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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.
Pilot Study: Simultaneous Daily Recording of Total Locomotor Activity and Heart Rate in Horses for Application in Precision Livestock Farming.
Animals : an open access journal from MDPI    April 22, 2025   Volume 15, Issue 9 1189 doi: 10.3390/ani15091189
Aragona F, Rizzo M, Arrigo F, Arfuso F, Fazio F, Giudice E, Pugliatti P, Piccione G, Giannetto C.Among physiological parameters, total locomotor activity (TLA) and heart rate (HR) are used as physiological indicators in animal welfare evaluations. The present study aimed to simultaneously record for 24 h the TLA and HR of ten clinically healthy horses housed in conventional individual boxes subjected to a natural photoperiod and temperature. An actigraphy-based data logger was placed on the headstall, and an equine HR monitor was placed around the chest to monitor TLA and HR, respectively. Activity was monitored with 5 min sampling intervals and HR with 5 s intervals. To make the data poi...
Effects of clodronate disodium on endocrine regulators of calcium in yearling horses.
Journal of animal science    April 22, 2025   Volume 103 doi: 10.1093/jas/skaf132
Conrad MB, Leatherwood JL, Paris BL, George JM, Martinez RE, Vergara-Hernandez FB, Nielsen BD, Colbath AC, Arnold CE, Glass KG, Welsh TH, Bradbery AN.Extra-label bisphosphonate (BP) use in juvenile horses is anecdotally reported, primarily for analgesic effects, despite the limited scientific understanding of biologic impacts on skeletally immature horses undergoing exercise. The objective of this study was to determine the effects of clodronate disodium (CD), a form of BP, on endocrine regulators of calcium. Thus, 32 yearling Quarter Horses were stratified by age (500 ± 13 d), body weight (BW; 336 ± 26 kg), sex (n = 16 geldings, n = 16 fillies), and initial bone optical density into 1 of 4 treatment groups for a 168-d tr...
Evaluation of the Effects of Detomidine on Equine Myoelectrical Activity Using Electrointestinography.
Journal of veterinary emergency and critical care (San Antonio, Tex. : 2001)    April 20, 2025   Volume 35, Issue 2 120-130 doi: 10.1111/vec.13464
Munsterman AS, Dias Moreira AS, Kottwitz J.To evaluate the effects of detomidine on equine intestinal slow-wave activity and frequency distribution measured by electrointestinography (EIG). Methods: Prospective, experimental study. Methods: University teaching hospital. Methods: A convenience sample of twelve 7- to 21-year-old clinically normal horses. Methods: Horses were randomly assigned to saline control (four horses) or detomidine treatment (eight). After obtaining a 30-min baseline EIG, a saline or detomidine bolus was administered, followed by a constant rate infusion, and another EIG was recorded. Ultrasonographic examinations ...
The application of the pyramidal training model for conditioning thoroughbred horses.
Journal of equine veterinary science    April 9, 2025   Volume 149 105567 doi: 10.1016/j.jevs.2025.105567
Davie A, Beavers R, Denham J.The purpose of this review was to analyze the scientific background and practical application of a very successful human training methodology, the pyramidal training model, to the Thoroughbred racehorse. Despite years of research and accumulated knowledge on human training methods to enhance endurance performance, Thoroughbred training programs have lacked significant change. This review introduces the pyramidal training model, explains the science that underpins this training concept and outlines one approach to translate this science into the equine field. It also discusses the importance of...
Physiological and behavioral parameters of pain and stress in mares during and after transvaginal ultrasound-guided follicular aspiration.
Frontiers in veterinary science    April 9, 2025   Volume 12 1574351 doi: 10.3389/fvets.2025.1574351
Van den Branden E, Salamone M, Broothaers K, Peere S, Polfliet E, Dewulf M, Van Steenkiste G, van Loon G, Smits K, Govaere J.The use of transvaginal ultrasound-guided follicle aspiration (TVA) for oocyte collection has become a widely used procedure in horses for embryo production (IVEP). The TVA procedure is characterized by various manipulations, which are physical restraint, perineal preparation, transrectal palpation, insertion of an intravaginal device, and needle punctures. These repeated transvaginal and transovarian punctures have raised concerns about their potential effects on mare welfare. Our study aimed to investigate the effects of TVA manipulations, and especially puncturing, on pain and stress in ma...
The effects of a plant-based and a plant- and marine-based n-3 oil supplement on behavioural reactivity, heart rate variability, and plasma fatty acid profile in young healthy horses.
Journal of animal science    April 8, 2025   skaf117 doi: 10.1093/jas/skaf117
Hartwig S, Rankovic A, McCrae P, Gagliardi K, Burron S, Ellis J, Ma DWL, Shoveller AK.Behavioural reactivity in horses poses a welfare and safety risk to both the horse and the handler, however, beneficial effects have been observed when dietary fat is increased in replacement of sugar. Supplementation with the fatty acids (FA) eicosapentaenoic (EPA) and docosahexaenoic acid (DHA) appear to improve negative behaviours in rodents and humans, but the effect of α-linolenic acid (ALA), EPA, and DHA, specifically, on reactivity in horses is unknown. The objective of this study was to evaluate the effects of camelina oil (CAM; ALA-enriched) and a mix of camelina and algal oil (ALG; ...
Running a genetic stop sign accelerates oxygen metabolism and energy production in horses.
Science (New York, N.Y.)    March 28, 2025   Volume 387, Issue 6741 eadr8589 doi: 10.1126/science.adr8589
Castiglione GM, Chen X, Xu Z, Dbouk NH, Bose AA, Carmona-Berrio D, Chi EE, Zhou L, Boronina TN, Cole RN, Wu S, Liu AD, Liu TD, Lu H, Kalbfleisch T....Horses are among nature's greatest athletes, yet the ancestral molecular adaptations fueling their energy demands are poorly understood. Within a clinically important pathway regulating redox and metabolic homeostasis (NRF2/KEAP1), we discovered an ancient mutation-conserved in all extant equids-that increases mitochondrial respiration while decreasing tissue-damaging oxidative stress. This mutation is a de novo premature opal stop codon in KEAP1 that is translationally recoded into a cysteine through previously unknown mechanisms, producing an R15C mutation in KEAP1 that is more sensitive to ...
Effects of Age, Sex, and Exercise on Measurement of Serum CTnI Levels and Some Parameters Related to the Cardiovascular Capacity of Caspian Horses.
Veterinary medicine and science    March 11, 2025   Volume 11, Issue 2 e70202 doi: 10.1002/vms3.70202
Mehrazin H, Sakha M, Safi S.Due to their high specificity and exclusive cardiac myocyte sensitivity, cardiac troponins T and I (cTnT, cTnI) are currently regarded as ideal biomarkers to identify cardiomyocyte damage, myocardial injury, myocardial infarction, and chronic heart failure. In fact, cTnI is considered the most reliable biomarker for diagnosing heart-related issues. This study aimed to investigate the effects of age, gender, and exercise training on serum cTnI levels and various parameters related to the cardiovascular capacity of Caspian horses. For this purpose, 50 adult Caspian horses over 3 years old, both ...
Absolute Quantitative Lipidomics Reveals Differences in Lipid Compounds in the Blood of Trained and Untrained Yili Horses.
Veterinary sciences    March 10, 2025   Volume 12, Issue 3 255 doi: 10.3390/vetsci12030255
Wang T, Meng J, Wang J, Ren W, Yang X, Adina W, Bao Y, Zeng Y, Yao X.The purpose of this study was to explore the relationship between blood lipid levels and the differences in cardiac structure and function of trained and untrained Yili horses as related to exercise performance. We utilized quantitative lipidomics technology to elucidate how the differences in lipid compounds in the blood influenced performance outcomes. Sixteen 18-month-old Yili horses were selected, ten of which received a 15-week training regimen, and six were kept as untrained controls. Cardiac structure and function were assessed by echocardiography, while plasma lipid metabolites were de...
Short communication: “Gene expression patterns of the four cardiac chambers in the Thoroughbred horse”.
Journal of equine veterinary science    March 9, 2025   105415 doi: 10.1016/j.jevs.2025.105415
Edling CE, Arevalo-Turrubiarte M, Premont A, Uribe MC, Forbes B, Kemp V, Weir J, Marr C, Lewis R, Jeevaratnam K.Athlete horses' contraction and conduction of the healthy heart influences racing performance. Gene expression patterns in the horse heart are not yet fully investigated. We aimed to evaluate the gene expression from the four chambers of the heart overall and with focus on genes involved in the electrophysiology of the heart in Thoroughbred racehorses with no clinical cardiac abnormalities. Tissue was collected from the left atrium (LA), right atrium (RA), left ventricle (LV), and right ventricle (RV). Total RNA was analysed by microarray technique. We compared gene expression in the heart cha...
How facial expressions reveal acute pain in domestic animals with facial pain scales as a diagnostic tool.
Frontiers in veterinary science    March 4, 2025   Volume 12 1546719 doi: 10.3389/fvets.2025.1546719
Mota-Rojas D, Whittaker AL, Coria-Avila GA, Martínez-Burnes J, Mora-Medina P, Domínguez-Oliva A, Hernández-Avalos I, Olmos-Hernández A....The growing interest in managing and recognizing pain in animals has led to the search for more sensitive methods to evaluate it, especially because some species conceal any visible changes associated with pain or are not easily assessed. Research has shown that an animal's facial expression changes when exposed to painful stimuli. Thus, developing several pain scales (grimace scales) in species such as horses, cattle, pigs, sheep, donkeys, rabbits, rats, mice, and cats has helped to improve the study of pain in veterinary medicine. The possibility of using facial expression as an indicator of...
Investigation of the impact of seasonal climate conditions on feed intake and body weight in horses.
International journal of biometeorology    February 26, 2025   doi: 10.1007/s00484-025-02881-z
Giannetto C, Aragona F, Fazio F, Piccione G, Giudice E, Arfuso F, Zumbo A.This study aimed to examine the impact of climatic conditions on feed intake, body weight, and rectal temperature in horses residing in the hot Summer Mediterranean climate zone. Eight gelding Italian Saddle horses, with an average body weight of 427.78 ± 54.55 kg and aged between 10 and 13 years, participated in the research. All horses were individually housed in a paddock (800 m/horse), and fed ad libitum. Food intake, body weight, and rectal temperature were recorded every 10 days at four 30-day collection periods in the year: January, April, July, and October. The formula adapted fr...
In vivo effects of cold therapy and bandaging on core temperatures of equine superficial and deep digital flexor tendons.
Veterinary surgery : VS    February 25, 2025   Volume 54, Issue 3 470-477 doi: 10.1111/vsu.14235
McCarthy RD, Ordóñez HJ, Semevolos SA.To compare core temperatures of superficial and deep digital flexor tendons (SDFT, DDFT) during application of a compression cooling system versus ice boots and during subsequent lower limb bandaging. Methods: Experimental study. Methods: Paired forelimbs of six sedated horses. Methods: Thermocouples were placed in the core of SDFT and DDFT of both front limbs of each horse. A compression cooling system was applied to one front limb and an ice boot was applied to the other front limb for 60 min and then removed for 60 min. Standing bandages were then applied to both front limbs for 60 mi...
Facial expressions during compound interventions of nociception, conspecific isolation, and sedation in horses.
Scientific reports    February 13, 2025   Volume 15, Issue 1 5373 doi: 10.1038/s41598-025-89329-x
Lundblad J, Rhodin M, Hernlund E, Bjarnestig H, Hidén Rudander S, Haubro Andersen P.Facial expressions in prey animals such as equines can convey information regarding their internal state and are therefore often used as cues for welfare and pain evaluation. The state of pain is commonly compared to a neutral state with little consideration given to other experiences that could affect the animal, although this situation is rare outside of experimental contexts. To evaluate the effect of managerial contexts on facial expressions from a nociceptive input, conspecific isolation and sedation with known physiological effects were compared to compound states of nociception. Using a...
Heart rate variability responses of horses and veterans with post-traumatic stress disorder to ground-based adaptive horsemanship lessons: a pilot study.
Translational animal science    February 12, 2025   Volume 9 txaf019 doi: 10.1093/tas/txaf019
Rankins EM, Faremi BE, Hartmann K, Quinn A, Posada-Quintero HF, McKeever KH, Malinowski K.Heart rate variability (HRV) can be measured as an indicator of autonomic nervous system (ANS) balance and thus, stress and affective arousal. Mixed results have been reported in the limited literature addressing the effects of equine-assisted services (EAS) on HRV in the human and horse participants. The aims of the present study were to determine the effects of ground-based adaptive horsemanship (AH) lessons on veterans' and horses' HRV during weekly lessons as well as veterans' resting HRV outside of lessons. Veterans with post-traumatic stress disorder (PTSD) were randomly assigned to 8 w...
Evaluation of the Effect of Intravenous Mannitol and Hypertonic Saline on Intraocular Pressure and Biometry Parameters in Anesthetized Horses With Experimentally Increased Intracranial Pressure.
Veterinary ophthalmology    February 11, 2025   doi: 10.1111/vop.70000
Bercovitz GR, Sullivan SN, Reed RA, Ryan CA, Diehl KA.To evaluate the effect of intravenous mannitol and hypertonic saline (HS) on intraocular pressure (IOP) and biometry parameters of horses with elevated intracranial pressure (ICP). Methods: Seven horses with normal ophthalmic exams were anesthetized and placed in lateral recumbency. A subarachnoid transducer was placed, and the head was lowered until ICP measured 40 mmHg +/- 2 mmHg. Five intravenous boluses of mannitol 20% (0.4 g/kg) or HS 7.2% (1 mL/kg) were administered, each 15 min apart. A 2-day washout period ensued, and the procedure was repeated with the other hyperosmotic age...
The inability of spectral Doppler ultrasonography to identify stallions with testicular dysfunction.
Journal of equine veterinary science    February 4, 2025   105367 doi: 10.1016/j.jevs.2025.105367
Pozor MA, Macpherson ML, Kelleman AA, Smith H, McNaughten JW.This retrospective study aimed to validate the usefulness of spectral Doppler ultrasonography in diagnosing testicular dysfunction in stallions. The first part of the study included a pony stallion with age-related testicular degeneration (18 - 21 yrs. old). Testicular dimensions, spectral Doppler parameters of testicular blood flow, and semen parameters were obtained numerous times from this pony over four consecutive years. While testicular volume decreased dramatically, and the stallion became azoospermic, there were no changes in any of the spectral Doppler parameters of testicular blood f...
Therapeutic Exercises for Rehabilitation of Muscle Injury.
The Veterinary clinics of North America. Equine practice    January 31, 2025   Volume 41, Issue 1 213-225 doi: 10.1016/j.cveq.2024.11.011
King MR, Colla S.Successful rehabilitation of muscle injury requires a comprehensive understanding of the injury process, healing phases, and resources to be employed. The initial phase is characterized by acute inflammatory signs, followed by the regenerative and remodeling phases. Therapeutic exercises can be utilized in all 3 phases, progressing from isometric exercises to aquatic therapies. The classification and phase of injury and the individual response to the applied therapies will guide the progression of the therapeutic exercises through the rehabilitation program.
Effect of an adjustable ceiling to prevent premature rising attempts after general anesthesia in healthy ponies and horses: A pilot study.
Veterinary surgery : VS    January 30, 2025   Volume 54, Issue 5 851-859 doi: 10.1111/vsu.14181
Lindqvist A, Nyman G, Rydén A, Wattle O.This study aimed to assess and compare the recovery of ponies and horses following general anesthesia in two different settings: a recovery box with an inflexible, adjustable ceiling, and free recovery without restraints. Our primary objective was to evaluate the effect of adjustable ceilings on the prevention of premature attempts to rise during recovery. The secondary aim was to compare the physiological stress indicators during recovery. Methods: Prospective, randomized, experimental study. Methods: Six healthy ponies and 10 healthy horses. Methods: This study used a crossover design with t...
Clinical Examination of the Muscle System.
The Veterinary clinics of North America. Equine practice    January 29, 2025   S0749-0739(24)00087-7 doi: 10.1016/j.cveq.2024.10.001
McKenzie E.The equine muscle system is complex and prone to a large range of hereditary and acquired diseases that often have overlapping clinical signs with orthopedic, neurologic, and other disorders. Obtaining a clinical history that fully outlines the client concerns and any known information about the disorder and following with a comprehensive physical examination and screening clinical pathology tests are fundamental to achieve definitive diagnosis. A methodical and comprehensive approach aids accurate diagnosis and development of an optimal treatment and management plan. This article describes th...
The myometrial transcriptome changes in mares with endometrosis.
Scientific reports    January 25, 2025   Volume 15, Issue 1 3173 doi: 10.1038/s41598-025-86742-0
Drzewiecka EM, Molcan T, Sadowska A, Piotrowska-Tomala K, Słyszewska M, Dias GF, Skarżyński DJ, Szóstek-Mioduchowska A.Mares with endometrosis exhibit histological changes not only in the endometrium but also in the myometrium that suggest possible functional impairment. The molecular background of these changes is not well understood. We hypothesize that the transcriptomic profile of the mare myometrium varies depending on the degree of endometrosis in mares. Myometria were collected from mares in the mid-luteal phase of the estrous cycle with endometrium categories I, IIA, IIB, and III (∑n = 23), according to Kenney and Doig´s histopathological classification. Myometrial RNA was isolated and subjected...
Pulse oximetry at two sensor placement sites in conscious foals.
Acta veterinaria Scandinavica    January 23, 2025   Volume 67, Issue 1 6 doi: 10.1186/s13028-025-00794-w
Rossi HS, Mykkänen AK, Junnila JJT, Hyytiäinen HK.Pulse oximetry has not been thoroughly evaluated for assessment of oxygenation in conscious foals. Compared with invasive arterial blood sampling, it is a painless and non-invasive method for real-time monitoring of blood oxygen saturation. The aim of this prospective clinical study was to evaluate the usability, validity, and reliability of pulse oximetry at two measuring sites (lip and caudal abdominal skin fold) for blood oxygen saturation measurement in conscious foals with and without respiratory compromise. Thirty-two foals under one month of age were used. Nineteen foals had normal resp...
Dynamic Adaptation of Hematological Parameters, Albumin, and Non-Esterified Fatty Acids in Saddlebred and Standardbred Horses During Exercise.
Animals : an open access journal from MDPI    January 21, 2025   Volume 15, Issue 3 300 doi: 10.3390/ani15030300
The response to exercise following a rest period may vary among horse breeds based on the importance of the hematological and hematochemical profiles of athletic horses. Ten Standardbred and ten Italian Saddlebred mares were subjected to a 4-week training program after a seasonal rest, and hematological parameters (red blood cells-RBCs; hemoglobin-Hb; hematocrit-Hct; platelets-PLTs; platelet aggregation-AG; aggregation slope-Slope; fibrinogen-Fb), as well as Albumin (Alb) and non-esterified fatty acids (NEFAs), were analyzed. Blood samples were obtained each week during the training program fo...
The Challenge of Defining Laterality in Horses: Is It Laterality or Just Asymmetry?
Animals : an open access journal from MDPI    January 21, 2025   Volume 15, Issue 3 288 doi: 10.3390/ani15030288
Haussler KK, le Jeune SS, MacKechnie-Guire R, Latif SN, Clayton HM.The defining characteristic of laterality is the dominance of one side of the brain controlling specific functions of paired organs or on one side of the body. Structural and functional asymmetries are ubiquitous in horses and range from anatomical features (e.g., the length of long bones) to the gathering of sensory information (e.g., which eye is used to observe unfamiliar scenes) and motor functions (e.g., left-right differences in locomotion). There is a common tendency to assign observed structural or functional asymmetries to lateralization, which often involves more than a simple left-r...
A structured exercise regimen enhances autonomic function compared to unstructured physical activities in geriatric horses.
Scientific reports    January 20, 2025   Volume 15, Issue 1 2493 doi: 10.1038/s41598-025-86679-4
Sanigavatee K, Poochipakorn C, Huangsaksri O, Vichitkraivin S, Pakdeelikhit S, Chotiyothin W, Wongkosoljit S, Wonghanchao T, Chanda M.Older horses often show reduced autonomic responses, affecting their well-being. While regular exercise can help maintain autonomic function, the impact of structured exercise on geriatric horses is not well understood. A study involving 27 geriatric horses examined their autonomic modulation over 12 weeks under different activity levels. Horses were divided into three groups: (1) sedentary (SEL), (2) those participating in unstructured activities (RAT), and (3) those following a structured exercise regimen (SER). Results showed that the minimum and average heart rates decreased in the structu...
Soft palate angle and basihyoid depth increase with tongue size and with body condition score in horses.
Equine veterinary journal    January 2, 2025   doi: 10.1111/evj.14445
Talbot AM, Shanks-Boon H, Baldwin CM, Barnes H, Maddox TW.Obesity has been associated with human obstructive sleep apnoea and canine brachycephalic obstructive airway syndrome. The effect of body condition score (BCS) on structures of the oropharynx, nasopharynx and upper airway of the horse has not been investigated. Objective: To investigate the effect of BCS on tongue measurements, soft palate angle and basihyoid depth in horses. Methods: Retrospective, analytical, cross-sectional. Methods: Computed tomographic (CT) images of the head of 58 horses were assessed. DICOM viewing software was used to measure head length, basihyoid-skin depth, soft pal...
Impact of musical rhythm on blood, physiological and welfare parameters in stabled horses.
Scientific reports    December 28, 2024   Volume 14, Issue 1 31311 doi: 10.1038/s41598-024-82637-8
The aim of this study was to evaluate the effects of two styles of classical music, based on different tempos (BPM), on the physiological and blood parameters of horses during social isolation and restriction of movements. First experiment was carried out using nine horses of no defined breed, distributed in Control, Slow-tempo music and Moderate-tempo music .For social isolation and restriction of movement, the animals were housed daily in individual stalls for two hours and exposed to the stimuli for 60 min, and eye temperature, heart rate, and respiratory rate were assessed. The second exp...
The rhythm of horse gaits.
Annals of the New York Academy of Sciences    December 28, 2024   doi: 10.1111/nyas.15271
Laffi L, Raimondi T, Ferrante C, Pagliara E, Bertuglia A, Briefer EF, Gamba M, Ravignani A.What makes animal gaits so audibly rhythmic? To answer this question, we recorded the footfall sound of 19 horses and quantified the rhythmic differences in the temporal structure of three natural gaits: walk, trot, and canter. Our analyses show that each gait displays a strikingly specific rhythmic pattern and that all gaits are organized according to small-integer ratios, those found when adjacent temporal intervals are related by a mathematically simple relationship of integer numbers. Walk and trot exhibit an isochronous structure (1:1)-similar to a ticking clock-while canter is characteri...
A novel placement method of a calibration-free pH capsule for continuous wireless measurement of intragastric pH in horses.
Journal of veterinary internal medicine    December 23, 2024   Volume 39, Issue 1 e17273 doi: 10.1111/jvim.17273
Hodgson E, Thirouin M, Narayanan P, Romano TR, Wise J, Bond S.Current methods to measure intragastric pH in horses have limitations. A wireless capsule has been designed for continuous esophageal pH monitoring in humans. Objective: To (1) determine the feasibility and describe the methodology of measuring intragastric pH wirelessly in horses; and (2) determine attachment duration of the capsules. Methods: Eleven healthy adult horses. Methods: Capsules were attached to squamous and glandular gastric mucosa under gastroscopic guidance, using suture loops and 1 to 4 hemostasis clips. pH was continuously recorded using a wireless recorder in both fed and fas...
The effect of lysophosphatidic acid on myometrial contractility and the mRNA transcription of its receptors in the myometrium at different stages of endometrosis in mares.
BMC veterinary research    December 19, 2024   Volume 20, Issue 1 571 doi: 10.1186/s12917-024-04384-2
Piotrowska-Tomala KK, Szóstek-Mioduchowska A, Jonczyk AW, Drzewiecka EM, Wrobel MH, Hojo T, Ferreira-Dias G, Skarzynski DJ.Endometrosis (chronic degenerative endometritis) results in morphological changes in the equine endometrium and impairs its secretory function. However, the effect of this condition on the myometrium remains unclear. Lysophosphatidic acid (LPA) may affect female reproductive function and embryo transport by influencing uterine contractility through its receptors (LPARs). The objective of this study was to determine myometrial LPAR1-6 mRNA transcription, and the effects of LPA on myometrial contractions in mares with endometrosis during the mid-luteal and follicular phases of the estrous cycle....