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Topic:Reproduction

Reproduction in horses encompasses the biological processes and mechanisms involved in the breeding and development of equine offspring. This includes the study of reproductive anatomy, physiology, and endocrinology in both mares and stallions. Key areas of interest include the estrous cycle, ovulation, conception, gestation, and parturition. Researchers also examine factors influencing fertility, reproductive technologies such as artificial insemination and embryo transfer, and management practices that impact reproductive success. This page compiles peer-reviewed research studies and scholarly articles that explore the physiological, genetic, and environmental aspects of equine reproduction.
The comparison of equine articular cartilage progenitor cells and bone marrow-derived stromal cells as potential cell sources for cartilage repair in the horse.
Veterinary journal (London, England : 1997)    October 2, 2011   Volume 192, Issue 3 345-351 doi: 10.1016/j.tvjl.2011.08.036
McCarthy HE, Bara JJ, Brakspear K, Singhrao SK, Archer CW.A chondrocyte progenitor population isolated from the surface zone of articular cartilage presents a promising cell source for cell-based cartilage repair. In this study, equine articular cartilage progenitor cells (ACPCs) and equine bone marrow-derived stromal cells (BMSCs) were compared as potential cell sources for repair. Clonally derived BMSCs and ACPCs demonstrated expression of the cell fate selector gene, Notch-1, and the putative stem cell markers STRO-1, CD90 and CD166. Chondrogenic induction revealed positive labelling for collagen type II and aggrecan. Collagen type X was not detec...
Ultrasound characteristics of experimentally induced luteinized unruptured follicles (LUF) and naturally occurring hemorrhagic anovulatory follicles (HAF) in the mare.
Theriogenology    September 29, 2011   Volume 77, Issue 3 514-524 doi: 10.1016/j.theriogenology.2011.08.026
Cuervo-Arango J, Newcombe JR.The development of hemorrhagic anovulatory follicles (HAF) involves luteinization and hemorrhage of the follicle. This is observed on ultrasound as an increase in the echogenicity of the granulosa layer and formation of echoic particles in the antrum. The inhibition of prostaglandin synthesis with flunixin meglumine (FM) during the periovulatory period induces ovulatory failure with development of luteinized unruptured follicles (LUF). These two types of anovulatory follicles appear to share similar ultrasound features but they have not been compared critically. The following endpoints: follic...
Ultrasonographic features of the mule embryo, fetus and fetal-placental unit.
Theriogenology    September 29, 2011   Volume 77, Issue 2 240-252 doi: 10.1016/j.theriogenology.2011.07.030
Paolucci M, Palombi C, Sylla L, Stradaioli G, Monaci M.The aim of this study was to establish baseline ultrasound data concerning the mule conceptus during gestation. Ten multiparous Trotter mares were artificially inseminated with chilled semen from an Amiatino jack donkey. Daily transrectal ultrasonography was carried out from the day of ovulation until Day 50 of gestation to determine the following: first detection of the embryonic vesicle (EV), mobility phase, EV diameter, day of EV fixation, changes in EV shape, date of yolk sac regression and embryo crown-rump length. Monthly ultrasonic assessments from Day 50 of gestation to term were carri...
Effects of nutritional cues on the duration of the winter anovulatory phase and on associated hormone levels in adult female Welsh pony horses (Equus caballus).
Reproductive biology and endocrinology : RB&E    September 29, 2011   Volume 9 130 doi: 10.1186/1477-7827-9-130
Salazar-Ortiz J, Camous S, Briant C, Lardic L, Chesneau D, Guillaume D.Mares have an annual reproductive rhythm, with a phase of inactivity in midwinter. The aim of this study was to determine the impact of food restriction on physiological and metabolic hallmarks of this rhythm. Methods: Over three successive years, 3 groups of 10 mares were kept under natural photoperiod. A 'well-fed' group was fed to maintain the mares in good body condition; a 'restricted' group received a diet calculated to keep the mares thin and a 'variable' group was fed during some periods like the 'restricted' group and during some other periods like the 'well-fed' group, with the aim o...
Uterine artery blood flow remains unchanged in pregnant mares in response to short-term administration of pentoxifylline.
Theriogenology    September 29, 2011   Volume 77, Issue 2 430-436 doi: 10.1016/j.theriogenology.2011.08.018
Bailey CS, Sper RB, Schewmaker JL, Buchanan CN, Beachler TM, Pozor MA, Whitacre MD.The objective of this study was to use Doppler ultrasound technology to determine whether pentoxifylline administration increased uterine blood flow in normal pregnant pony mares. Thirteen pregnant pony mares between 18 and 190 d of gestation (mean ± SEM, 101 ± 55) were utilized for the study during two trial periods. In each trial, pentoxifylline (17 mg/kg by mouth every 12h, diluted in syrup) was administered to half of the mares for 3 d, while the other mares were treated with syrup only. Doppler measurements were obtained from the right and left uterine arteries from each mare for 2 d pr...
Distribution of mast cells in vaginal, cervical and uterine tissue of non-pregnant mares: investigations on correlations with ovarian steroids.
Reproduction in domestic animals = Zuchthygiene    September 27, 2011   Volume 47, Issue 2 e29-e31 doi: 10.1111/j.1439-0531.2011.01897.x
Walter J, Klein C, Wehrend A.Mast cells constitute part of the cellular immune system of the genital tract. They play a potential role in cervical remodelling during parturition and are subject to the influence of ovarian steroids. In this study, the influence of oestradiol-17β and progesterone on the distribution of mast cells in equine vaginal, cervical and uterine tissue was evaluated. Genital tracts were retrieved from healthy mares at a local slaughterhouse. The cervix was divided along the longitudinal axis into five equivalent regions from cranial to caudal (R1 to R5). Toluidine blue staining was used for the dete...
The effects of an advanced uterine environment on embryonic survival in the mare.
Equine veterinary journal    September 25, 2011   Volume 44, Issue 4 432-439 doi: 10.1111/j.2042-3306.2011.00473.x
Wilsher S, Lefranc AC, Allen WR.During embryo transfer (ET) the equine embryo can tolerate a wide degree of negative asynchrony but positive asynchrony of >2 days usually results in embryonic death. There is still confusion over whether this is due to the inability of the embryo to induce luteostasis or to an inappropriate uterine environment. Objective: To assess embryo survival and development in an advanced uterine environment. Objective: Embryo-uterine asynchrony, not the embryo's inability to induce luteostasis, is responsible for embryonic death in recipient mares with a >2 days chronologically advanced uterus. Methods...
A pilot comparison of laser-assisted vs piezo drill ICSI for the in vitro production of horse embryos.
Reproduction in domestic animals = Zuchthygiene    September 24, 2011   Volume 47, Issue 1 e1-e3 doi: 10.1111/j.1439-0531.2011.01814.x
Smits K, Govaere J, Hoogewijs M, Piepers S, Van Soom A.Intracytoplasmic sperm injection (ICSI) is the method of choice for the in vitro production (IVP) of equine embryos. However, conventional ICSI has been associated with mechanical damage to the oocyte caused by the deformation of the zona pellucida (ZP) and exposure of the oolemma to negative pressure during injection. Introduction of the less traumatic and more efficient piezo drill-assisted ICSI (PDAI) yielded higher cleavage rates and more consistent results. Nevertheless, PDAI is also associated with disadvantages such as the use of mercury and possible DNA damage. This led us to explore a...
Comparison of transvaginal natural orifice transluminal endoscopic surgery (NOTES®) and laparoscopy for elective bilateral ovariectomy in standing mares.
Veterinary surgery : VS    September 23, 2011   Volume 40, Issue 8 998-1008 doi: 10.1111/j.1532-950X.2011.00877.x
Pader K, Freeman LJ, Constable PD, Wu CC, Snyder PW, Lescun TB.To compare surgical trauma, perioperative pain, surgical time, and complication rate in mares undergoing standing bilateral ovariectomy by natural orifice transluminal endoscopic surgery (NOTES) or laparoscopy. Methods: Experimental study. Methods: Healthy mares (n = 12). Methods: Ovariectomy was performed with a vessel-sealing device by transvaginal NOTES (n = 6) using a flexible endoscope and specialized instruments or by bilateral flank laparoscopy (n = 6). Preoperative and postoperative complete blood count (CBC), plasma fibrinogen concentration, serum amyloid A concentration, peritoneal n...
Expression of apoptotic genes in immature and in vitro matured equine oocytes and cumulus cells.
Zygote (Cambridge, England)    September 21, 2011   Volume 21, Issue 3 279-285 doi: 10.1017/S0967199411000554
Leon PM, Campos VF, Kaefer C, Begnini KR, McBride AJ, Dellagostin OA, Seixas FK, Deschamps JC, Collares T.The gene expression of Bax, Bcl-2, survivin and p53, following in vitro maturation of equine oocytes, was compared in morphologically distinct oocytes and cumulus cells. Cumulus-oocyte complexes (COC) were harvested and divided into two groups: G1 - morphologically healthy cells; and G2 - less viable cells or cells with some degree of atresia. Total RNA was isolated from both immature and in vitro matured COC and real-time reverse transcription polymerase chain reaction (qRT-PCR) was used to quantify gene expression. Our results showed there was significantly higher expression of survivin (P &...
Treatment with recombinant equine follicle stimulating hormone (reFSH) followed by recombinant equine luteinizing hormone (reLH) increases embryo recovery in superovulated mares.
Animal reproduction science    September 17, 2011   Volume 128, Issue 1-4 52-59 doi: 10.1016/j.anireprosci.2011.09.002
Meyers-Brown G, Bidstrup LA, Famula TR, Colgin M, Roser JF.The dynamics of ovarian follicular development depend on a timely interaction of gonadotropins and gonadal feedback in the mare. The development and efficacy of genetically cloned recombinant equine gonadotropins (reFSH and reLH) increase follicular activity and induce ovulation, respectively, but an optimum embryo recovery regimen in superovulated mares has not been established. The objective of this study was to determine if treatment with reFSH followed by reLH would increase the embryo per ovulation ratio and the number of embryos recovered after superovulation in mares. Sixteen estrous cy...
Liposomes as an alternative to egg yolk in stallion freezing extender.
Theriogenology    September 14, 2011   Volume 77, Issue 2 268-279 doi: 10.1016/j.theriogenology.2011.08.001
Pillet E, Labbe C, Batellier F, Duchamp G, Beaumal V, Anton M, Desherces S, Schmitt E, Magistrini M.Egg yolk is normally used as a protective agent to freeze semen of equine and other species. However, addition of egg yolk in extenders is not without disadvantages and the demand to find cryoprotective alternatives is strong. The objective of this study was to test the cryoprotective capacities of liposomes composed of egg yolk phospholipids. Two experiments were conducted: 1) the first to determine the optimal composition and concentration of liposomes to preserve post-thaw motility and membrane integrity of spermatozoa; 2) the second to assess in vivo the cryoprotective capacities of these ...
Standing ovariectomy in mares using a transvaginal natural orifice transluminal endoscopic surgery (NOTES®) approach.
Veterinary surgery : VS    September 7, 2011   Volume 40, Issue 8 987-997 doi: 10.1111/j.1532-950X.2011.00871.x
Pader K, Lescun TB, Freeman LJ.To develop and assess the feasibility of standing transvaginal Natural Orifice Transluminal Endoscopic Surgery (NOTES®) ovariectomy in the mare. Methods: Descriptive study. Methods: Mares (n=10). Methods: The technique was developed in 6 mares and then evaluated in a short-term study in 4 mares. Abdominal access was developed using controlled access vaginal cannula placement under endoscopic guidance. With viewing provided by a flexible endoscope, hemostasis, and transection of ovarian pedicles was performed using a customized bipolar vessel-sealing device. The ovaries were retrieved and the ...
Concentrations of circulating hormones during the interval between pulses of a PGF2α metabolite in mares and heifers.
Animal reproduction science    September 6, 2011   Volume 128, Issue 1-4 22-28 doi: 10.1016/j.anireprosci.2011.08.009
Ginther OJ, Rodriguez MB, Beg MA.The temporal relationship of several hormones to a metabolite of prostaglandin F2α (PGFM) was studied in mares and heifers from the beginning of the first PGFM pulse during luteolysis to the end of the second pulse. Mares (n=7) were selected with a 9-h interval between the peaks of the two pulses. In mares, estradiol-17β (estradiol) increased (P<0.05) within each PGFM pulse and plateaued for a mean of 6h between the pulses, resulting in a stepwise estradiol increase. Progesterone decreased linearly (P<0.0001) throughout the intra-pulse and inter-pulse intervals of PGFM. In heifers (n=6...
What is your diagnosis? Unilateral ovarian mass in a mare.
Veterinary clinical pathology    September 3, 2011   Volume 40, Issue 3 399-400 doi: 10.1111/j.1939-165X.2011.00345.x
Fernandes TR, Grandi F, Monteiro LN, Salgado BS, Rocha NS.No abstract available
Progesterone production in mares and echographic evaluation of the corpora lutea formed after follicular aspiration.
Reproduction in domestic animals = Zuchthygiene    August 30, 2011   Volume 47, Issue 2 288-292 doi: 10.1111/j.1439-0531.2011.01856.x
Mozzaquatro FD, Verstegen JP, Douglas RH, Troedsson MH, Delacorte FD, Silva CA, Rubin MI.Ultrasound-guided follicular aspiration was performed in 26 Criollo crossbred mares, followed by the evaluation of ultrasonographic images of the Corpus luteum (CL) that was formed after puncture of follicles of different diameters (Group 25-29 mm; Group 30-35 mm and Group >35 mm). Serum progesterone (P(4) ) concentrations were measured to determine CL function. The size of the CL was measured and the CL was classified based on the following echoscore: 1- anechoic tissue; 2- poorly defined luteal structure with low echogenicity; 3- echogenicity analogous to a luteal structure. The proport...
Artificial insemination and embryo transfer in mares.
Compendium (Yardley, PA)    August 27, 2011   Volume 33, Issue 7 E1-E5 
Scherzer J.Mares can be artificially inseminated with chilled or frozen semen to increase the revenue from their offspring. Embryo transfer can be used to produce more than one foal from a single mare per season. Recent advances in using equine follicle-stimulating hormone to induce superovulation in mares have stimulated research on preserving equine embryos. Equine embryos are usually collected on day 7 or 8 after ovulation, and younger (day 6.5) embryos are typically cryopreserved. Cryopreservation improves the ability of veterinary clinicians to preserve embryos for implantation in recipient mares an...
Temporal relationships of a pulse of prolactin (PRL) to a pulse of a metabolite of PGF2α in mares.
Theriogenology    August 26, 2011   Volume 77, Issue 1 99-107 doi: 10.1016/j.theriogenology.2011.07.021
Ginther OJ, Pinaffi FL, Silva LA, Beg MA.Hourly blood samples were collected from 10 mares during 24 h of each of the preluteolytic, luteolytic, and postluteolytic periods. The autocorrelation function of the R program was used to detect pulse rhythmicity, and the intra-assay CV was used to locate and characterize pulses of prolactin (PRL) and a metabolite of prostaglandin F2α (PGFM). Rhythmicity of PRL and PGFM concentrations was detected in 67% and 89% of mares, respectively. Combined for the three periods (no difference among periods), the PRL pulses were 5.2±0.4 h (mean±SEM) at the base, 7.5±1.5 h between nadirs of adjacent p...
Comparison of the cytobrush, cottonswab, and low-volume uterine flush techniques to evaluate endometrial cytology for diagnosing endometritis in chronically infertile mares.
Theriogenology    August 23, 2011   Volume 77, Issue 1 89-98 doi: 10.1016/j.theriogenology.2011.07.020
Cocchia N, Paciello O, Auletta L, Uccello V, Silvestro L, Mallardo K, Paraggio G, Pasolini MP.Endometritis is the most important cause of infertility in barren mares. The quick method of endometrial cytology (EC) has a relatively high reliability in diagnosing endometrial inflammation in the mare. For reliable cytological results, a collection technique that yields many well-preserved cells representative of a large uterine surface area without causing harm to the reproductive tract is required. The aim of the study was to compare three usually employed techniques for collection of endometrial and inflammatory cells (guarded cotton swab, uterine lavage, and cytobrush) in chronically in...
Morpho-functional studies regarding the fertility prognosis of mares suffering from equine endometrosis.
Theriogenology    August 23, 2011   Volume 76, Issue 7 1326-1336 doi: 10.1016/j.theriogenology.2011.06.001
Lehmann J, Ellenberger C, Hoffmann C, Bazer FW, Klug J, Allen WR, Sieme H, Schoon HA.The aim of the present study was to characterize the morpho-functional features of endometrosis in barren and foaling mares, using both conventional histopathological and immunohistochemical methods. Endometrial biopsy samples were collected during the physiological breeding season from 159 estrous, clinically healthy mares (mean age 12 years), and the quality and degree of endometrosis was histomorphologically defined. The mares were bred and those that foaled were put in the foaling group whereas those that did not foal were placed in the barren group. Foaling mares were then compared with b...
Production of live foals via intracytoplasmic injection of lyophilized sperm and sperm extract in the horse.
Reproduction (Cambridge, England)    August 16, 2011   Volume 142, Issue 4 529-538 doi: 10.1530/REP-11-0145
Choi YH, Varner DD, Love CC, Hartman DL, Hinrichs K.Work with lyophilized sperm helps delineate the factors required for successful fertilization. We investigated the use of lyophilized sperm in equine embryo production. In Experiment 1, sperm DNA fragmentation index was not affected by three freeze/thaw or lyophilization cycles. In Experiment 2, oocytes injected with lyophilized sperm or with sperm from a treatment in which lyophilized sperm were suspended in sperm cytoplasmic extract (SE) yielded blastocyst development rates of 0 and 28% respectively (P < 0.05). In Experiment 3, blastocyst development rate was 6-11% after injection of sper...
Administration of a gonadotropin-releasing hormone antagonist to mares at different times during the luteal phase of the estrous cycle.
Animal reproduction science    August 16, 2011   Volume 127, Issue 3-4 188-196 doi: 10.1016/j.anireprosci.2011.07.020
Evans MJ, Alexander SL, Irvine CH, Kitson NE, Taylor TB.The GnRH antagonist cetrorelix was given during the early (Days 1-5), mid (Days 6-10 or 5-12) or for the entire (Days 1-16) luteal phase of mares to inhibit the secretion of FSH and LH (Day 0=ovulation). Frequent blood sampling from Day 6 to Day 14 was used to determine the precise time-course of the suppression (cetrorelix given Days 6-10). Cetrorelix treatment caused a decrease in FSH and LH concentrations by 8 and 16 h, respectively, and an obliteration of the response to exogenous GnRH given 24h after treatment onset. Treatment never suppressed gonadotropin concentrations to undetectable l...
Relationship between progesterone, oestrone sulphate and cortisol and the components of renin angiotensin aldosterone system in Spanish purebred broodmares during pregnancy.
Theriogenology    August 10, 2011   Volume 76, Issue 8 1404-1415 doi: 10.1016/j.theriogenology.2011.06.009
Satué K, Domingo R, Redondo JI.The coordinated interaction of the components of the renin angiotensin aldosterone system (RAAS) with reproductive hormones such as progesterone, oestrogens and cortisol during pregnancy has been widely reported to play a vital role in foetal and placental development in various species, significantly influencing the proper achievement of pregnancy and foetal viability at birth. These interactions have not yet been clarified in mares. Thus, the purpose of the present research was to analyse the relationship between cortisol (CORT), progesterone (P4) and oestrone sulphate (OESTRONE), and the co...
Dissecting the molecular damage to stallion spermatozoa: the way to improve current cryopreservation protocols?
Theriogenology    August 10, 2011   Volume 76, Issue 7 1177-1186 doi: 10.1016/j.theriogenology.2011.06.023
Peña FJ, García BM, Samper JC, Aparicio IM, Tapia JA, Ferrusola CO.We review recent developments in the technology of freezing stallion sperm, paying special attention to the molecular lesions that spermatozoa suffer during freezing and thawing, such as osmotic stress, oxidative damage, and apoptotic changes. We also discuss the applicability of colloidal centrifugation in stallion sperm cryobiology. Increased knowledge about the molecular injuries that occur during cryopreservation may lead to improved protective techniques and thus to further improvements in fertility in the current decade.
Evaluation of two magnetic-bead-based viral nucleic acid purification kits and three real-time reverse transcription-PCR reagent systems in two TaqMan assays for equine arteritis virus detection in semen.
Journal of clinical microbiology    August 10, 2011   Volume 49, Issue 10 3694-3696 doi: 10.1128/JCM.01187-11
Miszczak F, Shuck KM, Lu Z, Go YY, Zhang J, Sells S, Vabret A, Pronost S, Fortier G, Timoney PJ, Balasuriya UB.This study showed that under specifically defined conditions with respect to nucleic acid extraction method and testing reagents, a previously described real-time reverse transcription-PCR (rRT-PCR) assay (T1 assay) provides sensitivity equal to or higher than that of virus isolation for the detection of equine arteritis virus in semen.
Review of seven cases of granulosa cell tumour of the equine ovary.
The Veterinary record    August 10, 2011   Volume 169, Issue 10 251 doi: 10.1136/vr.d4635
Crabtree J.This paper reviews the equine granulosa cell tumour (GCT) and describes the clinicopathological features, treatment and outcome in seven cases of GCT in mares. Mares were presented with unilateral ovarian enlargement during the 2007 to 2010 breeding seasons. The mean (sd) age of the mares was 11.7 (5.96) years. Three mares were multiparous barren, three were nulliparous and one was primigravida. Behaviour at presentation was 57 per cent anoestrus, 28 per cent with stallion-like behaviour and 14 per cent with persistent oestrus. All mares had unilateral ovarian enlargement. Six non-pregnant mar...
Concurrent lactation and pregnancy: pregnant domestic horse mares do not increase mother-offspring conflict during intensive lactation.
PloS one    August 10, 2011   Volume 6, Issue 8 e22068 doi: 10.1371/journal.pone.0022068
Bartošová J, Komárková M, Dubcová J, Bartoš L, Pluháček J.Lactation is the most energy demanding part of parental care in mammals, so parent-offspring conflict arises over milk provided by the mother. In some species females commonly become pregnant shortly after parturition of previous young. This further intensifies mother-offspring conflict due to concurrent pregnancy and lactation. In equids it has been well established that pregnant females wean their foals earlier than non-pregnant ones. Intensified mother-offspring conflict was presumed to associate with pregnancy also during the period of intensive lactation, i.e., before the weaning process ...
G6PDH-activity in equine oocytes correlates with morphology, expression of candidate genes for viability, and preimplantative in vitro development.
Theriogenology    August 5, 2011   Volume 76, Issue 7 1215-1226 doi: 10.1016/j.theriogenology.2011.05.025
Mohammadi-Sangcheshmeh A, Held E, Ghanem N, Rings F, Salilew-Wondim D, Tesfaye D, Sieme H, Schellander K, Hoelker M.Efficiencies for in vitro production of equine embryos are still low due to highly variable developmental competences of equine immature oocytes. In contrast to the equine, in vitro developmental competence of immature oocytes has been predicted successfully by the activity of glucose-6-phosphate dehydrogenase (G6PDH) indicated by brilliant cresyl blue (BCB) dye in a range of different species. Therefore, the aim of the present study was to test the association between G6PDH activity in equine oocytes with: (1) cumulus morphology and oocyte properties in terms of diameter and volume; (2) matur...
An update on cyathostomins: anthelmintic resistance and diagnostic tools.
Equine veterinary journal. Supplement    August 4, 2011   Issue 39 133-139 doi: 10.1111/j.2042-3306.2011.00397.x
Stratford CH, McGorum BC, Pickles KJ, Matthews JB.Cyathostomins represent a potential cause of equine morbidity and have become the main focus of endoparasite control in managed horses. All grazing horses are at risk of infection with cyathostomins; therefore, the application of appropriate management measures is essential. Anthelmintics currently comprise the main method of control for equine nematodes and the ready availability of these products in some countries has resulted in their use becoming dissociated from veterinary involvement. This is concerning given the levels of anthelmintic resistance that have been recorded in cyathostomin p...
The additional N-glycosylation site of the equine LH/CG receptor is not responsible for the limited cyclic AMP pathway activation by equine chorionic gonadotropin relative to luteinizing hormone.
Reproductive biology    August 2, 2011   Volume 11, Issue 2 157-164 doi: 10.1016/s1642-431x(12)60052-7
Saint-Dizier M, Foulon-Gauze F, Lecompte F, Combarnous Y, Chopineau M.In order to investigate the role of the unique seventh N23-glycosylation site of the equine LH/CG receptor (eLHCGR) in the cAMP pathway activation, COS-7 cells were transiently transfected with either the wild-type or the mutant eLHCGR(N23Q) cDNA and challenged with porcine LH and eCG for cAMP production. We showed that the N23-glycosylation site of the eLHCGR is not required for the functional coupling of the receptor with the cAMP pathway and is not responsible for the limited potency of eCG relative to pLH to activate this receptor.
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