Fetal development in horses refers to the progression of growth and differentiation that occurs from conception until birth. This process involves a series of complex stages, beginning with fertilization and continuing through embryonic and fetal stages. Key developmental milestones include organogenesis, skeletal formation, and maturation of physiological systems. The equine gestation period averages around 340 days, during which the fetus undergoes significant anatomical and functional changes. Understanding fetal development in horses is essential for optimizing breeding practices and ensuring the health of both mare and foal. This page compiles peer-reviewed research studies and scholarly articles that explore the stages, mechanisms, and factors influencing fetal development in equines.
Wilsher S, Allen WR.The normality of equine placentation is essential for fetal health and development. Substantial information exists on the gross morphological status of the placenta but few studies have addressed the problem of degenerative lesions that interfere with placental morphology and placental efficiency. Objective: Degenerative changes in the endometrium with increasing age and parity are reflected in the morphology and density of the placental microcotyledons. Objective: To assess placental efficiency on the basis of foal birthweight as a function of total microscopic area of fetomaternal contact. M...
Li X, Morris LH, Allen WR.This study investigated the basic conditions required for the production of horse embryos by the transfer of the nuclei of fetal and adult fibroblast cells to enucleated oocytes. Cumulus-oocyte complexes were recovered from abattoir ovaries and matured in vitro in groups of 20-30 for 28-30 h in tissue culture medium 199 containing 20% v:v fetal bovine serum in coculture with equine oviduct epithelial cells. Fetal fibroblast cells (FFC) were derived from a 32-day-old Thoroughbred x Pony fetus, and adult skin fibroblast cells (SFC) were obtained from subdermal biopsies recovered from a 4-yr-old ...
Allen WR, Wilsher S, Turnbull C, Stewart F, Ousey J, Rossdale PD, Fowden AL.The interacting influences of maternal size and fetal genotype on placental and fetal development in the mare were assessed by comparing conventional within-breed Thoroughbred (Tb-in-Tb, n = 7) and Pony (P-in-P, n = 7) control pregnancies established by artificial insemination (AI) with between-breed (Tb-in-P, n = 8; deprived in utero condition and P-in-Tb, n = 7; luxurious in utero condition) experimental pregnancies established by embryo transfer. All foals were born spontaneously and the mean (+/- SEM) duration of gestation in the two groups of control mares was significantly different (P <...
Allen WR, Wilsher S, Stewart F, Stewart F, Ousey J, Ousey J, Fowden A.Within-breed artificial insemination and between-breed embryo transfer were carried out in small pony (P) and large Thoroughbred (Tb) mares to create 4 types of horse pregnancy in which the fetus experienced spatial and nutritional deprivation (Tb-in-P; n=8), luxury (P-in-Tb; n=7) or normality (Tb-in-Tb; n=7 and P-in-P; n=7) in utero. Measurement of equine chorionic gonadotrophin (eCG), total conjugated oestrogens and progestagen concentrations in serial peripheral serum samples recovered from all the mares throughout gestation showed that the amount of eCG produced during the first half of ge...
Acker DA, Curran S, Bersu ET, Ginther OJ.To describe the gross and histologic changes that develop in the equine embryo proper (ie, the portion of the embryo that becomes the fetus) from days 17 to 40 after ovulation and to compare the external features of equine embryos with those of porcine, ovine, and human embryos. Methods: 34 embryos collected from mixed-breed pony mares. Methods: External features for each embryo proper, including length, number of branchial arches, growth of appendages, face and head features, and body features, were examined, using a dissecting microscope, for embryos collected on days 17 to 40. Internal feat...
Allen WR.The equine embryo takes 6 days to traverse the oviduct and, when it finally enters the uterus, it remains spherical in shape and moves continually throughout the uterine lumen until day 17 after ovulation to deliver its maternal recognition of pregnancy signal to the entire endometrium. Between day 25 and day 35 after ovulation, the trophoblast cells of a discrete annulate portion of the chorion multiply rapidly and acquire an invasive phenotype and, between day 36 and day 38, migrate deeply into the maternal endometrium to form the equine-unique endometrial protuberances known as endometrial ...
Wooding FB, Morgan G, Fowden AL, Allen WR.The placenta is the only channel for transport of nutrients to the conceptus and the fetal nutrient demands increase exponentially to term. The 9 kDa calcium binding protein (calbindin, 9CBP) and the iron binding protein uteroferrin (UF) are proving to be reliable markers for epithelia that mediate active transcellular calcium and iron transport and the glucose transporter proteins (GT1 and GT3) for glucose transport by facilitated diffusion. Light and electron microscope immunocytochemistry have been used on perfusion fixed resin embedded material to establish the distribution of 9CBP, UF, GT...
Enders AC, Liu IK.The free allantois and allantochorion of conceptuses from 17 mares between 20 and 90 days of gestation were examined to determine the manner in which the associated mesodermal derivatives differentiated. It was found that a robust basement membrane developed under the allantoic endoderm, and that this basement membrane was partially isolated from the vascular layer of the allantois by a mesothelial layer and an exocelom-like space. The exocelom-like space persisted until approximately the stage of villous formation, and remnants of the space persisted over larger allantoic vessels even later. ...
Betteridge KJ.The developmental changes in the equine conceptus, its maternal environment and their interaction during the first 4 weeks following fertilization are reviewed. Attention is drawn to species-specific events to show why the horse is such a valuable model in which to study early pregnancy.
Sharp DC.This paper will discuss development of the equine conceptus, especially from the perspective of the maternal environment in which it develops and to which it has considerable influence.
Soana S, Gnudi G, Bertoni G.The aim of this work was to study the ontogenetic process in teeth from their early appearance in the ossifying matrix of the mandible and maxilla, in different foetuses of scalar ages. Radiographic examinations of the skull and mandible hemisections were performed and the latero-medial (LM) and dorsoventral (DV) projections for the skull and mandible were analysed. A high-definition film-screen combination was used for this study. The exposure values ranged from 35 kV/6 mAs to 58 kV/10 mAs, according to the size of the skulls and their degree of ossification. The first dental germ observed wa...
Gerstenberg C, Allen WR, Stewart F.Placentation involves considerable growth and reorganization of both maternal and fetal tissues. In this investigation, immunohistochemical localization of the proliferation marker Ki-67 antigen was used to monitor cell division during placentation in mares. Endometrial biopsies were obtained from eight mares between day 14 and day 26 of pregnancy and from eight anoestrous mares that had been treated with various combinations of progesterone and oestrogen. Samples of endometrium and fetal membranes were obtained from 19 mares carrying normal horse conceptuses between day 30 and day 250 of gest...
Renaudin CD, Gillis CL, Tarantal AF, Coleman DA.Transrectal and transabdominal ultrasonography were performed on normal pregnant mares (n=10) at 2 week intervals from day 100 of gestation to parturition to evaluate fetal growth. Several fetal anatomical regions (head, eye, aorta, abdomen, rib, gonad, kidney and femur) were imaged and measured using standardized scan plans. The results of these analyses indicate that all of the biometric parameters correlate strongly with the day of gestation. Growth charts were developed, which demonstrate that the following variables have linear relationships with the day of gestation on which they were me...
Simpson KS, Adams MH, Behrendt-Adam CY, Baker CB, McDowell KJ.For development to proceed normally, the appropriate genes must be expressed in the correct tissues and in the correct time frame. Knowledge of gene expression during development provides information about the changes taking place within the conceptus as well as possible reasons for pregnancy failure. However, little is known about gene expression during development in the equine conceptus. In this study, we examined differences in gene expression between day 12 and day 15 equine conceptuses by suppression subtractive hybridization. This technique was used to isolate transcripts that are more ...
Bragulla H.The pre- and perinatal development of the hoof cartilage is described concerning the histological structure and surrounding vessels. Beginning in the third month of fetal development, the anlage of the hoof cartilage is still present in typical shape and location. It is built out of mesenchymal connective tissue. During further fetal development, the connective tissue cells will differentiate into two cell populations, fibroblasts and chondroblasts. Vessels, traversing the hoof cartilage, are surrounded by loose connective tissue, which will partially develop fibrocartilage. At birth, hoof car...
Bragulla H, Budras KD, Reilly JD.The fetal development of the white line (Zona alba) in the equine hoof is described. Its specific structure of lamellar and interlamellar horn, which in turn is composed of cap and terminal horn, is formed in the second half of the hoof's fetal development. In equine fetuses with a crown-rump length of less than 550 mm, the hoof capsule lacks a 'characteristic' white line since no borders between stratum medium, stratum internum and sole horn are discernible. In the hoof of an equine fetus with a crown-rump length of 550 mm, a narrow white line has taken shape. Its shallow lamellae are arrange...
Soana S, Gnudi G, Bertoni G, Botti P.The aim of this study is to point out the time of appearance of the carpal and tarsal bones in the fetal horse, considering an estimated fetal age, to follow their morphological development through to birth, and to characterize possible abnormal shape and/or delay of their ossification. The right carpal and tarsal region of 140 equine fetuses of both sexes (71 males, 69 females) and different ages (from 70 to 340 days of gestation) were examined radiographically in order to identify the sites of ossification from their earliest appearance. The times of appearance of the sites of ossification o...
Allen AL, Fretz PB, Card CE, Doige CE.A syndrome of congenital hypothyroidism and dysmaturity has been an important cause of reproductive loss and foal mortality in western Canada. The cause and pathogenesis of this syndrome is under investigation. One issue to be addressed is whether all the anomalies present in affected foals are produced concurrently by the same agent, or if affected foals are primarily hypothyroid in utero which induces the associated lesions. This study was designed to document the effects of fetal thyroidectomy, at about 215 days of gestation, on the growth and development of the equine fetus and to compare ...
Chavatte P, Holtan D, Ousey JC, Rossdale PD.Major progress on the endocrinology of the pregnant mare has been possible thanks to the catheterised equine fetal preparation developed by Marian Silver. In particular, these preparations led to the identification of the source of progestagens within the feto-placental unit and provided the impetus for further work on their biosynthesis and biological activities. The biosynthesis of the progestagens involves close interaction between the fetus, the endometrium and the placenta, and gives rise to some fundamental biochemical questions. The biological role of the progestagens is also discussed:...
Barber JA, Troedsson MH.A 12-year-old Arabian mare with a history of repeated early embryonic losses gave birth to a mummified fetus. The fetus was not the result of a pregnancy with twins. The mare had been given a progestogen throughout gestation and expelled the mummified fetus at about 325 days of gestation, 2 weeks after progestogen treatment was discontinued. We estimate that the size of the fetus was consistent with a fetal age of 5 months. The mare and mummified fetus illustrated that progestogen administration after 100 days of gestation can promote retention of a nonviable fetus. When the fetoplacental unit...
Bracher V, Mathias S, Allen WR.Placentation between 80 and 220 days of gestation was studied by measurement of fetal dimensions and gross, light and transmission and scanning electron microscopic examinations of the allantochorion and endometrium from 4 fertile mares with no, or very mild, endometrial histopathology and 6 subfertile Thoroughbred mares suffering varying degrees of age-related chronic degenerative endometritis (endometrosis). Spontaneously occurring twin gestation was observed in 3 animals. Several of the subfertile mares had endometrial cysts which showed 2 distinct features during pregnancy: those located n...
Schutzer WE, Holtan DW.The mare possesses unique steroid hormone metabolic activity during pregnancy in that peripheral 4-pregnene-3,20-dione (progesterone; P4) is undetectable by 220 days gestation. This study examines in vivo metabolism of progestins by the pregnant mare and in vitro metabolic activity of maternal and fetal tissues. Pregnant mares (n = 3) received intravenous infusions of 3 beta-hydroxy-5-pregnen-20-one (pregnenolone; P5), P4, 5 alpha-pregnane-3,20-dione (5 alpha-DHP), 3 beta-hydroxy-5 alpha-pregnan-20-one (3 beta-5 alpha), deuterium labeled (D4)-P5, D4-3 beta-5 alpha and vehicle. Anestrous mares ...
Engelhardt H, King GJ.The epitheliochorial placenta represents the least intimate association between maternal and fetal tissues. The best known examples of this form of placentation are the domestic livestock species. Current information on the nature and proposed functions of uterine lymphocyte populations in ruminants (sheep and cattle), horses and pigs is presented. In ruminants unusual gamma delta T cells may play a role in mid to late gestation. During normal horse pregnancy, fetally derived endometrial cup cells invade the uterine stroma and are destroyed by maternal leukocytes midway through gestation. Natu...
Lennard SN, Stewart F, Allen WR.In situ hybridization, Northern blotting, and immunohistochemical techniques were used to study the expression of transforming growth factor beta 1 (TGF beta 1) in the endometrium of the mare during the first 150 days of pregnancy (term = 330-340 days). In situ hybridization using an oligonucleotide (45mer) probe, based on a homologous region within all known mammalian TGF beta 1 DNA sequences, demonstrated TGF beta 1 mRNA accumulation in the glandular and lumenal epithelial cells of the endometrium from day 33 onwards which corresponds to the time of implantation (day 33-45). Expression in th...
Silver M, Fowden AL, Taylor PM, Knox J, Hill CM.Blood amino acids were measured in twelve chronically catheterized mares and fetuses between 250 and 310 days gestation. The concentrations of the majority of individual amino acids were similar in maternal and fetal arterial blood and no gestational changes were detected. Only methionine, phosphoserine, 3-methyl-histidine and glutamine were consistently higher in the fetus than the mare, whilst certain other amino acids were higher in the maternal blood. Fasting the mares for 36 h led to significant falls in plasma glucose and rises in urea and maternal free fatty acids (FFA). Small but signi...
Binanti D, Livini M, Riccaboni P, Sironi G.A 16-year-old primiparous mare aborted an apparently normal fetus at 240 days of gestation. A large, oval mass, measuring approximately 20 cm × 30 cm × 20 cm, was detected attached to the umbilical cord of the fetus. On the cut surface, the mass showed multifocal cystic structures, foci of mineralization, and diffuse hemorrhages. Histological examination of the mass revealed haphazardly arranged cartilage, bone, mesenchymal stroma, adipose tissue, vascular structures, smooth muscle, ciliated epithelium, squamous cornifying epithelium, and undifferentiated germ cells with areas of necrosis an...
Barber JA, Troedsson MH.A 12-year-old Arabian mare with a history of repeated early embryonic losses gave birth to a mummified fetus. The fetus was not the result of a pregnancy with twins. The mare had been given a progestogen throughout gestation and expelled the mummified fetus at about 325 days of gestation, 2 weeks after progestogen treatment was discontinued. We estimate that the size of the fetus was consistent with a fetal age of 5 months. The mare and mummified fetus illustrated that progestogen administration after 100 days of gestation can promote retention of a nonviable fetus. When the fetoplacental unit...
Orellana-Guerrero D, Uribe-Salazar JM, El-Sheikh Ali H, Scoggin KE, Ball B, Daels P, Finno CJ, Dini P.The placenta is a temporary organ that is essential for the survival of the fetus, with a lifelong effect on the health of both the offspring and the dam. The functions of the placenta are controlled by its dynamic gene expression during gestation. In this study, we aimed to investigate the equine placental DNA methylome as one of the fundamental mechanisms that controls the gene expression dynamic. Chorioallantois samples from four (4M), six (6M), and ten (10M) months of gestation were used to map the methylation pattern of the placenta. Globally, methylation levels increased toward the end o...
Yamauchi S.The horse fetal adrenal gland was shown to begin to increase in weight from about the end of the 4th month of pregnancy when the fetus has a crown-rump length of about 20 cm. Growth then proceeds steadily to term but, in contrast to the adult horse, the medulla remains thicker than the cortex throughout fetal life. The cortex also becomes established around 20 cm crown-rump length and at the same time the glomerular and fascicular zones become distinguishable. In contrast the reticular zone is not differentiated until around 50 cm crown-rump length. In the fetal adrenal cortex, the fascicular ...
Arvidson G, Astedt B, Ekelund L, Rossdale PD.Phospholipids in embryonic lung tissue, pulmonary washings and amniotic fluid were measured to study the development of lyng surfactant in the horse. A significant increase in the concentration of total phospholipids in lung tissue and a concomitant rise in the amount of dipalmitoyl lecithin in amniotic fluid between 100 and 150 days of gestation indicated the initial formation of surfactant in the fetal lung during this period.
Bunton TE.Developmental defects are rarely reported in the horse. Severe craniofacial and central nervous system defects in an equine foetus are described and their possible causation and pathogenesis are suggested.
Sweasey D, Patterson DS, Leadon DP.The lipid content of spinal cord, expressed as a percentage of adult values, was considerably higher for newborn foals than for several other species and traces of esterified cholesterol (type A) were only rarely present in horse fetal cord (from 270 days gestational age onwards). This suggested that, at birth, the spinal cord is neurochemically more 'mature' in the horse than in cattle, sheep and pigs. Data for premature foals revealed no lipid abnormality suggestive of myelin immaturity or degeneration.
Bragulla H.The pre- and perinatal development of the hoof cartilage is described concerning the histological structure and surrounding vessels. Beginning in the third month of fetal development, the anlage of the hoof cartilage is still present in typical shape and location. It is built out of mesenchymal connective tissue. During further fetal development, the connective tissue cells will differentiate into two cell populations, fibroblasts and chondroblasts. Vessels, traversing the hoof cartilage, are surrounded by loose connective tissue, which will partially develop fibrocartilage. At birth, hoof car...
Platt H.Many infections of the equine placenta and fetus result from ascending spread along the cervical canal. Most abortions due to infection occur during the later stages of pregnancy and the possible effects of intrauterine infection on the developing fetus and young foal are discussed.
Campos IS, de Souza GN, Gomes GM, Pinna AE, Ferreira AMR.The aim of the study was to correlate the spectral index of the right and left uterine arteries with equine placental development in mares with advanced pregnancies. We examined 32 multiparous Mangalarga Marchador mares with gestation of 150-240 days. During pregnancy, the pulsatility index (PI) and resistance index (RI) of the uterine arteries were obtained using spectral Doppler ultrasonography, and the combined uteroplacental thickness was obtained monthly using B-mode ultrasonography. The combined uteroplacental thickness correlated with gestational time of up to 13 years of age, and the s...
London WT, Levitt NH, Altshuler G, Curfman BL, Kent SG, Palmer AE, Sever JL, Houff SA.Fetal rhesus monkeys were inoculated intracerebrally with an attenuated strain of western equine encephalitis virus. All animals developed microcephaly. Twelve of sixteen monkeys developed ex vacuo hydrocephalus. All virus inoculated fetuses developed WEE virus antibody. Virus could not be recovered at the time of delivery. Monkeys with the highest WEE antibody titers showed the greatest degree of hydrocephalus.
Hashimoto M, Nambo Y, Kondo T, Watanabe K, Orino K.In mammal circulation, ferritin-binding proteins (FBPs) are thought to be involved in clearance of circulating ferritin after complex formation with it through receptor-mediated uptake. However, there is no report on fetal FBP in fetal circulation. Although iron concentrations of fetal horse plasma were higher than those of adult horse plasma, plasma ferritin concentrations and ferritin-binding activities were found to be significantly lower in fetus than in adult. FBPs were purified from fetal or adult horse plasma on horse spleen ferritin-Sepharose 4B affinity column. Partially affinity-puri...
Simpson KS, Adams MH, Behrendt-Adam CY, Baker CB, McDowell KJ.For development to proceed normally, the appropriate genes must be expressed in the correct tissues and in the correct time frame. Knowledge of gene expression during development provides information about the changes taking place within the conceptus as well as possible reasons for pregnancy failure. However, little is known about gene expression during development in the equine conceptus. In this study, we examined differences in gene expression between day 12 and day 15 equine conceptuses by suppression subtractive hybridization. This technique was used to isolate transcripts that are more ...
Read JE, Cabrera-Sharp V, Offord V, Mirczuk SM, Allen SP, Fowkes RC, de Mestre AM.Equine chorionic girdle trophoblast cells play important endocrine and immune functions critical in supporting pregnancy. Very little is known about the genes and pathways that regulate chorionic girdle trophoblast development. Our aim was to identify genes and signalling pathways active in vivo in equine chorionic girdle trophoblast within a critical 7-days window. We exploited the late implantation of the equine conceptus to obtain trophoblast tissue. An Agilent equine 44K microarray was performed using RNA extracted from chorionic girdle and chorion (control) from equine pregnancy days 27, ...
Loux S, Robles M, Chavatte-Palmer P, de Mestre A.Development and the subsequent function of the fetal membranes of the equine placenta require both complex and precise regulation of gene expression. Advancements in recent years in bioinformatic techniques have allowed more extensive analyses into gene expression than ever before. This review starts by combining publically available transcriptomic data sets obtained from a range of embryonic, placental and maternal tissues, with previous knowledge of equine placental development and physiology, to gain insights into key gene families relevant to placentation in the horse. Covering the whole o...
Joujou-Sisic K, Granérus M, Wetterling H, Wikström K, Engström W, Jeffcott L, Schofield PN, Welin A.The expression of the insulin like growth factor (IGF) II gene has been examined in the developing equine fetus. It was found that IGF II transcripts were present in abundant quantities in third trimester embryonic and extraembryonic tissues as for example the placenta. The expression of the IGF II gene was high in the fetal liver where two prominent transcripts--4.6 and 4.1--kB were produced. However, these transcripts could not be traced in the adult liver. Instead we found two different transcripts with the sizes of 4.0 and 2.9 kB in the adult liver. These findings taken together with the d...
Vieira GS, Paludo GR, Ramos AF, Pivato I, de Oliveira RA.For a long time, Pantaneiro and Campeiro breeds were raised only within their places of origin. Consequently, there are few of these horses; therefore, establishing reproductive and clinical standards for these animals is necessary to implant new biotechnologies for reproduction to preserve their genetics. This study aimed to perform a descriptive evaluation of fetal age determination by fetus ocular orbit measurement in mares of the Campeiro and Pantaneiro breeds. We also evaluated sequential changes in hematology and biochemistry for foals from birth to six months of life by counting red blo...
Robles M, Peugnet PM, Valentino SA, Dubois C, Dahirel M, Aubrière MC, Reigner F, Serteyn D, Wimel L, Couturier-Tarrade A, Chavatte-Palmer P.Following embryo transfer (ET), the size and breed of the recipient mare can affect fetal development and subsequent post natal growth rate and insulin sensitivity in foals. Objective: To investigate placental adaptation in pregnancies where increased or restricted fetal growth was induced through ET between Pony, Saddlebred and Draught horses. Methods: In vivo experiment. Methods: Control Pony (P, n = 21) and Saddlebred (S, n = 28) pregnancies were obtained by artificial insemination. Increased pregnancies were obtained by transferring Pony (P-D, n = 6) and Saddlebred (S-D, n = 8) emb...
Hanada M, Maeda Y, Oikawa MA.The secretions of the equine endometrial glands are essential for the survival, growth, and development of the conceptus in early pregnancy, and endometrial gland density is directly related to successful pregnancy outcome. Endometrial biopsy is routinely used to assess the reproductive potential of broodmares. Some previous studies have shown that equine endometrial glands are uniformly distributed throughout the uterus; however, other work has shown variation of the endometrial architecture between biopsy sites, suggesting that a single biopsy is not representative of the entire endometrium....
Aucamp J, van der Zwan H, Geldenhuys Z, Abera A, Louw R, van der Sluis R.In animal breeding, a species sex can influence the value of the animal. For example, in the horse breeding industry, mares are preferred as polo horses, while in wildlife breeding males with larger horns are more valuable. Therefore, the economic advantages of knowing the unborn fetus' sex are important to successful animal management. Ultrasonography is used to determine the sex of unborn fetuses, but this method places additional stress on the animal and require specialized equipment and expertise. Conversely, molecular-based sexing techniques require less invasive sampling and can determin...
Bragulla H, Budras KD, Reilly JD.The fetal development of the white line (Zona alba) in the equine hoof is described. Its specific structure of lamellar and interlamellar horn, which in turn is composed of cap and terminal horn, is formed in the second half of the hoof's fetal development. In equine fetuses with a crown-rump length of less than 550 mm, the hoof capsule lacks a 'characteristic' white line since no borders between stratum medium, stratum internum and sole horn are discernible. In the hoof of an equine fetus with a crown-rump length of 550 mm, a narrow white line has taken shape. Its shallow lamellae are arrange...
Renaudin C, Wensley F, Morgan J, Cassano J, Spriet M.Fetal age in Quarter Horses can be predicted within 2 weeks from 100- to 200- days of gestation using femur length, biparietal diameter (cranium diameter) and eye approximated volume. However, as pregnancy advances, the femur and cranium become too large to be imaged in their entirety using ultrasound and the corresponding biometric parameters can no longer be measured. In this longitudinal study, the proximal phalanx (P1) was evaluated as a novel biometric parameter for late gestation to predict fetal age and bone maturation. Transrectal ultrasound was performed in ten pregnant mares with kno...