Ovarian follicular dynamics in heifers: test of two-wave hypothesis by ultrasonically monitoring individual follicles

The two-wave hypothesis for follicular development during the bovine estrous cycle was tested by ultrasonically monitoring individual follicles in 10 heifers during an interovulatory interval. A dominant follicle was defined as one that reached a diameter of at least 11 mm. Subordinate follicles were defined as those that appeared to originate from the same follicular pool as a dominant follicle. A dominant follicle and its cohorts were defined as a wave. Two waves during an interovulatory interval were identified in 9 of 10 heifers. The first wave was first identified, retrospectively, on a mean of Day 0.2 +/- 0.1 (ovulation = Day 0) and gave origin to a dominant anovulatory follicle and a mean of 1.4 +/- 0.3 identified subordinates. The dominant follicle reached maximum diameter (mean, 15.8 +/- 0.8 mm) on an average of Day 7 and then decreased (P less than .04) by Day 11. The subordinate follicles increased in diameter for a few days and then regressed. The second wave was first identified on a mean of Day 10.0 +/- 0.4 and gave origin to the ovulatory follicle and a mean of 0.9 +/- 0.3 subordinates. One of the 10 heifers had 3 waves of follicular activity characterized by an anovulatory wave emerging on Day 0, another anovulatory wave emerging on Day 10, and an ovulatory wave emerging on Day 16. Results strongly supported the two-wave hypothesis but also indicated that a minority of interovulatory intervals in this heifer population may have 3 waves of follicular activity.     

Follicular and hormonal dynamics during the estrous cycle in goats

Transrectal ultrasonography of ovaries was performed daily in 6 goats for 3 consecutive estrous cycles. Blood samples collected daily were measured for concentrations of FSH, inhibin A, and estradiol-17beta. Follicular and hormonal data were analyzed for associations between the follicular waves and hormonal concentrations. During the interovulatory intervals, follicular growth and regression occurred in a wave like pattern (2-5 waves), and the predominant patterns were three and four follicular waves. In addition, there was no significant difference among the diameters of dominant follicles during the growth phase of the follicular waves. The number of 3 mm follicles peaked on days 0, 7, and 11 in interovulatory intervals that had three follicular waves and on days -1, 5, 11, and 15 in those that had four follicular waves. Plasma concentrations of FSH increased around the day of follicular wave emergence and declined with the growth of follicles. Circulating FSH increased again concomitant with regression of dominant follicles in the anovulatory wave, whereas FSH levels remained low in the ovulatory wave. Inhibin A was negatively correlated with FSH, while it was positively correlated with estradiol-17beta, suggesting that inhibin A is a product of healthy growing follicles and that it contributes to the suppression of FSH secretion. In conclusion, the growth of ovarian follicles in goats exhibits a wave-like pattern, and follicular dominance is less apparent in goats. Moreover, inhibin A may be a key hormone for regulation of the follicular wave through suppression of FSH secretion in goats.     

Relationship of fertility to ovarian follicular waves before breeding in dairy cows

Cows with two waves of follicular growth during the estrous cycle yield follicles that are older and larger at ovulation compared with cows having three waves. The objectives of the current research were 1) to compare fertility in cows with two or three follicular waves and 2) to examine associations between luteal function, follicular development, and fertility after breeding. Follicular waves were monitored by ultrasonography during the estrous cycle before insemination in 106 dairy cows. Fewer cows had three follicular waves before next estrus and ovulation than two waves (P < 0.01; 30% vs 68%, respectively), but pregnancy rate was higher (P = 0.058; 81 vs 63%, respectively). Cows with two waves had shorter estrous cycles (P < 0.01), with the ovulatory follicle being both larger (P < 0.05) and older (P < 0.01). In cows with three waves, luteal function was extended (P < 0.05) and the peak in plasma progesterone occurred later (P < 0.05) in the estrous cycle compared to two wave cows. Considering cows that became pregnant, luteal phase length was shorter (P < 0.05) during the estrous cycle preceding insemination than for nonpregnant cows. In conclusion, fertility was greater in lactating cows inseminated after ovulation of the third-wave follicle that had developed for fewer days of the estrous cycle as compared with two-wave cows.     

Circulating concentrations of immunoreactive inhibin and FSH during the estrous cycle of mares

A heterologous radioimmunoassay developed to measure inhibin in rat plasma was validated and used to characterize changes in peripheral concentrations of immunoreactive inhibin (ir-inhibin) in relation to follicle stimulating hormone (FSH) concentrations during the estrous cycle of mares. The primary antiserum used in the assay was developed against a synthetic porcine inhibin -subunit [(1-26)-Gly-Tyr] fragment. The same synthetic peptide was used for preparation of standards and tracer. Slopes of the dose-response curves for pooled estrus and diestrus mare plasma and equine follicular fluid were similar to the slopes for the porcine inhibin -subunit standard curve and porcine follicular fluid dose-response curve. Twelve mares were bled once daily beginning when diameter of the largest follicle reached ≤25 mm and continuing until 3 days after the end of an interovulatory interval (ovulation=Day 0). Each of the 12 interovulatory intervals were normalized to the mean length of the interovulatory interval (22.2 days; range, 19 to 26). There was an effect of day for concentrations of ir-inhibin (P<0.001) and FSH (P<0.006). Significant mean changes were as follows: 1) ir-inhibin decreased between Days 0 and 1, whereas FSH increased between Days 0 and 5; and 2) ir-inhibin increased between Days 7 and 12, whereas FSH decreased between Days 11 and 14. Mean concentrations of ir-inhibin and FSH were negatively correlated (r=-0.548; P<0.002). In conclusion, mean peripheral concentrations of ir-inhibin and FSH were inversely related during the estrous cycle of mares.     

Nutritionally induced anovulation in beef heifers: ovarian and endocrine function preceding cessation of ovulation.

Angus x Hereford heifers were used to determine endocrine and ovarian function preceding nutritionally induced anovulation. Six heifers were fed to maintain body condition score (M), and 12 heifers were fed a restricted diet (R) until they became anovulatory. Starting on d 13 of an estrous cycle, heifers were given PGF2alpha every 16 d thereafter to synchronize and maintain 16 d estrous cycles. Ovarian structures of M and R heifers were monitored by ultrasonography daily from d 8 to ovulation (d 1 of the subsequent cycle) until R heifers became anovulatory. Concentrations of LH and FSH were quantified in serum samples collected every 10 min for 8 h on d 2 and 15 (48 h after PGF2alpha), and estradiol and IGF-I were quantified in daily plasma samples from d 8 to 16 during the last ovulatory cycle (Cycle -2) and the subsequent anovulatory cycle (Cycle -1). During the last two cycles before anovulation, M heifers had 50% larger (P < .0001) ovulatory follicles than R heifers and 61% greater (P < .0001) growth rate of the ovulatory follicles. There was a treatment x cycle x day effect (P < .001) for concentrations of estradiol. The preovulatory increase in estradiol occurred in the R and M heifers during Cycle -2 but only in M heifers during Cycle -1. A treatment x cycle x day effect (P < .05) influenced LH concentrations. During Cycle -2, LH concentrations were similar for M and R heifers, but during Cycle -1, M heifers had greater LH concentrations than did R heifers. Concentrations of FSH were greater (P < .05) in R than M heifers after induced luteolysis when R heifers failed to ovulate. There was a treatment x cycle interaction (P < .05) for IGF-I concentrations, and M heifers had 4.7- and 8.6-fold greater IGF-I concentrations than did R heifers during Cycle -2 and -1, respectively. We conclude that growth rate and diameter of the ovulatory follicle, and concentrations of LH, estradiol, and IGF-I are reduced before the onset of nutritionally induced anovulation in beef heifers.     

Ovarian follicular growth, function and turnover in cattle: a review

Studies in cattle assessing changes in number and size of antral follicles, concentrations of estradiol, androgens and progesterone in serum and follicular fluid, and numbers of gonadotropin receptors per follicle during repetitive estrous cycles and postpartum anestrus are reviewed. The rate of growth of small follicles (1 to 3 mm) into larger follicles increases as the estrous cycle progresses from d 1 to 18 (d 0 = estrus). Size of the largest antral follicle present on the ovary also increases with advancement of the estrous cycle. Most large follicles (greater than 10 mm) persist on the ovarian surface for 5 d or more between d 3 and 13 of the bovine estrous cycle. After d 13, most of these large follicles are replaced more frequently by new growing follicles (turnover) with an increased probability for recruitment of the ovulatory follicle after d 18. More research is needed to determine the time required for growth of bovine follicles from small to large antral size and evoke recruitment of the ovulatory follicle. Factors that regulate selection of the ovulatory follicle are unknown but may involve increased frequency of LH pulses in blood, altered blood flow and(or) changes in intrafollicular steroids and proteins. Quantitative evaluation of ovarian follicles indicated occurrence of consistent short-term changes in fluid estradiol and numbers of luteinizing hormone receptors in cells of large follicles only during the pre-ovulatory period. Presumably, low concentrations of follicular estradiol found during most of the estrous cycle are not due to a lack of aromatizable precursor or follicle-stimulating hormone receptors. Follicular fluid concentrations of progesterone increase only near the time of ovulation. Little is known about changes in follicular growth, turnover and function during postpartum anestrus in cattle. However, preliminary data suggest that the steroidogenic capacity of large follicles changes markedly during the postpartum period.     

Effect of deslorelin acetate on gonadotropin secretion and ovarian follicle development in cycling mares

OBJECTIVE: To evaluate gonadotropin secretion and ovarian function after administration of deslorelin acetate to induce ovulation in mares. DESIGN: Randomized controlled trial. ANIMALS: 16 healthy mares with normal estrous cycles. PROCEDURE: 8 control mares were allowed to ovulate spontaneously, whereas 8 study mares received deslorelin to induce ovulation when an ovarian follicle > 35 mm in diameter was detected. Follicle development and serum concentrations of gonadotropins were monitored daily during 1 estrous cycle. Pituitary responsiveness to administration of gonadotropin-releasing hormone (GnRH) was evaluated 10 days after initial ovulation. RESULTS: Interovulatory intervals of mares treated with deslorelin (mean +/- SD, 25.6 +/- 2.6 days) were longer than those of control mares (22.9 +/- 1.8 days). Diameter of the largest follicle was significantly smaller during 2 days of the diestrous period after ovulation in deslorelin-treated mares than in control mares. Concentrations of follicle-stimulating hormone (FSH) were lower in deslorelin-treated mares on days 5 through 14 than in control mares. Concentrations of luteinizing hormone were not different between groups during most of the cycle. Gonadotropin release in response to administration of GnRH was lower in mares treated with deslorelin acetate than in control mares. CONCLUSIONS AND CLINICAL RELEVANCE: Administration of deslorelin was associated with reduction in circulating concentrations of FSH and gonadotropin response to administration of GnRH during the estrous cycle. Low concentration of FSH in treated mares may lead to delayed follicular development and an increased interovulatory interval.     

On the negative feedback regulation of gonadotropins in castrate and intact cattle with comparison of two FSH radioimmunoassays

Two homologous radioimmunoassays for bovine follicle stimulating hormone (bFSH) were utilized in comparing the differential regulation of FSH and luteinizing hormone (LH) in response to ovariectomy or administration of gonadal steroids in cattle. There appeared to be significant LH cross-reactivity in one of the bFSH systems (bFSH-HS-2-17), but not in the other (bFSH-BP3). Concentrations of FSH in plasma measured by these two systems suggested both qualitative and quantitative differences. Following ovariectomy in heifers, LH concentrations in plasma were increased by 7.5 h, while FSH (measured in the bFSH-BP3 system) was not significantly elevated until 18 h. Administration of 200 micrograms of estradiol-17 beta to ovariectomized heifers inhibited levels of FSH in plasma but large doses of testosterone (100 mg), androstenedione (400 mg) and dihydrotestosterone (800 mg) had no effect. Similarly, LH was not affected by the androgens, while estradiol induced LH surges, leading to increased mean LH concentrations. In contrast to the results in heifers, LH concentrations in plasma from steers were inhibited by administration of androgens as well as by estradiol. In steers, FSH (bFSH-BP3) was marginally inhibited by estradiol and not at all by the androgens. Differences in the secretory patterns of FSH and LH also occurred in intact heifers during the estrous cycle. The 72-h period preceding estrus (follicular phase) was characterized by rapidly declining serum progesterone concentrations, followed by concurrent increases in both LH and estradiol. The circulating levels of bFSH (BP3) tended to decline during this interval. Overall, during the estrous cycle, progesterone levels were positively correlated with bFSH-BP3 (r = .37) and negatively correlated with LH (r = -.39). The gonadotropins were not significantly related (r = -.15). These relationships are consistent with the concept that LH controls the final stages of follicular development in cattle and that FSH may exert only a permissive effect.     

The effect of active immunization against inhibin on gonadotropin secretions and follicular dynamics during the estrous cycle in cows

The hypothesis of the present study is that active immunization of cows against inhibin would neutralize endogenous inhibin, increase circulating levels of follicle stimulating hormone, and subsequently affect follicular dynamics and the ovulation rate during the estrous cycle. Thirteen cows were immunized against inhibin alpha-subunit and, 6 cows were immunized with a placebo. Both groups were given 4 booster immunizations 7, 14, 21, and 34 weeks after the primary injection. Ovaries were examined daily after the 2nd, 3rd, and 4th booster immunizations by transrectal ultrasonography for 25 days. After the 4th booster immunization, blood samples were collected daily for one complete estrous cycle to measure FSH and LH. The results showed that the immunized cows generated antibodies against inhibin, and that they had higher FSH levels compared with the controls. The number of follicular waves during the estrous cycle was higher in the immunized cows (3 or 4 waves) than in the controls (2 or 3 waves). Moreover, the immunized cows had a greater number of follicles during the estrous cycle compared with the control cows. The maximum number of follicles was 14.8 +/- 1.7 vs 5.4 +/- 0.2 in inhibin-immunized and control cows, respectively, during the first follicular wave and 13.9 +/- 1.9 vs 5.6 +/- 0.7, respectively, during the ovulatory wave. Multiple ovulations were increased in the immunized cows. However, the ovulation rate varied greatly in the immunized animals. In conclusion, immunization against inhibin increased FSH secretions during the estrous cycle in the cows. Moreover, the immunized cows had a greater number of follicular waves during the estrous cycle and a greater number of follicles, and this could be used as a potential source of oocytes for use in IVF/embryo transfer programs.     

Use of a small dose of estradiol benzoate during diestrus to synchronize development of the ovulatory follicle in cattle

We tested the hypothesis that a small dose of estradiol benzoate (EB) at the midstage of the estrous cycle in cattle would synchronize the subsequent pattern of ovarian follicular development, estrus, and ovulation. Nonlactating Friesian cows received either 1 mg of EB i.m. on d 13 of the estrous cycle (T; n = 12; estrus = d0) or served as untreated controls (C; n = 12). Their ovaries were examined daily with transrectal ultrasonography from d 7, and blood samples were collected 0, 2, 4, 8, 24, and 48 h after treatment on d 13. Plasma concentrations of estradiol-17beta were elevated to 12 pg/mL during the initial 24 h following treatment, compared with a baseline of 1 pg/mL in untreated controls (P < .001). Progesterone concentrations in cows of the T group declined between 24 and 48 h after treatment (-3.2 +/- .5 ng/mL) compared with little change in concentrations of progesterone in cows of the C group at this time (P < .01). This difference was coincident with an earlier time to regression of the corpus luteum in cows of the T group. Disregarding treatment groups, the second dominant follicle of the estrous cycle (DF2) emerged on d 10.6 +/- .3 and was 9.4 +/- .4 mm in diameter on d 13. Further growth of the DF2 was halted by EB treatment on d 13. Cessation of growth occurred irrespective of whether the DF2 was in the early or late growth phase, and a new follicular wave emerged 4.5 +/- .2 d later. The dominant follicle from this wave (DF3) ovulated 5 d after emergence in most cases. During the estrous cycle of every cow in the T group, there were three waves of follicular development (3-wave), whereas the ratio of 2:3 waves of follicular development in cows of the C group was 1:3. Consequently, the interval from emergence to ovulation of the ovulatory dominant follicle in cows of the C group ranged from 3 to 11 d. The dynamics of ovarian follicular wave development during the estrous cycle can be strategically manipulated by treating with a small dose of EB to synchronize proestrous development of the ovulatory follicle.     

Dynamics of ovarian follicular development in cattle during the estrous cycle, early pregnancy and in response to PMSG

Ovarian follicular dynamics of cattle were examined during the estrous cycle, early pregnancy and in response to PMSG. Number and size of follicles were monitored by ultrasonographic examinations. During the estrous cycle, distinct periods of follicular dominance (measured by the increase in difference in size between the largest and second largest follicle) occurred in both the luteal (Days 6-8) and proestrus (18-22) phases of the estrous cycle (two follicular waves). Associated with the well timed development of the first dominant follicle was a change in distribution of follicle numbers in small (less than 5 mm; increased on Days 2-4), medium (6-8 mm; increased on Days 3-5) and large (greater than or equal to 9 mm; increased on Days 6-9) follicular size classes. Follicular development was greater on the ovary bearing the CL for the period that the CL was present. The dominant follicle formed during the first follicular wave was capable of ovulating (6 of 8 heifers) following an injection of a synthetic analogue of prostaglandin F-2 alpha on Day 9 of the estrous cycle. During early pregnancy (Days 6-34), follicular development (size of largest follicle, number of follicles and total accumulated size of all follicles) on the ovary bearing the CL was suppressed between Days 24 and 34 of pregnancy. This was a local effect in that follicular development was sustained on the contralateral ovary. Therefore, the CL or conceptus may be regulating follicular development in a manner to help prevent luteolysis. Associated with the injection of PMSG was an initial increase in the number of small follicles followed by their recruitment into medium and large size classes leading to ovulation. Number of follicles greater than 5 mm on the Day of estrus was related (r = .97) to the number of subsequent embryos and oocytes collected. Ultrasonography is a valuable technique to monitor ovarian follicular dynamics in cattle, and can thereby be used to infer changes in physiological and endocrine states.     

Endocrinology of number of follicular waves per estrous cycle and contralateral or ipsilateral relationship between corpus luteum and preovulatory follicle in heifers

A 3-d extension of the luteal phase occurs in interovulatory intervals (IOIs) with a contralateral relationship between the corpus luteum (CL) and preovulatory follicle with 3 follicular waves (Contra-3W group). Concentrations of FSH, progesterone, LH, and estradiol-17β for the ipsilateral versus contralateral CL and/or follicle relationship and 2 versus 3 waves per IOI were studied in 14 heifers. Follicular waves and FSH surges were designated 1, 2, or 3, according to order of occurrence in the IOI. The day (day 0 = ovulation) of the FSH peak in surge 2 occurred earlier (P < 0.02) in 3-wave IOIs (day 6.3 ± 0.5) than in 2-wave IOIs (day 8.5 ± 0.5). Mean FSH was higher in 3-wave than in 2-wave IOI on 82% of the days in the IOI. Repeatability or individuality in FSH concentration was indicated by a correlation (r = 0.54, P < 0.04) in FSH concentrations between ovulations at the beginning and at the end of the IOI. Concentrations of LH and estradiol increased (P < 0.05) near the beginning of the luteolytic period in 2-wave IOI regardless of the CL and/or follicle relationship. In the Contra-3W group, LH and estradiol remained at basal concentrations concurrently with FSH surge 3 and extension of the luteal phase. The hypotheses were supported that FSH surge 2 occurs earlier in 3-wave IOIs than in 2-wave IOIs and that the development of 3-wave IOIs occurs in individuals with greater FSH concentrations. Extension of the luteal phase in the Contra-3W group was temporally associated with lower concentrations of LH and estradiol.     

Factors that affect ovarian follicular dynamics in cattle.

Studies of ovarian follicular dynamics in cattle may lead to methods for improving fertility, for synchronizing estrus with more precision, and for enhancing superovulatory responses. Within an estrous cycle, two or three large (> 10 mm) follicles develop during consecutive waves of follicular growth. The last wave provides the ovulatory follicle, whereas preceding wave(s) provide follicles that undergo atresia. The life span of large follicles seems to depend on the pulsatile secretion of LH; decreased frequency of LH pulses results in atresia of large follicles. Aromatase activity in the walls of the largest follicles is greatest during the first 8 d of the estrous cycle and decreases by d 12. Steroidogenesis of the largest and second-largest ovarian follicles differs on d 5, 8, and 12 of the estrous cycle. Follicular dynamics are altered by negative energy balance and lactation. The number of large follicles and concentration of estradiol during the preovulatory period differs between postpartum lactating and nonlactating cows. Dietary fats stimulate follicular growth when they are fed to increase energy balance. Administration of bovine somatotropin decreases energy balance and has a differential effect on ovarian follicular responses; growth of the largest follicle does not change, but growth of the second-largest follicle is stimulated by somatotropin. Studies of follicular dynamics in lactating cows demonstrate changes in ovarian function associated with energy balance that may be related to inefficient reproductive performance of cows producing high yields of milk.     

Effects of low-dose follicle-stimulating hormone administration on follicular dynamics and preovulatory follicle characteristics in dairy cows during the summer

The well-documented phenomenon of reduced conception rate in dairy cows during the hot season involves impaired functioning of the ovarian follicles and their enclosed oocytes. Three experiments were performed to examine the administration of low doses of follicle-stimulating hormone (FSH) to induce turnover of follicles that are damaged upon summer thermal stress and to examine whether this FSH administration has beneficial effects on preovulatory follicles. In experiment 1, synchronized heifers were treated with 100 mg of Folltropin-V (n = 7) or 4.4 mg of Ovagen (n = 6) on day 3 of the estrous cycle. Treatment with both FSH sources resulted in greater (P < 0.05) numbers of follicles than in control animals (n = 12) on day 6 of the estrous cycle, indicating that low doses of FSH can increase the number of emerging follicles in a follicular wave. In experiment 2, milking cows were assigned to a control group (n = 4) or treated with 2.2 mg (FSH-2.2; n = 6) or 4.4 mg (FSH-4.4; n = 5) Ovagen. Follicle-stimulating hormone was administrated on day 3 or 4 and day 10 or 11 of the estrous cycle, coinciding with emergence of the first and second follicular waves, respectively. The number of follicles emerging during the first wave tended to be higher (P < 0.1) in FSH-4.4-treated cows than in controls. The second-wave dominant follicles emerged 2 d later in the treated cows and were smaller in diameter (P < 0.05) than controls, 2 d before aspiration. Despite being younger, the preovulatory follicles of FSH-4.4 cows expressed a steroidogenic capacity that was similar to controls with a tendency toward greater insulin concentrations (P < 0.09). In experiment 3, milking cows were assigned to a control group (n = 6) or treated with 4.4 mg Ovagen (FSH-4.4; n = 6). Follicle-stimulating hormone was administrated on day 3 and day 12 or 13 of the estrous cycle. The number of emerging follicles was higher (P < 0.05) in the treated vs control cows. However, the features of the preovulatory follicle developed in the subsequent cycle did not differ between groups. In summary, low doses of FSH can efficiently induce follicular turnover accompanied by a modest effect on the preovulatory follicle of the treated cycle. It appears that the administration of low doses of FSH, precisely timed to synchronize with the emergence of follicular waves, might have a beneficial effect on the preovulatory follicle and its enclosed oocyte.     

Follicle Deviation in Ovulatory Follicular Waves with One or Two Dominant Follicles in Mares

The follicle and hormone aspects of diameter deviation and development of one dominant (≥28 mm) follicle (1DF) vs two dominant follicles (2DF) were studied in 32 ovulatory follicular waves in mares. Follicles were ranked each day as F1 (largest) to F3. The beginning of deviation was designated day 0 and preceded the first increase in the differences in diameter between F1 and F2 in the 1DF group and between a combination of F1 and F2 vs F3 in the 2DF group. One dominant follicle and 2DF developed in 21 (66%) and 11 (34%) waves, respectively. Double ovulations occurred in only one of the waves with 2DF. In 8/11 waves with 2DF, a second deviation occurred between F1 and F2 on 2.5 ± 0.4 days after the first deviation. On day 0, 1DF and 2DF waves were similar in number of days after ovulation, number of follicles, difference in diameter between F1 and F2, and plasma concentrations of LH, estradiol and immunoreactive inhibin. The interval from maximum FSH concentration to day 0 was longer (p < 0.05) and FSH concentration was lower (p < 0.05) on days -1 to 4 in the 2DF group. The similarities on day 0 in the characteristics of 1DF and 2DF waves despite the differences in the declining portions of the FSH profile indicated that a specific day of the FSH decline or a specific concentration were not factors in initiating deviation. Unlike reported results in heifers, the results in mares did not indicate a hormonal basis for the development of 2DF or two deviations.     

Ovarian follicular dynamics in Caspian mares

The characteristics of the major follicular waves (primary and secondary) throughout estrous cycle were studied in 7 healthy Caspian mares (age, 4-15 years; weight, 198.6 ± 0.9 kg) during the breeding season. Ovarian follicular dynamics were monitored by using an ultrasound scanner equipped with a 5-MHz, B-mode, linear-array, rectal transducer throughout 2 complete estrous cycles. The diameters of antral follicles (5 mm) were measured, averaging the narrowest and widest dimensions. To detect follicular wave emergence, the diameter profile of the 3 largest follicles per ovary of each mare was determined without considering day-to-day identity of follicles but with maintenance of distinction between left and right ovaries. The primary waves originated on day 6.4 ± 0.81 (ovulation = day 0) when the mean diameter of ovarian follicles was 9.6 ± 1.05 mm. Divergence between the dominant preovulatory follicle and subordinate follicles occurred on day 13.4 ± 0.81, when the dominant follicle was 18.1 ± 2.67 mm in diameter. The intervals from emergence to divergence and from divergence to ovulation were 7 ± 0.68 and 8.7 ± 0.68 days, respectively. Secondary major follicular waves were not observed during this study. In conclusion, only 1 major follicular wave was detected in a Caspian mare, confirming the data previously described in other equine breeds. It is also indicated that the occurrence of 1 major follicular wave per cycle is a more common phenomena in equine species.     

Dynamics of the Equine Preovulatory Follicle and Periovulatory Hormones: What's New?

Recent studies (2005–2008) on the interrelationships among the preovulatory follicle and periovulatory circulating hormones are reviewed. Close temporal and mechanistic relationships occur between estradiol/inhibin and follicle-stimulating hormone (FSH), between estradiol and luteinizing hormone (LH), and between progesterone and LH. Estradiol from the dominant follicle forms a surge that reaches a peak 2 days before ovulation. Estradiol, as well as inhibin, has a negative effect on FSH, and estradiol has a negative effect on LH. When estradiol decreases, the negative effect diminishes and accounts for the beginning of an FSH increase and a transition from a slow to rapid increase in LH on the day of the estradiol peak. The decrease in estradiol and the reduction or cessation in the growth of the preovulatory follicle beginning 2 days before ovulation are attributable to the development of a reciprocal negative effect of LH on follicle estradiol production when LH reaches a critical concentration. The LH decrease after the peak of the LH surge on the day after ovulation is related to a negative effect of a postovulatory increase in progesterone. Measurable repeatability within mares between consecutive estrous cycles occurs during the preovulatory period in diameter of the ovulatory follicle and concentrations of LH and FSH. Hormone-laden follicular fluid passes into the peritoneal cavity at ovulation and transiently alters the circulating concentrations of LH and FSH. Double ovulations are associated with greater estradiol concentrations and reduced concentrations of FSH.     

德州驴发情周期中卵泡动力学和激素变化规律的研究

本研究旨在揭示德州母驴发情季节的卵巢卵泡波变化特点以了解其卵泡的募集、选择、优势化、闭锁或者排卵,为驴人工授精和超数排卵提供可靠依据.本研究在德州驴发情季节(5～8月份)用直肠超声探查法每日检测5头德州母驴的10个排卵间隔的卵泡发育动态.同时每日采血,用放免法(RIA)检测E2、FSH、LH和IGF-1浓度.研究结果发现:(1)驴的排卵间隔平均为(23.22±0.28)d,每个周期中存在2个卵泡波,其中有4个展示主-主卵泡波,有6个展示次-主卵泡波.(2)第1波和第2波开始出现募集的时间分别在排卵后的第1和第(10.2±1.4)天,募集的卵泡数分别为(5.2±1.2)和(4.0±0.3)个,选择的卵泡数分别是(2.3±0.5)和(2.5±0.4)个.第1和第2卵泡波开始优势化的时间分别在排卵后的第(5.8±0.6)和第(18.1±1.4)天.波峰平均出现时间分别为第(7.67±0.24)和第(23.22±0.28)天.波峰时优势卵泡平均最大直径分别为(26.44±2.37)和(40.7±2.45)mm.第1和第2波最大优势卵泡每日平均生长率分别为(2.01±0.5)和(3.15±0.6)mm·d-1.(3)黄体(CL)最大直径平均为(37.61±1.58)mm,CL平均持续时间为(17.30±1.62)d.(4)E2浓度从排卵前3d至排卵后2d的发情期内均很高,平均维持在(46.47±9.53)Pg·mL-1,在排卵前2d达到峰值(62.84±4.89)Pg·mL-1.(5)由于E2的负反馈作用,FSH浓度到优势卵泡最后阶段(即第7天闭锁或第23或24天排卵时)大幅度下降.(6)LH在排卵前几天迅速升高,在排卵后第2天达到峰值(4.69±0.43)mIU·mL-1.(7)IGF-1的升降状况与E2基本相同,在排卵前1d达到峰值(47.9±3.91)ng·mL-1.通过上述卵泡和激素动态学变化,本研究得出下述结论:驴的卵巢卵泡发育是以卵泡波形式进行的,每个发情周期中最常出现的是2个卵泡波;排卵发生于发情结束前2 d;E2和IGF-1调节驴的发情活动;FSH调节小卵泡的募集;LH增高与排卵有关.     

Gonadotropin-releasing hormone-induced ovulation and luteinizing hormone release in beef heifers: effect of day of the cycle

The COSynch protocol has been used to synchronize ovulation and facilitate fixed-time AI in beef cattle. Establishment and maintenance of pregnancy was negatively affected, in previous studies, by GnRH-induced ovulation of small dominant follicles (/=10 mm) and increased ovulatory response after GnRH 2.     

Physiology and Endocrinology Symposium: Harnessing basic knowledge of factors controlling puberty to improve synchronization of estrus and fertility in heifers

The development of replacement heifers is a major economic investment for all beef and dairy operations. The costs associated with heifer development cannot be recovered if heifers do not conceive and remain productive in the herd; therefore, heifers need to conceive early in the breeding season or risk being culled. Previous research has reported up to a 21% increase in fertility from pubertal estrus to the third estrus of a heifer. The use of reproductive tract scores to determine pubertal status has demonstrated that peripubertal and pubertal heifers have increased pregnancy success to estrous synchronization compared with heifers that were prepubertal. The development of RIA has allowed accurate measurement of peripheral blood hormone concentrations associated with the pubertal process and luteal formation. This basic knowledge has increased our understanding of the mechanisms that control puberty in heifers. In addition, understanding the hormonal changes that occur during the estrous cycle has allowed for the development of estrous synchronization protocols that result in increased control of follicular growth, regression of luteal tissue, and ovulation. Transrectal ultrasonography has increased our understanding of follicular waves; this understanding led to research investigating the endocrine regulation of follicular waves and development of methods to synchronize follicular waves for purposes of fixed-time AI. Current topics of research include the effect of antral follicle count on fertility and the effect of maternal nutrition (on the fetus in utero) on subsequent reproductive potential of a heifer (i.e., fetal programming). Advancements in genomic technologies will likely provide a powerful tool for selecting heifers at birth that will have a greater probability of being reproductively successful if managed correctly. Therefore, knowledge gained through basic research on factors that control puberty has improved and will continue to improve heifer development and fertility.