal equivalent to the time of the oestradiol peak in ewes at mid breeding season, oestradiol concentrations were 6.70 ± 1.14 pg ml⁻¹ in ewes in transition (P < 0.05). In oestradiol treated ewes, peak serum oestradiol concentrations (24.8 ± 2.1 pg ml⁻¹) and time to peak (41.00 ± 0.05 h) did not differ between seasons (P > 0.05). During the breeding season, all six control ewes and four of six ewes given oestradiol showed oestrus with LH and FSH surges. The two ewes not showing oestrus did not respond to oestrus synchronisation and had persistently high serum concentrations of progesterone. During transition, three of six control ewes showed oestrus but only two had LH and FSH surges; all oestradiol treated ewes showed oestrus and gonadotrophin surges (P < 0.05). The timing and magnitude of LH and FSH surges did not vary with treatment or season. In blood samples collected every 12 min for 6 h, from 12 h after the start of oestradiol infusion, mean serum concentrations of LH and LH pulse frequency were lower in control ewes during transition than during mid breeding season (P < 0.05). Oestradiol treatment resulted in lower mean serum concentrations of LH in season and lower LH pulse frequency in transition (P < 0.05). We concluded that enhancing the height of the preovulatory peak in serum concentrations of oestradiol during the breeding season did not alter the timing or the magnitude of the preovulatory surge of LH and FSH secretion and that at transition into anoestrus, oestradiol can induce oestrus and the surge release of LH and FSH as effectively as during the breeding season.     

The introduction of rams induces an increase in pulsatile LH secretion in cyclic ewes during the breeding season

Application of the ram effect during the breeding season has been previously disregarded because the ewe reproductive axis is powerfully inhibited by luteal phase progesterone concentrations. However, anovulatory ewes treated with exogenous progestagens respond to ram introduction with an increase in LH concentrations. We therefore tested whether cyclic ewes would respond to ram introduction with an increase in pulsatile LH secretion at all stages of the estrous cycle. We did two experiments using genotypes native to temperate or Mediterranean regions. In Experiment 1 (UK), 12 randomly cycling, North of England Mule ewes were introduced to rams midway through a frequent blood-sampling regime. Ewes in the early (EL; n=3) [corrected] and late luteal (LL; n=6) phase responded to ram introduction with an increase in LH pulse frequency and mean and basal concentration [corrected] of LH (at least P<0.05). In Experiment 2 (Australia), the cycles of 32 Merino ewes were synchronised using intravaginal progestagen pessaries. Pessary insertion was staggered to produce eight ewes at each stage of the estrous cycle: follicular (F), early luteal (EL), mid-luteal (ML) and late luteal (LL). In all stages of the cycle, ewes responded to ram introduction with an increase in LH pulse frequency (P<0.01); EL, ML and LL ewes also had an increase in mean LH concentration (P<0.05). In conclusion, ram introduction to cyclic ewes stimulated an increase in pulsatile LH secretion, independent of ewe genotype or stage of the estrous cycle.     

The effects of ram exposure during progestagen oestrus synchronisation and time of ram introduction post progestagen withdrawal on fertility in ewes

Three experiments were undertaken to investigate the effect of a pre-mating ram exposure during progestagen synchronisation treatment on time of breeding, ovulation rate, embryo quality and fertility and any interaction with time of ram introduction for breeding post sponge withdrawal. Crossbred ewes in experiment 1a (n = 348), 1b (mule; n = 133) and 2 (n = 58) underwent a 12-14 days synchronisation protocol. Three days prior to sponge withdrawal ewes were divided into Control (ewes in continued isolation from rams) or +Ram (ram-exposed) groups. Rams were introduced to +Ram ewes and remained with ewes until sponge withdrawal. Ewes in experiments 1a and 2 received eCG at sponge withdrawal and were reintroduced to rams at either 36 or 48 h post sponge removal (PSR). In experiment 1b, ewes did not receive eCG and were reintroduced to rams at 24 h PSR. In experiments 1a and 1b time of breeding, date of lambing and litter size were recorded. In experiment 2, ewes were slaughtered 5 days post breeding, reproductive tracts flushed and corpora lutea, ova and embryos assessed. Fewer +Ram ewes were mated by 96 h PSR (P < 0.001) than Control ewes in experiment 1a but not when rams were introduced earlier in experiment 1b. In experiment 1a, ram introduction at 36 h PSR improved conception to first service compared to introduction at 48 h PSR (P < 0.01) in both +Ram and Control groups. In experiments 1a and 1b, +Ram ewes had reduced litter size caused by more single births (1a; P < 0.001, 1b; P < 0.01). In experiment 2, +Ram ewes had fewer corpora lutea than Control ewes (P < 0.001) but embryo quality was similar. However, more good embryos were produced when rams were introduced for breeding at 36 h compared to 48 h PSR (P < 0.001). We conclude that a pre-mating ram exposure during the synchronisation treatment reduced the number of ewes mated at and conceiving to the first service. This was partially overcome by introducing rams for breeding earlier (24 or 36 h compared to 48 h PSR) but the most dramatic decrease in fertility was due to a reduction in ovulation rate in the ram-exposed ewes.     

The influence of breeding intensity on above- and below-average sexual performance rams in single- and multiple-sire breeding environments

Two studies were conducted to evaluate the relationship between serving capacity scores and breeding performance of rams. The first study was conducted to determine whether rams with above or below mean serving capacity scores could perform equally in greater and lesser breeding intensity, single-sire mating schemes. The second study was conducted to determine whether rams with above and below mean serving capacity scores could perform equally well when only one or two ewes were in estrus daily in a multiple-sire breeding scheme (two rams/pen). Rams (n=68) were ranked according to average number of ejaculations recorded in serving capacity tests. Sixteen rams with the greatest scores (above-average) and 16 rams with least scores (below-average) were identified for breeding. Half of above-average and half of below-average rams were used in the two studies. For study 1, each ram was individually introduced to 23 estrus-synchronized ewes for 9 d to simulate high breeding intensity. Rams were given a 5-d rest before they were individually introduced to 23-24 naturally cyclic ewes for 17 d (low breeding intensity). For study 2, 16 rams were paired across ram types, and each pair competed for 20 ewes for 18 d (8 pens). For study 1, ewe fertility (ewes lambing/ewes present at lambing) and number of lambs born were greater (P<0.001) for above-average (0.67+/-0.03 and 27.6+/-1.2, respectively) than for below-average rams (0.39+/-0.07 and 15.3+/-2.7) with greater breeding intensity. Ewe fertility and lambs born did not differ for above-average (0.91+/-0.03 and 37.8+/-1.9, respectively) and below-average rams (0.86+/-0.03 and 39.0+/-1.9) with less breeding intensity. For study 2, number of ewes lambing (99+/-8.0 compared with 72+/-13.6; P=0.12) and number of lambs sired (149+/-18.5 compared with 101+/-22.8; P=0.14) did not differ between above- and below-average rams, respectively, in direct competition. Sexual classifications based on serving capacity tests are related to breeding performance of rams in certain breeding environments. When breeding intensity is greater, above-average rams impregnate more ewes and sire more lambs than below-average rams. When only a small number of ewes are in estrus daily, below-average rams for serving capacity scores perform as well as above-average rams in multiple-sire and single-sire breeding environments. We suggest that above-average rams should be used to reduce number of rams required when breeding intensity is greater.     

Induction of estrus with intramuscular injections of GnRH or PMSG in lactating goats (Capra hircus ) primed with a progestagen during seasonal anestrus

In Experiment 1, goats in seasonal anestrus (n=154) were treated with sponges impregnated with 1 of 2 types of progestagen (MAP or FGA) followed by PMSG (400 IU im) 48 h before sponge removal. The type of progestagen used had no effect on kidding, abortion, pseudogestation, multiple births, stillbirths, number of live births per doe or gestation length. In Experiment 2, lactating goats (n=24) in seasonal anestrus were treated with progestagen sponges (MAP). At sponge removal they received one of the following treatments: 1 injection of PMSG (400 IU im), 1 injection of GnRH (125 mug im; GnRH-1), or 2 injections of GnRH (125 mug/injection im; GnRH-2) at a 48 h interval. Serum samples were taken at 6-h intervals for 96 h, starting 12 h after sponge removal. Heterologous radioimmunoassays were validated for the measurements of goat FSH, LH, E(2) and P(4). The onset of estrus (P=0.004), mean doe receptivity (P=0.0006), maximum preovulatory E(2) concentrations (P=0.0001) and LH peak concentrations (P=0.08) occurred significantly later for GnRH-1 and GnRH-2 than for PMSG treatment. The PMSG treatment induced a preovulatory LH peak in a greater number of goats (P=0.05) and gave a higher gestation rate than GnRH-1 and GnRH-2 treatments (57 vs 0 vs 12%; P=0.03). It is likely that the GnRH treatments administered did not reactivate the hypothalamo-pituitary-gonadal axis. Thus, intramuscular injections of GnRH in lactating goats primed with a progestagen were not as effective in regulating reproductive performance during seasonal anestrus as were injections of PMSG.     

Seasonality of ovulation and estrus, and the ram effect in poll dorset ewes

The effects of month on the proportion of Poll Dorset ewes expressing estrus and ovulating and on their ovulation rate when continually exposed to vasectomised rams and the effects of isolating the ewes from rams in winter/early spring were examined during 15 months in New South Wales, Australia. The percentage of ewes ovulating and their ovulation rate varied from 23% and 1.10 in November to an average of 99% and 1.85 in April and June, and the proportion of ewes expressing estrus followed a similar trend. Some ewes (6.8%) ovulated throughout the 15 months, and the average breeding season was 294 (SE = 6.1) days. Isolating ewes from rams in late winter/early spring significantly decreased the proportion of ewes ovulating from September to November, increased the proportion in December, but did not significantly alter ovulation rate. The variability in reproductive measures during spring offers scope for selection; with the ram effect, improvements in spring joining results from Dorset ewes are possible.