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Topic:Physiology

The physiology of horses encompasses the study of the biological functions and processes that occur within the equine body. This includes the examination of various systems such as the cardiovascular, respiratory, musculoskeletal, digestive, and nervous systems. Understanding equine physiology is essential for comprehending how horses adapt to different environmental conditions, perform physical activities, and respond to health challenges. Research in this field often focuses on the mechanisms of energy metabolism, thermoregulation, and muscle function during exercise, as well as the physiological responses to stress and disease. This page compiles peer-reviewed research studies and scholarly articles that explore the diverse aspects of equine physiology, providing insights into the biological processes that support the health and performance of horses.
Distal interphalangeal articular cartilage assessment using low-field magnetic resonance imaging. Olive J.The suitability of low-field magnetic resonance (MR) imaging for assessment of articular cartilage has been questioned, based on insufficient image quality. The purposes of this study were to describe the MR anatomy of the normal distal interphalangeal (DIP) cartilage, and to evaluate the sensitivity and accuracy of low-field MR imaging for identification of cartilage erosions that were created ex vivo. Imaging sequences included sagittal and dorsal multiple-oblique T1-weighted gradient-recalled echo (GRE) and sagittal dual echo sequences. In the thickest regions, normal cartilage appeared as ...
Comparison of endoscopy, histology, and cytokine mRNA of the equine gastric mucosa.
Veterinary research communications    May 18, 2010   Volume 34 Suppl 1 S121-S124 doi: 10.1007/s11259-010-9401-6
Pietra M, Morini M, Perfetti G, Spadari A, Vigo P, Peli A.In recent years, gastric ulceration has been recognized as a common, possibly performance-limiting disease, of adult horses. The aim of this study was to compare endoscopic features, histological diagnosis, and mRNA levels of various cytokines (TNF-alpha, IL-1beta, and IL-13) from horse gastric biopsies. Eleven horses suffering from equine gastric ulcer syndrome and seven horses with normal histological gastric features were assessed. No correlation between endoscopic features and histology (i.e., the gold standard) was observed. Based on histological diagnosis, a significant (p < 0.05) inc...
On the search for in vitro in vivo correlations in the field of intra-articular drug delivery: administration of sodium diatrizoate to the horse.
European journal of pharmaceutical sciences : official journal of the European Federation for Pharmaceutical Sciences    May 16, 2010   Volume 41, Issue 1 10-15 doi: 10.1016/j.ejps.2010.05.006
Frost AB, Larsen F, Ostergaard J, Larsen SW, Lindegaard C, Hansen HR, Larsen C.Development of suitable in vitro release models for formulation development as well as quality control purposes has to be initiated in the early design phase of injectable depots. Optimally, construction of an in vitro release model may lead to the establishment of in vitro in vivo correlations. By using a model compound (sodium diatrizoate, DTZ), the purpose of this study was to investigate the possibility of establishing in vitro in vivo relations between the DTZ disappearance profile obtained from the donor compartment of the rotating dialysis cell model and the joint disappearance profile ...
Determination of (13)C/(12)C ratios of urinary excreted boldenone and its main metabolite 5beta-androst-1-en-17beta-ol-3-one.
Drug testing and analysis    May 15, 2010   Volume 2, Issue 5 217-224 doi: 10.1002/dta.124
Piper T, Geyer H, Gougoulidis V, Flenker U, Schänzer W.Boldenone (androsta-1,4-dien-17beta-ol-3-one, Bo) is an anabolic steroid known to have been used in cattle breeding or equine sport as a doping agent for many years. Although not clinically approved for human application, Bo or its main metabolite 5beta-androst-1-en-17beta-ol-3-one (BM1) were detected in several doping control samples. For more than 15 years the possibility of endogenous Bo production in human beings has been discussed. This is a challenging issue for doping control laboratories as Bo belongs to the list of prohibited substances of the World Anti-Doping Agency and therefore th...
Membrane lipids of the stallion spermatozoon in relation to sperm quality and susceptibility to lipid peroxidation.
Reproduction in domestic animals = Zuchthygiene    May 12, 2010   Volume 46, Issue 1 141-148 doi: 10.1111/j.1439-0531.2010.01609.x
García BM, Fernández LG, Ferrusola CO, Salazar-Sandoval C, Rodríguez AM, Martinez HR, Tapia JA, Morcuende D, Peña FJ.Lipids were extracted from ejaculated spermatozoa from seven individual stallions to distinguish neutral lipids (NL) and polar lipids (PL) and determine their variation among stallions and their relationship with sperm quality and sperm susceptibility to lipid peroxidation. The isolated fatty acids were correlated with sperm quality (membrane integrity, mitochondrial membrane potential (ΔΨm) and expression of active caspases) and the sensitivity of the sperm plasma membrane to LPO. The miristic (C14: 0), palmitic (C16: 0), stearic (C18: 0) and oleic (C18: 1n9) acids were predominant among th...
Selenium in blood, semen, seminal plasma and spermatozoa of stallions and its relationship to sperm quality.
Reproduction, fertility, and development    May 11, 2010   Volume 22, Issue 5 886-891 doi: 10.1071/RD10032
Bertelsmann H, Keppler S, Höltershinken M, Bollwein H, Behne D, Alber D, Bukalis G, Kyriakopoulos A, Sieme H.The essential trace element selenium is indispensable for male fertility in mammals. Until now, little data existed regarding the relationship between selenium and sperm quality in the stallion. Selenium, or selenium-dependent glutathione peroxidase activity, was determined in red blood cells, semen, seminal plasma and spermatozoa, and the percentages of spermatozoa with progressive motility (PMS), intact membranes (PMI), altered (positive) acrosomal status (PAS) and detectable DNA damage, determined by the sperm chromatin structure assay, were evaluated in 41 healthy stallions (three samples ...
Tachypnea and antipyresis in febrile horses after sedation with alpha-agonists.
Journal of veterinary internal medicine    May 11, 2010   Volume 24, Issue 4 1008-1011 doi: 10.1111/j.1939-1676.2010.0528.x
Kendall A, Mosley C, Bröjer J.Signs of tachypnea after sedation of febrile horses with alpha2-agonists have been noted previously but have not been further investigated. Objective: To examine the effects of xylazine and detomidine on respiratory rate and rectal temperature in febrile horses and to investigate if either drug would be less likely than the other to cause changes in these variables. Methods: Nine febrile horses and 9 healthy horses were included in the study. Methods: Horses were randomly assigned to sedation with xylazine 0.5 mg/kg or detomidine 0.01 mg/kg. Heart rate and respiratory rate were recorded before...
Inhibition of the mitochondrial permeability transition pore reduces “apoptosis like” changes during cryopreservation of stallion spermatozoa.
Theriogenology    May 10, 2010   Volume 74, Issue 3 458-465 doi: 10.1016/j.theriogenology.2010.02.029
Ortega Ferrusola C, González Fernández L, Salazar Sandoval C, Macías García B, Rodríguez Martínez H, Tapia JA, Peña FJ.In order to evaluate to what extent the changes that occur during cryopreservation involve the mitochondrial permeability transition pore (PT-pore), a specific inhibitor was used during the cryopreservation process of stallion spermatozoa. Four ejaculates from each of 7 stallions were frozen using a standard protocol. Before freezing, each ejaculate was split into three subsamples. The first was supplemented with 2.5 microM bongkrekic acid (BA) and the second with 5 microM BA. The third subsample served as control. The BA significantly reduced the percentage of spermatozoa depicting active cas...
A comparison of the moment arms of pelvic limb muscles in horses bred for acceleration (Quarter Horse) and endurance (Arab).
Journal of anatomy    May 10, 2010   Volume 217, Issue 1 26-37 doi: 10.1111/j.1469-7580.2010.01241.x
Crook TC, Cruickshank SE, McGowan CM, Stubbs N, Wilson AM, Hodson-Tole E, Payne RC.Selective breeding for performance has resulted in distinct breeds of horse, such as the Quarter Horse (bred for acceleration) and the Arab (bred for endurance). Rapid acceleration, seen during Quarter Horse racing, requires fast powerful muscular contraction and the generation of large joint torques, particularly by the hind limb muscles. This study compared hind limb moment arm lengths in the Quarter Horse and Arab. We hypothesized that Quarter Horse hind limb extensor muscles would have longer moment arms when compared to the Arab, conferring a greater potential for torque generation at the...
Concentrations of altrenogest in plasma of mares and foals and in allantoic and amniotic fluid at parturition.
Theriogenology    May 10, 2010   Volume 74, Issue 2 229-235 doi: 10.1016/j.theriogenology.2010.02.006
Palm FM, Schenk I, Neuhauser S, Schubert D, Machnik M, Schänzer W, Aurich C.Treatment with the progestin altrenogest is widely used in pregnant mares. The fact that foals born from healthy mares treated with altrenogest until term suffered from neonatal problems raises the question of direct effects of altrenogest on vital functions in the neonate. We have therefore investigated altrenogest concentrations in maternal and neonatal blood plasma and in fetal fluids. Pregnant mares were treated with altrenogest orally once daily (0,088 mg/kg bodyweight, n = 7) or left untreated (n = 8) from 280 d of gestation until foaling. Altrenogest concentration was determined in plas...
Horse prion protein NMR structure and comparisons with related variants of the mouse prion protein.
Journal of molecular biology    May 8, 2010   Volume 400, Issue 2 121-128 doi: 10.1016/j.jmb.2010.04.066
Pérez DR, Damberger FF, Wüthrich K.The NMR structure of the horse (Equus caballus) cellular prion protein at 25 degrees C exhibits the typical PrP(C) [cellular form of prion protein (PrP)] global architecture, but in contrast to most other mammalian PrP(C)s, it contains a well-structured loop connecting the beta2 strand with the alpha2 helix. Comparison with designed variants of the mouse prion protein resulted in the identification of a single amino acid exchange within the loop, D167S, which correlates with the high structural order of this loop in the solution structure at 25 degrees C and is unique to the PrP sequences of e...
In vivo administration of acepromazine or promethazine to horse decreases the reactive oxygen species production response of subsequently isolated neutrophils to stimulation with phorbol myristate acetate.
Journal of veterinary pharmacology and therapeutics    May 7, 2010   Volume 32, Issue 6 541-547 doi: 10.1111/j.1365-2885.2009.01077.x
Péters F, Franck T, Pequito M, de la Rebière G, Grulke S, Salccicia A, Verwilghen D, Chiavaccini L, Deby-Dupont G, Serteyn D.The previous experiments have shown that some phenothiazines have antioxidant and anti-inflammatory properties in vitro. In this study the inhibition of the production of reactive oxygen species (ROS) by neutrophils was studied in two groups of horses, which received a dose of 0.1 mg/kg of either acepromazine or promethazine intravenously. Blood samples were collected before (T0) and 0.5, 1, 3 and 5 h after drug administration. The chemiluminescence (CML) response of neutrophils was measured ex vivo in the presence of luminol for a period of 10 min and the maximum CML value (peak value) record...
Immunohistochemical analysis of MCT1 and CD147 in equine skeletal muscle fibres.
Research in veterinary science    May 4, 2010   Volume 89, Issue 3 432-437 doi: 10.1016/j.rvsc.2010.03.026
Mykkänen AK, Hyyppä S, Pösö AR, Ronéus N, Essén-Gustavsson B.Monocarboxylate transporter 1 (MCT1) and its ancillary protein CD147 facilitate efflux of lactate from the muscle. Expression of MCT1 and CD147 were studied with immunohistochemistry in type I, IIA, IIAB and IIB fibres of equine gluteal muscle. Staining intensity of MCT1 in the cytoplasm as well as in the membranes of fibre types decreased in the order I=IIA>IIAB>IIB and correlated with the oxidative capacity. Capillaries were pronounced in the MCT1 staining. CD147 antibody stained plasma membranes of all fibre types evenly, whereas the staining in the cytoplasm followed that of MCT1. In...
Expression of cyclooxygenase isoforms in ulcerated tissues of the nonglandular portion of the stomach in horses.
American journal of veterinary research    May 4, 2010   Volume 71, Issue 5 592-596 doi: 10.2460/ajvr.71.5.592
Rodrigues NL, Doré M, Doucet MY.To characterize the expression of the cyclooxygenase (COX)-1 and COX-2 isoforms in naturally occurring ulcers of the nonglandular portion of the stomach in horses. SPECIMEN POPULATION: 38 specimens from ulcerated stomachs and 10 specimens from healthy stomachs. Methods: Specimens were collected at an abbatoir; for each specimen of squamous gastric mucosa, 1 portion was fixed in neutral-buffered 10% formalin for immunohistochemical analysis and another was frozen at -70 degrees C for immunoblotting analysis. Immunoreactivity to 2 antibodies, MF241 (selective for COX-1) and MF243 (selective for ...
Mechanical loading of the distal end of the third metacarpal bone in horses during walking and trotting.
American journal of veterinary research    May 4, 2010   Volume 71, Issue 5 508-514 doi: 10.2460/ajvr.71.5.508
Merritt JS, Pandy MG, Brown NA, Burvill CR, Kawcak CE, McIlwraith CW, Davies HM.To assess the net mechanical load on the distal end of the third metacarpal bone in horses during walking and trotting. Methods: 3 Quarter Horses and 1 Thoroughbred. Methods: Surface strains measured on the left third metacarpal bone of the Thorough-bred were used with a subject-specific model to calculate loading (axial compression, bending, and torsion) of the structure during walking and trotting. Forelimb kinematics and ground reaction forces measured in the 3 Quarter Horses were used with a musculoskeletal model of the distal portion of the forelimb to determine loading of the distal end ...
Evaluation of the effect of phosphodiesterase on equine platelet activation and the effect of antigen challenge on platelet phosphodiesterase activity in horses with recurrent airway obstruction.
American journal of veterinary research    May 4, 2010   Volume 71, Issue 5 534-540 doi: 10.2460/ajvr.71.5.534
Dunkel B, Rickards KJ, Werling D, Page CP, Cunningham FM.To determine whether expression of equine platelet activation-dependent surface markers is influenced by phospodiesterase (PDE) isoenzyme activity and whether antigen challenge alters platelet PDE activity in horses with recurrent airway obstruction (RAO). Methods: 16 horses. Methods: 7 healthy horses were used for in vitro experiments, 6 horses with RAO were used for antigen challenge, and 6 healthy horses were used as control animals. Three of the healthy horses had also been used in the in vitro experiments. Effects of PDE inhibition and activation of adenylyl cyclase on CD41/61 and CD62P e...
Seasonal variation of plasminogen activator activity in spermatozoa and seminal plasma of boar, buck, bull and stallion.
Reproduction in domestic animals = Zuchthygiene    April 24, 2010   Volume 45, Issue 6 e440-e446 doi: 10.1111/j.1439-0531.2010.01597.x
Zervos IA, Lavrentiadou SN, Tsantarliotou MP, Georgiadis MP, Kokolis NA, Taitzoglou IA.Plasminogen activators (PA) are proteolytic enzymes present in the spermatozoa and seminal plasma of various species. They play a role in the binding of the spermatozoon and its penetration through the layers surrounding the oocyte. Plasminogen activator activity (PAA) is modulated by hormones that have a seasonal variation, such as testosterone and melatonin. The present study investigates the seasonal variation of PA activity in sperm extracts and seminal plasma of four farm animal species: boar, buck, bull and stallion. Semen samples were collected every second week during a 12-month period...
Heat shock protein 70 gene expression in equine blastocysts after exposure of oocytes to high temperatures in vitro or in vivo after exercise of donor mares.
Theriogenology    April 22, 2010   Volume 74, Issue 3 374-383 doi: 10.1016/j.theriogenology.2010.02.020
Mortensen CJ, Choi YH, Ing NH, Kraemer DC, Vogelsang MM, Hinrichs K.Heat above homeothermy can be detrimental to embryonic development, and cells may produce heat shock proteins to try to mitigate these effects. The authors examined the developmental competence of equine oocytes after a single heat exposure (42 degrees C, 2 or 4 h) during early or late stages of in vitro maturation. Rates of nuclear maturation, cleavage after intracytoplasmic sperm injection, and advanced embryonic development (morula or blastocyst) were compared to those for unexposed controls. Concentrations of heat shock protein 70 (HSPA1A) mRNA were determined by real-time RT-PCR in result...
Comparison of the antioxidant effects of synovial fluid from equine metacarpophalangeal joints with those of hyaluronic acid and chondroitin sulfate.
American journal of veterinary research    April 22, 2010   Volume 71, Issue 4 399-404 doi: 10.2460/ajvr.71.4.399
Müller AJ, Letelier ME, Galleguillos MA, Molina-Berríos AE, Adarmes HH.To evaluate the antioxidant effects of synovial fluid (SF) pooled from metacarpophalangeal joints of healthy horses and horses with various pathological conditions, and to compare then with the antioxidant effects of hyaluronic acid (HA) and chondroitin sulfate (CS). Methods: SF from 1 metacarpophalangeal joint was obtained from 42 horses immediately after humane slaughter. Samples were classified into 3 groups on the basis of origin: healthy joints or joints with chronically damaged cartilage or vascularly congested synovial membranes as detected via macroscopic evaluation. Methods: Antioxida...
Cellular localization of androgen synthesis in equine granulosa-theca cell tumors: immunohistochemical expression of 17alpha-hydroxylase/17,20-lyase cytochrome P450.
Theriogenology    April 22, 2010   Volume 74, Issue 3 393-401 doi: 10.1016/j.theriogenology.2010.02.022
Neto AC, Ball BA, Browne P, Conley AJ.Elevated blood testosterone concentrations, often accompanied by male-typical behaviors, is a common signalment of mares with granulosa-theca cell tumors (GCTCs), but no definitive information exists regarding the cellular differentiation of tumors associated with androgen secretion. This study was conducted to localize and thereby define the cellular expression of 17alpha-hydroxylase/17,20-lyase cytochrome P450 (P450c17), the enzyme most directly responsible for androgen synthesis, in 30 GTCTs and control tissues (gonads and adrenal glands) using immuno-histochemistry (IHC). Immuno-reactivity...
A pendulum test as a tool to evaluate viscous friction parameters in the equine fetlock joint.
Veterinary journal (London, England : 1997)    April 21, 2010   Volume 188, Issue 2 204-209 doi: 10.1016/j.tvjl.2010.03.016
Noble P, Lumay G, Coninx M, Collin B, Magnée A, Lecomte-Beckers J, Denoix JM, Serteyn D.An equine fetlock joint pendulum test was studied and the influence of post mortem time and intra-articular lipid solvent on the viscous frictional response examined. Fresh equine digits (group 1, n=6 controls; group 2, n=6 lipid solvent) were mounted on a pendulum tribometer. Assuming that pendular joint damping could be modelled by a harmonic oscillator fluid damping (HOFD), damping time (τ), viscous damping coefficient (c) and friction coefficient (μ) were monitored for 5h under experimental conditions (400N; 20°C). In all experiments, pendular joint damping was found to follow an expone...
Sex, science and satisfaction: a heady brew.
Animal reproduction science    April 21, 2010   Volume 121, Issue 1-2S S262-S278 doi: 10.1016/j.anireprosci.2010.04.178
Allen WR.Genus-specific reproductive processes and strategies displayed by equids, camelids and elephantids are compared and contrasted to illustrate the amazing diversity of reproductive physiology between genera and the equally surprising conservation of reproductive processes across a genus in the face of other dramatic phenotypic modifications and adaptation to prosper in the prevailing environment. From intrauterine conceptus mobility and chorionic gonadotrophic secretion by specialised invasive trophoblast cells in equids, through induced ovulation, an asymmetrical uterus and an almost absurdly s...
[Usual values in veterinary haematology].
Revue francophone des laboratoires : RFL    April 18, 2010   Volume 2010, Issue 420 27-42 doi: 10.1016/S1773-035X(10)70419-6
Bellier S, Cordonnier N.The goal of this article is to present the technical aspects and the applications of the blood cells examination and haemostasis in dogs and cats. Technical realisation of blood counts, cytology of normal and abnormal blood cells and significance of the pathologic cell counts are successively treated. Distinctive features of cytology and cell counts in herbivores, horses and cattle, are detailed in a separate chapter.Difficulties of veterinary haematology lie in qualitative and quantitative differences between blood cells of various animal species.Bone marrow cells exploration is not performed...
[The influence of pH on cholinesterase hydrolysis of alpha-naphthylacetate in the presence of some cationic detergents].
Ukrains'kyi biokhimichnyi zhurnal (1999 )    April 15, 2010   Volume 81, Issue 4 23-27 
Kuznetsova LP, Nikitina ER, Sochilina EE, Vasil'eva KA.The influence of some cationic detergents on the catalytic activity of the horse blood plasma cholinesterase in reaction of hydrolysis of alpha-naphthylacetate at different pH were investigated. It was shown, that in the absence of detergents in acid pH of the reaction medium the Km value increases, but V remain constant. In the range of pH from 8.5 to 5.0 in the presence of detergents the Km and V values are not practically changed. That is why the activation of cholinesterase hydrolysis of alpha-naphthylacetate in the presence of detergents is considerably higher than that of the neutral pH....
Effects of endotoxaemia and carbohydrate overload on glucose and insulin dynamics and the development of laminitis in horses.
Equine veterinary journal    April 14, 2010   Volume 41, Issue 9 852-858 doi: 10.2746/042516409x479027
Tóth F, Frank N, Chameroy KA, Bostont RC.Insulin resistance (IR) is a risk factor for pasture-associated laminitis in equids and alimentary carbohydrate overload may trigger laminitis. Whether glucose metabolism responses to carbohydrate overload are more pronounced in insulin-resistant horses requires further study. Objective: Horses pretreated with endotoxin to alter insulin sensitivity differ significantly in their glucose and insulin responses to carbohydrate overload. Methods: Horses (n=24) were divided into 3 groups. A lipopolysaccharide (LPS; n=8) group that received endotoxin as an 8 h 7.5 ng/kg bwt/h i.v. continuous rate inf...
Ultrasound B-scan image simulation, segmentation, and analysis of the equine tendon.
Medical physics    April 14, 2010   Volume 37, Issue 3 1038-1046 doi: 10.1118/1.3292633
Meghoufel A, Cloutier G, Crevier-Denoix N, de Guise JA.The hypothesis is that an imaging technique based on decompression and segmentation of B-scan images with morphological operators can provide a measurement of the integrity of equine tendons. Methods: Two complementary approaches were used: (i) Simulation of B-scan images to better understand the relationship between image properties and their underlying biological structural contents and (ii) extraction and quantification from B-scan images of tendon structures identified in step (i) to diagnose the status of the superficial digital flexor tendon (SDFT) by using the proposed imaging technique...
Metabolic evaluation of cooled equine spermatozoa.
Andrologia    April 14, 2010   Volume 42, Issue 2 106-111 doi: 10.1111/j.1439-0272.2009.00963.x
Vasconcelos AB, Santana MA, Santos AM, Santoro MM, Lagares MA.Microscopy has been used in the routine evaluation of sperm metabolism. Nevertheless, it has limited capacity to preview male fertility. As calorimetry may be used to evaluate directly the metabolic activity of a biological system, the aim of this study was to use microcalorimetry as an additive method for sperm metabolism evaluation of cooled equine semen. Two ejaculates of four stallions were collected and motility, viability (eosin 3%) and membrane functional integrity (hyposmotic swelling test) of spermatozoa were evaluated. Sperm samples were processed following different protocols and th...
Do horses suffer from irritable bowel syndrome?
Equine veterinary journal    April 14, 2010   Volume 41, Issue 9 836-840 doi: 10.2746/042516409x474284
Hunter JO.Irritable bowel syndrome (IBS) in man is not a single entity but has several causes. One of the most common forms has similarities with colic and laminitis in horses. Undigested food residues may pass from the small intestine into the colon where bacterial fermentation produces chemicals that lead to disease. In horses the consequences may be disastrous, but in healthy humans such malabsorption may not be harmful. After events such as bacterial gastroenteritis or antibiotic treatment, an imbalance of the colonic microflora with overgrowth of facultative anaerobes may arise, leading to malferme...
Acute laminitis: medical and supportive therapy.
The Veterinary clinics of North America. Equine practice    April 13, 2010   Volume 26, Issue 1 103-114 doi: 10.1016/j.cveq.2009.12.011
van Eps AW.Acute laminitis is a serious complication of many primary conditions in the horse. This article summarizes the most appropriate approach to management of the horse with acute laminitis, based on current information.
Carbohydrate alimentary overload laminitis.
The Veterinary clinics of North America. Equine practice    April 13, 2010   Volume 26, Issue 1 65-78 doi: 10.1016/j.cveq.2010.01.006
Pollitt CC, Visser MB.In acute laminitis, the suspensory apparatus of the distal phalanx fails at the lamellar dermal/epidermal interface. A grading system for the histopathology of laminitis is based on the consistent pattern of histologic changes to the secondary epidermal lamellae, basal cells, and basement membrane that occur as carbohydrate-induced laminitis develops. The actual trigger factors of carbohydrate-induced laminitis remain unidentified.