Developmental biology in horses involves the study of the processes by which horses grow and develop from a single fertilized egg into a fully formed organism. This field encompasses various stages, including embryonic development, fetal growth, and postnatal maturation. Researchers in this area examine cellular differentiation, gene expression, and morphogenetic movements that contribute to the formation of tissues and organs in equine species. Key topics include the molecular mechanisms that regulate developmental pathways, the influence of genetic and environmental factors on development, and the identification of developmental disorders. This page compiles peer-reviewed research studies and scholarly articles that explore the fundamental aspects of equine developmental biology, offering insights into the mechanisms driving normal and abnormal development in horses.
Allen WR, Skidmore JA, Stewart F, Antczak DF.Measurement of the concentrations of equine chorionic gonadotrophin (eCG) in the serum of pregnant mares and Jenny donkeys carrying normal intraspecies and hybrid interspecies pregnancies suggested that the production of this hormone may be influenced by parental gene imprinting. Specifically, a differential expression of maternal and paternal genes may control the size and secretory activity of the structures that secrete eCG, the fetal endometrial cups. However, bisection of an interspecies mule embryo followed by transfer of the resulting demi-embryos and other intact mule embryos to horse ...
Collins MH.The placenta provides many critical services to the developing foetus. Proper placental implantation, growth and function are necessary for normal foetal growth and development. Placental structure varies widely among species but all mammalian placentas have a convoluted materno-foetal interface that may be quite simple or highly complex; the more complicated interdigitations tend to characterise smaller placentas that have limited areas of contact between the placenta and the endometrium. The intimacy of the contact between maternal and foetal tissue varies from apposition only, as in the equ...
Sisken BF, Zwick M, Hyde JF, Cottrill CM.This review covers the development and maturation of the cerebellum of the horse and compares this developmental sequence with that of man, mouse and chicken. These comparisons attempt to correlate morphological and neurochemical features, developmental time and functional performance necessary for survival at birth. Although there is great disparity between these 3 species, the basic anatomical structures of the cerebellum are present as are specific cellular groups, synapses and neurochemical markers. In addition to this structural homogeneity, other attributes of the cerebellum are its easi...
Riggs CM, Lanyon LE, Boyde A.A novel technique for determining the collagen fibre orientation pattern of cross-sections of cortical bone was used to study mid-diaphyseal sections from the equine radius. Several in vivo strain gauge studies have demonstrated that this bone is loaded in bending during locomotion in such a way that the cranial cortex is consistently subjected to longitudinal tensile strains and the caudal cortex to longitudinal compressive strains. Twenty-three radii from 17 horses were studied. All the bones obtained from adult horses exhibited a consistent pattern of collagen fibre orientation across the c...
Knospe C, Budras KD.To answer the many open questions concerning the development of the horse's ovary, first the prenatal development was investigated. It resulted that follicles derive from the germinal epithelium and its cords, whereas the Leydig cells and the rete blastema originate from the mesonephros. In the second third of pregnancy the Leydig cells undergo an enormous proliferation, in the last third they degenerate. However this degeneration is not connected with the postnatal development of the ovulation groove.
Donaldson WL, Oriol JG, Plavin A, Antczak DF.Between days 36-38 of pregnancy equine trophoblastic cells of the chorionic girdle migrate and form endometrial cups. Just prior to invasion, the chorionic girdle cells express high levels of polymorphic, paternally inherited, major histocompatibility complex (MHC) class I antigens. Their descendents, the mature, invasive trophoblast cells of the endometrial cups, however, express low or undetectable levels of MHC class I antigens by day 44 of pregnancy. Experiments with MHC compatible pregnancies, the study of residual chorionic girdle cells that had failed to invade the endometrium and remai...
Fowden AL, Mijovic J, Ousey JC, McGladdery A, Silver M.The activities of glucose-6-phosphatase (G6P), fructose diphosphatase, phosphoenolpyruvate carboxykinase (PEPCK), aspartate and alanine transferases were measured in liver and kidney of fetal foals between 100-318 days of gestation (term approximately 335 days) and during the immediate postnatal period (0-48 h after birth). All 5 enzymes could be detected in the fetal liver and kidney at the youngest gestational age studied. Mean fetal activities were lower than those observed in their mothers and showed no change with gestational age for the majority of enzymes studied. However, renal PEPCK a...
O'Donohue DD, Smith FH, Strickland KL.A two part survey was carried out in Irish Thoroughbred horses in 1988 and 1989 to establish the incidence and prevalence of developmental skeletal problems, particularly possible manifestations of developmental orthopaedic disease (DOD). Survey One was a retrospective study based on a questionnaire involving the foal crops of 46 stud farms for 3 successive seasons; the 1711 animals initially documented represented 10.46% of Irish foal registrations. The second survey involved repeated monitoring of the 1988 foal crop from birth to 18 months of age on 17 stud farms. The 248 foals initially exa...
Freeman DA, Woods GL, Vanderwall DK, Weber JA.The hypothesis that equine embryos initiate oviductal transport in mares was tested by placing day 6 uterine embryos in the oviducts of day 2 (n = 10) or day 5 (n = 10) recipient mares and attempting to collect the embryos from the uterus 48 h later. To determine whether the surgical transfer procedure initiated oviductal transport, medium alone was placed in the oviducts of day 2 (n = 10) inseminated mares (sham transfer), and uterine embryo collections were attempted 48 h later. Embryos were transported through the oviduct of day 2 recipients by day 4 (instead of day 5 to 6) in six of ten ma...
Ball BA, Miller PG.In this study we examined the ability of equine oviductal epithelial cells (OEC) to support the development of four- to eight-cell equine embryos in vitro and investigated the ability of co-cultured embryos to continue normal development after transfer to synchronous recipient mares. Equine embryos obtained at Day 2 after ovulation were cultured with or without OEC for 5 days. Those OEC co-cultured embryos that reached the blastocyst stage and embryos recovered from the uterus at Day 7 were surgically transferred to synchronous recipient mares. Co-culture with OEC improved (P < 0.01) develo...
Horn F, Windle JJ, Barnhart KM, Mellon PL.The molecular mechanisms for the development of multiple distinct endocrine cell types in the anterior pituitary have been an area of intensive investigation. Though the homeodomain protein Pit-1/GHF-1 is known to be involved in differentiation of the somatotrope and lactotrope lineages, which produce growth hormone and prolactin, respectively, little is known of the transcriptional regulators important for the gonadotrope cell lineage, which produces the glycoprotein hormones luteinizing hormone and follicle-stimulating hormone. Using transgenic mice and transfection into a novel gonadotrope ...
Martínez-Galisteo A, Diz A, Agüera E, Vivo J.34 Andalusian foals of both sexes were divided into three age-groups (A = mean age 1 month, B = 7 months, C = 14 months). Samples of the right m. gluteus medius were stained for m-ATPase and NADH-TR in order to determine fibre type composition and size as well as the relative area occupied by each type. Results revealed no statistically significant variation in the proportion of type I fibre among the three age-groups. Significant differences were recorded, however, for type II fibres; an increase in the proportion of IIA fibres was accompanied by a decrease in IIB ones, the difference being m...
Enders AC, Liu IK.The structure of the equine chorionic girdle between days 28 and 42 of gestation was examined. Of particular interest were differentiation of trophoblastic cells within the girdle, adhesion between girdle and endometrium, invasion and displacement of the uterine epithelium, and the nature of the endometrium when girdle cells migrate into it to form endometrial cup cells. The chorionic girdle, identified initially as a band of tall columnar cells, becomes a stratified columnar epithelium indented by clefts and pits. Adhesion to and penetration through the endometrial luminal epithelium are rapi...
Freeman DA, Weber JA, Geary RT, Woods GL.The objectives of this study were 1) to determine the time of embryo transport through the mare oviduct, 2) to determine whether equine embryos increase in diameter prior to the time of oviductal transport, and 3) to assess the stage of equine embryonic development at the time of oviductal transport. The time of oviductal transport (interval from ovulation to uterine entry) was estimated by collecting embryos from the mare oviduct or uterus at 2-hour intervals from 120 to 168 h postovulation. The time of oviductal transport was 130 to 142 h, since 9 9 embryos were located in the oviduct from 1...
Freeman DA, Butler JE, Weber JA, Geary RT, Woods GL.Oviductal and uterine embryos were collected from mares at 5 to 7 days following ovulation 1) to evaluate the effects of oviductal tissue explants on in vitro growth and development of equine embryos and 2) to study the morphologic development of equine embryos in culture. Embryos were incubated for 5 days in a medium (control group) or in medium supplemented with oviductal tissue explants (co-culture group). Embryos were evaluated and the media changed daily. Following 5 days in culture, 10 10 (100%) control embryos and 27 29 (93%) co-cultured embryos had doubled in diameter. All embryos that...
Zicker SC, Spensley MS, Rogers QR, Willits NH.The concentrations of 23 amino acids in the plasma of 13 healthy foals were determined before suckling, when foals were 1 to 2 days old, 5 to 7 days old, 12 to 14 days old, and 26 to 28 days old. The ratio of the branched chain amino acids to the aromatic amino acids was also calculated at the 5 time points. Analysis of the concentrations at the 5 ages revealed a significant temporal relationship for each amino acid ranging from a polynomial order of 1 to 4 inclusively. There were significant differences between several concentrations of amino acids in plasma at specific sample times; however,...
Ball BA, Altschul M, Ellington JE.This study compared the in vitro development of Day-2 equine embryos co-cultured with either trophoblastic vesicles or oviductal explants. Embryos were collected surgically from the oviducts of pony mares 2 d after ovulation and assessed for stage of development. Culture medium was Ham's F12 and Dulbecco's Modified Eagle's Medium (50:50 v/v) in a humidified atmosphere of 5% CO(2) in air at 38.5 degrees C with either trophoblastic vesicles or oviductal explants. The quality score of embryos was assessed daily. After 4 d in culture, embryos were stained (Hoechst 33342) and evaluated with epifluo...
Nilson JH, Bokar JA, Clay CM, Farmerie TA, Fenstermaker RA, Hamernik DL, Keri RA.Expression of the glycoprotein hormone alpha-subunit gene occurs in the pituitary of all mammals but in placenta of only primates and horses. In humans, two different elements, termed upstream regulatory element (URE) and cAMP response element (CRE), are required for placenta-specific expression of the alpha-subunit gene. The URE binds a protein unique to placenta whereas the CRE binds a ubiquitous protein. Comparative analysis of the promoter-regulatory region of the alpha-subunit gene from a number of mammals indicates that a functional URE has been retained and suggests the potential for pl...
Steger DJ, Altschmied J, Büscher M, Mellon PL.Primate and equine species are thought to be unique among mammals in synthesizing placental gonadotropin glycoprotein hormones. Human chorionic gonadotropin (CG) and equine pregnant mare's serum gonadotropin (PMSG) are produced in placenta by the specific activation of a glycoprotein hormone alpha-subunit gene and a corresponding beta-subunit gene. The evolutionary mechanisms for the apparently independent acquisition of tissue specificity were investigated by cloning the 5' flanking region of the equine alpha-subunit gene and comparing the DNA elements and trans-acting factors involved in pla...
Hinrichs K, DiGiorgio LM.A technique was developed in which immature horse oocytes, obtained from slaughterhouse specimens, were transferred to the pre-ovulatory follicle of a mare in vivo, with resulting oocyte maturation, ovulation, fertilization and embryo development. Oocytes were collected from all follicles greater than 3 mm, and were classified as immature, maturing, expanded or denuded. The transfers were performed in the standing, tranquilized mare. The ovary containing the pre-ovulatory follicle was grasped per rectum. A trochar and cannula were placed through the abdominal wall in the flank area, ipsilatera...
Pruitt JA, Forrest DW, Burghardt RC, Evans JW, Kraemer DC.The viability and ultrastructure of equine embryos were assessed following culture in a static or perifusion system. The percentage change in diameter was greater (P less than 0.025) for embryos in the static treatment (71%) than in the perifusion treatment (33%). Fluorescein diacetate (FD) scores, the percentage of fluorescing cells (FC) and fluorescent intensity (FI), also were greater (P less than 0.05 and P less than 0.01) following static culture than for embryos cultured in the perifusion system. Four of 9 control embryos resulted in pregnancies but no embryos cultured in either system p...
Enders AC, Liu IK.The equine blastocyst becomes fixed in position in the uterus on approximately Day 16 of gestation, but allantochorionic villi are not formed until about Day 50. The purpose of this study was to examine evidence that the blastocyst is orientated during this time period, and to determine what morphological features might assist retention of the position of the blastocyst within the uterus. Implantation sites were collected on Days 10-42 of gestation, and the reproductive tracts perfused with fixative for light and electron microscopic examination. The conceptus is found at the bend of a uterine...
Brück I, Hyland JH.The contributions of 2 biochemical pathways to the total metabolism of glucose (the Embden-Meyerhof pathway [EMP] and the pentose phosphate pathway [PPP]), were assessed for equine embryos recovered on Day 4.5, 7.5 and 11.5 post ovulation. At all developmental stages studied, glucose was metabolized through both pathways. Through the EMP, the amounts of glucose metabolized per nl embryo volume per hour were 4.0, 9.9 and 3.1 pmol, whereas via the PPP, amounts were 0.9, 1.7 and 0.07 pmol for Day-4.5, -7.5 and -11.5 embryos, respectively. The ratio of EMP:PPP with age was 9.7 for Day -4.5 embryos...
Hinrichs K, Schmidt AL, Memon MA, Selgrath JP, Ebert KM.Embryos were recovered from the uteri of mares 5 d after ovulation. Six embryos, all morulae, were placed singly in 200-ul droplets of Ham's F-12 with 10% fetal calf serum and cultured at 37 degrees C in a 5% CO(2) atmosphere. The embryos expanded to form blastocysts by the third day of culture. The blastocysts hatched from their zona pellucida, rather than the zona thinning and flaking off, as occurs in vivo. Hatching from the zona pellucida began on the third day of culture and was complete in five of six embryos by the sixth day. The embryonic capsule, normally present in equine embryos aft...
Oikawa M, Yoshihara T, Kaneko M, Yoshikawa T.The morphology of the equine allantochorion at the tip of the pregnant horn was studied in the membranes of 14 mares. The findings in the allantochorion at the tip of the pregnant horn were of two types; one was growth retardation (hypoplastic villi, tunica adventitia of the vessels resembling embryonal connective tissue and the extended spaces of remnants of the extraembryonic coelom), the other was placental hypoxia or ischaemia (parakeratosis, stratified squamous metaplasia, necrosis of the trophoblasts, thickening of the basement membrane and fibrous hyperplasia of villous stroma). It seem...
Fenstermaker RA, Farmerie TA, Clay CM, Hamernik DL, Nilson JH.Expression of the glycoprotein hormone alpha-subunit gene occurs in the pituitaries of all mammals and in the placentas of primates and horses. In humans, tandem cAMP response elements (CREs), located in the proximal promoter-regulatory region of the alpha-subunit gene, act together with an adjacent upstream regulatory element to confer placenta-specific expression. Here, we report that the alpha-subunit genes of Old World Monkeys contain a single functional CRE. This suggests that tandem CREs are unique to higher primates and humans and are not absolutely required for placenta-specific expres...
Komosa M, Flisinska-Bojanowska A, Gill J.1. The development of diurnal rhythm activity of FDPA, AspAT and A1AT and in levels of cortisol, T3 and T4 was observed in the blood serum of six foals. 2. The studies began when a foal was 7 days old and were repeated every month until foals reached 1 year of age. Blood samples were taken every 4 hr for one day each month. 3. As a control group four barren mares were used, kept and examined in the same conditions. 4. In mature mares, diurnal rhythms in activity of A1AT (acrophase at 2200 hr), AspAt (2400 hr) and cortisol (0630 hr) but in T3 only in summer months (acrophase at 0100 hr) were ob...
Rao UR, Chapman MR, Singh RN, Mehta K, Klei TR.Transglutaminases (E.C. 2.3.3.13) are a family of Ca(2+)-dependent enzymes that stabilize protein structure by catalyzing the formation of isopeptide bonds. A novel form of transglutaminase has been identified and characterized that seem to play an important role in growth, development, and molting in adult and larval stages of filarial nematodes. The aim of this study was to identify the ubiquitous nature of this enzyme in other nematodes and to measure its significance to larval growth, molting, and development. For this purpose, equine Strongylus spp. were used. Activity of this enzyme was ...
Firth EC, Poulos PW.Selective perfusion and microangiographic examination of the radius and metacarpus of 30 foals were performed. Vessels of extraosseous origin supplying the peripheral part of the metaphysis were not demonstrated in young foals, but became more obvious in older animals. The development of this vessel system coincided with the reduction of transphyseal vessels from the epiphysis crossing the growth plate.
Roberts MC, Kidder DE, Hill FW.Two enzymes having lactase activity are present in the equine small intestine. The first, the digestive enzyme, neutral beta-galactosidase, declines in activity from birth to three years, disappearing completely between 3 and 4 years of age. The other, the soluble lysosomal enzyme, acid beta-galactosidase, having affinity for lactose and a synthetic beta-galactoside, shows a decrease in activity in the first three months of life and thereafter varies little in activity and represents the lactase enzyme in the adult horse. This pattern may parallel the development of lactase activity in many ot...
Silver M.THERE is no shortage of anatomical information on the equine foetus and its placenta, from the early work of Ruini in the 16th century to the recent studies of Steven and colleagues (Steven 1982); by contrast, knowledge of the physiology of the foal in utero is sparse. In other domestic animals there have been considerable ads ances in foetal and neonatal physiology and endocrinology in recent years due mainly to the develop-ment of the chronically catheterised foetal preparation in which sequential observations can he made in the conscious animal (Silver 1981). Some information about the deve...
Salamone D, Maserati M.Horse cloning by somatic cell nuclear transfer (SCNT) is an attractive scientific and commercial endeavor. Moreover, SCNT allows generating genetically identical animals from elite, aged, castrated, or deceased equine donors. Several variations in the horse SCNT method have been described, which may be useful for specific applications. This chapter describes a detailed protocol for horse cloning, thus including SCNT protocols using zona pellucida (ZP)-enclosed or ZP-free oocytes for enucleation. These SCNT protocols are under routine use for commercial equine cloning.
Breit S.Ten horses, younger than 1 year, were used to investigate the articulatio interphalangea distalis of the pelvic limb. The expansion of the dorsal recess was compared to those of adult horses. The recessus dorsalis always formed a shape like the letter 'm' or like a triangle with oblique angles. Consequently the great proximo-distal expansion on the medial side of the dorsal recess very probably has nothing to do with the age of horses.
Möstl E.In the pregnant mare two different groups of oestrogens are produced by the placenta. The precursor of "classical" oestrogens (oestrone, oestradiol-17 beta and oestradiol-17 alpha) is dehydroepiandrosterone which originates from the fetal gonads. The ring B unsaturated oestrogens (equilin and equilenin and their derivatives) derive from farnesyl pyrophosphate by a pathway not involving cholesterol.
Fischer B, Rose-Hellekant TA, Sheffield LG, Bertics PJ, Bavister BD.Preimplantation embryos of the pig (Days 11 to 15), cow (Days 14 to 16), sheep (Day 14) and pony (Day 16) bind epidermal growth factor (EGF) specifically. Binding was not detected in embryos of the rabbit at Day 5 or 6 or the hamster at Day 3. Transforming growth factor-alpha displaced [(125)I] EGF in pig, cow and pony embryos almost as much as unlabeled EGF. The binding affinities of EGF ranged from 12 to 233 pM in pig and cow embryos. The range of species and binding features indicate that the EGF family may play a significant role in mammalian preimplantation development.