The estrous cycle in horses is a recurring reproductive cycle that mares undergo, typically during the breeding season. This cycle is characterized by a series of hormonal changes that prepare the mare for potential conception. The cycle is divided into two main phases: estrus and diestrus. During estrus, the mare is receptive to mating, and ovulation occurs. In contrast, the diestrus phase is a period when the mare is not receptive to a stallion. The regulation of the estrous cycle involves a complex interplay of hormones such as estrogen, progesterone, luteinizing hormone, and follicle-stimulating hormone. Understanding the estrous cycle is important for effective breeding management and reproductive health in horses. This page compiles peer-reviewed research studies and scholarly articles that explore the physiology, hormonal regulation, and implications of the estrous cycle in equine reproduction.
Johnson AL.Studies were conducted to characterize circulating concentrations of prolactin, thyroxine (T4) and triiodothyronine (T3) relative to season and the estrous cycle, and to evaluate the ability of thyrotropin-releasing hormone (TRH) to induce prolactin secretion in the horse mare. The increase in serum prolactin following the iv injection of 0, 5, 50 and 500 micrograms TRH was dose-related, while all three doses of TRH induced a significant and comparable increase in serum T4. Seasonal variations in serum prolactin were directly correlated (P less than .001) with changes in photoperiod (r = .80) ...
Lauderdale JW.Puberty: For the gilt and filly, the first corpus luteum (CL) appears to have a normal lifespan. For both species, first CL usually is associated with estrus but can form in the absence of estrus. For the ewe and cow, a transient (1 to 4 d, ewe; 3 to 10 d, cow) rise and fall of ovarian derived progesterone (P4) is detected in peripheral blood (80% of ewes; 50% of heifers) prior to first "normal" CL. The first CL of apparent normal lifespan is not accompanied by estrus in the ewe. The first CL in the cow may or may not be accompanied by estrus; first estrus in the cow can be anovulatory. Data a...
Malinowski K, Johnson AL, Scanes CG.The ability of interrupted photoperiods to induce early estrus and ovulation was examined. Horse mares were exposed to long (16 h light) or short (10 h light), noninterrupted photoperiods, ambient light, or various interrupted photoperiod treatments from December 1 to April 15 (135 d). Follicular development was assessed by rectal palpation and estrous behavior was determined by teasing with a stallion. Serum concentrations of progesterone were used as an indicator of corpus luteum function. Differences among the light treatment groups were compared for the following behavioral and ovarian cha...
Hayes KE, Pierson RA, Scraba ST, Ginther OJ.The morphological changes in ultrasound images of the uterus at various times of the year were characterized in nonbred mares, using a linear-array scanner. The uterus was recorded as having an ultrasonic morphology characteristic of diestrus (uterine score 1, endometrial folds not visible), estrus (score 3, distinct endometrial folds), or an intermediate stage (score 2). In Experiment I, uterine scores for the first ovulatory period of the year were compared to scores for the second period in 23 pony mares. More mares (P<0.05) showed endometrial folding prior to the second ovulation of the...
Fitzgerald BP, I'Anson H, Legan SJ, Loy RG.To determine whether luteinizing hormone (LH) secretion during the first estrous cycle postpartum is characterized by pulsatile release, circulating LH concentrations were measured in 8 postpartum mares, 4 of which had been treated with 150 mg progesterone and 10 mg estradiol daily for 20 days after foaling to delay ovulation. Blood samples were collected every 15 min for 8 h on 4 occasions: 3 times during the follicular phase (Days 2-4, 5-7, and 8-11 after either foaling or end of steroid treatment), and once during the luteal phase (Days 5-8 after ovulation). Ovulation occurred in 4 mares 13...
Widders PR, Stokes CR, David JS, Bourne FJ.The effect of cycle stage on immunoglobulin and albumin levels in serum, follicular fluid, oviductal, uterine and vaginal secretions was measured. There was no variation in serum immunoglobulin levels during the oestrous cycle, although IgM levels were elevated in cyclic mares compared to non-cyclic (immature and anoestrous) animals. Similarly, there was no cyclical variation in follicular or oviductal protein concentrations. In the uterus, IgG and IgA levels relative to total protein were higher in oestrogenic than in progestagenic secretions, while the trend in relative IgM concentrations wa...
Brockschmidt LD, Loch WE, Sikes JD.An experiment was conducted to test the effect of repeated transcervical (non-surgical) uterine flushing and a prostaglandin analogue (PG) on the estrous cycle of pony mares. Uteri in group A were trancervically flushed for embryos 7 to 9 days post ovulation. In addition, group B mares were given 5 ml of PG by intramuscular injection on the day of flushing. Group C served as controls and were not flushed or given PG but were allowed to cycle normally. All mares (except controls) were bred A.I. every other day during estrus. There was no effect on embryo recovery rate from repeated flushing or ...
Squires EL, Voss JL, Maher JM, Shideler RK.The effect of prior treatment with anabolic steroids was studied in 46 three-year-old mares. In the preceding year, these mares had been assigned to 1 of 4 treatment groups and had received the manufacturer's recommended dosage of 1.1 mg of boldenone undecylenate (BU)/kg of body weight, 4.4 mg of boldenone undecylenate (4 BU)/kg, 1.1 mg of nandrolone decanoate (ND)/kg, or 0.04 ml of sesame oil/kg (control, C). Mares had received an injection every 3 weeks for 54 weeks for a total of 19 injections, with the final injection in December. In the following breeding season, fewer (P less than 0.05) ...
Penzhorn BL, Gilbert RO.A 17-year-old Nooitgedacht mare was presented in mid-summer after failing to conceive during the previous 2 breeding seasons. The mare conceived to service during a PG-induced oestrus when synthetic GnRH was used to induce ovulation.
Aguado LI, Ojeda SR.During the days preceding the first ovulation the ovary of the rat exhibits a remarkable increase in estradiol (E2) and progesterone (P) release in response to gonadotropins. No such increase is observed in the case of androgens (A, testosterone + dihydrotestosterone). The present experiments were undertaken to examine the possibility of reproducing these developmental events by stimulating the ovary with a gonadotropin that has substantial FSH-like activity. In vivo administration of pregnant mare serum gonadotropin (PMSG) to juvenile 29-day-old rats greatly increased the in vitro E2 and A re...
Strzemienski PJ, Do D, Kenney RM.Luminal fluid from the mare uterus was used to investigate its relation to antibacterial defenses. Uterine flushings were collected at Day 3 of estrus, Day 8 postovulation and Day 15 postovulation. Uterine proteins were concentrated by ultrafiltration, dialyzed and examined for chemotactic activity to neutrophils and for antibacterial properties. Serum taken at the time of flushing was dialyzed and studied in a similar manner. Neutrophil migration in response to serum from Day 3 estrus and Day 8 postovulation was increased (P less than 0.05) above controls. Uterine protein from Day 8 postovula...
Zavy MT, Vernon MW, Sharp DC, Bazer FW.Comparisons of estrone, 17 beta-estradiol, and plasma progestin concentrations were made in uterine fluid and peripheral blood of nonpregnant and pregnant pony mares. Concentrations of these steroids were also measured within yolk sac fluid from blastocysts on days 12, 14, 16, and 18 of pregnancy to obtain more complete analyses of the uterine environment (uterine fluid plus yolk sac fluid) of early pregnancy. Thirty mares were randomly assigned to six treatment groups (n = 5/group), and uterine fluid and peripheral blood samples were obtained on days 8, 12, 14, 16, 18, and 20 postovulation. A...
Blue HB, Blue MG, Kenney RM, Merritt TL.Forty uterine fluid samples were obtained from 4 mares classified as resistant to uterine bacterial infection. The uterus of each mare was flushed with 50 ml of saline solution during estrus and diestrus of successive estrous cycles. Bacteria or fungi were isolated from 4 samples, and 7 additional samples were obtained from a mare with active intrauterine infection. Fluid volumes obtained during estrus (means = 40.3 +/- 11 ml) tended to be greater than those recovered during diestrus (means = 36.8 +/- 7.9 ml), but the difference was not significant. Concentrations and yields of protein in reco...
Brook D.A guarded, sterile swab is used to obtain samples for uterine culture. With the mare in stocks, the tail bandage and the perineum washed, the culture rod is introduced into the vagina with a gloved hand. After the rod is guided through the cervix, the guard cap is dislodged and the swab is rubbed along the endometrium, after which the rod is extracted. Samples for uterine culture should only be obtained during full estrus. Swabs should be directly plated onto agar within 2 hours of collection. Blood agar is appropriate for initial screening, but use of specialized types of agar expedites ident...
Driancourt MA, Palmer E.An experiment was carried out on pony mares to establish the time of the oestrous cycle at which ovarian follicles are recruited for ovulation. In one group (n=7), the cycle was interrupted at the preovulatory stage by removing the preovulatory follicle; in another group (n=13) the cycle was interrupted at day 6 of the luteal phase by inducing luteolysis with a prostaglandin injection (PG). In a subgroup (n=7) of those given PG, the ovary not bearing the corpus luteum was removed at the time of injection. A further group (n=6) served as surgical controls. The interval to the next ovulation and...
Thompson DL, Voelkel SA, Reville-Moroz SI, Godke RA, Derrick DJ.Effects of testosterone propionate (TP) treatment on plasma concentrations of luteinizing hormone (LH) and follicle-stimulating hormone (FSH) before and after an injection of gonadotropin releasing hormone (GnRH) were studied using ovariectomized cows and pony mares. An initial injection of GnRH (1 microgram/kg of body weight) was followed by either TP treatment or control injections for 10 (cows) or 11 (ponies) d. A second GnRH injection was administered 1 d after the last TP or oil injection. Concentrations of LH and FSH were determined in samples of plasma taken before and after each GnRH i...
Zavy MT, Vernon MW, Asquith RL, Bazer FW, Sharp DC.Two experiments were conducted to assess the effect of exogenous hormone treatment on uterine luminal prostaglandin F (PGF). In the first experiment ovariectomized pony mares received either corn oil (21 days, n = 3), estradiol valerate (21 days, n = 3), progesterone (21 days, n = 3) or estradiol valerate (7 days) followed by progesterone (14 days, n = 4). Progesterone treated mares had higher (P less than .01) uterine luminal PGF compared with all other groups, and no differences were detected between other treatment comparisons. In Experiment II, uterine fluid was collected from 4 ovariectom...
Thompson DL, Reville SI, Derrick DJ, Walker MP.Eight seasonally anestrous mares were administered intravaginal polyurethane sponges on December 15 and then weekly thereafter until February 1. Control mares received no sponges or genital contact. Sponge insertion caused an immediate surge in follicle-stimulating hormone (FSH) concentrations in jugular plasma in 50% of treated mares whereas no control mares had surges in FSH (P less than .05). The effect of treatment on luteinizing hormone (LH) concentrations was much less dramatic and only three treated mares appeared to have positive responses. Sponge-treated mares exhibited positive respo...
Asa CS, Goldfoot DA, Ginther OJ.Pregnant mares (N = 12) were observed with a stallion from early gestation until parturition for sexual behavior. Observations were conducted for 20 min per day for 5 days each month from September until July. No mares exhibited full estrus and no intromissions or ejaculations occurred at any time during the study. Social interactions such as male approach and female/female mutual grooming occurred with greater mean frequency or duration in some months of spring and fall than in winter months. (P less than 0.05). None of the recorded behaviors differed by month of gestation. In a comparison of...
Asa CS, Robinson JA, Ginther OJ.Daily blood samples from four mares were assayed for cortisol through a total of eight ovulatory cycles. Mean cortisol concentrations on days -14, -13, -10, -9 and -8 before ovulation (dioestrus) were greater than on days -5 to -1 (oestrus). The highest mean (+/- S.E.M) value of cortisol occurred on day -10 (260 +/-28 nmol/l) and the lowest on day -2 (142 +/- 14 nmol/l). A single episode on a day in late dioestrus characterized the maximum cortisol value per cycle for five of eight cycles. Extraction of plasma samples with petroleum ether or chromatography before assay, to eliminate interferen...
Thompson DL, Reville-Moroz SI, Derrick DJ, Walker MP.Sixteen intact cyclic mares were treated on the fourth day of estrus and then every other day for a total of six injections with 1) testosterone propionate, 2) dihydrotestosterone (DHT) benzoate, 3) estradiol (E2) benzoate or 4) safflower oil. Mares were given gonadotropin releasing hormone (GnRH) on Day 3 of estrus (pretreatment) and again 24 h after the last steroid or oil injection. Treatment with testosterone propionate resulted in a greater (P less than 0.05) follicle-stimulating hormone (FSH) response to the second injection of GnRH compared with all other treatments. Treatment with DHT ...
YoungLai EV, Jarrell JF.Granulosa cells were harvested from mares at various stages of the oestrous cycle and incubated in Krebs-Ringer bicarbonate buffer with 1 beta,2 beta[3H]androstenedione as substrate. The release of 3H2O expressed as CPM/h/mg protein varied from 44000 to 768000 in follicles from 7 mares. The release of 3H2O was not significantly altered by luteinizing hormone, follicle stimulating hormone or pregnant mare's serum gonadotrophin. There was a significant negative correlation between the release of 3H2O and the concentration of progesterone in the follicular fluid. Based on the assumption that the ...
Withrow JM, Sargent GF, Scheffrahn NS, Kesler DJ.Two pony mares were administered 150 mg of testosterone propionate every other day for 20 days (ten injections) and every ten days there-after. An additional two mares and one stallion were not treated and served as controls. Testosterone propionate was dissolved in absolute ethanol and administered subcutaneously. Sex behavior tests were conducted 26 and 40 days after the first injection. Control mares exhibited very little male sex behavior. Both testosterone propionatetreated mares, however, exhibited mounting, sniffing, flehmen, biting and vocalization behavior in the presence of an estrou...
Ginther OJ, Scraba ST, Nuti LC.Pony mares (n=480) and 16 stallions were assigned to four herds of 60 mares and one stallion (large herds) and to 12 herds of 20 mares and one stallion (small herds). The stallions remained with the herds continuously for all of the large herds and seven of the small herds. In the five remaining small herds the stallion was put into a herd for three hours every two days for 12 observation periods. Pregnancy rates and day of ovulation were estimated by size of embryonal enlargements. Mean pregnancy rates of 51% and 54% were obtained in the small herds and 42% in the large herds during a 48-day ...
Howey WP, Jochle W, Barnes WJ.Alfaprostal (K 11941), a novel prostaglandin F2 alpha analogue was clinically tested in 24 dioestrous mares, 40 anoestrous mares and 31 postpartum mares, all being given 2 or 3 mg intramuscularly. Blood samples were taken for the determination of plasma progesterone levels and the objective confirmation of luteolytic drug effects. Two hundred and thirty-six mares at the same location were used for comparisons of the rates of pregnancy, early embryonic loss and foaling. Alfaprostol was found to act as a potent luteolytic agent with good oestrus induction, follicular development and normal conce...
Thompson DL, Reville SI, Walker MP, Derrick DJ, Papkoff H.To study the possible role of ovarian androgens in regulation of follicle stimulating hormone (FSH) secretion in the cycling mare, five mature, intact mares were treated with testosterone (20 micrograms/kg of body weight) daily during estrus; five control mares received safflower oil on the same schedule. Mares were teased for estrus and samples of jugular blood were drawn daily through one full estrous cycle. Concentrations of FSH in plasma were measured by a newly developed radioimmunoassay based on anti-ovine FSH serum and radioiodinated equine FSH. Testosterone treatment during estrus had ...
Thompson DL, Godke RA, Squires EL.Twelve mares fed altrenogest for 14 d were used to study the effects of a single injection of testosterone propionate on concentrations of follicle-stimulating hormone (FSH) during diestrus, and to relate the normal and perturbed patterns of FSH secretion to subsequent estrous characteristics and fertility. Seven of 12 mares received testosterone propionate at 200 micrograms/kg of body weight on d 5 of progestogen feeding. Mares were teased and blood samples were drawn daily; all mares were artificially inseminated at the first estrus after progestogen treatment. Testosterone propionate treatm...
Adams GP, Bosu WT.This article reviews the reproductive events in the nonpregnant mare with emphasis on recent advances. The discussion is restricted to the salient features of puberty (prenatal and prepubertal events), seasonality (gonadotropins, photoperiod, and other modifying factors), and the estrous cycle (hormones, estrus, diestrus, and the control of cyclicity) in the nonpregnant mare.
Vivrette SL, Irvine CH.Acyclic mare given oestradiol for 3 days to simulate the preovulatory plasma oestradiol surge showed a non-significant 37% decrease in plasma LH during treatment. When GnRH analogue injections were given with oestradiol on Days 1--3, oestradiol had no effect on each GnRH-induced LH increase, but LH increases were more prolonged following subsequent GnRH injections on Days 4--7 when oestradiol was no longer being given. A much greater prolongation of LH release occurred when the course of GnRH injections was commenced after oestradiol treatment ceased; the LH response was almost identical to th...
Strzemienski PJ, Do D, Kenney RM.Luminal fluid from the mare uterus was used to investigate its relation to antibacterial defenses. Uterine flushings were collected at Day 3 of estrus, Day 8 postovulation and Day 15 postovulation. Uterine proteins were concentrated by ultrafiltration, dialyzed and examined for chemotactic activity to neutrophils and for antibacterial properties. Serum taken at the time of flushing was dialyzed and studied in a similar manner. Neutrophil migration in response to serum from Day 3 estrus and Day 8 postovulation was increased (P less than 0.05) above controls. Uterine protein from Day 8 postovula...
Belonje PC, van Niekerk CH.Attention is drawn to the beneficial effect of improved nutrition during winter and early spring on the ovarian activity of mares. Furthermore, the necessity of an adequate plane of nutrition during early pregnancy to prevent embryonic resorption is stressed.
Montavon S.Few scientists predicted the impact that echography would have on the equine breeding industry. The aim of this practical review is to give the practitioner the ability to be familiar with the ovarian echographic exam. He should be able to recognize different stages of estrous cycle in the mare, status of preovulatory follicles and to predict precisely the time of ovulation. Various specific criteria and details regarding the equine echography are studied and illustrated, using the last scientific data published after the Fifth International Symposium on Equine Reproduction in Deauville.
Stecco R, Paccamonti D, Gutjahr S, Pinto CR, Eilts B.Oxytocin is released in response to teasing during both estrus and diestrus in mares, and at least during estrus, teasing results in an increase in electromyographic activity in the uterus. Exogenous oxytocin causes an increase in intrauterine pressure and prior studies have shown that this response is correlated to the day of the estrous cycle. To determine if teasing causes an increase in intrauterine pressure and if this response varies by day of the cycle, intrauterine pressure was measured while mares were teased with a stallion 2 days before ovulation, on the day ovulation was detected a...
Vogelsang SG, Sorensen AM, Potter GD, Burns SJ, Kraemer DC.Embryos were collected nonsurgically on Day 7 or 8 after ovulation from 7 Quarter horse mares using a modified 30-ml Foley catheter to flush the uterine horn ipsilateral to the recent ovulation with 500 ml TCM-199 containing Hepes buffer. After collection, the uteri were infused with nitrofurazone to reduce the chances of infection due to the procedure. Eleven collections from 7 mares resulted in recovery of 9 embryos and nonsurgical transfer of 4 of these resulted in the birth of one foal. After collections, 8 oestrous cycles averages 22.75 days and 2 extended oestrous cycles were 43 and 59 d...
Watson ED, Dixon CE.The distribution of T lymphocytes and of cells bearing MHC Class II antigens in the endometrium of the mare was studied using an avidin-biotin-peroxidase staining method. The cells within the endometrium which expressed MHC Class II were macrophages, lymphocytes, monocytes, dendritic cells, epithelial cells and endothelial cells. MHC Class II expression increased significantly (P < 0.05) in the luminal epithelium and tended (P = 0.0573) to increase in the subepithelial layers during oestrus. Numbers of T lymphocytes did not differ between oestrus and dioestrus. MHC Class II expression and T...
Gillon A, Ho ENM, Chan GHM, Kauff A, Hughes G, Lund RA, Ashley Z, Wan TSM, Heather AK.Altrenogest is a commonly used progestogen for the suppression of oestrus and associated distracting behaviours that interfere with training and performance of female racehorses. The steroid is derived from 19-nor testosterone and is structurally similar to the anabolic androgenic steroid, trenbolone. In this study, the relative androgen potency of altrenogest was determined by a kidney (HEK293) cell androgen bioassay. The HEK293 bioassay shows that in its pure form, altrenogest has a high relative potency compared with testosterone but is not as strong as β-trenbolone. Our results also show ...
Nitschelm D, van der Horst CJ.It appeared that in mares suffering from endometritis the concentrations of pregnenolone, of the compound "5.4", and of progesterone in the blood during the first 2 days of the heat period were significantly higher than in normal cycling mares, in which the concentrations of all steroids were less than or equal to 1 ng/ml. In five endometritis mares the "5.4" and progesterone concentrations decreased after the first days of the heat period, and at about two days before ovulation the values were comparable to those of normal cycling mares. Eight mares suffering from a severe endometritis did no...
Allen WE, Newcombe JR.The condition of anoestrus in the mare is described and the various causes of this syndrome are discussed. A regimen for examining systematically the clinically anoestrus mare is proposed, and methods of treatment are suggested.
Jeannerat E, Janett F, Sieme H, Wedekind C, Burger D.The theory of ejaculate economics was mainly built around different sperm competition scenarios but also predicts that investments into ejaculates depend on female fecundity. Previous tests of this prediction focused on invertebrates and lower vertebrate, and on species with high female reproductive potential. It remains unclear whether the prediction also holds for polygynous mammals with low female reproductive potential (due to low litter size and long inter-birth intervals). We used horses (Equus caballus) to experimentally test whether semen characteristics are adjusted to the oestrous cy...
Ignácio FS, Bergfelt DR, Mendes D, Orlandi C, Araújo GH, Oliveira JV, Montechiesi DF, Carvalho LR, Meira C.The primary objective of this study was to examine the follicular and ovulatory responses following treatment with pFSH in association with ablation-induced or spontaneous follicular wave emergence or follicle deviation during diestrus in crossbred (Mangalarga×Arabian) and Brazilian Warmblood mares with a propensity for spontaneous multiple ovulations; secondary considerations were given to the collection of embryos. In Experiment 1, crossbred mares were administered (im) saline (control, n=7) or pFSH (25mg) when the largest follicle of the ablation-induced follicular wave reached ≥13mm (n=...
Campbell MLH, Hampshire D, Hamstead LE, Rose BV, Smith KC, de Mestre AM.Intrauterine infusion of peanut oil at Day 10 post-ovulation has been reported to prolong dioestrus in mares. However, the effects of peanut oil treatment on the endometrium and whether the technique is painful have not been assessed. The objectives of this study were, (i) to determine the effect of intrauterine infusion of peanut oil on endometrial health, (ii) to determine whether use of intrauterine peanut oil is painful and (iii) to confirm that peanut oil causes prolonged dioestrus. Six mares aged 3-12 years old were used in a cross-over design with each mare administered both 1 ml of in...
King SS, Evans JW.Follicular growth, circulating estradiol concentrations and endometrial prostaglandin F2 alpha (PGF2 alpha) production were measured to determine whether there is an interrelationship among these factors associated with luteolysis. Follicular growth was monitored by rectal palpation every other day during diestrus in 16 mares. Plasma estradiol was determined for daily samples during all estrous cycles. Endometrial tissue was removed for PGF2 alpha analysis by radioimmunoassay on d 10, 12, 14 or 16 during several normal cycles and after d 30 during spontaneously prolonged cycles. Circulating es...