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Topic:Tissue

Tissue in horses refers to the various types of biological material that make up the body of the animal, including muscle, connective, epithelial, and nervous tissues. Each type of tissue has specific functions and characteristics, contributing to the overall physiology and health of the horse. Muscle tissue is responsible for movement and locomotion, connective tissue provides structural support and protection, epithelial tissue covers body surfaces and lines cavities, and nervous tissue is involved in transmitting signals for communication and coordination within the body. The study of equine tissue encompasses aspects such as growth, repair, and response to injury or disease. This page compiles peer-reviewed research studies and scholarly articles that explore the structure, function, and pathology of different tissue types in horses, offering insights into their role in equine health and disease management.
Quantitative Assessment of Tendon Hierarchical Structure by Combined Second Harmonic Generation and Immunofluorescence Microscopy.
Tissue engineering. Part C, Methods    May 5, 2020   Volume 26, Issue 5 253-262 doi: 10.1089/ten.TEC.2020.0032
Durgam S, Singh B, Cole SL, Brokken MT, Stewart M.Histological evaluation of healing tendons is primarily focused on monitoring restoration of longitudinal collagen alignment, although the elastic property of energy-storing flexor tendons is largely attributed to interfascicular sliding facilitated by the interfascicular matrix (IFM). The objectives of this study were to explore the utility of second harmonic generation (SHG) imaging to objectively assess cross-sectional tendon fascicle architecture, to combine SHG microscopy with elastin immunofluorescence to assess the ultrastructure of collagen and elastin in longitudinal and transverse se...
On-site testing of sutured organs: An experimental set up to cyclically tighten sutures.
Journal of the mechanical behavior of biomedical materials    May 1, 2020   Volume 109 103803 doi: 10.1016/j.jmbbm.2020.103803
Pascoletti G, Pressanto MC, Putame G, Terzini M, Audenino AL, Zanetti EM.A number of surgical practices are aimed to compensate for tissue relaxation or weakened/atrophied muscles by means of suture prostheses/thread lifts. The success rate of these procedures is often very good in the short term, while it is quite variable among subjects and techniques in the middle-long term. Middle-long term failures are mostly related to suture distraction, loosening or wear, coming from repeated loading cycles. In this work, an experimental device to perform ex vivo tests on prosthetic sutures has been set up. An equine laryngoplasty has been used as a benchmark, being represe...
Data from cyclic tensile tests on sutured organs to evaluate creep behaviour, distraction, and residual thread strength.
Data in brief    April 30, 2020   Volume 30 105644 doi: 10.1016/j.dib.2020.105644
Pascoletti G, Pressanto MC, Putame G, Terzini M, Franceschini G, Zanetti EM.A number of applications in the surgical practice are based on tensile sutures aimed to keep soft tissues in place and compensate the exit of neuropathies, prolapses or general tissue relaxation. Long-term behaviour of these constructs need to be carefully examined in order to define tensile forces to be applied and to compare different suture anchors. Data here reported refer to equine laryngoplasties, where a suitable loading system has been designed in order to be able to test sutures in-sito, applying known forces ("On-site testing of sutured organs: an experimental set up to cyclically ti...
Equine Hoof Canker: Bovine Papillomavirus Infection Is Not Associated With Impaired Keratinocyte Differentiation.
Veterinary pathology    April 29, 2020   Volume 57, Issue 4 525-534 doi: 10.1177/0300985820921820
Apprich V, Licka T, Freiler S, Gabriel C.Impaired keratinocyte differentiation has recently been suggested as a key event in equine hoof canker development. Koilocytotic appearance of keratinocytes, one of the most characteristic morphological alterations in hoof canker tissue, is also a common marker for papillomavirus (PV) infection, and bovine PV-1 and/or -2 (BPV-1/2) has previously been detected in equine canker patients. Therefore, the present study aimed to correlate the frequency and severity of koilocytotic keratinocytes with BPV detection in hoof canker samples. Hoof tissue of 5/18 canker-affected horses and 2/6 control hors...
Evidence that human and equine erythrocytes could have significant roles in the transport and delivery of amino acids to organs and tissues.
Amino acids    April 21, 2020   Volume 52, Issue 5 711-724 doi: 10.1007/s00726-020-02845-0
Thorn B, Dunstan RH, Macdonald MM, Borges N, Roberts TK.Erythrocytes have a well-defined role in the gaseous exchange of oxygen and carbon dioxide in the mammalian body. The erythrocytes can contain more than half of the free amino acids present in whole blood. Based on measures showing that venous erythrocyte levels of amino acids are much less than arterial erythrocyte levels, it has previously been proposed that erythrocytes also play a role in the delivery of amino acids to tissues in the body. This role has been dismissed because it has been assumed that to act as an amino acid transport vehicle, the erythrocytes should release their entire a...
Multi-differentiation potential is necessary for optimal tenogenesis of tendon stem cells.
Stem cell research & therapy    April 9, 2020   Volume 11, Issue 1 152 doi: 10.1186/s13287-020-01640-8
Rajpar I, Barrett JG.Tendon injury is a significant clinical problem due to poor healing and a high reinjury rate; successful treatment is limited by our poor understanding of endogenous tendon stem cells. Recent evidence suggests that adult stem cells are phenotypically diverse, even when comparing stem cells isolated from the same tissue from the same individual, and may in fact exist on a spectrum of proliferation and differentiation capacities. Additionally, the relationships between and clinical relevance of this phenotypic variation are poorly understood. In particular, tenogenic capacity has not been studie...
Preliminary Notes on Equine Tissue Transglutaminase Serology and A Case of Equine Gluten-Sensitive Enteropathy and Dermatitis in an 11-Year-Old Dutch Warmblood Horse.
Journal of equine veterinary science    April 2, 2020   Volume 90 102999 doi: 10.1016/j.jevs.2020.102999
van Proosdij R, Mulder C, Reijm M, Bontkes H, von Blomberg M, van der Kolk H.It has been suggested that gluten may play a role in equine inflammatory small bowel disease (ISBD). Previous work showed an association between equine gluten-sensitive enteropathy and IgA antibodies to tissue transglutaminase (TGA) in serum. The purpose of this study is to investigate the prevalence of IgA antibodies to TGA in a group of healthy non-gluten-free sport ponies and to present a case of tentative gluten-sensitive enteropathy and dermatitis in a horse. Blood samples were obtained from 40 healthy jumping ponies. The ponies comprised 12 mares, 8 stallions, and 20 geldings with an ave...
Cyclical strain improves artificial equine tendon constructs in vitro.
Journal of tissue engineering and regenerative medicine    March 23, 2020   Volume 14, Issue 5 690-700 doi: 10.1002/term.3030
Atkinson F, Evans R, Guest JE, Bavin EP, Cacador D, Holland C, Guest DJ.Tendon injuries are a common cause of morbidity in humans. They also occur frequently in horses, and the horse provides a relevant, large animal model in which to test novel therapies. To develop novel cell therapies that can aid tendon regeneration and reduce subsequent reinjury rates, the mechanisms that control tendon tissue regeneration and matrix remodelling need to be better understood. Although a range of chemical cues have been explored (growth factors, media etc.), the influence of the mechanical environment on tendon cell culture has yet to be fully elucidated. To mimic the in vivo e...
Detection of Age-Related Changes in Tendon Molecular Composition by Raman Spectroscopy-Potential for Rapid, Non-Invasive Assessment of Susceptibility to Injury.
International journal of molecular sciences    March 20, 2020   Volume 21, Issue 6 2150 doi: 10.3390/ijms21062150
Yin NH, Parker AW, Matousek P, Birch HL.The lack of clinical detection tools at the molecular level hinders our progression in preventing age-related tendon pathologies. Raman spectroscopy can rapidly and non-invasively detect tissue molecular compositions and has great potential for in vivo applications. In biological tissues, a highly fluorescent background masks the Raman spectral features and is usually removed during data processing, but including this background could help age differentiation since fluorescence level in tendons increases with age. Therefore, we conducted a stepwise analysis of fluorescence and Raman combined s...
Decellularization of Extracellular Matrix from Equine Skeletal Muscle.
Journal of equine veterinary science    March 19, 2020   Volume 90 102962 doi: 10.1016/j.jevs.2020.102962
Miranda CMFC, Therrien J, Leonel LCPC, Smith OE, Miglino MA, Smith LC.Equine represents an attractive animal model for musculoskeletal tissue diseases, exhibiting much similarity to the injuries that occur in humans. Cell therapy and tissue bioengineering have been widely used as a therapeutic alternative by regenerative medicine in musculoskeletal diseases. Thus, the aim of this study was to produce an acellular biomaterial of equine skeletal muscle and to evaluate its effectiveness in supporting the in vitro culture of equine induced pluripotency stem cells (iPSCs). Biceps femoris samples were frozen at -20°C for 4 days and incubated in 1% sodium dodecyl s...
Identification and Quantification of Transient Receptor Potential Vanilloid 1 (TRPV1) in Equine Articular Tissue.
Animals : an open access journal from MDPI    March 18, 2020   Volume 10, Issue 3 506 doi: 10.3390/ani10030506
Braucke AFGV, Frederiksen NL, Berg LC, Aarsvold S, Müller FC, Boesen MP, Lindegaard C.Joint pain and osteoarthritis (OA) are some of the most common causes of lameness in horses, and most of the available treatments focus on symptomatic relief without a disease-modifying effect. TRPV1 is a potential target for treating joint diseases, including OA, and the present study aims to investigate if the TRPV1 receptor is present in equine articular tissue and determine whether the number of receptors is upregulated in joint inflammation. Metacarpo/metatarsophalangeal (MCP/MTP) joints from 15 horses euthanised for reasons unrelated to this study were included. Based on synovial fluid a...
Molecular and cellular evidence of natural Venezuelan equine encephalitis virus infection in frugivorous bats in Colombia.
Veterinary world    March 16, 2020   Volume 13, Issue 3 495-501 doi: 10.14202/vetworld.2020.495-501
Guzmán C, Calderón A, Oviedo T, Mattar S, Castañeda J, Rodriguez V, Moraes Figueiredo LT.Venezuelan equine encephalitis virus (VEEV) is an alphavirus that causes encephalitis with a high impact on public health in Latin America. However, only in Guatemala, Trinidad and Tobago, and Mexico have found antibodies in VEEV in bats, using immunohistochemistry, the sensitivity and specificity are improved; thus, it is better for demonstrating natural infection in bats as potential hosts. This study aimed to determine the presence of VEEV in tissues of frugivorous bats. Methods: A prospective descriptive cross-sectional study with a non-probabilistic sampling was carried out in 12 localiti...
Sub- and Supramolecular X-Ray Characterization of Engineered Tissues from Equine Tendon, Bovine Dermis, and Fish Skin Type-I Collagen.
Macromolecular bioscience    March 12, 2020   Volume 20, Issue 5 e2000017 doi: 10.1002/mabi.202000017
Terzi A, Gallo N, Bettini S, Sibillano T, Altamura D, Madaghiele M, De Caro L, Valli L, Salvatore L, Sannino A, Giannini C.Collagen represents one of the most widely used biomaterial for scaffolds fabrication in tissue engineering as it represents the mechanical support of natural tissues. It also provides physical scaffolding for cells and it influences their attachment, growth, and tissue regeneration. Among all fibrillary collagens, type I is considered one of the gold standard for scaffolds fabrication, thanks to its high biocompatibility, biodegradability, and hemostatic properties. It can be extracted by chemical and enzymatic protocols from several collagen-rich tissues, such as tendon and skin, of differen...
Coronary corium, a new source of equine mesenchymal stromal cells.
Veterinary research communications    March 4, 2020   Volume 44, Issue 2 41-49 doi: 10.1007/s11259-020-09771-2
da Silva LL, Silveira MD, da Costa Garcia CAS, Grudzinski PB, Martins CF, Nardi NB.Mesenchymal stromal cells (MSCs) have attracted great attention for therapeutic applications. Since cells derived from different tissues have different properties, using the right tissue source may impact their efficiency in regenerative medicine. This study describes for the first time the isolation and characterization of MSCs derived from the equine coronary corium, which may be useful for treating diseases such as laminitis. Seven coronary corium samples were used for isolation of cells (ccMSCs). Adherent cells were characterized for morphology, immunophenotype, proliferation and different...
Histologic Comparison of the Dura Mater among Species.
Comparative medicine    February 3, 2020   Volume 70, Issue 2 170-175 doi: 10.30802/AALAS-CM-19-000022
Kinaci A, Bergmann W, Bleys RL, van der Zwan A, van Doormaal TP.The biocompatibility, biodegradation, feasibility, and efficacy of medical devices like dural sealants and substitutes are often evaluated in various animal models. However, none of these studies explain the rationale for choosing a particular species, and a systematic interspecies comparison of the dura is not available. We hypothesized that histologic characteristics of the dura would differ among species. We systematically investigated basic characteristics of the dura, including thickness, composition, and fibroblast orientation of the dura mater, in 34 samples representing 10 animal speci...
The Potential of Mesenchymal Stem Cells to Treat Systemic Inflammation in Horses.
Frontiers in veterinary science    January 21, 2020   Volume 6 507 doi: 10.3389/fvets.2019.00507
MacDonald ES, Barrett JG.One hallmark of mesenchymal stem cells (MSCs) is the ability to differentiate into multiple tissue types which assists in tissue regeneration. Another hallmark of MSCs is their potent anti-inflammatory and immunomodulatory properties and the potential to treat inflammatory, immune-mediated, and ischemic conditions. In equine practice, MSCs have shown efficacy in the treatment of musculoskeletal disorders such as tendinopathy, meniscal tears and cartilage injury. However, there are many equine disease processes and conditions that may benefit from the immunomodulatory properties of MSCs. Exampl...
Prognostic value of colonic and peripheral venous lactate measurements in horses with large colon volvulus.
Veterinary surgery : VS    January 9, 2020   Volume 49, Issue 3 472-479 doi: 10.1111/vsu.13361
Orr KE, Baker WT, Lynch TM, Hughes FE, Clark CK, Slone DE, Fogle CA, Gonzalez LM.To determine the prognostic value of (1) colonic venous lactate or peripheral lactate values obtained before and after manual correction of a large colon volvulus and (2) a combination of variables including pelvic flexure biopsy. Methods: Prospective clinical study. Methods: Forty adult horses in which large colon volvulus was diagnosed intraoperatively. Methods: Colonic venous, peripheral venous, and arterial blood samples were collected to measure lactate values before and after manual correction. Mucosal biopsy samples were obtained in cases that underwent enterotomy or colonic resection a...
Therapeutic mesenchymal stromal stem cells: Isolation, characterization and role in equine regenerative medicine and metabolic disorders.
Stem cell reviews and reports    December 5, 2019   Volume 16, Issue 2 301-322 doi: 10.1007/s12015-019-09932-0
Al Naem M, Bourebaba L, Kucharczyk K, Röcken M, Marycz K.Mesenchymal stromal cells (MSC) have become a popular treatment modality in equine orthopaedics. Regenerative therapies are especially interesting for pathologies like complicated tendinopathies of the distal limb, osteoarthritis, osteochondritis dissecans (OCD) and more recently metabolic disorders. Main sources for MSC harvesting in the horse are bone marrow, adipose tissue and umbilical cord blood. While the acquisition of umbilical cord blood is fairly easy and non-invasive, extraction of bone marrow and adipose tissue requires more invasive techniques. Characterization of the stem cells a...
Pododermal angioarchitecture in the equine hoof wall: A light and scanning electron microscopic study of the wall proper.
Clinical hemorheology and microcirculation    November 28, 2019   Volume 74, Issue 1 21-44 doi: 10.3233/CH-199233
Sievers H, Hiebl B, Hünigen H, Hirschberg RM.Blood supply is an important factor for the normal function of the equine hoof, but earlier studies present conflicting data on functional characteristics of its angioarchitecture. Objective: Emphasis was laid on demonstration of the microvascularisation in the different hoof wall regions, aiming at assessment of specialised vascular structures, e.g. vascular sphincter mechanisms and arteriovenous anastomoses. Methods: The angioarchitecture of the adult pododerma in the equine hoof wall was examined by scanning electron microscopy of micro-corrosion casts assisted by exemplary histological and...
Endotoxin-induced changes in expression of cyclooxygenase isoforms in the lamellar tissue of extracorporeally haemoperfused equine limbs.
Anatomia, histologia, embryologia    November 27, 2019   Volume 49, Issue 5 597-605 doi: 10.1111/ahe.12520
Patan-Zugaj B, Egerbacher M, Licka TF.Angiogenesis and sepsis-related equine laminitis have several features in common. Both events can be induced by endotoxin (lipopolysaccharide- LPS) and both are associated with increased expression of the enzyme cyclooxygenase (COX), of which two isoforms (COX-1 and COX-2) exist. To examine the causal relationship between LPS exposure and COX expression and to investigate the tissue distribution of COX in the LPS-exposed tissue, the technique of extracorporeal haemoperfusion of isolated equine forelimbs was utilized. Perfusion was performed for 10 hr under physiological conditions (control-pe...
Comparison of the Chondrogenic Potential of Mesenchymal Stem Cells Derived from Bone Marrow and Umbilical Cord Blood Intended for Cartilage Tissue Engineering.
Stem cell reviews and reports    November 21, 2019   Volume 16, Issue 1 126-143 doi: 10.1007/s12015-019-09914-2
Contentin R, Demoor M, Concari M, Desancé M, Audigié F, Branly T, Galéra P.Osteoarthritis (OA) remains incurable in humans or horses and mesenchymal stromal/stem cells (MSCs) represent an attractive solution for producing a neocartilage substitute. However, the best MSC source still needs to be identified. This study compared the chondrogenic potential of equine MSCs derived from bone marrow (BM) and umbilical cord blood (UCB), at their undifferentiated status to check if one cell source is better proned, and after chondrogenic-induced differentiation. Chondrogenesis was induced by culture in collagen scaffold with BMP-2 + TGF-ß1 in hypoxia or normoxia. MSCs cho...
The Equine Hoof: Laminitis, Progenitor (Stem) Cells, and Therapy Development.
Toxicologic pathology    November 19, 2019   Volume 49, Issue 7 1294-1307 doi: 10.1177/0192623319880469
Yang Q, Lopez MJ.The equine hoof capsule, composed of modified epidermis and dermis, is vital for protecting the third phalanx from forces of locomotion. There are descriptions of laminitis, defined as inflammation of sensitive hoof tissues but recognized as pathologic changes with or without inflammatory mediators, in the earliest records of domesticated horses. Laminitis can range from mild to serious, and signs can be acute, chronic, or transition from acute, severe inflammation to permanently abnormal tissue. Damage within the intricate dermal and epidermal connections of the primary and secondary lamellae...
Dimethyl sulfoxide maintains structure and function of cryopreserved equine endometrial explants.
Cryobiology    October 15, 2019   Volume 91 90-96 doi: 10.1016/j.cryobiol.2019.10.006
Thompson RE, Johnson AK, Prado TM, Premanandan C, Brown ME, Whitlock BK, Pukazhenthi BS.Availability of viable frozen-thawed endometrial tissues could facilitate detailed studies into physiologic and disease processes influencing the endometrium. This study was designed to investigate the cryosurvival of equine endometrial tissue. Previous studies in the human and horse have focused on cryopreservation of dissociated endometrial cells. To our knowledge, there are no studies on cryopreservation of endometrial explants. Our objectives were to 1) determine the influence of differing concentrations of the permeating cryoprotectant dimethyl sulfoxide (MeSO) on viability, structural in...
Multiple excisions of an equine sarcoid and the repercussions.
The Canadian veterinary journal = La revue veterinaire canadienne    October 11, 2019   Volume 60, Issue 10 1115-1117 
Melinyshyn H.A 13-year-old Morgan crossbred mare was presented in May 2018 with a 10-cm mass on the medial aspect of the left hind fetlock. In September and November 2018 and June 2019 the mass was incompletely excised and continued to grow. A diagnosis of spindle cell tumor, consistent with equine sarcoid, was made based on histopathology from the last excision. . Une jument Morgan croisée âgée de 13 ans a été présentée en mai 2018 avec une masse de 10 cm sur l’aspect médial du boulet arrière gauche. En septembre et novembre 2018 ainsi qu’en juin 2019, la masse avait été excisée de manièr...
Equine Fetal, Adult, and Embryonic Stem Cell-Derived Tenocytes Are All Immune Privileged but Exhibit Different Immune Suppressive Properties In Vitro.
Stem cells and development    October 8, 2019   Volume 28, Issue 21 1413-1423 doi: 10.1089/scd.2019.0120
McClellan A, Paterson YZ, Paillot R, Guest DJ.In horses and humans, tendon injuries are a significant problem. Not only can they occur in both athletes and nonathletes, they require lengthy periods of recuperation and undergo poor natural regeneration, which leads to high reinjury rates. Embryonic stem cells (ESCs) may provide a renewable source of allogeneic cells to use in clinical applications to aid tissue regeneration. Equine ESCs can undergo tenocyte differentiation in vivo and in vitro, but the immune properties of tenocytes isolated from either ESCs or tissues have not previously been characterized. Here, we demonstrate that equin...
An MRI study of solute transport in the intervertebral disc.
Magma (New York, N.Y.)    October 4, 2019   Volume 33, Issue 2 299-307 doi: 10.1007/s10334-019-00781-z
Palfrey RM, Summers IR, Winlove CP.Quantitative magnetic resonance imaging was used to determine partition coefficients and characteristic time constants for diffusion of MRI contrast agents in disc tissue. Methods: Twenty-two excised equine intervertebral discs were exposed to a range of contrast agents: six to manganese chloride, eight to Magnevist (gadopentetate dimeglumine) and eight to Gadovist (gadobutrol), and uptake into the disc was quantified in T-weighted images. Results: Diffusion for all contrast agents was approximately 25% faster in the nucleus than in the outer annulus; disc-average time constants ranged from (2...
Differential Gene Expression in Articular Cartilage and Subchondral Bone of Neonatal and Adult Horses.
Genes    September 25, 2019   Volume 10, Issue 10 745 doi: 10.3390/genes10100745
Kemper AM, Drnevich J, McCue ME, McCoy AM.Skeletogenesis is complex and incompletely understood. Derangement of this process likely underlies developmental skeletal pathologies. Examination of tissue-specific gene expression may help elucidate novel skeletal developmental pathways that could contribute to disease risk. Our aim was to identify and functionally annotate differentially expressed genes in equine neonatal and adult articular cartilage (AC) and subchondral bone (SCB). RNA was sequenced from healthy AC and SCB from the fetlock, hock, and stifle joints of 6 foals (≤4 weeks of age) and six adults (8-12 years of age). There w...
Insulin-like growth factor binding protein (IGFBP6) is a cross-species tendon marker.
European cells & materials    September 24, 2019   Volume 38 123-136 doi: 10.22203/eCM.v038a10
Turlo AJ, Mueller-Breckenridge AJ, Zamboulis DE, Tew SR, Canty-Laird EG, Clegg PD.The main challenge in tendon injury management is suboptimal tissue healing that fails to re-establish original tendon function. Tissue bioengineering is a promising approach for tendon therapy, with potential to improve its functional outcomes. However, evaluation criteria for tissue-engineered tendon are unclear due to the lack of specific markers of differentiated tendon. The study aim was to identify a panel of genes that characterised tendons in comparison to cartilage or muscles and validate those genes, both in human and key species used as models for tendon diseases. Gene expression pr...
Cyclical cervical function in the mare involves remodelling of collagen content, which is correlated with modification of oestrogen receptor 1 abundance.
Animal reproduction science    September 12, 2019   Volume 210 106192 doi: 10.1016/j.anireprosci.2019.106192
Campbell MLH, Peachey L, Callan L, Wathes DC, de Mestre AM.This study was conducted to elucidate mare cervical dilation mechanisms by testing two hypotheses: (i) the proportion of collagen staining in histological samples of mare cervices and (ii) the abundance of hormone receptors in the equine cervix differ with stage of the oestrous cycle and site within the cervix. Tissues and jugular vein blood samples were collected from 15 mares. Collagen content was assessed using Masson's Trichome staining. Receptor abundance was assessed using RT-PCR, qRT-PCR and immunohistochemistry. In sub-epithelial stroma, there was less collagen during the follicular th...
Comparison of tensile properties of xenopericardium from three animal species and finite element analysis for bioprosthetic heart valve tissue.
Artificial organs    September 1, 2019   Volume 44, Issue 3 278-287 doi: 10.1111/aor.13552
Rassoli A, Fatouraee N, Guidoin R, Zhang Z.Bioprosthetic heart valves still have poor long-term durability due to calcification and mechanical failure. The function and performance of bioprostheses is known to depend on the collagen architecture and mechanical behavior of the target tissue. So it is necessary to select an appropriate tissue for such prostheses. In this study, porcine, equine, and bovine pericardia were compared histologically and mechanically. The specimens were analyzed under light microscopy. The planar biaxial tests were performed on the tissue samples by applying synchronic loads along the axial (fiber direction) a...
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