Veterinary research in horses encompasses the study of diseases, health management, and medical treatments specific to equine species. This field investigates various aspects of horse health, including infectious diseases, metabolic disorders, and musculoskeletal conditions. Researchers focus on understanding the pathophysiology of equine ailments, developing diagnostic tools, and evaluating therapeutic interventions. The study of horse health also involves examining preventive measures such as vaccination protocols and nutritional management to promote overall well-being. This page collects peer-reviewed research studies and scholarly articles that explore the diverse areas of veterinary science related to horses, providing insights into disease mechanisms, treatment strategies, and advancements in equine healthcare.
Galuppo LD, Stover SM, Willits NH.To compare the biomechanical properties, in full limb preparations, of intact second phalanx and a simulated comminuted second phalangeal fracture stabilized with either two bone plates or a custom Y-plate. Methods: In vitro biomechanical assessment of intact limbs and of paired limbs with a simulated second phalangeal fracture stabilized by one of two fixation methods. Animal Population-Thirteen pairs of equine cadaveric forelimbs. Methods: A comminuted second phalangeal fracture was created in six paired cadaveric limbs. For each limb pair, the fracture was stabilized with two plates in one ...
Shiue Y-L , Millon LV, Skow LC, Honeycutt D, Murray JD, Bowling AT.The hypothesis that the conservation of sex-chromosome-linked genes among placental mammals could be extended to the horse genome was tested using the UCDavis horse-mouse somatic cell hybrid (SCH) panel. By exploiting the fluorescence in-situ hybridization (FISH) technique to localize an anchor locus, X-inactivation-specific transcript (XIST) on the horse X chromosome, together with the fragmentation and translocation of the X- and Y-chromosome fragments in a somatic cell hybrid panel, we regionally assigned 13 type I and 13 type II (microsatellite) markers to the horse X- and Y-chromosomes. T...
Chism PN, Latimer FG, Patton CS, Rohrbach BW, Blackford JT.To evaluate the tensile strength and wound morphology of the equine linea alba at intervals over 6 months after ventral median celiotomy. Methods: Linea alba tensile strength and wound morphology were determined at 2, 4, 8, 16, and 24 weeks postoperatively. Linea alba samples from 3 unoperated horses were used as controls. Methods: Eighteen adult horses, weighing 400 to 500 kg, 4 to 15 years old. Methods: Tensile strength and thickness of incised linea alba samples collected at 2, 4, 8, 16, and 24 weeks after ventral median celiotomy were compared with control linea alba samples. Additional sa...
McDï¿® LA, Stover SM, Taylor KT.To determine cyclic biomechanical properties of gap osteotomized adult equine tibiae stabilized with an equine interlocking nail (EIN). Methods: In vitro experimental biomechanical investigation. Methods: Thirteen adult equine cadaveric tibiae. Methods: Adult equine tibiae with transverse, midshaft, 1-cm gap osteotomies, stabilized with an equine interlocking nail, underwent cyclic biomechanical testing in vitro under axial compression, 4-point bending, and torsion. Different specimens were subjected to different load levels that represented estimated in vivo loads at 2 Hz for 740,000 cycles. ...
Sechin A, Deschamps JC, Lucia T, Aleixo JA, Bordignon V.We evaluated the effect of PMSG on the weaning-to-first service interval, total litter size and born alive litter size in swine. Four doses of PMSG (0, 500, 750 and 1,000 IU) were administered intramuscularly after weaning to sows at 3 different farms, grouped by parities (1, 2 and 3 or higher) and 2 distinct time periods. The associations among main effects and response variables were assessed by analysis of variance. Polynomial orthogonal terms were used to adjust the estimates of weaning-to-first service interval, total litter size and born alive litter size for the interaction effect of pa...
Cadario ME, Merritt AM, Archbald LF, Thatcher WW, LeBlanc MM.Intrauterine pressure was measured in 4 reproductively normal mares and 4 mares with delay in uterine clearance after administration of oxytocin to determine if intrauterine pressure varied between dosage and group. Changes in intrauterine pressure were measured during estrus, when a follicle was > or =35 mm, using a Millar "Mikro-tip" catheter that had 3 discrete pressure sensors/channels. Mares received 4 different treatments of 10, 5, 2.5 or 0 IU (vehicle) of oxytocin. The protocol for each treatment consisted of a 10-min baseline recording, administration of treatment and measurement of...
Bedford SJ, Gowdy HL, Hinrichs K.This study was designed to compare the effects of different media and containers on longevity of motility of spermatozoa during in vitro incubation at 38 degrees C in either air or 5% CO2 atmosphere. Three ejaculates were collected from each of 4 stallions. The media tested were skim milk-glucose, modified Krebs/Ringer and Hank's salts solution for incubation in an air atmosphere, and modified Krebs/Ringer and Brackett and Oliphant (BO) defined medium for incubation in a 5% CO2 atmosphere. All samples were incubated in 5-mL borosilicate glass tubes filled with 3 mL of extended spermatozoa, 5-m...
Davies PC.An 11-year-old mare presented with neuromuscular deficits and what resembled shivering in the left hind limb. On necropsy, there was no evidence of denervation atrophy of the left hind gastrocnemius muscle. The spinal cord had a small, right-sided lesion at C3-C4 and C4-C5. Tests for equine herpesvirus-1 and Sarcocystis spp. were negative.
Sasaki N, Yoshihara T.Seven Thoroughbred horses were laparotomized and Force Transducers were fixed on the proximal jejunal and cecal serosa. After observation of the digestive tract motility in consciousness, cisapride (0, 0.5, 0.75 or 1 mg/kg) was orally administered. In horses treated with 0.75 mg/kg or 1.0 mg/kg cisapride, the migrating contraction (MC) of the jejunum was significantly increased in frequency.
Leblond A, Villard I, Leblond L, Sabatier P, Sasco AJ.Epidemiological studies should allow comparisons to be made of the prevalence of disease in populations from different countries, but the population characteristics and health problems in French horses are not well established. We have conducted a retrospective evaluation of the causes of death and vital characteristics of insured horses in France for the year 1995, with a view to comparison with published data from other countries. Files on 448 deceased horses were provided by nine insurance companies. Most of the animals were used for breeding (60%), followed by leisure (20%), eventing and r...
Provitera P, Bouamr F, Murray D, Carter C, Scarlata S.Human immunodeficiency virus (HIV) and equine infectious anemia virus (EIAV) are closely related lentiviruses that infect immune cells, but their pathogenesis differ. Localization to the cytosolic leaflet of the plasma membrane is critical for replication of both viruses. This localization is accomplished through the matrix (MA) domain of the Gag precursor protein. In HIV-1, association of MA to anionic membranes appears to be primarily driven by a linear cluster of basic residues in the MA domain and an N-myristoylation signal. Interestingly, the MA protein of EIAV does not contain either of ...
Rosenstein DS, Bowker RM, Bartlett PC.To describe the vascular anatomy of the palmar digital artery and its major branches in the equine foot and to quantify the diameter of these vessels by use of digital angiograms. Sample Population-6 thoracic limbs obtained from 6 horses. Methods: Distal portions of each limb were perfused with aerated Krebs-Henseleit solution. Digital angiograms were acquired in standing and lateral recumbent positions, following an intra-arterial injection of iopamidol. Select vessels were measured on radiographic views, and values were corrected for magnification. Results: The palmar digital artery tapered ...
Goodrich LR, Moll HD, Crisman MV, Lessard P, Bigbie RB.To evaluate healing of pinch-grafted wounds on the distal aspect of the limbs of ponies bandaged with equine amnion or a standard nonadherent wound dressing material. Methods: 6 ponies. Methods: A 2.5x2.5-cm full-thickness section of skin was removed from the dorsal aspect of each limb at the midpoint of the metacarpus or metatarsus. Six days later, wounds were grafted with partial-thickness pinch grafts. Half the wounds were bandaged with amnion, and the other half were bandaged with a nonadherent dressing. Bandages were changed every 3 days until wound healing was complete. At each bandage c...
McDï¿® LA, Stover SM, Coleman K.To determine a range of limb loading activity for healthy adult horses confined to box stalls in an equine veterinary teaching hospital and determine the effects of hospital environmental factors on load rates and daily limb loading patterns. Methods: 6 mature healthy horses of various ages, breeds, and sexes, and 1 horse with a repaired metatarsal fracture. Methods: Step monitors were placed on 2 limbs of adult horses confined to box stalls. Relocation steps and weight shifts were recorded, as loading events, for 24 hours. Influence of forelimb versus hind limb and environmental factors on lo...
Asperio RM, Marzola P, Sbarbati A, Osculati F, Addis F.To analyze the tendon of the deep digital flexor (TDDF) muscle of the forelimb in horses by use of a contrast radiographic agent (gadopentate dimeglumine [Gd-DTPA/Dimeg]) and magnetic resonance imaging (MRI) and to determine the concentration of water protons in the tendons by use of MRI. Methods: 8 TDDF harvested from the forelimbs of 6 horse cadavers. Methods: Examinations were performed on the same portion of each tendon. Tendons were examined by use of two techniques: MRI before and after treatment with Gd-DTPA/Dimeg as well as scanning electron microscopy. Results: Tendons did not have de...
Watson JL, Jackson KA, King DP, Stott JL.Expression of the low-affinity IgE receptor (CD23) on the surface of mononuclear cells is a critical event in the development of IgE-mediated immunologic responses. Using PCR and cDNA library screening, a 2.7kb cDNA encoding equine CD23 and a 627bp PCR fragment of cattle CD23 were sequenced and characterized. The equine CD23 sequence encodes a complete and continuous open reading frame of 327 amino acids, while the shorter cattle fragment encodes 209 amino acids corresponding to nucleotides 325-1098 of the equine CD23 transcript. In addition to high identities in their nucleotides and translat...
Reinemeyer CR, Scholl PJ, Andrews FM, Rock DW.A 3 m, video gastroscope was used to screen 47 horses suspected of being naturally infected with equine bot larvae. 17 of 47 (36.2%) candidate horses harbored Gasterophilus nasalis larvae in the proximal duodenum and 46 of 47 (97.9%) had G. intestinalis larvae in the stomach. All horses infected with G. nasalis had concurrent infections with G. intestinalis. 14 horses with dual infections were allocated randomly to two treatment groups. Seven horses in Group 1 received 2% moxidectin oral gel once at a dosage of 0.4 mg/kg bodyweight (BW), and seven horses in Group 2 were untreated controls. 14 ...
Sandin A, Andrews FM, Nadeau JA, Nilsson G.Nervous excitation was induced by various means in horses provided with a gastric cannula. Insulin hypoglycaemia profoundly inhibited the basal acid output and volume secreted from the stomach. No clear effect on acid secretion was noted after administration of bethanechol, as the acid output was covered by the copious secretion of saliva. Atropine almost abolished the basal acid output. Sensoric stimulation by teasing caused a slight but not significant increase in the total acid output. These data suggest that cholinergic excitation might play a role in the stimulation of both volume and aci...
Stephen JO, Baptiste KE, Townsend HG.To describe clinical signs and clinicopathologic findings in donkeys with hypothermia. Methods: Retrospective study. Methods: 10 hypothermic donkeys. Methods: Information on signalment, history, physical examination findings, results of diagnostic tests, treatments, and necropsy findings was extracted from medical records of all donkeys with hypothermia between 1988 and 1998 and compared with information from medical records of all normothermic donkeys and hypothermic horses admitted to the hospital during the same period. Results: Donkeys were more likely to be hypothermic than horses. The me...
Molenkamp R, Rozier BC, Greve S, Spaan WJ, Snijder EJ.Equine arteritis virus (EAV), the type member of the family Arteriviridae, is a single-stranded RNA virus with a positive-stranded genome of approximately 13 kb. EAV uses a discontinuous transcription mechanism to produce a nested set of six subgenomic mRNAs from which its structural genes are expressed. We have generated the first documented arterivirus defective interfering (DI) RNAs by serial undiluted passaging of a wild-type EAV stock in BHK-21 cells. A cDNA copy of the smallest DI RNA (5.6 kb) was cloned. Upon transfection into EAV-infected BHK-21 cells, transcripts derived from this clo...
van Herwerden L, Gasser RB, Blair D.This study investigated sequence heterogeneity in the first internal transcribed spacer (ITS-1) of ribosomal DNA within and among species and strains of Echinococcus. Different ITS-1 sequence variants exist in Echinococcus granulosus and Echinococcus multilocularis, which represent at least four evolutionary lineages: (1) a sheep strain-lineage of E. granulosus, (2) a sister lineage of a cervid and camel E. granulosus ITS-1 variants, (3) a lineage including the ITS-1 variants representing horse, bovine and camel strains of E. granulosus, as well as variants from E. multilocularis, Echinococcus...
Vatistas NJ, Snyder JR, Carlson G, Johnson B, Arthur RM, Thurmond M, Zhou H, Lloyd KL.Although gastric ulcers have been identified relatively frequently in racing Thoroughbreds, there have been no large scale studies to determine their effect on health and performance. Two hundred and two Thoroughbred horses in active race training were selected by the attending veterinarians for gastro-endoscopic examination. Images of the stomach mucosa were stored in a digitised format for subsequent evaluation. The number of ulcers and a score of severity were determined. Gastric ulceration of the squamous mucosa was identified in 82% of horses. Seventy-three (39%) horses displayed clinical...
Argenzio RA.Ulceration of the nonglandular, stratified squamous mucosa of the equine and porcine stomach is a common event in both species, although in pigs the fatality rate is significant and it is an economically important disease. Because the barrier function of this mucosa in horses and pigs appears similar, it is probable that similar pathophysiological mechanisms may be responsible for the initiating lesions and reparative events. Recent studies of ulcer pathogenesis in the pig have shown that feed preparation or prolonged fasting can result in disruption of the normal stratification of gastric con...
Vatistas NJ, Sifferman RL, Holste J, Cox JL, Pinalto G, Schultz KT.Gastric ulceration is a prevalent condition of racehorses. A number of models of gastric ulceration have been described, but none mimic the conditions of a horse in training. The objectives of this study were to determine whether gastric ulcers could be induced and maintained in a group of horses in simulated race training. In addition, serum cortisol was measured on a weekly basis to investigate the possibility that stress may be important in the pathogenesis of gastric ulceration. Thirty horses used in the trial were fed Bermuda grass hay and 6 kg of a concentrate diet, and exercised 6 days/...
Aggarwal N, Holmes MA.Despite the importance of IgG Fc receptors in the regulation of various immunological mechanisms, these receptors have not been well characterised in the domesticated animals including equines. This paper describes the production of two monoclonal antibodies (CVS 59 and CVS 61) that recognise equine IgG Fc receptors. Fusions were conducted using BALB/c mice hyperimmunised with equine peripheral blood mononuclear cells. Hybridoma supernatants were screened on the basis of their ability to inhibit the rosetting of equine antibody coated sheep erythrocytes with equine peripheral blood mononuclear...
Becker AC, Stock KF, Distl O.Detailed movement evaluations of warmblood foals and mares were performed in connection with regular breeding events of the Oldenburg horse breeding societies in 2009 and 2010. Unfavourable movement characteristics considered indicative for impaired balance were noted by a special judge (SJ) and the regular judges of the breeding events (RJ) and served as the basis for definition of new movement traits. Detailed movement information on 3374 foals and 2844 mares showed that more severe findings like irregular motion pattern in hind legs or irregularity in general motion pattern occurred only sp...
Earley ET, Reiswig JD.Odontoplasty (floating and occlusal equilibration) is the most commonly performed procedure in equine dentistry. From an anatomic perspective, an irregular occlusal surface, prominent cingula, transverse ridges, and enamel points all contribute to the function, form, and longevity of the equine cheek tooth. With limited reserve crown available and an average functional life range between 18 and 25Â years, removal of tooth structure should be conservative. The authors consider a quality oral examination to be the most important dental procedure performed in the horse. Individual tooth evaluatio...
Aboashia FA, Alatrag F, Elmarimi A.Hematological and biochemical values are widely used in veterinary clinics for disease prognosis, nutritional and therapeutic monitoring, as well as in understanding the disease process in farm animals, including equines. This study aims to assess the alterations in hematological and biochemical parameters in pure Arabian horses infested with internal parasites. Samples of feces and blood were collected from 20 adult mares. Fecal samples were proceeded by flotation test. The blood samples were analyzed for hematological and biochemical parameters to determine their means ± standard error (M Â...
Collier MA, Brighton CT, Norrdin R, Twardock AR, Rendano VT.The effect of a 20-microA direct-current implantable bone growth stimulator (BGS) on bone production with a "gap healing" model in the horse was evaluated. The right and left 4th metatarsal bones (Mt-4) were used in 7 adult horses to create the "gap healing" model. A 4-mm section of the Mt-4 bone was resected bilaterally in each horse. The BGS was surgically placed into the 7 left Mt-4 defects. The 7 right Mt-4 defects served as controls. Six horses survived the 16-week experimental period. Signs of pain, decreased range of limb motion, or lameness was not observed in any animal during the 16 ...
Kirkpatrick JF, Liu IM, Turner JW, Bernoco M.Twenty-six free-roaming feral mares were immunized against porcine zonae pellucidae (PZP) between February and May, 1988. Eight sexually mature mares received 2 inoculations 2 weeks apart, and 18 mares received 3 inoculations at intervals of 2 and 4 weeks. Analysis of urinary oestrone conjugates (E1C) and non-specific progesterone metabolites (iPdG) in samples collected in October, 1988, revealed that none of the 18 mares that received 3 and only 1 of the 6 mares that received two inoculations were pregnant, whereas 3 of 6 sham-injected control mares and 5 of 11 untreated mares were pregnant. ...
Hamm D, Willeford KO, White G, Reed SM, Hamm J.Suppurative lower airway disease is a common debilitating disease in performance horses and, while rarely fatal, is often recalcitrant to conventional therapy. A variety of treatments have been used to combat this condition and we conducted two types of studies to determine if caprine serum fraction--immunomodulator (CSFI), a nonspecific immunomodulator, improved recovery from lower respiratory disease. Two dose response studies were performed to ascertain the efficacy of CSFI. Horses were maintained daily on conventional antibiotic therapy. Respiratory tract exudate, nasal discharge, dyspnoea...
Hubbell JA, Muir WW.The rate of rise of arterial partial pressure of carbon dioxide (PaCO2) was determined in 49 apneic halothane-anesthetized horses following controlled ventilation. Drugs given for induction of anesthesia did not affect the rapid rate of rise of PaCO2 during the first minute after controlled ventilation, the PaCO2 at 1 minute after controlled ventilation, or the PaCO2 at which spontaneous ventilation began. Horses given xylazine-ketamine for induction of anesthesia had a significantly (P less than 0.05) faster rate of rise of PaCO2 after 1 minute following controlled ventilation than did horses...
Grabowska A, Kozdrowski R.Progesterone plays a crucial role in the maintenance of pregnancy from conception to about 100-120 days of gestation when placenta becomes the main source of gestagens. The aim of the study was to test progesterone concentration 14 days after ovulation in pregnant mares and relate it to peak estral endometrial edema and the presence of intrauterine fluid (IUF) after artificial insemination (AI), the number of treatments against IUF, and the time from AI to the day when the uterus was found free of fluid. Results: Mares were divided into two groups: group A (n = 13; age 10.8 ± 4.5 y...
Prosser CG, McLaren RD.Ligand blotting analysis of serum from the horse using radiolabelled IGF-I revealed a protein at 96 kDa which was not present in serum from goat, cow, sheep, deer or donkey. These latter species all displayed five labelled bands in the range 24 to 41 kDa. Conversely, these were only weakly labelled in serum from the horse. Size exclusion chromatography of horse serum pre-incubated with radiolabelled IGF-I revealed reduced binding in the 130-kDa peak compared with goat plasma, and ligand blotting analysis indicated the 96-kDa protein was present in this peak. The 96-kDa protein from horse serum...
Haywood PE, Teale P, Moss MS.Thoroughbred geldings were fed racehorse cubes containing a predetermined concentration of theobromine in the form of cocoa husk. They were offered 7 kg of cubes per day, divided between morning and evening feed, and food consumption was monitored. Urinary concentrations of theobromine were determined following the consumption of cubes containing 11.5, 6.6, 2.0 and 1.2 mg per kg of theobromine, to verify whether or not such concentrations would produce positive urine tests. Pre-dose urine samples were collected to verify the absence of theobromine before each experiment. It became apparent fro...
Bowker RM, Rockershouser SJ, Linder K, Vex KB, Sonea IM, Caron JP.The innervation of the navicular bone (os sesamoideum distale) and its suspensory ligaments (ligamenta sesamoidea collateralia) (CSL) or proximal suspensory ligament and the ligamentum sesamoideum distale impar or the distal sesamoidean impar ligament (DS-impar ligament) was examined using combined anatomical techniques of silver impregnation and immunocytochemistry. Silver impregnation studies revealed an abundance of nerve fibres present in both the CSL and DS-impar ligament with the latter having relatively more nerve fibres. These silver-impregnated nerves coursed parallel to and were asso...
Sobral GG, Neto OCG, da Silva AM, Carneiro GF.The aim of the present study is to evaluate the correlation between the total protein measured by an optical refractometer and the concentration of IgG by radial immunodiffusion (RID) to determine the performance of the optical refractometer to diagnose the failure of passive transfer of immunity (FPTI) in 12-hour-old foals. Blood was collected from foals (n = 30) 12 hours after birth. A study was carried out to measure the serum IgG concentration by RID test and measure total protein (TP) by optical refractometer. The correlation coefficient was measured between the TP concentration and the...
De Silva M, Tagliavia C, Galiazzo G, Gifuni G, Caiazza M, Chiocchetti R, Grandis A.The morphometric studies of the atrioventricular valves are still limited in the horse. Objective: To investigate the anatomy of the atrioventricular valves in the horse, focusing on the morphometric features of the valvular leaflets and the tendinous cords. We hypothesised that accessory leaflets occur commonly and exist as independent structures in the atrioventricular valves of the horse. Methods: Descriptive anatomical study. Methods: Twenty normal hearts from slaughtered half-bred horses were used. The cardiac weight and circumference were recorded. The atrioventricular valves were expose...
Kitzman JV, Wilson RC, Booth NH, Hendricks HL, Bush PB.The pharmacokinetics of 4-aminopyridine (4-AP), a drug capable of antagonizing nondepolarizing neuromuscular blocking drugs, as well as several classes of injectable sedative and anesthetic agents, were studied in 6 intact, awake horses. Plasma samples were assayed for 4-AP over a frequent sampling schedule for 8 hours after IV administration. The plasma 4-AP vs time data best fit a 2-compartment pharmacokinetic model. Distribution half-life was 7.4 minutes, elimination half-life was 259 minutes, volume of the central compartment was 0.89 L/kg, volume of distribution (area) was 1.98 L/kg, volu...
Pezzanite LM, Griffenhagen GM, Bass L, Okudaira M, Larson B, Hendrickson DA.Liposomal local anaesthetic solutions may provide extended-duration analgesia postoperatively but have not been assessed following intra-peritoneal local infiltration in any species. Objective: To evaluate two doses of 1.33% liposomal bupivacaine (LB) versus 0.75% bupivacaine HCL (BHCl) for analgesia following laparoscopic ovariectomy in mares. Methods: Prospective cohort study. Methods: Fifteen healthy Quarter Horse mares (age 2-20 years) with normal bilateral ovarian palpation and appearance were enrolled. Horses were restrained in standing stocks and administered an α-2 agonist, butorpha...
Vatistas NJ, Wright IM, Dyson SJ.Forty-eight horses with osteochondritic lesions of the femoropatellar joint were treated, 23 of them by an arthrotomy and 25 by arthroscopy. Follow-up information was obtained from either the owner or the referring veterinarian by telephone inquiry. There were no statistical differences between the groups of horses undergoing the two procedures with regard to age, sex, breed, the involvement of one or both limbs, the size of the lesion, and the duration of either the surgery or anaesthesia. However, the horses treated by an arthrotomy spent 14.5 days in hospital after the operation whereas tho...
Kwak ML, Schubert J.Although forensic parasitology remains in its infancy compared with more developed fields like forensic entomology, parasites can be useful forensic indicators. We present the case of an individual who disobeyed animal health legislation and crossed animal quarantine boundaries and two state lines with a cattle tick (Rhipicephalus australis) infested horse. Following recognition of the infestation by animal health officers, the individual asserted that the infestation had occurred either during transport or upon arrival at the site of discovery, and that they had not contravened quarantine leg...
Weers JG, Maki AH.Triplet-singlet energy transfer has been studied in the complex formed between auramine O (AO) and horse liver alcohol dehydrogenase with optically detected magnetic resonance (ODMR) spectroscopy. The results show that Trp-15 and Tyr residues transfer triplet energy mainly by a trivial process, whereas Trp-314 transfers triplet energy by a Förster process with two observed lifetimes at 77 K of 170 and 50 ms. The different Förster energy-transfer lifetimes are ascribed either to quenching of the two Trp-314 residues of the dimer by a single asymmetrically bound AO or to two distinct conformat...