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Topic:Equine Health

Equine health encompasses the study and management of diseases, disorders, and overall well-being of horses. It involves understanding various physiological systems, preventive care, and treatment strategies to maintain optimal health in equine populations. Common areas of focus include nutrition, infectious diseases, orthopedic conditions, and reproductive health. Research in equine health aims to advance knowledge on diagnostic methods, therapeutic interventions, and management practices that improve horse welfare and performance. This page collects peer-reviewed research studies and scholarly articles that explore the diverse aspects of equine health, offering insights into current findings and advancements in the field.
The effect of time of insemination with fresh cooled transported semen and natural mating relative to ovulation on pregnancy and embryo loss rates in the mare.
Reproduction in domestic animals = Zuchthygiene    November 30, 2010   Volume 46, Issue 4 678-681 doi: 10.1111/j.1439-0531.2010.01728.x
Newcombe JR, Cuervo-Arango J.One hundred and fifty-four mares were inseminated with fresh semen either during the pre- or post-ovulatory periods at different intervals relative to ovulation: 36-24 h (n = 17) and 24-0 h (n = 30) before ovulation; 0-8 h (n = 21), 8-16 h (n = 24), 16-24 h (n = 48) and 24-32 h (n = 14) h after ovulation. All mares received the same routine post-mating treatment consisting of an intrauterine infusion with 1 litre of saline and antibiotics followed 8 h later by an intravenous administration of oxytocin. Artificial inseminations (AI) from 36 h before ovulation up to 16 h post-ovulation were perf...
Equine influenza virus: a jumping virus that races with Thoroughbred horses and greyhounds.
Veterinary journal (London, England : 1997)    November 30, 2010   Volume 189, Issue 1 3-4 doi: 10.1016/j.tvjl.2010.10.023
Murcia PR, Wood JL.No abstract available
Borna disease virus antibody and RNA from peripheral blood mononuclear cells of race horses and jockeys in Korea.
Psychiatry investigation    November 27, 2010   Volume 8, Issue 1 58-60 doi: 10.4306/pi.2011.8.1.58
Song JW, Na KS, Tae SH, Kim YK.During the last two decades, Borna disease virus (BDV) has received much attention as a possible zoonotic agent, particularly as a cause of psychiatric disease. Although several studies have shown that BDV is present in Asia, BDV has not been detected in Korea. This study was designed to further investigate the presence of BDV infection in Korea. Methods: Blood samples were taken from 39 race horses and 48 jockeys. Antibody to BDV was detected by indirect immunofluorescence antibody test and RNA of BDV by real time reverse transcriptase PCR (rRT-PCR). Results: No evidence of BDV was detected i...
Use of pressor therapy in 34 hypotensive critically ill neonatal foals.
Australian veterinary journal    November 26, 2010   Volume 88, Issue 12 472-477 doi: 10.1111/j.1751-0813.2010.00652.x
Dickey EJ, McKenzie H, Johnson A, Furr MO.Arginine vasopressin (AVP) is used in human medicine in the management of vasodilatory shock and cardiac arrest, but it is not widely used in equine neonatal intensive care because of concerns about potential side effects and suboptimal efficacy. This retrospective study reports the clinical use of AVP and norepinephrine (NE) in foals with refractory hypotension. Objective: To report the cardiovascular responses and fluid balance in critically ill, hypotensive foals receiving either NE or AVP. Methods: The medical records of neonatal foals (<7 days of age) from 2000 to 2007 admitted to the ...
Comparative human-horse sequence analysis of the CYP3A subfamily gene cluster.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 72-79 doi: 10.1111/j.1365-2052.2010.02111.x
Schmitz A, Demmel S, Peters LM, Leeb T, Mevissen M, Haase B.Cytochrome P450 enzymes (CYP450s) represent a superfamily of haem-thiolate proteins. CYP450s are most abundant in the liver, a major site of drug metabolism, and play key roles in the metabolism of a variety of substrates, including drugs and environmental contaminants. Interaction of two or more different drugs with the same enzyme can account for adverse effects and failure of therapy. Human CYP3A4 metabolizes about 50% of all known drugs, but little is known about the orthologous CYP450s in horses. We report here the genomic organization of the equine CYP3A gene cluster as well as a compara...
Microarray analysis after strenuous exercise in peripheral blood mononuclear cells of endurance horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 166-175 doi: 10.1111/j.1365-2052.2010.02129.x
Capomaccio S, Cappelli K, Barrey E, Felicetti M, Silvestrelli M, Verini-Supplizi A.It is known that moderate physical activity may have beneficial effects on health, whereas strenuous effort induces a state resembling inflammation. The molecular mechanisms underlying the cellular response to exercise remain unclear, although it is clear that the immune system plays a key role. It has been hypothesized that the physio-pathological condition that develops in athletes subjected to heavy training is caused by derangement of cellular immune regulation. The purpose of the present study was to obtain information on endurance horse gene transcription under strenuous conditions and t...
Molecular heterogeneity of XY sex reversal in horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 41-52 doi: 10.1111/j.1365-2052.2010.02101.x
Raudsepp T, Durkin K, Lear TL, Das PJ, Avila F, Kachroo P, Chowdhary BP.Male-to-female 64,XY sex reversal is a frequently reported chromosome abnormality in horses. Despite this, the molecular causes of the condition are as yet poorly understood. This is partially because only limited molecular information is available for the horse Y chromosome (ECAY). Here, we used the recently developed ECAY map and carried out the first comprehensive study of the Y chromosome in XY mares (n=18). The integrity of the ECAY in XY females was studied by FISH and PCR using markers evenly distributed along the euchromatic region. The results showed that the XY sex reversal condition...
Genome-wide association analysis of osteochondrosis of the tibiotarsal joint in Norwegian Standardbred trotters.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 111-120 doi: 10.1111/j.1365-2052.2010.02117.x
Lykkjen S, Dolvik NI, McCue ME, Rendahl AK, Mickelson JR, Roed KH.Osteochondrosis (OC), a disturbance in the process of endochondral ossification, is by far the most important equine developmental orthopaedic disease and is also common in other domestic animals and humans. The purpose of this study was to identify quantitative trait loci (QTL) associated with osteochondrosis dissecans (OCD) at the intermediate ridge of the distal tibia in Norwegian Standardbred (SB) using the Illumina Equine SNP50 BeadChip whole-genome single-nucleotide polymorphism (SNP) assay. Radiographic data and blood samples were obtained from 464 SB yearlings. Based on the radiographi...
Genetics of swayback in American Saddlebred horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 64-71 doi: 10.1111/j.1365-2052.2010.02108.x
Cook D, Gallagher PC, Bailey E.Extreme lordosis, also called swayback, lowback or softback, can occur as a congenital trait or as a degenerative trait associated with ageing. In this study, the hereditary aspect of congenital swayback was investigated using whole genome association studies of 20 affected and 20 unaffected American Saddlebred (ASB) Horses for 48,165 single-nucleotide polymorphisms (SNPs). A statistically significant association was identified on ECA20 (corrected P=0.017) for SNP BIEC2-532523. Of the 20 affected horses, 17 were homozygous for this SNP when compared to seven homozygotes among the unaffected ho...
Morphological variation in the horse: defining complex traits of body size and shape.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 159-165 doi: 10.1111/j.1365-2052.2010.02127.x
Brooks SA, Makvandi-Nejad S, Chu E, Allen JJ, Streeter C, Gu E, McCleery B, Murphy BA, Bellone R, Sutter NB.Horses, like many domesticated species, have been selected for broad variation in skeletal size. This variation is not only an interesting model of rapid evolutionary change during domestication, but is also directly applicable to the horse industry. Breeders select for complex traits like body size and skeletal conformation to improve marketability, function, soundness and performance in the show ring. Using a well-defined set of 35 measurements, we have identified and quantified skeletal variation in the horse species. We collected measurements from 1215 horses representing 65 breeds of dive...
Systemic treatment with high dose of flunixin-meglumine is able to block ovulation in mares by inducing hemorrhage and luteinisation of follicles.
Theriogenology    November 26, 2010   Volume 75, Issue 4 707-714 doi: 10.1016/j.theriogenology.2010.10.011
Cuervo-Arango J, Domingo-Ortiz R.Prostaglandins play an obligatory role during the process of ovulation in mammals. Ovulation can be blocked by intrafollicular administration of non-steroidal anti-inflammatory drugs (NSAIDs) in several domestic species including the mare as well as by systemic administration of these drugs in women. In the mare, the effect of systemic NSAIDs treatment on ovulation has not been critically studied. The objectives of this study were: a) to determine whether high dose of flunixin-meglumine (FM) administered systemically to mares during the periovulatory period was able to block ovulation; and b) ...
Refinement of quantitative trait loci on equine chromosome 10 for radiological signs of navicular disease in Hanoverian warmblood horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 36-40 doi: 10.1111/j.1365-2052.2010.02096.x
Lopes MS, Diesterbeck U, Machado Ada C, Distl O.Navicular disease is characterized by a progressive degenerative alteration of the equine podotrochlea. In this study, we refined a previously identified quantitative trait locus (QTL) on horse chromosome 10 for the abnormal development of canales sesamoidales (DCS) of the navicular bone in Hanoverian warmblood horses. Genotyping was done in 192 Hanoverian warmblood horses from 17 paternal half-sib groups. The whole marker set comprised 45 markers including seven newly developed microsatellites and 13 single nucleotide polymorphisms (SNPs) within positional candidate genes. Chromosome-wide sig...
Identification of equine major histocompatibility complex haplotypes using polymorphic microsatellites.
Animal genetics    November 26, 2010   Volume 41 Suppl 2, Issue Suppl 2 150-153 doi: 10.1111/j.1365-2052.2010.02125.x
Tseng CT, Miller D, Cassano J, Bailey E, Antczak DF.A system for identifying equine major histocompatibility complex (MHC) haplotypes was developed based on five polymorphic microsatellites located within the MHC region on ECA 20. Molecular signatures for 50 microsatellite haplotypes were recognized from typing 353 horses. Of these, 23 microsatellite haplotypes were associated with 12 established equine leucocyte antigen (ELA) haplotypes in Thoroughbreds and Standardbreds. Five ELA serotypes were associated with multiple microsatellite subhaplotypes, expanding the estimates of diversity in the equine MHC. The strong correlations between serolog...
A survey of aged horses in Queensland, Australia. Part 2: Clinical signs and owners’ perceptions of health and welfare.
Australian veterinary journal    November 26, 2010   Volume 88, Issue 12 465-471 doi: 10.1111/j.1751-0813.2010.00638.x
McGowan TW, Pinchbeck G, Phillips CJ, Perkins N, Hodgson DR, McGowan CM.To describe the prevalence and risk factors for clinical signs of disease and owner-reported health or welfare issues of aged horses in Queensland, Australia. Methods: Owners of horses were contacted via an equestrian organisation in Queensland and asked to complete a questionnaire about their horses aged 15 years and older, to gain information about clinical signs observed in the horse and disease history. Owners were asked to identify health or welfare issues they felt were important in aged horses. Results: Owners were able to identify many clinical signs of disease, with 83% of horses havi...
IgE, IgGa, IgGb and IgG(T) serum antibody levels in offspring of two sires affected with equine recurrent airway obstruction.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 131-137 doi: 10.1111/j.1365-2052.2010.02122.x
Scharrenberg A, Gerber V, Swinburne JE, Wilson AD, Klukowska-Rötzler J, Laumen E, Marti E.Equine recurrent airway obstruction (RAO) is a chronic lower airway disease of the horse caused by hypersensitivity reactions to inhaled stable dust, including mould spores such as Aspergillus fumigatus. The goals of this study were to investigate whether total serum IgE levels and allergen-specific IgE and IgG subclasses are influenced by genetic factors and/or RAO and whether quantitative trait loci (QTL) could be identified for these parameters. The offspring of two RAO-affected sires (S1: n=56 and S2: n=65) were grouped by stallion and disease status, and total serum IgE levels and specifi...
Identification of the myostatin locus (MSTN) as having a major effect on optimum racing distance in the Thoroughbred horse in the USA.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 154-158 doi: 10.1111/j.1365-2052.2010.02126.x
Binns MM, Boehler DA, Lambert DH.One hundred and eighty-nine Thoroughbred horses that had won Graded Stakes races in North America were genotyped with the Illumina Equine SNP50 bead chip. Association tests using PLINK to determine whether any SNPs were associated with optimum racing distance (7 furlongs and under compared to 8-10 furlongs) identified a locus on ECA18 that was statistically significant (-log 10 EMP2=1.63) at the genome-wide level following permutation analysis (10,000 permutations). Bioinformatic analysis revealed that the two ECA18 SNPs with the highest statistical significance spanned the MSTN (myostatin) lo...
Fine mapping of a quantitative trait locus for osteochondrosis on horse chromosome 2.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 87-90 doi: 10.1111/j.1365-2052.2010.02113.x
Dierks C, Komm K, Lampe V, Distl O.In this study, we refine a quantitative trait locus for equine osteochondrosis (OC) on horse chromosome (ECA) 2 to a genome-wide significant interval at 20.08-30.94 Mb. The marker set contained 27 newly developed microsatellites equidistantly distributed over ECA2 and 44 nucleotide polymorphisms, located in 16 positional candidate genes for OC. Genotyping was performed in 211 Hanoverian horses from 14 paternal half-sib groups. A NCDN-associated SNP and haplotype were significantly associated with OC in fetlock and/or hock joints. This study is a further step towards the identification of genes...
Genome-wide SNP association-based localization of a dwarfism gene in Friesian dwarf horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 2-7 doi: 10.1111/j.1365-2052.2010.02091.x
Orr N, Back W, Gu J, Leegwater P, Govindarajan P, Conroy J, Ducro B, Van Arendonk JA, MacHugh DE, Ennis S, Hill EW, Brama PA.The recent completion of the horse genome and commercial availability of an equine SNP genotyping array has facilitated the mapping of disease genes. We report putative localization of the gene responsible for dwarfism, a trait in Friesian horses that is thought to have a recessive mode of inheritance, to a 2-MB region of chromosome 14 using just 10 affected animals and 10 controls. We successfully genotyped 34,429 SNPs that were tested for association with dwarfism using chi-square tests. The most significant SNP in our study, BIEC2-239376 (P(2df)=4.54 × 10(-5), P(rec)=7.74 × 10(-6)), is lo...
Transcription of LINE-derived sequences in exercise-induced stress in horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 23-27 doi: 10.1111/j.1365-2052.2010.02094.x
Capomaccio S, Verini-Supplizi A, Galla G, Vitulo N, Barcaccia G, Felicetti M, Silvestrelli M, Cappelli K.A large proportion of mammalian genomes is represented by transposable elements (TE), most of them being long interspersed nuclear elements 1 (LINE-1 or L1). An increased expression of LINE-1 elements may play an important role in cellular stress-related conditions exerting drastic effects on the mammalian transcriptome. To understand the impact of TE on the known horse transcriptome, we masked the horse EST database, pointing out that the amount is consistent with other major vertebrates. A previously developed transcript-derived fragments (TDFs) dataset, deriving from exercise-stimulated hor...
Structural annotation of equine protein-coding genes determined by mRNA sequencing.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 121-130 doi: 10.1111/j.1365-2052.2010.02118.x
Coleman SJ, Zeng Z, Wang K, Luo S, Khrebtukova I, Mienaltowski MJ, Schroth GP, Liu J, MacLeod JN.The horse, like the majority of animal species, has a limited amount of species-specific expressed sequence data available in public databases. As a result, structural models for the majority of genes defined in the equine genome are predictions based on ab initio sequence analysis or the projection of gene structures from other mammalian species. The current study used Illumina-based sequencing of messenger RNA (RNA-seq) to help refine structural annotation of equine protein-coding genes and for a preliminary assessment of gene expression patterns. Sequencing of mRNA from eight equine tissues...
Linkage disequilibrium and historical effective population size in the Thoroughbred horse.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 8-15 doi: 10.1111/j.1365-2052.2010.02092.x
Corbin LJ, Blott SC, Swinburne JE, Vaudin M, Bishop SC, Woolliams JA.Many genomic methodologies rely on the presence and extent of linkage disequilibrium (LD) between markers and genetic variants underlying traits of interest, but the extent of LD in the horse has yet to be comprehensively characterized. In this study, we evaluate the extent and decay of LD in a sample of 817 Thoroughbreds. Horses were genotyped for over 50,000 single nucleotide polymorphism (SNP) markers across the genome, with 34,848 autosomal SNPs used in the final analysis. Linkage disequilibrium, as measured by the squared correlation coefficient (r(2)), was found to be relatively high bet...
Mitochondrial DNA insertions in the nuclear horse genome.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 176-185 doi: 10.1111/j.1365-2052.2010.02130.x
Nergadze SG, Lupotto M, Pellanda P, Santagostino M, Vitelli V, Giulotto E.The insertion of mitochondrial DNA in the nuclear genome generates numts, nuclear sequences of mitochondrial origin. In the horse reference genome, we identified 82 numts and showed that the entire horse mitochondrial DNA is represented as numts without gross bias. Numts were inserted in the horse nuclear genome at random sites and were probably generated during the repair of DNA double-strand breaks. We then analysed 12 numt loci in 20 unrelated horses and found that null alleles, lacking the mitochondrial DNA insertion, were present at six of these loci. At some loci, the null allele is prev...
A genome-wide association study for racing performances in Thoroughbreds clarifies a candidate region near the MSTN gene.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 28-35 doi: 10.1111/j.1365-2052.2010.02095.x
Tozaki T, Miyake T, Kakoi H, Gawahara H, Sugita S, Hasegawa T, Ishida N, Hirota K, Nakano Y.Using 1400 microsatellites, a genome-wide association study (GWAS) was performed to identify genomic regions associated with lifetime earnings and performance ranks, as determined by the Japan Racing Association (JRA). The minimum heritability (h(2) ) was estimated at 7-8% based on the quantitative trait model, suggesting that the racing performance is heritable. Following GWAS with microsatellites, fine mapping led to identification of three SNPs on ECA18, namely, g.65809482T>C (P=1.05E-18), g.65868604G>T (P=6.47E-17), and g.66539967A>G (P=3.35E-14) associated with these performance ...
RT-qPCR comparison of mast cell populations in whole blood from healthy horses and those with laminitis.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 16-22 doi: 10.1111/j.1365-2052.2010.02093.x
Brooks SA, Bailey E.Inflammatory damage to the digital laminae, a structure responsible for suspension of the distal skeleton within the hoof capsule, results in a painful and often life-threatening disease in horses called laminitis. There can be many diverse causes of laminitis; however, previous work in the horse has suggested that in each case, the inflammation and resulting tissue damage is consistent with the action of mediators released from mast cells (MC), as well as the downstream consequences of their activation. The recent development of molecular genetics tools to characterize cells based on their tr...
Interspersed repeats in the horse (Equus caballus); spatial correlations highlight conserved chromosomal domains.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 91-99 doi: 10.1111/j.1365-2052.2010.02115.x
Adelson DL, Raison JM, Garber M, Edgar RC.The interspersed repeat content of mammalian genomes has been best characterized in human, mouse and cow. In this study, we carried out de novo identification of repeated elements in the equine genome and identified previously unknown elements present at low copy number. The equine genome contains typical eutherian mammal repeats, but also has a significant number of hybrid repeats in addition to clade-specific Long Interspersed Nuclear Elements (LINE). Equus caballus clade specific LINE 1 (L1) repeats can be classified into approximately five subfamilies, three of which have undergone signifi...
A genome-wide scan for tying-up syndrome in Japanese Thoroughbreds.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 80-86 doi: 10.1111/j.1365-2052.2010.02112.x
Tozaki T, Hirota K, Sugita S, Ishida N, Miyake T, Oki H, Hasegawa T.Tying-up syndrome, also known as recurrent exertional rhabdomyolysis in Thoroughbreds, is a common muscle disorder for racehorses. In this study, we performed a multipoint linkage analysis using LOKI based on the Bayesian Markov chain Monte Carlo method using 5 half-sib families (51 affected and 277 nonaffected horses in total), and a genome-wide association study (GWAS) using microsatellites (144 affected and 144 nonaffected horses) to map candidate regions for tying-up syndrome in Japanese Thoroughbreds. The linkage analysis identified one strong L-score (82.45) between the loci UCDEQ411 and...
Genomic structure, polymorphism and expression of ACCN1 and ACCN3 genes in the horse.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 138-144 doi: 10.1111/j.1365-2052.2010.02123.x
Mata X, Ducasse A, Vaiman A, Diribarne M, Fraud AS, Guérin G.A category of cation gate proteins was shown to be present in sensory neurons and act as receptors of protons present in tissues such as muscles. The Amiloride-sensitive Cation Channel, Neuronal (ACCN) gene family is known to play a role in the transmission of pain through specialized pH sensitive neurons. Muscles from horses submitted to strenuous exercises produce lactic acid, which may induce variable pain through ACCN differential properties. The sequences of the equine cDNAs were determined to be 2.6 kb in length with an open reading frame of 1539 bp for ACCN1 and 2.1 kb in length with an...
Estimated prevalence of the Type 1 Polysaccharide Storage Myopathy mutation in selected North American and European breeds.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 145-149 doi: 10.1111/j.1365-2052.2010.02124.x
McCue ME, Anderson SM, Valberg SJ, Piercy RJ, Barakzai SZ, Binns MM, Distl O, Penedo MC, Wagner ML, Mickelson JR.The GYS1 gene mutation that is causative of Type 1 Polysaccharide Storage Myopathy (PSSM) has been identified in more than 20 breeds of horses. However, the GYS1 mutation frequency or Type 1 PSSM prevalence within any given breed is unknown. The purpose of this study was to determine the frequency of the GYS1 mutation and prevalence of genetic susceptibility to Type 1 PSSM in selected breeds from Europe and North America. The GYS1 mutation was detected in 11 breeds, including, in order of increasing allele frequency, Shires, Morgans, Appaloosas, Quarter Horses, Paints, Exmoor Ponies, Saxon-Thu...
A conserved segmental duplication within ELA.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 186-195 doi: 10.1111/j.1365-2052.2010.02137.x
Brinkmeyer-Langford CL, Murphy WJ, Childers CP, Skow LC.The assembled genomic sequence of the horse major histocompatibility complex (MHC) (equine lymphocyte antigen, ELA) is very similar to the homologous human HLA, with the notable exception of a large segmental duplication at the boundary of ELA class I and class III that is absent in HLA. The segmental duplication consists of a ∼ 710 kb region of at least 11 repeated blocks: 10 blocks each contain an MHC class I-like sequence and the helicase domain portion of a BAT1-like sequence, and the remaining unit contains the full-length BAT1 gene. Similar genomic features were found in other Perissod...
True stress and Poisson’s ratio of tendons during loading.
Journal of biomechanics    November 26, 2010   Volume 44, Issue 4 719-724 doi: 10.1016/j.jbiomech.2010.10.038
Vergari C, Pourcelot P, Holden L, Ravary-Plumioën B, Gerard G, Laugier P, Mitton D, Crevier-Denoix N.Excessive axial tension is very likely involved in the aetiology of tendon lesions, and the most appropriate indicator of tendon stress state is the true stress, the ratio of instantaneous load to instantaneous cross-sectional area (CSA). Difficulties to measure tendon CSA during tension often led to approximate true stress by assuming that CSA is constant during loading (i.e. by the engineering stress) or that tendon is incompressible, implying a Poisson's ratio of 0.5, although these hypotheses have never been tested. The objective of this study was to measure tendon CSA variation during qua...