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.     

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.     

Exogenous hormonal manipulation of ovarian activity in cattle

To achieve precise control of the oestrous cycle in cattle it is necessary to control both the life span of the corpus luteum and the follicle wave status at the end of the treatment. Antral follicle growth in cattle occurs in distinct wavelike patterns during the ovarian cycle and the postpartum anoestrous period. The emergence of each new wave is stimulated by a transient increase in FSH. Each follicle wave has an inherent life span of 7-10 days as it progresses through the different stages of development, viz., emergence, selection, dominance and atresia or ovulation. The dominant follicle (DF) is distinguishable from other subordinate follicles by its enhanced capacity to produce oestradiol, maintenance of low intrafollicular concentrations of insulin-like growth factor binding proteins-2, -4 and -5 and follistatin and an increase in free intrafollicular concentrations of IGF-I as well as an increase in size. Three approaches can be taken to control ovarian activity and regulate the oestrous cycle in cattle: (i) use of the luteolytic agent prostaglandin F2alpha (PGF2alpha) alone or one of its potent analogues, (ii) administration of exogenous progesterone-progestagen treatments combined with the use of exogenous oestradiol or gonadotrophin releasing hormone (GnRH) to control new follicle wave emergence and shorten the life span of the corpus luteum, and (iii) prior follicle wave synchrony followed by induced luteolysis. A number of different oestrous synchronisation regimens, viz., PGF2alpha-based only, short-term progesterone with prior follicle wave synchrony using oestradiol or GnRH have been developed but the problem of obtaining good follicle wave synchrony and CL regression limit their widespread application. GnRH-prostaglandin-GnRH regimens have recently been developed for beef and dairy cows. However, their success is variable. A better understanding of the hormonal control of follicle growth is a prerequisite in order to obtain more precise control the oestrous cycle allowing one AI at a predetermined time giving high pregnancy rates without recourse to detection of oestrus.     

Follicle Development in Mares

The mare provides a unique experimental model for studying follicle development in monovular species. Development of antral follicles in horses is characterized by the periodic growth of follicular waves which often involve the selection of a single dominant follicle. If properly stimulated, the dominant follicle will complete development and eventually ovulate a fertile oocyte. Regulation of follicular wave emergence and follicle selection involves an interplay between circulating gonadotropins and follicular factors that ensures that individual follicles are properly stimulated to grow (or to regress) at any given stage of follicular wave development. Periodic development of follicular waves continuously occurs during most of post-natal life in the mare and is influenced by factors such as stage of oestrous cycle, season, pregnancy, age, breed and individual so that different types of follicular waves (minor or major, ovulatory or anovulatory) and different levels of activity within waves may develop under different physiological conditions. Changes in gonadotropin levels and/or in the sensitivity of follicles to circulating gonadotropins seem to account largely for these physiological variations in follicle development.     

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.     

The localization and expression of epidermal growth factor and epidermal growth factor receptor in bovine ovary during oestrous cycle

Epidermal growth factor (EGF) is one of the important regulatory factors of EGF family. EGF has been indicated to effectively inhibit the apoptosis of follicular cells, to promote the proliferation of granulosa cells and the maturation of oocytes, and to induce ovulation process via binding to epidermal growth factor receptor (EGFR). However, little is known about the distribution and expression of EGF and EGFR in cattle ovary especially during oestrous cycle. In this study, the localization and expression rule of EGF and EGFR in cattle ovaries of follicular phase and luteal phase at different time points in oestrous cycle were investigated by using IHC and real-time qPCR. The results showed that EGF and EGFR in cattle ovary were mainly expressed in granulosa cells, cumulus cells, oocytes, zona pellucida, follicular fluid and theca folliculi externa of follicles. The protein and mRNA expression of EGF/EGFR in follicles changed regularly with the follicular growth wave both in follicular and in luteal phase ovaries. In follicular phase ovaries, the protein expression of EGF and EGFR was higher in antral follicles than that of those in other follicles during follicular growth stage, and the mRNA expression of EGFR was also increased in stage of dominant follicle selection. However, in luteal phase ovaries, the growth of follicles was impeded during corpus luteum development under the action of progesterone secreted by granular lutein cell. The mRNA and protein expressions of EGF and EGFR in ovarian follicles during oestrous cycle indicate that they play a role in promoting follicular development in follicular growth waves and mediating the selection process of dominant follicles.     

Ovarian follicular dynamics in lines of sheep (Finn, Merinos) selected on ovulation rate

Ewes from selected lines of sheep from each of two breeds, Finns (high ovulation rate, low ovulation rate and control lines with respective ovulation rates of 5.4, 2.7 and 3.3) and Merinos (T Merinos selected for increased ovulation rate and control Merinos with respective ovulation rates of 1.9 and 1.2) were used to examine how selection to alter ovulation rate had altered follicle development. Ovarian antral follicles were counted, measured, classified as nonatretic or atretic (more than five pyknotic bodies). The growth of ovulatory follicles in vivo, followed by repeated follicle ink marking, also was compared in the three lines of Finns. Regardless of breed, ewes selected for high ovulation rate had a similar number of antral follicles and a similar extent of atresia compared with their controls. Alterations induced by selection were located in the last stages of folliculogenesis. T Merinos exhibited a lower proportion of atretic follicles among follicles greater than 3 mm and a larger diameter of the largest healthy follicle when preovulatory follicles were excluded. High-line Finn ewes recruited more follicles, which produced smaller preovulatory follicles, each containing a smaller number of granulosa cells compared with either the low- or control-line ewes. Hence, physiological selection for high ovulation rate raised it by different methods in Merino than in Finn ewes.     

Physiological mechanisms of ovarian follicular growth in pigs--a review

Follicular growth after antrum formation is determined by follicle-stimulating hormone (FSH). Only two ways are possible for recruited follicles, continuing development or atresia. In gilts, intensive ovarian follicular growth begins between 60 and 100 days of age, and fluctuations of the ovarian morphological status last about 20 days; however, at that time there are no really large follicles. Final follicular development is under luteinising hormone (LH) control; this is why the attainment of puberty is related to an increase in serum oestradiol to a level that causes a preovulatory surge of this gonadotropin. The pool of follicles at the beginning of the oestrous cycle is about 30-40, most of which are small (< 3 mm) and growing. Then, the pool of follicles increases to about 80 in the mid-luteal phase but about 50 of them are small and 30 are medium sized (3-6.9 mm). Some of these follicles are in the growing phase, but some are atretic. Between days 7 and 15 of the oestrous cycle the percentage of atretic follicles fluctuates between 12 and 73%. At that time there are no large (> 7 mm) follicles because of the suppressing effect of progesterone. The number of small follicles declines after luteolysis. From the pool of medium follicles, large follicles are selected under the influence of LH, but about 70% of the medium-sized follicles become atretic. Because of the long-lasting selection process there is a significant heterogeneity in the diameter of large follicles in oestrus. However, the number of follicles correlates with the number of corpora lutea after ovulation. Individual follicular development and the relationship between follicles are still poorly known. The use of ultrasonography may give a closer insight into these phenomena.     

Mechanisms for Dominant Follicle Selection in Monovulatory Species: A Comparison of Morphological, Endocrine and Intraovarian Events in Cows, Mares and Women

The selection of a single ovarian follicle for further differentiation and finally ovulation is a shared phenomenon in monovulatory species from different phylogenetic classes. The commonality of dominant follicle (DF) development leads us to hypothesize that mechanisms for DF selection are conserved. This review highlights similarities and differences in follicular wave growth between cows, mares and women, addresses the commonality of the transient rises in FSH concentrations, and discusses the follicular secretions oestradiol and inhibin with their regulatory roles for FSH. In all three species, rising FSH concentrations induce the emergence of a follicle wave and cohort attrition occurs during declining FSH concentrations, culminating in DF selection. Cohort secretions are initially responsible for declining FSH, which is subsequently suppressed by the selected DF lowering it below the threshold of FSH requirements of all other cohort follicles. The DF acquires relative FSH-independence in order to continue growth and differentiation during low (cow, mare) or further declining FSH concentrations (women), and thus may be the one cohort follicle with the lowest FSH requirement due to enhanced FSH signalling. In all three monovulatory species a transition from FSH- to LH-dependence is postulated as the mechanism for the continued development of the selected DF. In addition, FSH and IGF enhance each other's ability to stimulate follicle cell function and access of IGF-I and -II to the type 1 receptor is regulated by IGF binding proteins that are in turn regulated by specific proteases; all of which have been ascribed a role in DF development. No fundamental differences in DF selection mechanisms have been identified between the different species studied. Thus functional studies of the selection of DFs in cattle and mares are also valuable for identifying genes and pathways regulating DF development in women.     

Follicular development in cattle; changes in their count and size during the ovarian cycle

Follicular development was examined by transrectal ultrasound scanning in 12 heifers during 51 oestrous cycles. Internal diameters of largest and second largest follicles and the number of smaller ovarian vesicles were determined. Diameters of dominant follicles showed inverse growth pattern to the second largest follicles and numbers of smaller follicles (greater than or equal to 5 mm). There was an increase in diameters of the largest follicles from beginning of dioestrous to day 9 and from time of luteolysis to ovulation, which was coincident which a decrease in diameters of the second largest follicles and numbers of smaller ovarian vesicles. Smaller follicles increased in count and diameter, when the dominant follicle achieved its largest dimension and started to regress. The cyclic corpus luteum had no local influence on diameters of the largest and second largest follicles in the ovary bearing the corpus luteum versus the contralateral ovary. Internal diameters of oestrous follicles measured 14.7 +/- 2.6 mm in heifers and 15.3 +/- 2.9 mm in cows at the day of oestrous (p greater than 0.05; t-test). Dioestrous follicles with similar size were detected during various stages of the oestrous cycle. The diameter of the dominant follicle is not an accurate criterion for determining the stage of the oestrous cycle.     

Association of Fertility with Numbers of Antral Follicles within a Follicular Wave During the Oestrous Cycle in Beef Cattle

The association between conception rate at first service and numbers of follicles developed during a follicular wave was examined in 102 suckled beef cows and 14 heifers. Follicular development was monitored using ultrasonography for either two (trial 1) or three (trial 2) consecutive oestrous cycles (pre-breeding, breeding and post-breeding equivalent). Animals were examined on alternate days from day 6 after first oestrus (day 0) until ovulation and from day 6 after insemination until next ovulation or day 24 of pregnancy and were observed for oestrus twice daily and inseminated artificially at either the second (trial 1) or third oestrus (trial 2). Cows were classified as having two or three waves of follicular development for each oestrous cycle. Numbers of follicles >or=4 mm per wave were determined, and based on the maximum diameter they attained, were classified as small (4-6 mm), medium (7-10 mm) or large (>or=11 mm) follicles. Total numbers of follicles, and primarily numbers of small and medium follicles, were affected by trial and within trial by cow, oestrous cycle and follicular wave. Heifers had more small and total numbers of follicles, but fewer large follicles than cows in trial 1 (p < 0.05). The average number of antral follicles per wave in the breeding cycle or post-breeding period did not affect conception rates, which averaged 84%. Repeatability of the total numbers of antral follicles between and among oestrous cycles and follicular waves ranged from 0.01 to 0.97. In conclusion, fertility was not affected by the numbers of antral follicles >or=4 mm in diameter in a single follicular wave.     

The final stages of dominant follicle selection in cattle

The final stages of ovarian follicle growth in cattle are typically characterized by the ultrasound-detectable emergence of a cohort of small (3-5mm in diameter) antral follicles, followed by a selection process during which the number of follicles continuing to grow decreases. Finally, only one follicle (the dominant follicle; DF) shows an enhanced growth rate and estradiol synthesis when it attains 8.5mm compared to its closest competitor (the largest subordinate follicle; SF). Cohort emergence is caused by a transient FSH rise, while DF selection occurs during declining FSH indicating differential FSH dependence of DF and SF. In order to elucidate the mechanisms underlying DF survival or SF atresia, this review aims to (i) describe follicular changes in the local production and regulation of members of the inhibin family of proteins and the insulin-like growth factor (IGF) system in relation to FSH deprivation leading to DF selection, and (ii) develop a model for DF selection outlining the putative involvement of inhibins, activin and follistatin on the one hand, and bioavailable IGFs regulated by IGF binding proteins (IGFBPs) and IGFBP proteases on the other hand. It is concluded, that the first indications of differential FSH dependence are seen within 33h of the FSH peak, and high amounts of precursor forms of inhibin and free activin, and low amounts of the lower molecular weight (MW) IGFBPs are related to follicle survival in terms of enhanced growth and estradiol synthesis, and suppression of granulosa cell apoptosis. In addition, maintenance of low amounts of intrafollicular IGFBP4 may constitute an important mechanism in the future DF to attain FSH independence.     

Recruitment and selection of ovarian follicles for determination of ovulation rate in the pig

Gonadotropins determine the follicle selection and ovulation rate. Follicle growth is independent of gonadotropins until antrum formation, at which time recruitment occurs. Once recruited, follicles will continue to grow or degenerate. In gilts, visible surface follicles are classified as small (<3mm), medium (3-6.9 mm) and large (> or =7.0mm). At estrus (day 0), there are approximately 15 small and medium follicles, and approximately 15 large follicles. By day 3, there may be approximately 30 small, 5 medium and no large follicles. During the remainder of the luteal phase, the pool of follicles increases and peaks at day 11-13 with approximately 50 small, and 30 medium, but with no large follicles observed. By the start of the follicular phase at day 15, numbers of small and medium follicles rapidly decline, while a pool of medium follicles is selected for the ovulation. The size of large follicles at estrus is heterogeneous (6.5-10.0 mm) but their number is reflective of the subsequent number of corpora lutea found following the ovulation. However, the time of medium follicle selection for ovulation is variable during the late luteal and early follicular phases. Suppression of FSH before and at the time of luteolysis reduces medium and large follicles but does not reduce the ovulation rate. In contrast, suppression of FSH for 3 days or unilateral ovariectomy after 3 days of the follicular phase prevents full ovulatory compensation. Therefore, FSH appears to be involved in the maintenance of a pool of medium follicles that can be selected by LH to mature and ovulate.     

Postpartum follicular and luteal activity in Holstein-Friesian cows genetically selected for high or low mature bodyweight: Relationships with follicle stimulating hormone,insulin, insulin-like growth factor-1 and growth hormone

AIM: To investigate ovarian follicular and luteal activity during the postpartum period of cows genetically selected for high or low mature bodyweight, in relation to metabolic and reproductive endocrine parameters, to determine whether there are differences between strains that could affect fertility outcomes.

METHODS: The presence of follicles ≥5 mm diameter and luteal structures was mapped in the ovaries of 12 high (heavystrain) and 12 low (light-strain) mature bodyweight cows by daily trans-rectal ultrasonography from Day 7 postpartum until the end of their first normal oestrous cycle. Blood samples were collected daily, for measurement of concentrations of follicle stimulating hormone (FSH), progesterone, growth hormone (GH), insulin-like growth factor-1 (IGF-1), and insulin. Intervals to first ovulation were calculated from ultrasonography data.

RESULTS: Heavy-strain cows had shorter intervals than light-strain cows from calving to the emergence of the first (9.0 (SE 0.9) vs 12.4 (SE 1.3) days) and second (16.4 (SE 1.8) vs 20.6 (SE 1.6) days) dominant follicles (p<0.05). Concentrations of FSH in heavy-strain cows prior to the emergence of the second, third and fourth dominant follicles were higher than in light-strain cows (p<0.05). Heavy-strain cows were more likely to have a large (>15 mm diameter) follicle earlier than light-strain cows (p<0.01). Concentrations of insulin and IGF-1, but not those of GH, were higher in heavy- than light-strain cows during the postpartum period (p=0.01 and p=0.02, respectively), and concentrations of both on Day 6 were inversely related to the time of emergence of the first dominant follicle (p>0.01).

Concentrations of progesterone were similar in both strains of cow until Day 10 of the first oestrous cycle, but thereafter were higher in light- than heavy-strain cows until Day 16. Progesterone concentrations in heavy-strain cows declined earlier and more rapidly than in their lighter counterparts.

CONCLUSION: These results indicate that there is a rapid postpartum resumption of follicular activity in both heavy-and light-strain cows, but that there is an earlier emergence of dominant follicles and ovulation in the former. Differences in luteal function, in terms of lower dioestrus progesterone concentrations and an earlier onset of luteolysis, in heavy- than light-strain cows might be sufficient to impair the fertility of the former.     

Control of ovarian follicles activity in the ewe.

During the ovine estrous cycles, three waves of follicular growth, closely associated with the FSH secretion pattern, were observed. The parameters of these follicular waves and the ability of follicles to produce steroids in vitro were studied in various conditions. In vivo, the follicular events were similar between the breeding season and the anestrus, except for the lack of ovulation; but at the end of the breeding season and in anestrus, the follicles lose a big part of their aromatization ability. In ewes carrying the Booroola fecundity gene or Cambridge fecundity gene, the reduction in follicular atresia seems to be one of the main follicular features implicated in the control of high ovulation rate. In vitro, the most relevant difference is an early acquisition of estrogen production ability of small follicles in Booroola fecundity gene barring ewes. Fluoro-gestone-acetate (FGA) pessaries reduced the number of growing follicles; despite this effect disappearing after the sponge withdrawal, the ovulation rate is significantly reduced. But an equine chorionic gonadotrophin (eCG) treatment restores the ovulation rate (OR) by reducing the atresia rate of pre-ovulatory follicles. In similar conditions, a pretreatment of the ewes with melatonin again reduced the atresia rate of large follicles and resulted in an increased ovulation rate. In vitro, FGA blocked aromatization ability, and melatonin inhibited both androstenedione and estradiol production, but a further treatment with eCG partly restores the steroid secretion. Immunization against androstenedione leads to a higher OR, owning to a reduced atresia of large follicles. Daily growth hormone injections for a hole cycle resulted in an increased follicular population and ovulation rate, while FSH plasma levels decreased and the follicle sensitivity to gonadotrophins was reduced.     

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.     

Characterization of immunoreactive IGF-I pattern during the peri-ovulatory period of the oestrous cycle of thoroughbred mares and its relation to other hormones

The aim of this study was to characterize ir-IGF-I pattern and its relation to other hormones during the oestrous cycle in mares. Nine non-pregnant non-lactating pluriparous thoroughbred mares were used. The studied mares were examined ultrasonically and bled daily to follow the ovarian changes and the hormonal milieu for a complete Interovulatory interval (IOI). Two (minor and major) follicular waves were characterized per IOI in thoroughbred mares. The largest follicle of the first follicular wave (DF1) was firstly detected at D - 1.75 ± 0.47 with a growth rate of 2.78 ± 0.14mm/day and maximum diameter of 22.45 ± 0.75mm on day 6.65 ± 0.82. The largest follicle of the second follicular wave (DF2) had a growth rate of 2.15 ± 0.29 mm/day, reached a maximum diameter of 42.70 ± 2.63 mm on D 19.25 ± 0.43. Ir-IGF-I increased significantly prior to ovulation and had a similar pattern to oestrogen (r = 0.84, p < 0.05), suggesting that the ovarian follicles are the main source of circulating ir-IGF-I during the oestrous cycle of mares and that ir-IGF-I may be a crucial factor in follicular differentiation and maturation. In conclusion, this study demonstrated that ir-IGF-I is secreted during the oestrous phase of the cycle concomitant with the development of the future ovulatory dominant follicle, and it may act in synergy with other hormones for the selection and differentiation of the dominant follicle.     

Use of Equine Chorionic Gonadotropin to Control Reproduction of the Dairy Cow: A Review

Equine chorionic gonadotropin (eCG) is a member of the glycoprotein family of hormones along with LH, FSH and thyroid‐stimulating hormone. In non‐equid species, eCG shows high LH‐ and FSH‐like activities and has a high affinity for both FSH and LH receptors in the ovaries. On the granulosa and thecal cells of the follicle, eCG has long‐lasting LH‐ and FSH‐like effects that stimulate oestradiol and progesterone secretion. Thus, eCG administration in dairy cattle results in fewer atretic follicles, the recruitment of more small follicles showing an elevated growth rate, the sustained growth of medium and large follicles and improved development of the dominant and pre‐ovulatory follicle. In consequence, the quality of the ensuing CL is improved, and thereby progesterone secretion increased. Based on these characteristics, eCG treatment is utilized in veterinary medicine to control the reproductive activity of the cow by i) improving reproductive performance during early post‐partum stages; ii) increasing ovulation and pregnancy rates in non‐cyclic cows; iii) improving the conception rate in cows showing delayed ovulation; and finally, iv) eCG is currently included in protocols for fixed‐time artificial insemination since after inducing the synchrony of ovulation using a progesterone‐releasing device, eCG has beneficial effects on embryo development and survival. The above effects are not always observed in cyclic animals, but they are evident in animals in which LH secretion and ovarian activity are reduced or compromised, for instance, during the early post‐partum period, under seasonal heat stress, in anoestrus animals or in animals with a low body condition score.     

FSH and LH concentrations preceding post-partum ovulation in the mare

Equine follicle stimulating hormone (FSH) and luteinizing hormone (LH) were measured in the serum obtained from pregnant mares, bled daily, from up to 17 days before parturition until the first ovulation after parturition. LH was at baseline levels until 2 days before ovulation when it started, and continued, to rise until after ovulation. FSH surges occurred at approximately 24 and 14 days before this post-partum ovulation (12 and 2 days before parturition), thus showing a similar pattern to the cyclic mare, consistent with the hypothesis that 2 surges at these times prepare and prime follicles for subsequent ovulation. The high plasma oestradiol levels that occurred during and immediately before these FSH surges did not show a negative feedback, or the inverse relationship between FSH and oestradiol which occurs in the cyclic mare, and in other species.