The effects of gonadotrophin releasing hormone administration in early postpartum dairy cows on hormone concentrations, ovarian activity and reproductive performance: a review

Gonadotrophin releasing hormones have become widely used hormonal compounds in veterinary medicine, particularly with respect to bovine reproduction. The character and physiological actions of gonadotrophin releasing hormone are briefly reviewed and its clinical applications are summarized. The endocrinological research concerned with the use of gonadotrophin releasing hormone in the early postpartum period is discussed. Field trials which have been conducted to assess the effects of postpartum gonadotrophin releasing hormone administration on reproductive performance have varied widely in both design and interpretation of results. These experiments are reviewed, including the clinical trials using normal cows as well as those on cows with retained placenta.     

Effects of gonadotrophin releasing hormone on reproductive performance of dairy cows with retained placenta.

The effects of gonadotrophin releasing hormone (GnRH) on the reproductive performance of dairy cows with retained placenta were studied. Three hundred and seventy-eight cows diagnosed as having retained placenta received intramuscular injections of either 2 mL sterile water or 200 micrograms of GnRH in 2 mL sterile water between day 8 and day 14 postpartum. Rectal palpation was performed at the time of treatment and ten to 20 days after treatment in order to determine the rate of uterine involution. Thereafter, monthly rectal examinations were carried out until insemination. Pregnancy diagnosis was made by rectal palpation at 40 days or more after breeding. Using the entire experimental population, there were no significant differences between GnRH-treated and control cows for the rate of uterine involution, the occurrence of reproductive problems, the interval from parturition to first observed estrus, the interval from parturition to first insemination, the interval from parturition to conception, the number of services per conception, the total number of services per cow regardless of conception and the incidence of culling for infertility. When the data for herds in which breeding began earlier in the postpartum period (herds having a mean less than or equal to 80 days from parturition to first service for retained placenta cows) were considered, the GnRH treatment resulted in a significantly shorter (p less than or equal to 0.01) calving to conception interval as compared to control cows. Also, there was a significant reduction (p less than or equal to 0.05) in the total number of services per cow regardless of conception and a significant reduction in the interval from parturition to first service.(ABSTRACT TRUNCATED AT 250 WORDS)     

Improved knowledge about conception rates influences the decision to stop insemination in dairy cows

The conception rate in dairy cows is dependent on a number of cow factors such as days in milk and insemination number. Unfortunately, some of these factors were not accounted for in optimal insemination and replacement decision models. By using wrong estimates of the conception rate, the calculated optimal insemination and replacement policy might differ from the real optimal insemination and replacement policy. The objective of this study was to evaluate different sets of conception rates with an increasing level of accuracy to determine the insemination policy. An existing dynamic program for optimal insemination and replacement was used to compare three different scenarios in the estimation of conception rates, based on the reproductive performance of Dutch dairy cattle: (i) constant conception rate throughout lactation, (ii) conception rate dependent on parity and months in milk, and (iii) conception rate dependent on parity, months in milk and insemination number. The time step of the model was 30.4 days (1 month). The discounted future cash flow of culling a cow at each time step (replace a heifer immediately) was compared with keeping that cow under optimal future decisions. The difference between immediate culling and optimal decisions is defined as the retention pay-off. The insemination value was calculated as the difference between the future cash flow between immediate insemination of a cow and waiting one time-step. The results show that the difference in the insemination values and the optimal time to stop insemination depend on parity, lactation stage and the relative milk yield. In older cows with equal milk yields and at the same months in milk, the insemination value was lower than in younger cows. Within a parity, the insemination value was higher for cows with a higher milk yield. On individual cow level, using more accurate conception rate as input in the optimal insemination and replacement model might reduce miscalculation of the economic consequences for at least of €20-€38 per cow per year. Basing insemination decisions on less accurate input of the probabilities of conception, however, did not have an economic consequence at the herd level. In conclusion, using the appropriate conception rate as input in the optimal insemination and replacement model would increase the precise decision for the optimal time to stop insemination and hence improve the reproductive management efficacy.     

Synchronisation of oestrus and reproductive performance of dairy cows following administration of oestradiol benzoate or gonadotrophin releasing hormone during a synchronised pro-oestrus

OBJECTIVE: To compare the use of gonadotrophin releasing hormone (GnRH) and oestradiol benzoate (ODB) administered following a synchronised pro-oestrus on reproductive performance of lactating dairy cows and the submission rates of non-pregnant cows following resynchronisation. DESIGN: Cohort study. PROCEDURE: Lactating Holstein cows enrolled in a controlled breeding program were first treated with an intravaginal progesterone releasing insert (IVP4) for 8 days, 2.0 mg of ODB intramuscular (i.m.) at device insertion (Day 0), an analogue of PGF2alpha at device removal and either 1.0 mg of ODB i.m., 24 h after device removal (ODB group, n = 242), or 0.25 mg of a GnRH agonist (GnRH group, n = 152) injected i.m. approximately 34 h after device removal. Every cow was artificially inseminated between 49 and 56 h after removal of its insert (Day 10). Cows detected in oestrus 1 day after artificial insemination (AI) that were not detected in oestrus on the previous day were re-inseminated on that day. All cows treated on Day 0 were resynchronised for reinsemination by insertion of a used IVP4 device on Day 23. Oestradiol benzoate at a dose of 1.0 mg was administered i.m. at the time of device insertion. Inserts were removed 8 days later (Day 31) and 1.0 mg of ODB was injected i.m. 24 h later. Those cows detected in oestrus between Days 31 and 35 were artificially inseminated. On Day 46 these cows were resynchronised for a third round of AI by insertion of an IVP4 device, used previously to synchronise cows for the first and second rounds of AI, and administration of 1.0 mg of ODB i.m.. Eight days later inserts were removed. Cows detected in oestrus between Days 54 and 58 were artificially inseminated. Bulls were run with the herd between rounds of AI and removed after 21 weeks of mating. RESULTS: Treatment with ODB or GnRH at the first synchronised pro-oestrus did not significantly alter the reproductive performance over three rounds of AI or over a 21-week breeding period. Treatment also did not alter submission rates at the second round of AI or the proportion of non-pregnant and non-return cows ('phantom' cows) detected and did not result in significant differences in concentrations of progesterone in plasma 10 and 18 days after removal of inserts at the first round of AI. Treatment with GnRH reduced the proportion of cows detected in oestrus at the first round of AI (36.2 vs 97.5%; P < 0.001). CONCLUSION: Administration of GnRH compared to ODB at a synchronised pro-oestrus results in similar reproductive performance. Treatment with GnRH reduced the proportion of cows detected in oestrus following treatment. This may offer advantages to the way AI is managed by enabling insemination at a fixed-time and removing the need for the detection of oestrus.     

Evolution of the corpus luteum volume determined ultrasonographically and its relation to the plasma progesterone concentration after artificial insemination in pregnant and non-pregnant dairy cows

The aim of this study was to assess the relationship of the evolution of the corpus luteum (CL) volume that was determined ultrasonographically with the pregnancy status in lactating dairy cows during early pregnancy. Ultrasound examinations were carried out on 76 cows following artificial insemination (AI). Plasma concentrations of progesterone were determined from blood samples collected at each ultrasound examination. Conception was confirmed by ultrasonography on day 30 after AI. Around day 14 post-insemination (p.i.), the CL volume tended to decrease in pregnant and non-pregnant cows, and, after day 19 p.i., both groups differed significantly, indicating the luteal regression in non-pregnant cows. Reaching signification on day 20. The diminution in CL volume was also reflected in the plasma progesterone concentration. However, the patterns of CL volume, estimated by ultrasonography, differed more evidently and earlier between both groups (around 1 week p.i., at day 9 p.i. P < 0.05, whereas progesterone started to differ around 2 weeks p.i., at day 14 p.i, P < 0.05). These results indicate that the estimation of the CL volume by ultrasonography could be useful for assessing the presence of a functional CL.     

Reproductive performance in dairy cows following postpartum treatment with gonadotrophin releasing hormone and/or prostaglandin: a field trial.

Three hundred and five Holstein Friesian cows were given either 250 micrograms gonadotrophin releasing hormone (GnRH) or saline on day 15 postpartum followed by 500 micrograms cloprostenol or saline on day 24 postpartum. Four treatment groups were formed using random allocation: Group I -- placebo (Day 15)/placebo (Day 24), Group II -- GnRH (Day 15)/placebo (Day 24), Group III -- placebo (Day 15)/cloprostenol (Day 24), Group IV -- GnRH (Day 15)/cloprostenol (Day 24). Double blind techniques were used during the follow-up period. Rectal palpation, to assess uterine involution and ovarian activity was performed just prior to each treatment and again at 28 days postpartum. In addition blood samples were collected at 15, 24 and 28 days postpartum for measurement of plasma progesterone. There were no significant differences among treatment groups with respect to services per conception, number of heats detected before first service and culling for infertility. Cows treated only with GnRH had an increased calving to first estrus and calving to first breeding interval, and tended to have an increased calving to conception interval. Treatment with cloprostenol significantly decreased calving to conception and calving to first observed estrus intervals. Treatment with GnRH on day 15 postpartum resulted in a significant increase in the subsequent incidence of pyometra and prebreeding anestrus. On the other hand, cloprostenol treatment on day 24 postpartum resulted in a decreased incidence of pyometra, regardless of GnRH treatment and a decreased incidence of prebreeding anestrus in GnRH treated cows compared to cows receiving only GnRH at day 15 postpartum.(ABSTRACT TRUNCATED AT 250 WORDS)     

Effects of progesterone and human chorionic gonadotrophin administration five days postinsemination on plasma and milk concentrations of progesterone and pregnancy rates of normal and repeat breeder dairy cows.

Treatment with a progesterone-releasing intravaginal device between days 5 and 12 after estrus elevated (p less than 0.05) plasma progesterone concentrations between days 6 and 8 in comparison with controls. Treatment with injectable progesterone (200 mg) on days 5, 7, 9 and 11 postestrus did not increase plasma progesterone concentrations over controls. The administration of 1500 IU human chorionic gonadotrophin (hCG) on day 5 after estrus resulted in a sustained increase (p less than 0.01) in plasma progesterone concentrations from day 8 until day 20 when measurements ceased. Pregnancy rates, as a result of artificial insemination (AI) at the pretreatment estrus, in these treatments (n = 12-14 each), were unaffected by any of the treatments and ranged from 57.1 to 75.0% at 45-60 days post-AI. In a field trial, of 36 repeat breeder cows treated with 1500 IU hCG 5.5 days after insemination, 47.2% were pregnant at 60 days, whereas 39.5% of saline-treated controls were diagnosed pregnant. Treatment with hCG significantly (p less than 0.05) increased milk progesterone concentrations over controls on days 14 and 20 after insemination.     

Comparison of effects of GnRH and prostaglandin in combination, and prostaglandin on conception rates and time to conception in dairy cows

OBJECTIVE: To assess the effectiveness of a gonadotrophin-releasing hormone (GnRH) / prostaglandin program (GnRH-PG-GnRH, Ovsynch) on conception rates and time to conception of lactating dairy cows compared with a PG program (double prostaglandin injection). DESIGN: A randomised multi-centre cohort study was conducted with 778 cows from nine dairy herds. Cows at different stages of lactation were randomly assigned, after matching for days open at the time of treatment, to either the PG or Ovsynch program. PROCEDURE: Cows on the PG program received two intramuscular injections of prostaglandin (2 mL, Prosolvin) 11 days apart. The Ovsynch program consisted of two intramuscular injections of GnRH (1 mL, Fertagyl) 9 days apart, separated by one injection of prostaglandin 40 h before the second GnRH injection. Milk samples were taken at the time of artificial insemination and assayed for progesterone by radioimmunoassay. RESULTS: The Ovsynch program was not significantly different to PG in achieving conception, with overall conception rates of 37.6% and 41.4%, respectively, for each program. There was, however, a significant interaction between the effects of parity and treatment (P = 0.03), because conception rates were higher in older cows (parity 5 or more) on the PG program than for older cows on the Ovsynch program. There was no significant effect of treatment (P > 0.5) on time to conception after treatment, but older cows were slower to conceive (P < 0.0001). Conception rates differed (P < 0.0001) among herds. CONCLUSION: The median days to conception for both groups was 22 and mean days from treatment to conception were 36.3 +/- 3.3 and 31.6 +/- 2.7 for the Ovsynch and PG programs respectively, indicating that reproductive performance of cows was not significantly different with Ovsynch program or PG program. There appears to be a need to evaluate causes of reproductive failure in older cows.     

Short-term changes in serum luteinizing hormone, ovarian response and reproductive performance following gonadotrophin releasing hormone treatment in postpartum dairy cows with retained placenta.

Thirty-six postpartum Holstein cows consisting of eighteen cows that shed the placenta soon after calving (NRP) and eighteen cows that retained placenta greater than 24 h (RP) were used. There were four treatment groups. Group I consisted of 9 NRP cows which received intramuscular injection of sterile saline on day 15 postpartum. The second group consisted of 9 NRPN cows which received 100 micrograms of gonadotrophin releasing hormone on day 15 postpartum (NRPT). The third group consisted of 9 RP cows which received saline on day 15 postpartum (RPN) and the fourth group consisted of 9 RP cows which received 100 micrograms of gonadotrophin releasing hormone on day 15 postpartum (RPT). Blood samples were collected once daily during the first month and once every other day during the second month postpartum. In addition fourteen cows (RPT, n = 7; NRPT, n = 7), were used to study short-term changes in serum luteinizing hormone concentrations following gonadotrophin releasing hormone treatment on day 15 postpartum. Blood samples were obtained from these cows every 15 min during 1 h before and 6 h after gonadotrophin releasing hormone administration. Sera from all samples were assayed for progesterone and luteinizing hormone concentrations. Starting from four days after calving rectal palpations and ultrasound examinations of the ovaries were carried out once every four days until day 28 postpartum in order to monitor ovarian changes. All cows were inseminated on the first estrus after day 60 postpartum and examined for pregnancy between 35 and 42 days after insemination.(ABSTRACT TRUNCATED AT 250 WORDS)     

奶牛胚胎移植中生殖激素含量变化及黄体发育状况与移植效率的关系

本研究通过测定奶牛血中相关激素水平,针对胚胎移植中对受体母牛生理状况判断时卵巢触诊法的主观性和盲目性,以相关激素水平为指标来判定受体牛的生理状态,研究胚胎移植受体黄体和生理状况及妊娠之间的关系.通过试验证实,利用黄体触诊法选择受体牛虽然有非量化的弊端,但是实际生产操作中仍然是可行的.     

Time of onset of post-partum luteal function and conception in dairy cows.

The following reproductive parameters were studied in 40 Icelandic dairy cows on 2 farms with free stall and tie stall housing: The time of onset of post-partum luteal activity; the time of first post-partum artificial insemination (AI); the time of conception; conception rate to first post-partum AI; the number of AI's per conception; length of gestation; and the number of days between 2 consecutive calvings. Milk for progesterone analysis was sampled 3 times a week from calving until first AI on farm 1, but every day for 90-100 days from calving on farm 2. In this study the average onset of luteal function was 21 days post-partum, the cows were inseminated 62 days and conceived 74 days post-partum. Conception rate to first AI was 71% and number of AI's per conception was 1.6. Significant differences were not found between the 2 farms. The reproductive performance of the cows in this study was greater than found in earlier studies of the Icelandic dairy cows and indicate that this animal has reproductive capabilities comparable to high producing foreign dairy cows.     

Thyroid-stimulating hormone responses after single administration of thyrotropin-releasing hormone and combined administration of four hypothalamic releasing hormones in beagle dogs

Thyrotropin (TSH) responses were determined in eight healthy male beagle dogs after a single administration of thyrotropin-releasing hormone (TRH) and the combined administration of four hypothalamic releasing hormones, i.e., corticotropin-releasing hormone, growth hormone-releasing hormone, gonadotropin-releasing hormone, and TRH. In both tests, TRH was administered in a dose of 10 μg/kg. Basal TSH concentrations ranged form 0.07 to 0.27 μg/1(mean ± SE, 0.14 ± 0.02 μg/1). The administration of TRH, alone or in the combined test, resulted in a prompt and significant increase in TSH with mean (±SE) plasma TSH peaks of 1.26 ± 0.22 μg/1 at 10 min and 0.85 ± 0.17 μg/1 at 30 min, respectively. The area under the curve (0–120 min) was significantly lower in the combined test than in the single TRH test, whereas the increments were not significantly different. It is concluded that measurements of TSH responses to TRH alone and in combination with other releasing hormones can be used for the assessment of pituitary thyrotropic cell function. In the combined test, the TSH response is slightly lower than that in the single test.     

Influence of cornual insemination on conception in dairy cattle

The objective of this study was to compare conception to artificial insemination (AI) services in dairy cattle when semen was deposited into the uterine body or into both uterine horns (cornual insemination). Nine herdsman inseminators (HI) in four commercial dairy herds in Washington constituted the experimental units. Herds ranged in size from 393 cows to 964 cows. The duration of the experiment was 12 mo in three herds and 18 mo in the fourth herd. At the beginning of the experiment all inseminators were trained to deposit semen in the body of the uterus. Inseminators were instructed to use this method for 6 mo. Following employment of body deposition, the same inseminators were retrained to deposit one-half of the semen into the right uterine horn and one-half into the left uterine horn. Cornual inseminations were performed for 6 mo. A total of 4,178 services constituted the data set. Milk samples were collected from cows on the day of insemination and later were assayed for progesterone (P4). There was variation (P less than .01) in conception associated with month of insemination and insemination method (P less than .001). The monthly variation was not associated with season of the year. Least squares means for conception when semen was deposited in the uterine body was 44.7%, compared with 64.6% when cornual insemination was employed. The insemination treatment X inseminator interaction was not significant. Results suggest that cornual insemination provides an alternative to deposition of semen in the uterine body.     

持久黄体性不孕奶牛用复方缩宫素乳剂处理后血清中孕酮和雌二醇水平的变化

选择患持久黄体的不孕奶牛14头进行复方缩宫素乳剂的临床治疗试验,并利用放射免疫方法对部分患牛治疗前后血清孕酮（P     

Formation of a secondary corpus luteum after ultrasound-guided follicular aspiration in cows

This paper reports the observed formation of a secondary corpus luteum (CL) in the presence of the cyclic corpus luteum, on the ovaries of a cow after ultrasound-guided follicular aspiration for oocyte recovery. The secondary structure, although smaller and lighter (4.97 g vs. 6.02 g) than the natural one, had the typical macroscopic appearance of a corpus luteum. Histological examination of the structure using electron microscopy revealed typical structural features of a natural CL. Mean tissue progesterone concentration was significantly lower in the secondary CL (31.15 +/- 3.11 compared with 58.29 +/- 6.32 micrograms/g tissue of the cyclic CL) and oestradiol-17 beta significantly higher than in the natural CL (108 +/- 11.6 compared with 74.2 +/- 7.81 pg/g tissue). P450scc and P450(17 alpha) mRNA was detected in both structures while P450arom and full-length mRNA FSH receptor were detected only in the secondary structure.     

The comparison of treating Holstein dairy cows with progesterone, CIDR and GnRH after insemination on serum progesterone and pregnancy rates

This experiment was conducted to investigate the effects of different exogenous progesterone sources administrated after artificial insemination (AI) on serum progesterone (P(4) ) concentration and pregnancy rates in Holstein lactating cows. Sixty-four lactating Holstein dairy cows were allocated to four different treatments (n = 16 per treatment): the cows 1) were injected with physiological saline on days 5 and 13 after AI (control group); 2) were injected with progesterone on days 5 and 13 after AI (P group); 3) received controlled internal drug releasing device (CIDR) for a period from day 5 to 19 after AI (CIDR group); and 4) were injected with gonadotropin-releasing hormone (GnRH) agonist on days 5 and 13 after AI (GnRH group). Blood samples were collected on days 0 (AI day), 5, 13, 16 and 19 after AI to determine serum P(4) concentration. The results revealed a significant difference among treatment groups for serum P(4) concentration on days 13, 16 and 19 with the lowest concentration of serum P(4) for the control group. The pregnancy rate was also positively affected by all the treatments with CIDR having the greatest effect on pregnancy rate. Overall, the results indicated that CIDR has the greatest effect on serum P(4) concentration and pregnancy rate, although the administration of P and GnRH during days after AI increased serum P(4) concentration in lactating dairy cows as well.     

Relationship between rectal findings of corpus luteum and whole milk progesterone levels in postpartum dairy cows

The reliability of clinical ovarian findings was assessed as an indicator of luteal function in primiparous dairy cows. The postpartum period of 103 cows following their first parturition was studied by thrice weekly rectal palpation of ovaries and whole milk progesterone assay from 1 week after parturition to the first insemination. The relationship between milk progesterone levels and 1101 ovarian findings was compared during the follicular phases, short luteal phases and during the early, mid and late thirds of normal luteal phases. The compatibility between elevated progesterone and palpable corpus luteum was 71%, and between low progesterone and lack of corpus luteum 77%. In 10% of all rectal examinations the finding was unspecified; i.e. the clinician could not differentiate between luteal and follicular activity. During the acyclic period prior to the initiation of luteal function, the proportion of false corpus luteum findings was 11%. The corpora lutea of the short oestrous cycles were more difficult to palpate than those of normal cycles. During early dioestrus the corpus luteum was significantly more difficult to palpate than during the rest of dioestrus. The percentage of unspecified findings was highest during early dioestrus.The paper discusses the reliability of rectal examination as a method of diagnosing cyclicity and of evaluating the responsiveness of a cow to prostaglandin treatment.     

Failure of an intramuscular injection of an analogue of gonadotrophin-releasing hormone 11 to 13 days after insemination to increase pregnancy rates in dairy cattle

At 11 to 13 d after insemination, 1028 dairy cows on 19 farms were given an intramuscular injection of 10 micrograms of the GnRH analogue buserelin. Their reproductive performance was compared to 1022 control cows receiving a placebo. The GnRH analogue-treated cows had fewer short interoestrous intervals compared to the control cows, but there were no significant differences in pregnancy rates to either the insemination preceding or following treatment, in calving to conception interval, or to percent pregnant by the end of mating. There were no differences between treatments in herds of high, average or low pregnancy rate, in 2-year-old cows or cows greater than 2-years-old and in cows calved 40 d or less, or greater than 40 d. The use of 10 micrograms of the GnRH analogue, buserelin, 11 to 13 d after insemination, cannot be recommended as a method of improving reproductive performance in dairy herds.     

Effect of administration of human chorionic gonadotropin after artificial insemination on concentrations of progesterone and conception rates in beef heifers

The objective of this study was to determine whether administration of hCG approximately 5 d after AI would increase plasma progesterone concentrations and conception rates in beef heifers. Heifers from two locations (Location 1: n = 347, BW = 367 +/- 1.72 kg; Location 2: n = 246, BW = 408 +/- 2.35 kg) received melengestrol acetate (0.5 mg.heifer(-1).d(-1)) for 14 d and an injection of PGF2alpha (25 mg i.m.) 19 d later. Heifers were observed for estrus continuously during daylight from d 0 to 4.5 after PGF2alpha and artificially inseminated approximately 12 h after the onset of estrus. Half of the heifers inseminated at Location 1 were assigned randomly to receive an injection of hCG (3,333 IU i.m.) 8 d after PGF2alpha, and a blood sample was collected from all heifers 14 d after PGF2alpha for progesterone analysis. Half of the heifers inseminated at Location 2 were administered hCG on d 9 after PGF2alpha, and a blood sample was collected from all heifers 17 d after PGF2alpha. Heifers at Location 1 had a 94% synchronization rate, exhibited estrus 2.45 +/- 0.03 d after PGF2alpha, and received hCG 5.55 +/- 0.03 d after AI. Heifers at Location 2 had an 85% synchronization rate, exhibited estrus 2.69 +/- 0.03 d after PGF2alpha, and received hCG 6.31 +/- 0.03 d after AI. Progesterone concentrations were greater (P < 0.01) for hCG-treated heifers than for controls at both locations (8.6 vs. 4.6 ng/mL for treatment vs. control at Location 1, and 11.2 vs. 5.6 ng/mL for treatment vs. control at Location 2). Pregnancy status was determined by ultrasound approximately 50 d after AI. Conception rates (65 vs. 70% for treatment vs. control, respectively) did not differ at Location 1. Conception rates tended (P = 0.10) to be increased with hCG treatment at Location 2 (61 vs. 50% for treatment vs. control, respectively). A second experiment was conducted with 180 heifers at a third location to determine the effects of hCG administration 6 d after timed insemination at approximately 60 h after PGF2alpha in heifers synchronized as in Exp. 1. Pregnancy rate to timed AI did not differ between hCG-treated (62%) and control heifers (59%). Final pregnancy rate after timed AI and bull exposure (92%) was not affected by treatment. In summary, administration of hCG 5 to 6 d after AI did not improve conception or pregnancy rates at two out of three locations evaluated, suggesting insufficient progesterone is not a major factor contributing to early pregnancy failure in beef heifers.