Changes in the peripheral concentrations of inhibin, follicle-stimulating hormone, luteinizing hormone, progesterone and estradiol-17beta during turnover of cystic follicles in dairy cows with spontaneous follicular cysts

Several studies have clarified that the follicular cysts degenerate and are replaced by newly growing follicles that develop into new follicular cysts without ovulation, i.e., turnover of ovarian follicular cysts in cows. However, the relativity of endocrinological changes, including the inhibin profile during turnover of spontaneous follicular cysts in dairy cows, is still unclear. In the present study, the relationship between turnover of follicular cysts and changes in the peripheral blood concentrations of progesterone (P), estradiol-17beta (E(2)), luteinizing hormone (LH), follicle stimulating hormone (FSH) and inhibin were examined in lactating dairy cows. Five cows diagnosed with follicular cysts (follicles of more than 25 mm in diameter in the absence of a corpus luteum) were investigated. Their ovarian dynamics were monitored using ultrasonography, and blood samples were collected at 2- or 3- day intervals throughout the experiment. The day when a follicle fated to become a follicular cyst reached more than 8 mm in diameter was defined as the start of a cystic follicular wave. Four of the 5 cows exhibited a similar patterns of cystic follicular changes and hormone profiles. The data from the 4 cows was used for analysis of the relationships between turnover of cystic follicles and the hormone profiles. Two or three new cystic follicular waves occurred in each cow during the experimental period. The mean diameter of the cystic follicles was more than 25 mm 13 to 15 days after the start of the cystic follicular wave, and it began to decrease 1 to 6 days before the start of the subsequent cystic follicular wave. The levels of E(2) and inhibin tended to decrease for 7 to 9 days before the start of a new cystic follicular wave and to increase concomitantly with new follicular cyst growth. The levels of FSH rose for 1 to 3 days before the start of a new cystic follicular wave. The present study clarified the relationship between FSH and inhibin during turnover of spontaneous follicular cysts in dairy cows and found that it was very similar to previous results for cows. The present results suggest that an increase in FSH secretion following a reduction in inhibin secretion triggers turnover of cystic follicles in cows with spontaneous follicular cysts.     

Passive immunoneutralization of endogenous inhibin increases ovulation rate in miniature Shiba goats

We reported previously that passive immunization against inhibin enhances follicular growth and increases the ovulation rate. However, the ovulation rate was not comparable to the number of follicles. Therefore, the aim of this study was to attempt to increase the ovulation rate by increasing the interval between inhibin immunization and PGF2alpha injection. Five miniature Shiba goats were treated with 10 ml inhibin antiserum (inhibin-AS) developed against [Tyro30]-inhibin alpha (1-30). A control group (n=5) was treated with normal goat serum. All animals were injected intramuscularly with 125 microg PGF2alpha 72 h after treatment to induce estrus and ovulation. Blood samples were collected for hormonal assay and the ovulation rate was determined by laparotomy. In contrast to the control group, there was a significant increase in plasma concentrations of FSH in the immunized group. After luteolysis, plasma concentrations of estradiol-17beta increased markedly to a preovulatory peak about 2 folds higher (P<0.01) than that of controls. In addition, the ovulation rate was greater in the immunized group (14.4 +/- 2.2) than in the control group (2.2 +/- 0.6), and the mean number of follicles > or = 4 mm in diameter was 10.0 +/- 0.8 in the inhibin-AS group compared with 2.4 +/- 0.3 in control group. The present results demonstrate that immunoneutralization of endogenous inhibin increased FSH secretions in miniature shiba goats. The increased FSH secretion enhanced follicular growth and increased the ovulation rate. Additionally, increasing the interval between inhibin-AS and PGF2alpha injections (to 72 h) resulted in a greater ovulation rate compared with the previous protocol (48 h). Therefore, inhibin-AS treatment proved to be an effective alternative to exogenous gonadotropin methods for induction of superovulation in goats.     

Ovarian follicular dynamics and hormones throughout the estrous cycle in Thai native (Bos indicus) heifers

Relatively few studies have been reported regarding the reproductive physiology of female Thai native cattle. Therefore, the objective of the present study was to evaluate the follicular dynamics and concentrations of follicle stimulating hormone (FSH), estradiol (E2) and progesterone (P4) during the estrous cycle in Thai native heifers (TNH) and to compare obtained results with those of European and Indian cattle breeds previously reported. For the detection of estrus, ovaries of all 20 heifers were examined twice daily (12 h intervals) by ultrasonography for three consecutive estrous cycles. From data of 60 estrous cycles (n = 60 estrous cycles from 20 heifers), it was found that 14 (70%) and 6 heifers (30%) had two (42 estrous cycles collected from 14 heifers) and three follicular waves (18 estrous cycles collected from 6 heifers), respectively. The days when estrus was detected, interovulatory intervals, life‐spans of corpus lutea (CL), and days for growing and regression of CLs were shorter in the two follicular waves than those in the three follicular waves (P < 0.05). In both two and thre follicular waves, larger maximum diameters and higher growth rates of the dominant follicle (DF) in an ovulatory wave were observed than those of the preceding waves without ovulation (P < 0.05). There was a progressive increase in follicular size and FSH and E2 production during follicular growth in each follicular wave. In addition, the FSH and E2 peak concentrations during the ovulatory wave were higher than those of the anovulation waves (P < 0.05). Moreover, although the ovarian follicular dynamic patterns in Thai native heifers were similar to those previously reported for European and Indian cattle breeds, the diameter of the largest preovulatory follicle (OF), subordinate follicles (SF) and CLs were smaller than those in European and Indian cattle breeds. In conclusion, when compared with European and some breeds of Indian cattle, the length of interovulatory intervals was shorter, and the sizes of dominant SF and CLs were smaller in Thai native heifers.     

Plasma concentrations of inhibin A in cattle with follicular cysts: relationships with turnover of follicular waves and plasma levels of gonadotropins and steroid hormones

We investigated the profiles of circulating levels of inhibin A and total inhibin in beef cows with follicular cysts in relation to the patterns of follicular development and circulating gonadotropins and steroid hormones. Turnover of follicular waves was monitored in five cows every 2 days for 70 days from 10 days after detection of estrus without ovulation. The mean interwave intervals were 19.6 ± 1.0 days (n = 18 waves with cysts from the five cows). Circulating levels of inhibin A were approximately 170 pg/ml before emergence of follicular waves with cysts and increased (P < 0.05) concomitantly with follicle emergence. High concentrations of inhibin A (greater than 300 pg/ml) were noted for 7 days during the growth phase of cystic follicles, but inhibin A levels decreased gradually when development of the cysts reached a plateau. This profile of inhibin A was similar to those of total inhibin and estradiol, but was inversely related to the changes in plasma FSH concentrations. LH pulse frequency and mean concentrations of LH in cows with cysts were higher than those observed in the luteal phase of normal cyclic cows. These results indicate that the capacity to secrete inhibin, as well as estradiol, is maintained in cystic follicles, the growth of which is extended by LH secretion at levels greater than those seen in the normal luteal phase. Inhibin A plays an important role in the extension of interwave intervals by suppressing recruitment of a new cohort of follicles.     

Dynamic changes in plasma concentrations of gonadotropins, inhibin, estradiol-17beta and progesterone in cows with ultrasound-guided follicular aspiration

To elucidate the effects of ultrasound-guided transvaginal follicular aspiration, plasma concentrations of FSH, LH, inhibin, estradiol-17beta and progesterone, and folliculogenesis were examined in Holstein cows. Four clinically healthy cows with regular estrous cycles were scanned by ultrasound per rectum once a week for 9 weeks before the commencement of follicular aspiration. All visible follicles were divided into 3 categories based on their sizes (2 < or = small < 5 mm; 5 < or = medium < 10 mm, large > or = 10 mm). The follicular aspiration was started at random during the estrous cycle and conducted under epidural anesthesia induced with 5 ml of 2% lidocaine once a week for 6 weeks. The average number of total visible follicles > or = 2 mm in diameter at 7 days after aspiration (21.7 +/- 7.4, n = 24) was similar to that before starting aspiration (26.7 +/- 10.5, n = 36). Plasma inhibin and estradiol-17beta declined and fell to a trough on 1.5 days and returned to pre-aspiration values by 5 days after aspiration. Plasma concentrations of FSH increased and reached peak levels between 1 and 1.5 days after aspirations. Plasma concentrations of LH also increased and reached peak levels between 0.5 and 1.5 days after aspirations. Both plasma FSH and LH had returned to pre-aspiration levels by 5 days after aspirations. Plasma concentrations of progesterone did not change with the follicular aspiration. These results demonstrate that follicular aspiration decreases plasma concentrations of inhibin and estradiol-17beta, which in turn leads to a rise in plasma concentrations of FSH and LH. It is suggested that marked increases in plasma concentrations of FSH and LH after the aspiration stimulate the development and maturation of a new cohort of follicles within one week in cows.     

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.     

Effect of Suppression of FSH with a GnRH Antagonist (Acyline) Before and During Follicle Deviation in the Mare

A GnRH antagonist (Acyline) was used to study the role of FSH in early development of a follicular wave in 61 mares. In Experiment 1, a single dose of 3 mg per mare, compared with 0 and 1 mg, suppressed both the FSH and follicle responses to exogenous GnRH. In Experiment 2, high concentrations of FSH were induced by two successive ablations of all follicles ≥ 6 mm on days 10 and 13 (day 0 = ovulation). A single treatment with Acyline resulted in significantly greater suppression of plasma concentrations of FSH than a single treatment with charcoal-extracted follicular fluid (source of inhibin) or oestradiol. Suppression of FSH was not significantly different between the group treated with Acyline alone and a group treated with a combination of Acyline, inhibin and oestradiol. In Experiment 3, all follicles were ablated on day 10 to induce an FSH surge and a new follicular wave. Acyline treatment on day 10 resulted in an immediate decrease in FSH, without a significant effect on day of emergence of a new wave or growth of follicles from 7 to 11 mm on days 11–13. Treatment on day 15, a day before expected follicle deviation and after the peak of the wave-stimulating FSH surge, resulted in an immediate decrease in FSH and cessation of follicle growth. Results indicated that growth of follicles for about 2 days after wave emergence was independent of FSH. In contrast, during the decline in the wave-stimulating FSH surge and before follicle deviation, growth of follicles was dependent on FSH.     

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.     

Follicular turnover and hormonal association in postpartum goats during early and late lactation

To clarify the effect of lactation period on ovarian follicular activity and associated hormonal levels in goats, six goats were monitored daily by ultrasonographic examination with blood sampling during early (Days 5 to 25; Day 0 was the day of kidding) and late (Days 40 to 60) lactation. While the presence of a corpus luteum of pregnancy retarded follicular growth in the ipsilateral ovary until Days 11-13 postpartum, the total follicular number (TFN) and area (TFA) increased during late lactation due to the significant increase in the number of medium- and large-sized follicles and decrease in the number of small follicles. Four goats showed a similar pattern of follicular development during the period studied characterized by the emergence of five and six waves during the early and late lactation, respectively. The largest follicle diameter of the first three waves monitored during early lactation was significantly smaller as compared with the diameter of those existing during late lactation. TFN showed a positive correlation with FSH but showed a negative correlation with immunoreactive (ir-) inhibin and estradiol during the postpartum period. TFA was positively correlated with ir-inhibin, estradiol and PRL and negatively correlated with FSH during the monitored periods. The plasma levels of ir-inhibin and progesterone were significantly higher during late lactation compared with the levels recorded during early lactation. Ir-inhibin levels showed a significant positive correlation with LH and estradiol during early and late lactation but showed a negative correlation with FSH during the whole lactation period. LH was positively correlated with estradiol and PRL during early and late lactation, respectively. These results suggest that the lactation period has a detrimental effect on ovarian activity during the early postpartum period in goats.     

Effect of active immunization of pony mares against recombinant porcine inhibin alpha subunit on ovarian follicular development and plasma steroids and gonadotropins

Two pony mares were immunized against recombinant porcine inhibin alpha subunit three times with 39 day intervals. Clinical findings and endocrinological changes before immunization were taken as the control. The first significant rise in the anti-inhibin titre (P<0.05) in the circulation was found 27 days after the first injection. Maximum binding activity was reached by the 12th day after the second booster dose. The number of small, medium and large sized follicles had increased significantly compared to before immunization (11.75 +/- 4.30, 2.75 +/- 0.69 and 2.51 +/- 0.63 vs 6.50 +/- 1.43, 1.83 +/- 0.44 and 1.33 +/- 0.38, respectively), but the ovulation rate remained unchanged after immunization. The average plasma concentration of FSH and estradiol-17beta during the estrous cycle increased significantly (P<0.05) after immunization. These results suggest that immunization against inhibin is a useful tool to increase the number of ovarian follicles during the estrous cycle of pony mares. Moreover, the present study supported the concept that inhibin plays a major role in the control of follicular growth through its inhibitory effect on FSH secretion synergistically with steroid hormones.     

Follicle-stimulating hormone isoforms and plasma concentrations of estradiol and inhibin A in dairy cows with ovulatory and non-ovulatory follicles during the first postpartum follicle wave

Following parturition, all cows display a wave of ovarian follicular growth, but a large proportion fail to generate a preovulatory rise in estradiol, and hence fail to ovulate. Follicle-stimulating hormone (FSH) exists as multiple isoforms in the circulation depending on the type and extent of glycosylation, and this has pronounced effects on its biological properties. This study examined differences in plasma FSH, estradiol, and inhibin A concentrations, and the distribution of FSH isoforms in cows with ovulatory or atretic dominant follicles during the first postpartum follicle wave. Plasma FSH isoform distribution was examined in both groups during the period of final development of the dominant follicle by liquid phase isoelectric focusing. Cows with an ovulatory follicle had higher circulating estradiol and inhibin A concentrations, and lower plasma FSH concentrations. The distribution of FSH isoforms displayed a marked shift toward the less acidic isoforms in cows with ovulatory follicles. A higher proportion of the FSH isoforms had a pI>5.0 in cows with ovulatory follicles compared to those with atretic follicles. In addition, cows with ovulatory follicles had greater dry matter intake, superior energy balance, elevated circulating concentrations of insulin and insulin-like growth factor-I, and lower plasma nonesterified fatty acids. The shift in FSH isoforms toward a greater abundance of the less acidic isoforms appears to be a key component in determining the capability for producing a preovulatory rise in estradiol, and this shift in FSH isoforms was associated with more favorable bioenergetic and metabolic status.     

Physiological roles of inhibin in regulation of FSH secretion and follicular development during early pregnancy in goats

The aim of the current study was to clarify the physiological role of inhibin in controlling FSH secretion and follicular development during the early pregnancy in goats. Eight goats investigated sonographically on Days 19-21 (Day 0=day of mating) for pregnancy were assigned into control (n=3) and treated (n=5) groups. The ovaries of all animals were daily scanned with ultrasound for follicles 2mm or more in diameter from 1 day before to 5 days after treatment. On Day 25 postbreeding; animals received either 10 ml, of normal goat serum or antiserum against [Tyr (30)]-inhibin alpha (1-30). Jugular blood samples were collected every 6 h starting 24 h before and until 120 h after treatment. The plasma concentration of FSH increased at 6 h and remained at significantly high levels until 120 h in treated vs. control group. The plasma concentrations of estradiol showed a marked increased at 66 h, with peak levels at 120 h after treatment of antiserum. The basal concentrations of LH and the pattern of plasma concentrations of progesterone were not significantly different between the two groups. The number of medium size (3.5-5.0 mm) follicles increased considerably from Day 2, whereas small (3.5 mm or less) and large (5 mm or more) follicles increased noticeably from Day 3, as compared with pre-treatment and controls. These results clearly indicated that inhibin is a key hormone in regulation of follicular development through regulation of endogenous FSH secretion during early pregnancy in goats.     

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.     

Regulatory role of inhibin in follicle-stimulating hormone secretion and folliculogenesis in the guinea pig.

The effects of unilateral and bilateral ovariectomy and passive immunization against inhibin on follicle-stimulating hormone (FSH) secretions and follicular development in the guinea pig were investigated. Bilateral ovariectomy decreased plasma immunoreactive (ir-) inhibin rapidly and increased plasma FSH significantly. Unilateral ovariectomy decreased plasma ir-inhibin and increased plasma FSH temporarily, and doubled the number of ova released from the remaining ovary at the subsequent ovulation in guinea pigs. Injection of 1.0 ml inhibin antiserum significantly increased concentrations of plasma FSH at 6 hr onwards and the number of small follicles (100-200 microm in diameter) at 48 hr after the injection in guinea pigs bearing progesterone-containing implants. In vitro bioassay showed that inhibin antiserum could neutralize the suppression of ovarian homogenate on FSH secretion from cultured rat anterior pituitary cells. These results confirm the evidence that the ovary is the main source of inhibin secretion and both in vitro bioassay and passive immunization against inhibin show that the inhibin is a major regulator in the follicular development through FSH secretion in guinea pigs.     

Immunization of goats against inhibin increased follicular development and ovulation rate

In the present study, two experiments were conducted to induce superovulation in goats using passive and active immunization against inhibin. In the first experiment, two groups of goats were given an intravenous injection of either 10 ml normal goat serum (control; n=6) or inhibin antiserum developed against [Tyro30]-inhibin alpha (1-30) (passively immunized; n=6) 48 h before treatment with PGF2alpha. In the second experiment, two groups of goats were immunized with inhibin vaccine (actively immunized; n=5) or Freund's adjuvant (control; n=5) followed by three booster immunizations at 4 week intervals. Blood samples were collected for determination of FSH, LH, estradiol-17beta, and progesterone. Ultrasonography was used to determine ovarian activity at PGF2alpha injection and ovulation rate one week after estrus. In both experiments, there was a significant increase in plasma FSH concentration compared with the controls. However, the pattern of the FSH levels was different between the passively and actively immunized goats. The numbers of follicles in passively and actively immunized goats (22.4 +/- 2.3 and 18.6 +/- 2.1, respectively) were significantly greater than those in the controls (2.6 +/- 0.4 and 2.3 +/- 0.4, respectively). In addition, the ovulation rate was greater in the immunized animals compared with the controls. Therefore, either passive or active immunization against inhibin could be used to induce superovulation in goats.     

Hormonal Response of Female Goats to Active Immunization against a Recombinant Human Inhibin α-subunit, and establishment of an Enzyme-linked Immunosorbent Assay for Caprine Follicle-stimulating Hormone

The effect of selective immunosuppression of endogenous inhibin in goats on FSH, LH, progesterone and estradiol-17β profiles was studied during the breeding and nonbreeding seasons. Eighteen adult female Boer goats were immunized against the recombinant human inhibin α-subunit (hINH-α). With the exception of estradiol, which was determined by radio-immunoassay (RIA), all plasma hormone concentrations were determined by ELISA. The ELISA for FSH presented in this paper was established in the authors' laboratory, based on an existing RIA. Mean basal concentrations of FSH were not affected by immunosuppression of endogenous inhibin, nor was there a difference in the amplitude of the pre-ovulatory FSH surge. Immunization against inhibin appears to eliminate the slight secondary rise of FSH occurring 12–20 h after the major surge associated with ovulation. The LH profiles of the immunized goats were characterized by lower basal concentrations both before and after the pre-ovulatory LH surge which itself was reduced by 50% in immunized does. By contrast, concentrations of circulating estradiol were significantly elevated after inhibin-immunization. Progesterone profiles were not affected. Extending immunization into the anoestrous season by a booster injection of hINH-α, implicating oestrus induction with a progestagen and eCG, produced no discernible differences in FSH and LH profiles in comparison with nonimmunized control goats. The findings suggest that in goats, paracrine factors may play a more significant role in controlling follicular activity than a feedback mechanism acting via the pituitary.     

Characteristics of Ovarian Follicle Development in Domestic Animals

In most domestic animals the later stages of follicle development occurs in a wave‐like pattern during oestrous cycles (cattle, sheep, goats, horses and buffalo) or periods of reproductive activity (llamas and camels). A follicle wave is the organized development of a cohort of gonadotrophin‐dependent follicles all of which initially increase in size, but most of which subsequently regress and die by atresia (subordinate follicles). The number of remaining (dominant) follicles is specific to the species and is indicative of litter size. Follicle waves develop during both luteal and follicular phases and it is the dominant follicle(s) of the last follicular wave that ovulates. However, there are cases where dominant follicles from the last two follicle waves can ovulate (sheep and goats). There are exceptions to the organized wave‐like pattern of follicle growth where follicle development is apparently continuous (pigs and chickens). In these animals many follicles develop to intermediate diameters and at specific times follicles that are destined to ovulate are selected from this pool and continue growing to ovulation. Understanding the pattern of follicle development in different species is increasingly important for designing improved methods to manipulate reproduction in domestic animals.     

Follicle Wave Growth in Cattle

Ovarian follicle growth in cattle culminates in the selection of a single dominant follicle which attains the ability for final maturation and ovulation once or twice during the luteal phase and at the end of the oestrous cycle, as well as during other reproductive states. This review will describe in detail the first follicle wave of the cycle leading to selection of the first wave dominant follicle, indicating the specific gonadotrophin dependencies of cohort and dominant follicles, and relating follicle fate to steroidogenesis. As a differential gonadotrophin response of growing antral follicles during the follies‐stimulating hormone (FSH) decline may determine which follicle becomes selected, first wave follicles are also characterized in relation to intrafollicular growth factors, which may modify the gonadotrophin response, such as inhibins and members of the insulin‐like growth factor (IGF) family. Subsequently, the follicular control of the transient FSH rise and decline so crucial to dominant follicle selection will be discussed. It is concluded that successful hormonal manipulation of follicle wave growth and dominant follicle selection will depend on our detailed understanding of the gonadotrophin requirements of differentiating wave follicles.     

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.     

Monitoring follicular development in cattle by real-time ultrasonography: a review.

The application of real-time ultrasonography to monitoring ovarian function in mammals has advanced the understanding of follicular dynamics and its regulation. Follicular development is a wave-like sequence of organised events. The waves consist of the synchronous growth of small (4 to 5 mm) antral follicles, followed by the selection and growth of one dominant follicle which achieves the largest diameter and suppresses the growth of the subordinate follicles. In the absence of luteal regression, the dominant follicle eventually regresses (becomes atretic) and a new follicular wave begins. The dominant follicle regulates the growth of the subordinate follicles, because the appearance of the next wave is accelerated if the dominant follicle is ablated, and delayed if the lifespan of the dominant follicle is prolonged. During bovine oestrous cycles, two or three successive waves emerge, on average, on the day of ovulation (day 0) and day 10 for two-wave cycles, and on days 0, 9 and 16 for three-wave cycles. During the oestrous cycle there are thus two or three successive dominant follicles, and the last of these ovulates. Ovarian folliculogenesis is a complex process involving interactions between pituitary gonadotrophins, ovarian steroids and non-steroidal factors. Subtle changes in the hormonal milieu regulate folliculogenesis and the emergence of a follicular wave is preceded by a small increase in the concentration of plasma follicle-stimulating hormone. The mechanisms that promote the selection of a dominant follicle have not been elucidated, but considerable progress has been made in understanding follicular development and its regulation. Most treatments designed to control the development of follicular waves have been based on the physical or hormonal removal of the suppressive effect of the dominant follicle, and the consequent controlled induction of the emergence of a new follicular wave. The studies reviewed here describe current methods for regulating the bovine ovarian cycle, interesting models for future studies, and information that may be used for improving reproductive efficiency.