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Topic:Testes

The testes in horses are reproductive organs responsible for the production of sperm and the secretion of hormones such as testosterone. Located in the scrotum, the testes play a vital role in male fertility and reproductive behavior. They consist of seminiferous tubules where spermatogenesis occurs, and interstitial cells, also known as Leydig cells, which produce testosterone. The function and health of equine testes can be influenced by factors such as age, season, and overall health status. This page compiles peer-reviewed research studies and scholarly articles that explore the anatomy, physiology, and pathology of the testes in horses, providing insights into their role in equine reproduction and breeding management.
Dexamethasone downregulates expression of several genes encoding orphan nuclear receptors that are important to steroidogenesis in stallion testes.
Journal of biochemical and molecular toxicology    February 24, 2019   Volume 33, Issue 6 e22309 doi: 10.1002/jbt.22309
Valdez R, Cavinder CA, Varner DD, Welsh TH, Vogelsang MM, Ing NH.Glucocorticoids impair testosterone synthesis by an unknown mechanism. Stallions treated with the synthetic glucocorticoid dexamethasone had testes collected at 6 or 12 hours postinjection. The testicular expression of selected genes encoding nuclear receptors and steroidogenic enzymes was measured. At 6 hours, dexamethasone treatment decreased levels of NR0B2, NR4A1, NR5A1, and NR5A2 messenger RNAs (mRNAs) and NR5A2 mRNA levels remained depressed at 12 hours. In contrast, dexamethasone increased levels of NFKBIA mRNA at both time points. At 6 hours, dexamethasone did not alter levels ...
The serine protease testisin is present on the surface of capacitated stallion spermatozoa and interacts with key zona pellucida binding proteins.
Andrology    December 13, 2018   Volume 7, Issue 2 199-212 doi: 10.1111/andr.12569
Swegen A, Smith ND, Gibb Z, Curry BJ, Aitken RJ.Serine proteases are emerging as important players in the spermatozoon's acquisition of functional competence. This study aimed to characterize the serine protease testisin (PRSS21) in stallion spermatozoa, examining its surface expression, possible origins in the testis and epididymis, and changes in response to capacitation and acrosome reaction, as well as its capacity to form high molecular weight complexes and interact with other proteins. The role of serine proteases in spontaneous capacitation and acrosome reaction of stallion spermatozoa was established using the serine protease inhibi...
Inhibin-A and Inhibin-B in stallions: Seasonal changes and changes after down-regulation of the hypothalamic-pituitary-gonadal axis.
Theriogenology    September 27, 2018   Volume 123 108-115 doi: 10.1016/j.theriogenology.2018.09.036
Ball BA, Davolli GM, Esteller-Vico A, Fleming BO, Wynn MAA, Conley AJ.The biological function of inhibin is mediated by two heterodimers, inhibin-A and inhibin-B. The relative importance of inhibin-A and -B in male reproductive function varies considerably across species with inhibin-B predominating in many species, whereas inhibin-A appears relatively more important in rams. Research reported to date in stallions has examined total or immunoreactive (ir) inhibin which does not distinguish the two heterodimers. Therefore, the objective of this study was to characterize changes in inhibin-A and inhibin-B concentrations in stallions: 1) across season for a period ...
Determination of the activity and relative abundance of mRNA for antioxidant enzymes in stallion testicular and epididymal tissues: A comparison between two breeding seasons.
Animal reproduction science    August 11, 2018   Volume 196 230-238 doi: 10.1016/j.anireprosci.2018.08.015
Koziorowska-Gilun M, Gilun P, Mietelska K, Kordan W.The key prerequisite for successful insemination is sperm characterized to have positive values for morphological and biological variables which are determined by, among others, effective antioxidant protection during the lifespan of sperm cells. This study evaluated the activity and relative abundance of mRNA for antioxidant enzymes in stallion testicular and epididymal tissues during breeding (n = 5) and non-breeding (n = 5) seasons. The activity of superoxide dismutase (SOD) was greater (P < 0.05) during the breeding season, in particular in the testes and the caput epididymi...
Horse Y chromosome assembly displays unique evolutionary features and putative stallion fertility genes.
Nature communications    July 27, 2018   Volume 9, Issue 1 2945 doi: 10.1038/s41467-018-05290-6
Janečka JE, Davis BW, Ghosh S, Paria N, Das PJ, Orlando L, Schubert M, Nielsen MK, Stout TAE, Brashear W, Li G, Johnson CD, Metz RP, Zadjali AMA....Dynamic evolutionary processes and complex structure make the Y chromosome among the most diverse and least understood regions in mammalian genomes. Here, we present an annotated assembly of the male specific region of the horse Y chromosome (eMSY), representing the first comprehensive Y assembly in odd-toed ungulates. The eMSY comprises single-copy, equine specific multi-copy, PAR transposed, and novel ampliconic sequence classes. The eMSY gene density approaches that of autosomes with the highest number of retained X-Y gametologs recorded in eutherians, in addition to novel Y-born and transp...
Source and localization of ovulation-inducing factor/nerve growth factor in male reproductive tissues among mammalian species.
Biology of reproduction    July 10, 2018   Volume 99, Issue 6 1194-1204 doi: 10.1093/biolre/ioy149
Bogle OA, Carrasco RA, Ratto MH, Singh J, Adams GP.The objectives of the study were to compare the presence and localization of ovulation-inducing factor (OIF)/nerve growth factor (NGF) in male reproductive organs and determine the abundance in ejaculates of species representative of both spontaneous and induced ovulators. We hypothesized that the protein is a widely conserved component of semen among mammals, but is most abundant in camelids. Immunohistochemical analysis was performed on tissues from the male reproductive system of llamas, rats, cattle, bison, elk, and white-tailed deer (n = 2 males/species), and the abundance of OIF/NGF in...
Equine Chorionic Gonadotropin (eCG) improves bucks’ semen quality during the nonbreeding season.
Reproduction in domestic animals = Zuchthygiene    June 8, 2018   Volume 53, Issue 5 1096-1102 doi: 10.1111/rda.13209
Beracochea F, Viera MN, Acevedo L, Santiago-Moreno J, Ungerfeld R.In most goat breeds, testosterone serum concentration and semen quality decrease during the nonbreeding season. However, bucks reproductive activity may be stimulated with the administration of equine chorionic gonadotropin (eCG). Therefore, the aim of this study was to determine whether the repeated administration of eCG stimulates the reproductive status of bucks during the nonbreeding season. The study was performed with 19 bucks that were assigned to a group that was treated with eCG (GeCG) and an untreated control group (GCon). The GeCG bucks received an initial dose of 800 IU of eCG (Da...
Effects of implants containing the GnRH agonist deslorelin on testosterone release and semen characteristics in Shetland stallions.
Animal reproduction science    May 26, 2018   Volume 195 230-241 doi: 10.1016/j.anireprosci.2018.05.027
Gautier C, Schmidt K, Aurich J, Aurich C.The hypothesis in this study was continuous treatment of stallions with the GnRH agonist deslorelin inhibits reproductive functions. A 2-week pre-experimental period was followed by an 11-week deslorelin implant treatment. Stallions received 4.7 (D1, n = 7), or 18.8 mg deslorelin (D2, n = 5) or remained untreated (C, n = 5). Libido, sperm motility, membrane integrity, DNA fragmentation, estrogen receptors, basal plasma testosterone and Anti Muellerian hormone (AMH) concentrations were evaluated once weekly during the treatment period. The testosterone response to the GnRH agonist buserelin a...
Anti-Müllerian hormone as a biomarker for acute testicular degeneration caused by toxic insults to stallion testes.
Theriogenology    May 17, 2018   Volume 116 95-102 doi: 10.1016/j.theriogenology.2018.05.009
Pozor M, Conley AJ, Roser JF, Nolin M, Zambrano GL, Runyon SP, Kelleman AA, Macpherson ML.Recently, anti-Müllerian hormone (AMH) was validated as a reliable marker of testicular damage caused by various chemotherapy drugs in humans and in mice. In horses, the reference values of AMH concentrations in normal stallions, during different seasons of a year, have been recently reported. However, this hormone was not evaluated in subfertile or infertile stallions with testicular damage. Therefore, the objective of this study was to investigate the effects of experimentally induced testicular degeneration on the concentration of AMH in stallions. Severe but transient testicular degenerat...
Anti-Müllerian hormone profiling in prepubertal horses and its relationship with gonadal function.
Theriogenology    May 15, 2018   Volume 117 72-77 doi: 10.1016/j.theriogenology.2018.05.012
Scarlet D, Wulf M, Kuhl J, Köhne M, Ille N, Conley AJ, Aurich C.Anti-Müllerian hormone (AMH) has gained increasing interest as a biomarker for assessment of gonadal activity. The ability to predict the ovarian follicular reserve of prepubertal female horses (fillies) or to identify stallions with testicular pathologies already during their prepubertal life has not been analyzed so far. Both would help to select fertile horses and reduce costs associated with keeping animals. The objectives of the present study were to (1) assess AMH, LH, FSH, progesterone (females) and testosterone (males) dynamics in prepubertal horses from birth onwards and (2) determin...
Endocrine and metabolic profile of peripubertal Standardbred colts.
Theriogenology    May 14, 2018   Volume 117 78-84 doi: 10.1016/j.theriogenology.2018.05.006
Ellerbrock RE, Canisso IF, Lima FS, Shipley CF, Thompson DL, Conley AJ, Kline KH.The objectives of this study were to determine the concentrations of reproductive and metabolic hormones during the peripubertal period and to assess their relationship with testicular development and body fat deposition. Blood samples were collected from 23 healthy Standardbred colts every four weeks for twelve months. Colts were weighed monthly, and percent of body fat and testicular volume estimated by ultrasound. Onset of puberty was determined as the month when testosterone was two standard deviations above the previous mean. Plasma FSH, LH, leptin, estradiol-17β, androstenedione, IGF-1,...
Blastocyst production after intracytoplasmic sperm injection with semen from a stallion with testicular degeneration.
Reproduction in domestic animals = Zuchthygiene    March 1, 2018   Volume 53, Issue 3 814-817 doi: 10.1111/rda.13153
Roels K, Smits K, Ververs C, Govaere J, D'Herde K, Van Soom A.In horse breeding, intracytoplasmic sperm injection (ICSI) has gained interest to obtain offspring from subfertile individuals. This paper presents a case report of a stallion with severe testicular degeneration. Semen analysis showed very low motility and 83.5% of detached heads. Histology of a testicular biopsy showed severely decreased spermatogenesis, while transmission electron microscopy of the sperm cells revealed no significant abnormalities. A total of 39 oocytes were fertilized by ICSI with frozen-thawed spermatozoa of this stallion: 25 oocytes with intact spermatozoa and 24 with det...
What Is Your Diagnosis? Testicular tumor in a horse.
Veterinary clinical pathology    February 14, 2018   Volume 47, Issue 1 166-167 doi: 10.1111/vcp.12577
Leidinger E, Springler G, Furman E, Wallner A.No abstract available
Maternal obesity increases insulin resistance, low-grade inflammation and osteochondrosis lesions in foals and yearlings until 18 months of age.
PloS one    January 26, 2018   Volume 13, Issue 1 e0190309 doi: 10.1371/journal.pone.0190309
Robles M, Nouveau E, Gautier C, Mendoza L, Dubois C, Dahirel M, Lagofun B, Aubrière MC, Lejeune JP, Caudron I, Guenon I, Viguié C, Wimel L....Obesity is a growing concern in horses. The effects of maternal obesity on maternal metabolism and low-grade inflammation during pregnancy, as well as offspring growth, metabolism, low-grade inflammation, testicular maturation and osteochondrotic lesions until 18 months of age were investigated. Twenty-four mares were used and separated into two groups at insemination according to body condition score (BCS): Normal (N, n = 10, BCS ≤4) and Obese (O, n = 14, BCS ≥4.25). BCS and plasma glucose, insulin, triglyceride, urea, non-esterified fatty acid, serum amyloid A (SAA), leptin and adiponect...
Maternal parity affects placental development, growth and metabolism of foals until 1 year and a half.
Theriogenology    December 21, 2017   Volume 108 321-330 doi: 10.1016/j.theriogenology.2017.12.019
Robles M, Dubois C, Gautier C, Dahirel M, Guenon I, Bouraima-Lelong H, Viguié C, Wimel L, Couturier-Tarrade A, Chavatte-Palmer P.Primiparous mares are known to produce smaller foals than multiparous mares. This difference seems to be partly explained by the reduced exchange surface and volume of the placental villi in primiparous compared to multiparous placentas. The effect of maternal parity on foals' post-natal growth, metabolism and sexual maturation, however, has been given little consideration. The objectives of this work were to analyse placental biometry and structure at term, growth of foals and yearlings, their metabolism and testicular maturation at one year of age. Twenty multiparous mares (M), aged over 6 y...
Early histone H4 acetylation during chromatin remodeling in equine spermatogenesis.
Biology of reproduction    December 1, 2017   Volume 98, Issue 1 115-129 doi: 10.1093/biolre/iox159
Ketchum CC, Larsen CD, McNeil A, Meyer-Ficca ML, Meyer RG.Chromatin remodeling during spermatogenesis culminates in the exchange of nucleosomes for transition proteins and protamines as an important part of spermatid development to give rise to healthy sperm. Comparative immunofluorescence analyses of equine and murine testis histological sections were used to characterize nucleoprotein exchange in the stallion. Histone H4 hyperacetylation is considered a key event of histone removal during the nucleoprotein transition to a protamine-based sperm chromatin structure. In the stallion, but not the mouse, H4 was already highly acetylated in lysine residu...
Characterization of equine GST A3-3 as a steroid isomerase.
The Journal of steroid biochemistry and molecular biology    November 24, 2017   Volume 178 117-126 doi: 10.1016/j.jsbmb.2017.11.011
Lindström H, Peer SM, Ing NH, Mannervik B.Glutathione transferases (GSTs) comprise a superfamily of enzymes prominently involved in detoxication by making toxic electrophiles more polar and therefore more easily excretable. However some GSTs have developed alternative functions. Thus, a member of the Alpha class GSTs in pig and human tissues is involved in steroid hormone biosynthesis, catalyzing the obligatory double-bond isomerization of Δ-androstene-3,17-dione to Δ-androstene-3,17-dione and of Δ-pregnene-3,20-dione to Δ-pregnene-3,20-dione on the biosynthetic pathways to testosterone and progesterone. The human GST A3-3 is the ...
Morcellation for testes extraction in horses undergoing standing laparoscopic cryptorchidectomy.
The Canadian veterinary journal = La revue veterinaire canadienne    November 2, 2017   Volume 58, Issue 11 1215-1220 
Sassot LN, Ragle CA, Farnsworth KD, Lund CM.The major objective of this study was to describe the use of morcellation for standing laparoscopic cryptorchidectomy in 30 client-owned horses. A second objective was to describe a laparoscopic-assisted standing scrotal technique for removal of a descended testis in unilateral cryptorchids. Morcellation for extraction of abdominal testes is effective, efficient, and well-tolerated. Morcellation enabled safe and straightforward extraction of testes from the abdomen through an approximately 13- to 20-mm incision using a 2-portal technique. No incisional or morcellator-related complications occu...
Equine fetal adrenal, gonadal and placental steroidogenesis.
Reproduction (Cambridge, England)    September 8, 2017   Volume 154, Issue 4 445-454 doi: 10.1530/REP-17-0239
Legacki EL, Ball BA, Corbin CJ, Loux SC, Scoggin KE, Stanley SD, Conley AJ.Equine fetuses have substantial circulating pregnenolone concentrations and thus have been postulated to provide significant substrate for placental 5α-reduced pregnane production, but the fetal site of pregnenolone synthesis remains unclear. The current studies investigated steroid concentrations in blood, adrenal glands, gonads and placenta from fetuses (4, 6, 9 and 10 months of gestational age (GA)), as well as tissue steroidogenic enzyme transcript levels. Pregnenolone and dehydroepiandrosterone (DHEA) were the most abundant steroids in fetal blood, pregnenolone was consistently higher bu...
Horse spermatogonial stem cell cryopreservation: feasible protocols and potential biotechnological applications.
Cell and tissue research    August 22, 2017   Volume 370, Issue 3 489-500 doi: 10.1007/s00441-017-2673-1
Costa GMJ, Avelar GF, Lacerda SMSN, Figueiredo AFA, Tavares AO, Rezende-Neto JV, Martins FGP, França LR.The establishment of proper conditions for spermatogonial stem cells (SSCs) cryopreservation and storage represents an important biotechnological approach for the preservation of the genetic stock of valuable animals. This study demonstrates the effects of different cryopreservation protocols on the survival rates and phenotypic expression of SSCs in horses. The cells were enzymatically isolated from testes of eight adult horses. After enrichment and characterization of germ cells in the suspension, the feasibility of several cryopreservation protocols were evaluated. Three different cryomedia...
Testicular volumetry and prediction of daily sperm output in stallions by orchidometry and two- and three-dimensional sonography.
Theriogenology    August 19, 2017   Volume 104 149-155 doi: 10.1016/j.theriogenology.2017.08.015
Pricking S, Bollwein H, Spilker K, Martinsson G, Schweizer A, Thomas S, Oldenhof H, Sieme H.Accurate determination of the testes volume and prediction of the daily sperm output (DSO) is valuable information for reproductive management of a stallion. The aim of this study was to compare different methods for measuring the testes volume, including caliper, 2D and 3D ultrasound. Special emphasis was on feasibility of 3D volume analysis. First, 22 castrated testes were measured and derived volumes were compared with volumes determined via volume displacement in a graded cylinder with saline solution. Then, during the breeding season, testes sizes of 52 stallions were measured in vivo an...
Relationship between echotextural and histomorphometric characteristics of stallion testes.
Theriogenology    June 1, 2017   Volume 99 134-145 doi: 10.1016/j.theriogenology.2017.05.031
Pozor M, Morrissey H, Albanese V, Khouzam N, Deriberprey A, Macpherson ML, Kelleman AA.The goal of this study was to investigate correlations between objective measures of testicular echotexture and histomorphometric attributes related to the histological composition of stallion testes. Fifty-four scrotal testes were obtained from three groups of stallions during routine castrations: colts  5 yrs old (n = 9). In addition, two scrotal testes with degeneration, 16 retained inguinal and 10 retained abdominal testes were surgically obtained. Cross-sectional and longitudinal ultrasonograms were obtained for each testis. Mean numerical pixel values (NPVs) as well as pixel standard...
Pulse Doppler ultrasound as a tool for the diagnosis of chronic testicular dysfunction in stallions.
PloS one    May 30, 2017   Volume 12, Issue 5 e0175878 doi: 10.1371/journal.pone.0175878
Ortiz-Rodriguez JM, Anel-Lopez L, Martín-Muñoz P, Álvarez M, Gaitskell-Phillips G, Anel L, Rodríguez-Medina P, Peña FJ, Ortega Ferrusola C.Testicular function is particularly susceptible to vascular insult, resulting in a negative impact on sperm production and quality of the ejaculate. A prompt diagnosis of testicular dysfunction enables implementation of appropriate treatment, hence improving fertility forecasts for stallions. The present research aims to: (1) assess if Doppler ultrasonography is a good tool to diagnose stallions with testicular dysfunction; (2) to study the relationship between Doppler parameters of the testicular artery and those of sperm quality assessed by flow cytometry and (3) to establish cut off values ...
Castration in the standing horse combining laparoscopic and conventional techniques.
Equine veterinary journal    April 19, 2017   Volume 49, Issue 6 776-779 doi: 10.1111/evj.12681
Rijkenhuizen ABM, van der Harst MR.Standing laparoscopic castration without orchidectomy has been shown to be a reliable method of castration in animals with abdominal cryptorchidism, but it cannot be recommended as a method of castration in those with inguinal cryptorchidism or normal males. An alternative laparoscopic method for the castration of horses with descended testes was developed. Objective: To report the results of laparoscopy-assisted abdominal transection of the spermatic cord and a modified open method in the standing horse for castration of descended testes in unilateral abdominal cryptorchids. Methods: Descript...
Laparoscopic cryptorchidectomy with a vessel-sealing device in dorsal recumbent horses: 43 cases.
Veterinary surgery : VS    March 3, 2017   Volume 46, Issue 4 559-565 doi: 10.1111/vsu.12624
Bracamonte JL, Thomas KL.To document laparoscopic cryptorchidectomy with a vessel-sealing device in dorsal recumbent horses. Methods: Retrospective study. Methods: Forty-three male horses. Methods: Food was withheld for 36 hours before horses were placed under general anesthesia and positioned in dorsal recumbency. One laparoscopic portal and 2 instrument portals were placed for the laparoscopic procedure. A third instrument portal was created in bilateral cryptorchid horses. A vessel-sealing device was used for hemostasis and transection of retained testes. Descended testes were removed via closed castration with pri...
Androgen Insensitivity Syndrome in a Family of Warmblood Horses Caused by a 25-bp Deletion of the DNA-Binding Domain of the Androgen Receptor Gene. Welsford GE, Munk R, Villagómez DA, Hyttel P, King WA, Revay T.Testicular feminization, an earlier term coined for describing a syndrome resulting from failure of masculinization of target organs by androgen secretions during embryo development, has been well documented not only in humans but also in the domestic horse. The pathology, actually referred to as androgen insensitivity syndrome (AIS), has been proposed to follow an X-linked recessive pattern of inheritance in some horse breeds already investigated. Affected individuals are characterized by a female phenotype but with a stallion genotype of 64,XY SRY+ constitution. We identified a Warmblood hor...
Xenografting of isolated equine (Equus caballus) testis cells results in de novo morphogenesis of seminiferous tubules but not spermatogenesis.
Andrology    February 3, 2017   Volume 5, Issue 2 336-346 doi: 10.1111/andr.12308
Zeng W, Alpaugh W, Stefanovski D, Schlingmann K, Dobrinski I, Turner RM.The study of spermatogenesis in the horse is challenging because of the absence of an in vitro system that is capable of reproducing efficient spermatogenesis and because of the difficulties and costs associated with performing well-controlled studies in vivo. In an attempt to develop novel methods for the study of equine spermatogenesis, we tested whether cells from enzymatically digested pre-pubertal equine testicular tissue were capable of de novo tissue formation and spermatogenesis following xenografting under the back skin of immunocompromised mice. Testes were obtained from normal pre...
Testosterone serum profile, semen characteristics and testicular biometry of Mangalarga Marchador stallions in a tropical environment.
Reproduction in domestic animals = Zuchthygiene    January 23, 2017   Volume 52, Issue 2 335-343 doi: 10.1111/rda.12918
Waddington B, Penitente-Filho JM, Neves J, Pinho RO, Chaya AY, Maitan PP, Silveira CO, Neves MG, Guimarães S, de Carvalho GR, Guimarães JD.This study was conducted to characterize the daily profile of testosterone secretion and its mean concentrations in the four seasons as well as to evaluate the semen characteristics and testicular biometry of Mangalarga Marchador stallions throughout the year in a tropical region. Three stallions were submitted to semen collections and evaluation of testicular biometry every 14 days along a year. Blood samples were collected once at the middle of each season, in a 20-min interval during 24 hr in order to evaluate the testosterone secretion profiles among seasons. Testosterone concentrations ...
Undifferentiated embryonic cell transcription factor 1 (UTF1) and deleted in azoospermia-like (DAZL) expression in the testes of donkeys.
Reproduction in domestic animals = Zuchthygiene    January 20, 2017   Volume 52, Issue 2 264-269 doi: 10.1111/rda.12889
Lee YS, Jung HJ, Yoon MJ.Putative markers for each specific germ cell stage can be a useful tool to study the fate and functions of these cells. Undifferentiated embryonic cell transcription factor 1 (UTF1) is a putative marker for undifferentiated spermatogonia in humans, rats and horses. The deleted in azoospermia-like (DAZL) protein is also expressed by differentiated spermatogonia and primary spermatocytes in several species. However, whether the expression patterns of these molecular markers are identical and applicable to donkeys remains to be elucidated. The objective of this study was to investigate the expres...
Maternal Nutrition during Pregnancy Affects Testicular and Bone Development, Glucose Metabolism and Response to Overnutrition in Weaned Horses Up to Two Years.
PloS one    January 12, 2017   Volume 12, Issue 1 e0169295 doi: 10.1371/journal.pone.0169295
Robles M, Gautier C, Mendoza L, Peugnet P, Dubois C, Dahirel M, Lejeune JP, Caudron I, Guenon I, Camous S, Tarrade A, Wimel L, Serteyn D....Pregnant mares and post-weaning foals are often fed concentrates rich in soluble carbohydrates, together with forage. Recent studies suggest that the use of concentrates is linked to alterations of metabolism and the development of osteochondrosis in foals. The aim of this study was to determine if broodmare diet during gestation affects metabolism, osteoarticular status and growth of yearlings overfed from 20 to 24 months of age and/or sexual maturity in prepubertal colts. Methods: Twenty-four saddlebred mares were fed forage only (n = 12, group F) or cracked barley and forage (n = 12, group ...