Complex interrelationships among CL,preovulatory follicle,number of follicular waves,and right or left ovaries in heifers

The diameter of the dominant follicle (DF) of wave 1 was studied on Days 9 to 17 (Day 0 = ovulation) in a survey of the ipsilateral and contralateral relationships between the location of the DF and CL, and number of follicular waves per interovulatory interval (IOI). For contralateral relationships, regardless of number of waves the diameter of the DF of wave 1 decreased (P < 0.03) between Days 11 and 13 when referenced to the follicle–CL relationship of wave 1 and decreased (P < 0.008) between Days 9 and 11 when referenced to the preovulatory follicle (PF)–CL relationship. For wave 2 in two-wave IOIs, the CL ovary of ipsilateral relationships had more (P < 0.05) follicles that reached at least 6 mm than the non-CL ovary. In three-wave IOIs, frequency of IOIs with the DF in the CL ovary was greater (P < 0.02) for wave 2 than for wave 3. In wave 3, the preovulatory and the largest subordinate follicles were located more frequently (P < 0.005) in the contralateral ovary. Ovulation in two-wave IOIs occurred more frequently (P < 0.0009) from the right ovary. In three-wave IOIs with a contralateral relationship ovulation occurred more frequently (P < 0.003) from the left ovary; a negative intraovarian effect of the CL on location of the PF may account for more ovulations from the left ovary and a reported greater frequency of the contralateral relationship. The hypothesis was supported that the ipsilateral versus contralateral relationship between the PF and CL is affected by the DF–CL relationship during the previous follicular waves and by the number and identity of waves per IOI.     

Relationship between days of the luteolytic period and locations of the preovulatory follicle and CL in interovulatory intervals with two or three follicular waves in heifers

Diameter of follicles was determined every 12 hours and progesterone (P4), FSH, and LH concentrations were determined every 6 hours from Day 12 (Day 0 = ovulation) to the ovulation at the end of the interovulatory interval (IOI). Groups were assigned on the basis of an ipsilateral (Ipsi) versus contralateral (Contra) relationship between the preovulatory follicle and CL and two follicular waves (2W) versus three waves (3W) per IOI. Numbers of IOIs were Ipsi-2W (n = 6), Ipsi-3W (n = 6), and Contra-3W (n = 8). Normalization to the end of luteolysis (day that P4 was closest to 1.0 ng/mL) indicated for the first time that concentrations of P4 and FSH were greater (P < 0.05) in 3W IOIs than in 2W IOIs for the 3 days before the beginning of a P4 decrease. The beginning of a P4 decrease occurred about 5 days and 6 hours after emergence of the preovulatory wave at 6 mm in 2W and 3W IOIs, respectively. On the day of diameter deviation between the future dominant and largest subordinate follicles in wave 3 of 3W IOIs, the future dominant follicle had the following characteristics: (1) distribution of diameters differed (P < 0.01) from unimodality; (2) diameter was greater (P < 0.05) in the Contra-3W group (9.8 ± 0.4 mm) than in the Ipsi-3W group (8.8 ± 0.3 mm); (3) diameter was similar to the diameter at the beginning of the P4 decrease (9.6 ± 0.9 mm); and (4) diameter was as small or smaller than diameter of the largest subordinate in seven of 14 heifers compared with zero of seven heifers in wave 2 of 2W IOIs. The differences involving deviation may be related to a reported greater frequency of the Contra-3W group than Ipsi-3W group. Results supported the hypothesis that emergence of the ovulatory wave occurs well before the beginning of luteolysis in 2W IOIs and near the beginning of luteolysis in 3W IOIs.     

Hormone concentrations temporally associated with contralateral and ipsilateral relationships between the CL and preovulatory follicle during the third follicular wave in heifers

Concentrations of circulating hormones after Day 14 (Day 0 = ovulation) were determined daily in 87 interovulatory intervals (IOIs) in heifers. The IOIs were grouped into four permutations according to an ipsilateral (Ipsi) or contralateral (Contra) relationship between the CL and the preovulatory follicle and two (2W) or three (3W) follicular waves per IOI. The number of IOIs per group differed (P < 0.005) from equality among the Ipsi-2W (n = 27), Contra-2W (n = 31), Ipsi-3W (n = 9), and Contra-3W (n = 20) groups. A continuous decrease in progesterone (luteolysis) began later (P < 0.05) in the Contra-3W group (Day 18.0 ± 0.4) than in each of the Ipsi-2W (15.4 ± 0.2), Contra-2W (15.6 ± 0.2), and Ipsi-3W (16.2 ± 0.5) groups. Concentrations of LH and estradiol began to increase near the beginning of luteolysis in each group. A minor FSH surge that did not stimulate a major follicular wave developed in about 50% of the IOIs in each group, except that none were detected in the Ipsi-3W group. The minor FSH surge reached a peak about 4 days before ovulation and several days after wave 3 had emerged. The hypothesis that luteolysis begins earliest in two-wave IOIs, intermediate in three-wave IOIs with an ipsilateral CL/follicle relationship, and latest in three-wave IOIs with a contralateral relationship was supported. The hypothesis that a minor FSH surge occurs most frequently in association with three follicular waves was not supported.     

Effect of intraovarian proximity between dominant follicle and corpus luteum on dimensions and blood flow of each structure in heifers

An intraovarian positive physiologic coupling between the extant CL and the ipsilateral preovulatory follicle (PF) or the future or established postovulatory dominant follicle (DF) was studied in 26 heifers. Ovaries were scanned by ultrasonic imaging from Day 16 (Day 0 = ovulation) of the preovulatory period until Day 6 of the postovulatory period. Hemodynamics of the follicles and CL were assessed by color-Doppler ultrasonography. When the PF and CL were ipsilateral compared with contralateral, blood-flow resistance in wall of the PF was lower (P < 0.04) on Days –2 and –1, and percentage blood-flow signals in the CL approached being greater (P < 0.08) on Days –4 to –1. During the postovulatory period, percentage of DF wall with blood-flow signals (44.1 ± 1.2% vs. 31.4 ± 2.8%) and percentage of CL with blood-flow signals (51.8 ± 1.2% vs. 42.5 ± 3.1%) were each greater (P < 0.05) when the two ipsilateral structures were adjacent (distance between antrum and CL wall, ≤ 3 mm) than when separated. On Day 0, the distance between follicle and ipsilateral CL was less (P < 0.02) for the future DF than for the future largest subordinate. Growth rate between Days 0 and 2 averaged over all growing follicles was greater (P < 0.01) when the follicles were ≤3 mm from the CL (1.1 ± 0.1 mm/day) than when farther from the CL (0.9 ± 0.1 mm/day). Results supported the hypotheses that (1) a positive intraovarian coupling occurs between the PF or postovulatory DF and the extant CL and (2) the coupling is enhanced when the ipsilateral DF and CL are in close proximity.     

Minor FSH surge,minor follicular wave,and resurgence of preovulatory follicle several days before ovulation in heifers

Blood samples were collected and follicle diameters were determined daily beginning on Day 12 (Day 0 = ovulation) in 35 interovulatory intervals (IOIs) in heifers. A minor follicular wave with maximal diameter (6.0 ± 0.3 mm) on Day −4 was detected in six of seven IOIs that were scanned for follicles 4 mm or greater. The number of IOIs with a CV-identified minor FSH surge toward the end of the IOI was greater (P < 0.03) in two-wave IOIs (10/17) than in three-wave IOIs (4/18). The 17 two-wave IOIs were used for study of the temporal relationships among preovulatory follicle, FSH, LH, and estradiol. Daily growth rate of the preovulatory follicle was maximum on Days −11 to −7, minimum (P < 0.05) on Days −7 to −4, and increased (resurged, P < 0.05) on Days −4 to −3. A transient increase in FSH was maximum on mean Day −4, and the peak of a minor FSH surge occurred on Day −4.5 ± 0.2. Concentration of LH and estradiol increased between Days −5 and −4. Results demonstrated resurgence of the preovulatory follicle apparently for the first time in any species. Resurgence seemed more related temporally to the minor FSH surge than to the LH increase, but further study is needed. Results supported the novel hypotheses that a minor FSH surge near the end of the IOI is temporally associated with (1) the emergence of a minor follicular wave and (2) the resurgence in growth rate of the preovulatory follicle.     

Ovarian follicular and luteal dynamics in wapiti during the estrous cycle

The reproductive tracts of 13 mature hinds were examined daily by transrectal ultrasonography and blood samples were taken daily from October to January to characterize follicular, luteal, and endocrine dynamics in wapiti during the estrous season. Follicle development occurred in waves characterized by regular, synchronous development of a group of follicles in temporal succession to a surge in serum FSH concentration. The mean interovulatory interval was 21.3 +/- 0.1 d, but was shorter in hinds exhibiting two follicular waves than in hinds exhibiting three and four waves (P < 0.05). The interwave interval was similar among waves in two-wave cycles and the first wave of three-wave cycles. All other interwave intervals in three- and four-wave cycles were shorter (P < 0.05). The maximum diameter of the dominant follicle of the first wave was similar among two-, three-, and four-wave cycles. For all other waves in three- and four-wave cycles, the maximum diameter was smaller (P < 0.05). Corpus luteum diameter and plasma progesterone concentrations were similar between two- and three-wave cycles, but the luteal phase was longer (P < 0.05) in four-wave cycles. The dominant follicle emerged at a diameter of 4 mm at 0.4 +/- 0.1 and 0.8 +/- 0.1 d before the largest and second largest subordinate follicles, respectively. The follicle destined to become dominant was larger (P < 0.05) than the largest subordinate follicle one day after emergence, which coincided with the first significant decrease in serum FSH concentration. We concluded that the estrous cycle in wapiti is characterized by two, three, or four waves of follicular development (each preceded by a surge in circulating FSH), that there is a positive relationship between the number of waves and the duration of the cycle, and an inverse relationship between the number of waves and the magnitude of follicular dominance (diameter and duration of the dominant follicle).     

Ultrasonic characteristics of preovulatory follicle and ovulation in Caspian mares

The Caspian breed of horses is believed to be the direct descendant of the earliest equine animals. Some special characteristics of Caspian horse differentiate this breed of horses from other breeds. In the current study the ultrasonically observed characteristics of a preovulatory dominant follicle and the lengths of estrus, diestrus as well as some related parameters were studied during 42 interovulatory intervals in 11 healthy Caspian mares. The preovulatory dominant follicle deviated from subordinate follicles and became the largest follicle in the ovaries at Day −8.7±0.53 (Day 0=ovulation). Every mare was a single ovulator with ovulations more frequent from the left ovary than from the right (65% versus 35%). Mean length of estrus, diestrus, and interovulatory interval were 8.3±0.86, 13.8±0.59, and 22.1±0.40 days, respectively. The time interval from ovulation until the time in which the mares were no longer in estrus was 1.9±0.42 days.     

Developmental pattern of small antral follicles in the bovine ovary

The study was designed to characterize the developmental pattern of 1- to 3-mm follicles and to determine the stage at which the future dominant follicle first attains a size advantage among its cohorts. In experiment 1, heifers (n = 18) were examined every 24 h by transrectal ultrasonography for one interovulatory interval (IOI). In experiment 2, cows (n = 9) were examined every 6 h from 5 to 13 days after ovulation to monitor precisely the diameter changes of individual follicles >/=1 mm during emergence of wave 2. Results revealed a change over days (P < 0.05) in the number of 1- to 3-mm follicles, with a maximum (P < 0.05) 1 or 2 days before wave emergence (conventionally defined as the time when the dominant follicle is first detected at 4 mm), followed 3-4 days later by a maximum (P < 0.05) in the number of >/=4-mm follicles. The profiles of small (1-3 mm) and large (>/=4-mm) follicles were inversely proportional (r = -0.79; P = 0.01). The profile of the number of 1- to 3-mm follicles during wave emergence was similar (P = 0.63) between waves in two-wave IOI, but differed (P < 0.01) among waves in three-wave IOI as a result of a greater number of follicles in the ovulatory wave (P < 0.04). As well, the number of follicles in the ovulatory wave tended to be greater (P < 0.06) in three-wave IOI than in two-wave IOI. The future dominant follicle was first identified at a diameter of 1 mm and emerged 6-12 h earlier than the first subordinate follicle (P < 0.01). After detection of the dominant follicle at 1 mm (0 h), its diameter differed from that of the first and second subordinate follicles at 24 h (P = 0.04) and 12 h (P = 0.01), when the dominant follicle was 2.4 +/- 0.17 mm and 1.7 +/- 0.14 mm, respectively. The growth rate of the dominant follicle differed from that of the first and second subordinate follicles at 120 h (P = 0.03) and 108 h (P = 0.02), when the dominant follicle was 9.5 +/- 0.30 mm and 8.8 +/- 0.49 mm, respectively. Emergence of the future dominant (r = 0.71), first (r = 0.73), and second (r = 0.76) subordinate follicles was temporally associated (P < 0.01) with a rise in circulating concentrations of FSH. Transient, nocturnal elevations in plasma FSH concentration were followed within 6 h by an increase in the growth rate of 1- to 3-mm follicles. We conclude that 1) 1- to 3-mm follicles develop in a wave-like manner in association with surges in plasma concentrations of FSH, 2) 1- to 3-mm follicles are exquisitely responsive to transient elevations in FSH, and 3) selection of the dominant follicle is manifest earlier than previously documented and is characterized by a hierarchical progression over a period encompassing the entire FSH surge (5 days).     

Effects of a dominant follicle on ovarian follicular dynamics during the oestrous cycle in heifers

Two hypotheses were tested: (1) a dominant follicle causes regression of its subordinate follicles, and (2) a dominant follicle during its growing phase suppresses the emergence of the next wave. Cyclic heifers were randomly assigned to one of four groups (6 heifers/group): cauterization of the dominant follicle of Wave 1 or sham surgery (control) on Day 3 or Day 5 (day of ovulation = Day 0). Ultrasonic monitoring of individually identified follicles was done once daily throughout the interovulatory interval. The onset of regression (decreasing diameter) of the largest subordinate follicle of Wave 1 was delayed (P less than 0.01) by cauterization of the dominant follicle of Wave 1 on Day 3 compared to controls (mean onset of regression, Days 10.8 +/- 2.1 vs 4.3 +/- 0.4). Cauterization of the dominant follicle of Wave 1 on Days 3 or 5 caused early emergence (P less than 0.01) of Wave 2 when compared to controls (Day-3 groups: Days 5.5 +/- 0.4 vs 9.6 +/- 0.7; Day-5 groups: Days 7.0 +/- 0.3 vs 9.1 +/- 0.4). The results supported the two hypotheses. In addition, cauterization of the dominant follicle of Wave 1 on Days 3 or 5 increased the incidence of 3-wave interovulatory intervals.     

Differential blood flow changes between the future dominant and subordinate follicles precede diameter changes during follicle selection in mares

Diameter deviation during a follicular wave is characterized by the continued growth of the developing dominant follicle and reduced growth and regression of the subordinate follicles. This study considered the hypothesis that reduced blood flow in the future largest subordinate follicle precedes the beginning of diameter deviation. The hypothesis was tested by quantifying the daily changes in blood-flow velocities and blood-flow area within the wall of follicles before and during diameter deviation in mares (n = 7). The blood-flow end points were quantified daily by transrectal color Doppler ultrasonography. Follicles were identified retrospectively by rank as F1 (largest) and F2 according to the maximum attained diameter. Follicles were grouped into nine F1 diameter ranges of 3.0 mm each (equivalent to 1 day's growth) centered on 6.5, 9.5, 12.5, 15.5, 18.5, 21.5, 24.5, 27.5, and 30.5 mm. Diameter deviation began in the 24.5-mm group, as indicated by a smaller (P < 0.05) difference between F1 and F2 in the 24.5-mm group than in the 27.5-mm group. Based on a similar approach, peak systolic velocity and time-averaged maximum velocity of blood flow began to deviate between F1 and F2 in the 18.5-mm group (P < 0.04) and blood flow area began to deviate in the 21.5-mm group (P < 0.009). Thus, differential blood flow area between F1 and F2 began an average of 3.0 mm (equivalent to 1 day) and differential blood-flow velocities began an average of 6.0 mm before the beginning of diameter deviation. The results demonstrated that deviation between F1 and F2 in the blood flow of the follicle walls occurred 1 or 2 days before deviation in follicle diameter during follicle selection in mares.     

Follicular populations during the estrous cycle in heifers. II. Influence of right and left sides and intraovarian effect of the corpus luteum

Ovarian follicles ≥2 mm were studied in 22 Holstein heifers by daily ultrasound examinations. Data were partitioned by right vs. left ovary and corpus luteum bearing ovary vs. the contralateral ovary. There were significantly more (P < 0.03) follicles 4–6 mm, > 13mm and total ≥2 mm in the right ovary, regardless of the presence of a corpus luteum. Significantly more (P < 0.05) follicles 2–3 mm, > 13 mm and total ≥2 mm were observed in the ovary bearing the corpus luteum. Interactions between day and corpus luteum appeared to be due to a greater number of follicles in the ovary bearing the corpus luteum during the first part of the interovulatory interval. There was also a day by right side vs. left side interaction for the number of follicles > 13 mm. Interpretation of the interactions was that the presence of a corpus luteum was conducive to the development of more anovulatory diestrous follicles > 12 mm. However, as regression of the corpus luteum progressed, there was an apparent proclivity for preovulatory follicular development in the right ovary. There was no apparent pattern of alternating sides of ovulation or of alternating sides of development of anovulatory diestrous follicles and preovulatory follicles in heifers observed for more than one interovulatory interval. There was not a significant difference in the maximum diameter attained by the anovulatory diestrous follicle or preovulatory follicle between ovaries ipsilateral or contralateral to the corpus luteum; however, the maximum diameter attained by the preovulatory follicle was greater (P < 0.05) than that attained by the anovulatory diestrous follicle.     

The effect of a single high dose of PGF2α administered to dairy cattle 3.5 days after ovulation on luteal function,morphology, and follicular dynamics

A single treatment with PGF2α is assumed to have no luteolytic effect on cows with corpora lutea < 5 days old. The objective of this study was to determine the effect of a single high dose of PGF2α administered to dairy cattle on the morphology and function of the early CL. The study followed a crossover design with a treatment cycle in which 50 mg of dinoprost were administered 3.5 days postovulation and a control untreated cycle. Ultrasound examination and blood samples were performed during the two consecutive cycles. Corpus luteum (CL) diameter, progesterone concentration, and follicular dynamics characteristics were compared between control and treated cycles. Two of nine cows (22%) developed full luteolysis. The remaining seven cows (78%) had partial luteolysis with a decrease (P < 0.05) in progesterone concentration and CL diameter for two and 12 days post-treatment, respectively. The interovulatory interval of treated cycles (19.7 ± 2.4 days) was not different (P > 0.05) from that of controls (23.8 ± 0.9 days). The transient reduction in progesterone of cows with partial luteolysis had no effect on the proportion of cows with two or three follicular waves, follicle growth rate, or preovulatory diameter (P > 0.05). Two cows developed ovarian cystic degeneration during the PGF2α-induced cycle. In conclusion, the treatment of cows with a high dose of PGF2α 3.5 days postovulation induced some degree of luteolysis in all treated cows. This resulted in partial luteolysis in 78% of treated animals and in full luteolysis in the remaining 22%.     

Temporal associations among ovarian events in cattle during oestrous cycles with two and three follicular waves

For 18 two-wave interovulatory intervals in heifers, the follicular waves were first detected on Days -0.2 +/- 0.1 and 9.6 +/- 0.2, and for 4 three-wave intervals on Days -0.5 +/- 0.3, 9.0 +/- 0.0 and 16.0 +/- 1.1 (ovulation is Day 0). The day-to-day mean diameter profile of the dominant follicle of the 1st wave and the day of emergence of the 2nd wave were not significantly different between 2-wave and 3-wave intervals. There were no indications, therefore, that events occurring during the first half of the interovulatory interval were associated with the later emergence of a 3rd wave. The dominant ovulatory follicle differed significantly (P less than 0.05 at least) between 2-wave and 3-wave intervals in day of emergence (Day 9.6 +/- 0.2 and 16.0 +/- 1.1), length of interval from emergence of follicle to ovulation (10.9 +/- 0.4 and 6.8 +/- 0.6 days), and diameter on day before ovulation (16.5 +/- 0.4 and 13.9 +/- 0.4 mm). The mean length of 2-wave interovulatory intervals (20.4 +/- 0.3 days) was shorter (P less than 0.01) than for 3-wave intervals (22.8 +/- 0.6 days). The mean day of luteal regression for 2-wave and 3-wave intervals was 16.5 +/- 0.4 and 19.2 +/- 0.5 (P less than 0.01). For all intervals, luteal regression occurred after emergence of the ovulatory wave, and the next wave did not emerge until near the day of ovulation at the onset of the subsequent interovulatory interval. In conclusion, the emergence of a 3rd wave was associated with a longer luteal phase, and the viable dominant follicle present at the time of luteolysis became the ovulatory follicle.     

Follicular populations during the estrous cycle in heifers. I. Influence of day

Ovarian follicles ⩾ 2 mm were studied in 22 Holstein heifers by daily ultrasound examinations. There were significant differences (P < 0.0001) among days of the estrous cycle for diameter of the largest and second largest follicles and in the numbers of follicles 2–3 mm, 4–6 mm, 7–10 mm, 11–13 mm, > 13 mm, and total number of follicles ⩾2 mm. Patterns of the mean profiles for all follicular endpoints except the number of follicles 4–6 mm and total number of follicles ⩾ 2 mm were bimodal. The days encompassed by the first and second portions, respectively, of the bimodal profiles were approximately: diameter of largest follicle, Days 0–14 and 15–21 (ovulation); diameter of second largest follicle, Days 0–7 and 8–20; number of follicles 2–3 mm, Days 1–11 and 12–20; number of follicles 7–10 mm, Days 0–6 and 7–18; number of follicles 11–13 mm, Days 0–8 and 9–20; and number of follicles > 13 mm, Days 2–14 and 16–21. Data for the various categories were recombined to demonstrate relationships between the numbers of follicles 2–3 mm and ⩾ 4 mm during the interovulatory interval. There were significant differences (P < 0.0001) among days in both 2–3 mm and ⩾ 4 mm follicular categories. Differences appeared due to periods of higher mean numbers of follicles 2–3 mm which began between Days 2 or 3 and Days 15 or 16 and reached maximum levels on Day 7 and Day 19, respectively. There was an inverse relationship between the number of follicles 2–3 mm vs ⩾ 4 mm and between the diameters of largest and second largest follicles. The process of selection of the follicle destined to ovulate appeared to become manifest as selective growth of the preovulatory follicle with concurrent decrease in diameter of the second largest follicle and regression of the other follicles in the various follicular categories. A similar process apparently occurred early in the interovulatory interval. There was apparently selective growth of a follicle to preovulatory size by Day 6, coincident decrease in diameter of the second largest follicle, and apparent regression of other follicles in the ultrasonically detectable pool. The only apparent difference was that the follicle which attained preovulatory diameter early in the interovulatory interval remained in the ovary for 5 or 6 days, then regressed, while the follicle which attained preovulatory diameter at approximately Day 18–20 ovulated.     

The effect of a single high dose of PGF2α administered to dairy cattle 3.5 days after ovulation on luteal function, morphology, and follicular dynamics

A single treatment with PGF2α is assumed to have no luteolytic effect on cows with corpora lutea < 5 days old. The objective of this study was to determine the effect of a single high dose of PGF2α administered to dairy cattle on the morphology and function of the early CL. The study followed a crossover design with a treatment cycle in which 50 mg of dinoprost were administered 3.5 days postovulation and a control untreated cycle. Ultrasound examination and blood samples were performed during the two consecutive cycles. Corpus luteum (CL) diameter, progesterone concentration, and follicular dynamics characteristics were compared between control and treated cycles. Two of nine cows (22%) developed full luteolysis. The remaining seven cows (78%) had partial luteolysis with a decrease (P < 0.05) in progesterone concentration and CL diameter for two and 12 days post-treatment, respectively. The interovulatory interval of treated cycles (19.7 ± 2.4 days) was not different (P > 0.05) from that of controls (23.8 ± 0.9 days). The transient reduction in progesterone of cows with partial luteolysis had no effect on the proportion of cows with two or three follicular waves, follicle growth rate, or preovulatory diameter (P > 0.05). Two cows developed ovarian cystic degeneration during the PGF2α-induced cycle. In conclusion, the treatment of cows with a high dose of PGF2α 3.5 days postovulation induced some degree of luteolysis in all treated cows. This resulted in partial luteolysis in 78% of treated animals and in full luteolysis in the remaining 22%.     

Selection of the dominant follicle in cattle: role of two-way functional coupling between follicle-stimulating hormone and the follicles

The functional coupling between the declining portion of the FSH surge and the growing follicles of a wave was studied by treating heifers with a minimal dose of estradiol to decrease FSH concentrations without an associated change in LH concentrations. Estradiol treatment when the largest follicle reached >/= 6.0 mm (Hour 0) resulted in depression of both FSH concentrations and diameter of the largest follicle by Hour 8. The smaller follicles were also inhibited. These results supported the hypothesis that FSH continues to be needed by the growing follicles even when the FSH concentrations are decreasing during the declining portion of the FSH surge. Estradiol treatment when the largest follicle was >/= 8.5 mm (expected time of follicular deviation) also resulted in a transient decrease in both FSH concentrations and diameter of the largest follicle, but the diameters of the smaller follicles were not affected. These results supported the hypothesis that the low concentrations of FSH at the expected time of deviation, although inadequate for the smaller follicles, were required for continued growth of the largest follicle. In another study, ablation (Hour 0) of the largest follicle was done at >/= 7.5 mm vs. >/= 8.5 mm. The mean FSH concentrations for the 8.5-mm groups were greater for the ablation group than for the control group at Hours 8 and 12, but there was no difference between the 7.5-mm groups at any hour. These results supported the hypothesis that by the time the largest follicle reaches the expected beginning of deviation it has developed a greater capacity for suppressing FSH. It is postulated that the essence of the selection of a dominant follicle is a close two-way functional coupling between changing FSH concentrations and follicular growth.     

Temporal interrelationships among luteolysis, FSH and LH concentrations and follicle deviation in mares

The effect of altered LH concentrations on the deviation in growth rates between the 2 largest follicles was studied in pony mares. The progestational phase was shortened by administration of PGF2alpha on Day 10 (Day 0=ovulation; n=9) or lengthened by daily administration of 100 mg of progesterone on Days 10 to 30 (n=11; controls, n=10). All follicles > or = 5 mm were ablated on Day 10 in all groups to initiate a new follicular wave. The interovulatory interval was not altered by the PGF2alpha treatment despite a 4-day earlier decrease in progesterone concentrations. Time required for growth of the follicles of the new wave apparently delayed the interval to ovulation after luteolysis. The FSH concentrations of the first post-ablation FSH surge were not different among groups. A second FSH surge with an associated follicular wave began by Day 22 in 7 of 11 mares in the progesterone group and in 0 of 19 mares in the other groups, indicating reduced functional competence of the largest follicle. A prolonged elevation in LH concentrations began on the mean day of wave emergence (Day 11) in the prostaglandin group (19.2 +/- 2.2 vs 9.0 +/- 0.7 ng/mL in controls; P<0.05), an average of 4 d before an increase in the controls. Concentrations of LH in the progesterone group initially increased until Day 14 and then decreased so that by Day 18 the concentrations were lower (P<0.05) than in the control group (12.9 +/- 1.6 vs 20.2 +/- 2.6 ng/mL). Neither the early and prolonged increase nor the early decrease in LH concentrations altered the growth profile of the second-largest follicle, suggesting that LH was not involved in the initiation of deviation. However, the early decrease in LH concentrations in the progesterone group was followed by a smaller (P<0.05) diameter of the largest follicle by Day 20 (26.9 +/- 1.7 mm) than the controls (30.3 +/- 1.7 mm), suggesting that LH was necessary for continued growth of the largest follicle after deviation.     

Two-way coupling between FSH and the dominant follicle in heifers

Follicular Wave 1 and 2 and the associated FSH Surge 1 and 2 were used to designate the first two waves and surges of the interovulatory interval in two experiments in heifers. In experiment 1, a group with early (group E, N = 9) and late (group L, N = 5) development of the dominant follicle of Wave 1 were used as natural models to study FSH/follicle coupling. The day of wave emergence and the day of deviation in diameters between the two largest follicles were not different between groups. Emergence of Wave 2 and maximal FSH concentration in Surge 2 was approximately 1 day later (P < 0.03) in group L. Diameter of the dominant follicle of wave 1 (13.8 ± 0.3 mm vs. 12.0 ± 0.3 mm) and FSH concentrations in Surge 2 (0.29 ± 0.02 ng/mL vs. 0.21 ± 0.03 ng/mL) were first greater (P < 0.05) in group E than in group L at 4 and 5 days, respectively, after wave emergence. In experiment 2, treatment with estradiol (N = 8) when the dominant follicle of Wave 1 was ≥11 mm (Hour 0) resulted in a decrease (P < 0.02) in FSH and slower (P < 0.05) growth rate of the follicle between Hours 0 and 4. Results supported the following hypotheses: (1) the FSH surge that stimulates emergence of a follicular wave is associated with final growth of the dominant follicle of the previous anovulatory wave; and (2) suppression of FSH Surge 2 when the dominant follicle of Wave 1 is ≥11 mm is associated with a decrease in diameter. It is concluded for the first time that two-way FSH/follicle coupling in heifers continues during final growth of the dominant follicle of Wave 1 and that Surge 2 is the FSH source.     

The mare: a 1000-pound guinea pig for study of the ovulatory follicular wave in women

The mare is a good comparative model for study of ovarian follicles in women, owing to striking similarities in follicular waves and the mechanism for selection of a dominant follicle. Commonality in follicle dynamics between mares and women include: (1) a ratio of 2.2:1 (mare:woman) in diameter of the largest follicle at wave emergence when the wave-stimulating FSH surge reaches maximum, in diameter increase of the two largest follicles between emergence and the beginning of deviation between the future dominant and subordinate follicles, in diameter of each of the two largest follicles at the beginning of deviation, and in maximum diameter of the preovulatory follicle; (2) emergence of the future ovulatory follicle before the largest subordinate follicle; (3) a mean interval of 1 day between emergence of individual follicles of the wave; (4) percentage increase in diameter of follicles for the 3 days before deviation; (5) deviation 3 or 4 days after emergence; (6) 25% incidence of a major anovulatory follicular wave emerging before the ovulatory wave; (7) 40% incidence of a predeviation follicle preceding the ovulatory wave; (8) small but significant increase in estradiol and LH before deviation; (9) cooperative roles of FSH and insulin-like growth factor 1 and its proteases in the deviation process; (10) age-related effects on the follicles and oocytes; (11) approximate 37-hour interval between administration of hCG and ovulation; and (12) similar gray-scale and color-Doppler ultrasound changes in the preovulatory follicle. In conclusion, the mare may be the premier nonprimate model for study of follicle dynamics in women.     

Selection of the dominant follicle in cattle: establishment of follicle deviation in less than 8 hours through depression of FSH concentrations

Deviation in follicle diameter in cattle is characterized by continued growth of the largest follicle of a follicular wave and a reduction or cessation of growth of the smaller follicles. Deviation begins when the largest follicle reaches about 8.5 mm. Two experiments were done to test the hypothesis that the deviation mechanism is established in < 8 h, as indicated by the temporal relationships between follicle removal and an increase in FSH concentrations (Experiment 1) and between a decrease in FSH concentrations and follicle inhibition (Experiment 2). In Experiment 1, the role of the first follicle to reach 8.5 mm was studied by follicle ablation (Hour 0). The combined mean FSH concentrations for the control group (n = 8) and ablation group before ablation (n = 7) progressively decreased (P < 0.02) over two 8-h intervals before the largest follicle reached > or = 8.5 mm (Hour-16, 1.77 +/- 0.11 ng/mL; Hour 0, 1.49 +/- 0.08 ng/mL). In controls, the concentrations continued to decrease (P < 0.02) until Hour 10 (1.21 +/- 0.09 ng/mL). Ablation of the largest follicle at > or = 8.5 mm resulted in increased (P < 0.02) circulating FSH concentrations between Hours 5 (1.34 +/- 0.04 ng/mL) and 8 (1.61 +/- 0.09 ng/mL). Growth rate of the second-largest follicle between Hours 0 and 8 was greater (P < 0.05) in the ablation group than in the controls, and the second largest follicle became dominant in 7 of 7 heifers following ablation of the largest follicle. In Experiment 2, a minimal single injection of a depressant of FSH concentrations (4.4 mL of steroid-reduced follicular fluid) was given when the largest follicle was a mean of 8.4 mm (Hour 0; controls, n = 4; treated, n = 4). An interaction of group and hour (P < 0.005) for FSH concentrations was attributable to an FSH decrease (P < 0.002) by Hour 6 and an increase (P < 0.002) between Hours 9 and 12 in the treated group. The growth rate of the largest follicle between Hours 0 and 12 was less (P < 0.05) in the treated group (0.2 +/- 0.2 mm/12 h) than in the control group (1.2 +/- 0.4 mm/12 h). The reduced diameter was recorded within 6 h after suppression of FSH concentrations, supporting the hypothesis. Our preferred interpretation is that when the largest follicle reaches a critical diameter of about > or = 8.5 mm, FSH concentrations continue to decrease and become lower than required by the smaller follicles but not the largest follicle. The results further indicate that a close temporal coupling between a change in FSH concentrations and the follicular response could establish the deviation mechanism in < 8 h or before the second largest follicle reaches a similar critical diameter.