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58-426: A tendon or sinew is a tough band of dense fibrous connective tissue that connects muscle to bone . It sends the mechanical forces of muscle contraction to the skeletal system, while withstanding tension . Tendons, like ligaments , are made of collagen . The difference is that ligaments connect bone to bone, while tendons connect muscle to bone. There are about 4000 tendons in the adult human body. A tendon

116-482: A human stride, the Achilles tendon stretches as the ankle joint dorsiflexes. During the last portion of the stride, as the foot plantar-flexes (pointing the toes down), the stored elastic energy is released. Furthermore, because the tendon stretches, the muscle is able to function with less or even no change in length , allowing the muscle to generate more force. The mechanical properties of the tendon are dependent on

174-487: A linear stress-strain curve until it begins to fail. The mechanical properties of tendons vary widely, as they are matched to the functional requirements of the tendon. The energy storing tendons tend to be more elastic, or less stiff, so they can more easily store energy, whilst the stiffer positional tendons tend to be a little more viscoelastic, and less elastic, so they can provide finer control of movement. A typical energy storing tendon will fail at around 12–15% strain, and

232-475: A stress in the region of 100–150 MPa, although some tendons are notably more extensible than this, for example the superficial digital flexor in the horse , which stretches in excess of 20% when galloping. Positional tendons can fail at strains as low as 6–8%, but can have moduli in the region of 700–1000 MPa. Several studies have demonstrated that tendons respond to changes in mechanical loading with growth and remodeling processes, much like bones . In particular,

290-521: A study showed that disuse of the Achilles tendon in rats resulted in a decrease in the average thickness of the collagen fiber bundles comprising the tendon. In humans, an experiment in which people were subjected to a simulated micro-gravity environment found that tendon stiffness decreased significantly, even when subjects were required to perform restiveness exercises. These effects have implications in areas ranging from treatment of bedridden patients to

348-405: A tendon's collagen is type I collagen , many minor collagens are present that play vital roles in tendon development and function. These include type II collagen in the cartilaginous zones, type III collagen in the reticulin fibres of the vascular walls, type IX collagen, type IV collagen in the basement membranes of the capillaries , type V collagen in the vascular walls, and type X collagen in

406-436: A well-developed rough endoplasmic reticulum and they are responsible for synthesis and turnover of tendon fibres and ground substance . Tendon cells form a connecting epithelial layer between the muscle and shell in molluscs . In gastropods , for example, the retractor muscles connect to the shell via tendon cells. Muscle cells are attached to the collagenous myo-tendon space via hemidesmosomes . The myo-tendon space

464-474: Is suan bao niu jin , in which the tendon is marinated in garlic. It is also sometimes found in the Vietnamese noodle dish phở . In some organisms, notably birds , and ornithischian dinosaurs , portions of the tendon can become ossified. In this process, osteocytes infiltrate the tendon and lay down bone as they would in sesamoid bone such as the patella. In birds, tendon ossification primarily occurs in

522-464: Is a major functional component of tendons , ligaments and aponeuroses , and is also found in highly specialized organs such as the cornea . Elastic fibers , made from elastin and fibrillin , also provide resistance to stretch forces. They are found in the walls of large blood vessels and in certain ligaments, particularly in the ligamenta flava . In hematopoietic and lymphatic tissues, reticular fibers made by reticular cells provide

580-676: Is a mix of fibrous and areolar tissue . Fibromuscular tissue is made up of fibrous tissue and muscular tissue . New vascularised connective tissue that forms in the process of wound healing is termed granulation tissue . All of the special connective tissue types have been included as a subset of fascia in the fascial system , with blood and lymph classed as liquid fascia . Bone and cartilage can be further classified as supportive connective tissue . Blood and lymph can also be categorized as fluid connective tissue , and liquid fascia . Membranes can be either of connective tissue or epithelial tissue . Connective tissue membranes include

638-492: Is also recommended in survival guides as a material from which strong cordage can be made for items like traps or living structures. Tendon must be treated in specific ways to function usefully for these purposes. Inuit and other circumpolar people utilized sinew as the only cordage for all domestic purposes due to the lack of other suitable fiber sources in their ecological habitats. The elastic properties of particular sinews were also used in composite recurved bows favoured by

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696-399: Is beneficial to have longer than average Achilles tendon and a shorter calf muscle . Tendon length is determined by genetic predisposition, and has not been shown to either increase or decrease in response to environment, unlike muscles, which can be shortened by trauma, use imbalances and a lack of recovery and stretching. In addition tendons allow muscles to be at an optimal distance from

754-406: Is bound by an epitenon , which is a sheath of dense irregular connective tissue . The whole tendon is enclosed by a fascia . The space between the fascia and the tendon tissue is filled with the paratenon , a fatty areolar tissue . Normal healthy tendons are anchored to bone by Sharpey's fibres . The dry mass of normal tendons, which is 30–45% of their total mass, is made of: Although most of

812-459: Is characterized by collagen fibers arranged in an orderly parallel fashion, giving it tensile strength in one direction. Dense irregular connective tissue provides strength in multiple directions by its dense bundles of fibers arranged in all directions. Special connective tissue consists of cartilage , bone , blood and lymph . Other kinds of connective tissues include fibrous, elastic, and lymphoid connective tissues. Fibroareolar tissue

870-454: Is controlled by the tenocytes and their surrounding extracellular matrix. The three main stages of tendon healing are inflammation, repair or proliferation, and remodeling, which can be further divided into consolidation and maturation. These stages can overlap with each other. In the first stage, inflammatory cells such as neutrophils are recruited to the injury site, along with erythrocytes . Monocytes and macrophages are recruited within

928-492: Is made of dense regular connective tissue , whose main cellular components are special fibroblasts called tendon cells (tenocytes). Tendon cells synthesize the tendon's extracellular matrix , which abounds with densely-packed collagen fibers . The collagen fibers run parallel to each other and are grouped into fascicles. Each fascicle is bound by an endotendineum , which is a delicate loose connective tissue containing thin collagen fibrils and elastic fibers. A set of fascicles

986-480: Is then attached to the base of the tendon cells via basal hemidesmosomes, while apical hemidesmosomes, which sit atop microvilli , attach the tendon cells to a thin layer of collagen . This is in turn attached to the shell via organic fibres which insert into the shell. Molluscan tendon cells appear columnar and contain a large basal cell nucleus . The cytoplasm is filled with granular endoplasmic reticulum and sparse golgi . Dense bundles of microfilaments run

1044-553: Is well known to promote angiogenesis and to induce endothelial cell proliferation and migration, and VEGF mRNA has been shown to be expressed at the site of tendon injuries along with collagen I mRNA. Bone morphogenetic proteins (BMPs) are a subgroup of TGF-β superfamily that can induce bone and cartilage formation as well as tissue differentiation, and BMP-12 specifically has been shown to influence formation and differentiation of tendon tissue and to promote fibrogenesis. In animal models, extensive studies have been conducted to investigate

1102-662: The meninges (the three membranes covering the brain and spinal cord ) and synovial membranes that line joint cavities. Mucous membranes and serous membranes are epithelial with an underlying layer of loose connective tissue. Fiber types found in the extracellular matrix are collagen fibers , elastic fibers , and reticular fibers . Ground substance is a clear, colorless, and viscous fluid containing glycosaminoglycans and proteoglycans allowing fixation of Collagen fibers in intercellular spaces. Examples of non-fibrous connective tissue include adipose tissue (fat) and blood . Adipose tissue gives "mechanical cushioning" to

1160-420: The stroma —or structural support—for the parenchyma (that is, the bulk of functional substance) of the organ. Mesenchyme is a type of connective tissue found in developing organs of embryos that is capable of differentiation into all types of mature connective tissue. Another type of relatively undifferentiated connective tissue is the mucous connective tissue known as Wharton's jelly , found inside

1218-486: The umbilical cord . This tissue is no longer present after birth, leaving only scattered mesenchymal cells throughout the body. Various types of specialized tissues and cells are classified under the spectrum of connective tissue, and are as diverse as brown and white adipose tissue , blood , cartilage and bone . Cells of the immune system—such as macrophages , mast cells , plasma cells , and eosinophils —are found scattered in loose connective tissue, providing

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1276-519: The ECM during the healing process after a tendon injury. Certain MMPs including MMP-1, MMP-2, MMP-8, MMP-13, and MMP-14 have collagenase activity, meaning that, unlike many other enzymes, they are capable of degrading collagen I fibrils. The degradation of the collagen fibrils by MMP-1 along with the presence of denatured collagen are factors that are believed to cause weakening of the tendon ECM and an increase in

1334-566: The actin cytoskeleton can activate integrins, which mediate "outside-in" and "inside-out" signaling between the cell and the matrix. G-proteins , which induce intracellular signaling cascades, may also be important, and ion channels are activated by stretching to allow ions such as calcium, sodium, or potassium to enter the cell. Sinew was widely used throughout pre-industrial eras as a tough, durable fiber . Some specific uses include using sinew as thread for sewing, attaching feathers to arrows (see fletch ), lashing tool blades to shafts, etc. It

1392-452: The body, among other functions. Although there is no dense collagen network in adipose tissue, groups of adipose cells are kept together by collagen fibers and collagen sheets in order to keep fat tissue under compression in place (for example, the sole of the foot). Both the ground substance and proteins (fibers) create the matrix for connective tissue. Type I collagen is present in many forms of connective tissue, and makes up about 25% of

1450-464: The collagen fiber diameter and orientation. The collagen fibrils are parallel to each other and closely packed, but show a wave-like appearance due to planar undulations, or crimps, on a scale of several micrometers. In tendons, the collagen fibres have some flexibility due to the absence of hydroxyproline and proline residues at specific locations in the amino acid sequence, which allows the formation of other conformations such as bends or internal loops in

1508-477: The collagen fibrils at specific locations. The proteoglycans are interwoven with the collagen fibrils – their glycosaminoglycan (GAG) side chains have multiple interactions with the surface of the fibrils – showing that the proteoglycans are important structurally in the interconnection of the fibrils. The major GAG components of the tendon are dermatan sulfate and chondroitin sulfate , which associate with collagen and are involved in

1566-484: The collagen fibrils. When decorin molecules are bound to a collagen fibril, their dermatan sulfate chains may extend and associate with other dermatan sulfate chains on decorin that is bound to separate fibrils, therefore creating interfibrillar bridges and eventually causing parallel alignment of the fibrils. The tenocytes produce the collagen molecules, which aggregate end-to-end and side-to-side to produce collagen fibrils. Fibril bundles are organized to form fibres with

1624-436: The deciding factor regarding actual and potential muscle size. For example, all other relevant biological factors being equal, a man with a shorter tendons and a longer biceps muscle will have greater potential for muscle mass than a man with a longer tendon and a shorter muscle. Successful bodybuilders will generally have shorter tendons. Conversely, in sports requiring athletes to excel in actions such as running or jumping, it

1682-399: The design of more effective exercises for astronauts . Tendons are subject to many types of injuries. There are various forms of tendinopathies or tendon injuries due to overuse. These types of injuries generally result in inflammation and degeneration or weakening of the tendons, which may eventually lead to tendon rupture . Tendinopathies can be caused by a number of factors relating to

1740-689: The effects of mechanical strain in the form of activity level on tendon injury and healing. While stretching can disrupt healing during the initial inflammatory phase, it has been shown that controlled movement of the tendons after about one week following an acute injury can help to promote the synthesis of collagen by the tenocytes, leading to increased tensile strength and diameter of the healed tendons and fewer adhesions than tendons that are immobilized. In chronic tendon injuries, mechanical loading has also been shown to stimulate fibroblast proliferation and collagen synthesis along with collagen realignment, all of which promote repair and remodeling. To further support

1798-462: The elongated tenocytes closely packed between them. There is a three-dimensional network of cell processes associated with collagen in the tendon. The cells communicate with each other through gap junctions , and this signalling gives them the ability to detect and respond to mechanical loading. These communications happen by two proteins essentially: connexin 43 , present where the cells processes meet and in cell bodies connexin 32 , present only where

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1856-443: The fibres of the tendon. More recently, tests carried out in vivo (through MRI ) and ex vivo (through mechanical testing of various cadaveric tendon tissue) have shown that healthy tendons are highly anisotropic and exhibit a negative Poisson's ratio ( auxetic ) in some planes when stretched up to 2% along their length, i.e. within their normal range of motion. After this 'toe' region, the structure becomes significantly stiffer, and has

1914-413: The fibril assembly process during tendon development. Dermatan sulfate is thought to be responsible for forming associations between fibrils, while chondroitin sulfate is thought to be more involved with occupying volume between the fibrils to keep them separated and help withstand deformation. The dermatan sulfate side chains of decorin aggregate in solution, and this behavior can assist with the assembly of

1972-454: The fibril to elongate and decrease in diameter under tension. However, the proteoglycans may also have a role in the tensile properties of tendon. The structure of tendon is effectively a fibre composite material, built as a series of hierarchical levels. At each level of the hierarchy, the collagen units are bound together by either collagen crosslinks, or the proteoglycans, to create a structure highly resistant to tensile load. The elongation and

2030-421: The fibrils become aligned in the direction of mechanical stress. The final maturation stage occurs after ten weeks, and during this time there is an increase in crosslinking of the collagen fibrils, which causes the tissue to become stiffer. Gradually, over about one year, the tissue will turn from fibrous to scar-like. Matrix metalloproteinases (MMPs) have a very important role in the degradation and remodeling of

2088-472: The fibrils then assemble further to form fascicles, which are about 10 mm in length with a diameter of 50–300 μm, and finally into a tendon fibre with a diameter of 100–500 μm. The collagen in tendons are held together with proteoglycan (a compound consisting of a protein bonded to glycosaminoglycan groups, present especially in connective tissue) components including decorin and, in compressed regions of tendon, aggrecan , which are capable of binding to

2146-458: The first 24 hours, and phagocytosis of necrotic materials at the injury site occurs. After the release of vasoactive and chemotactic factors, angiogenesis and the proliferation of tenocytes are initiated. Tenocytes then move into the site and start to synthesize collagen III. After a few days, the repair or proliferation stage begins. In this stage, the tenocytes are involved in the synthesis of large amounts of collagen and proteoglycans at

2204-413: The ground for starting inflammatory and immune responses upon the detection of antigens . There are many types of connective tissue disorders, such as: Tenocytes Tendon cells , or tenocytes, are elongated fibroblast type cells. The cytoplasm is stretched between the collagen fibres of the tendon. They have a central cell nucleus with a prominent nucleolus . Tendon cells have

2262-417: The healing process. IGF-1 increases collagen and proteoglycan production during the first stage of inflammation, and PDGF is also present during the early stages after injury and promotes the synthesis of other growth factors along with the synthesis of DNA and the proliferation of tendon cells. The three isoforms of TGF-β (TGF-β1, TGF-β2, TGF-β3) are known to play a role in wound healing and scar formation. VEGF

2320-699: The hindlimb, while in ornithischian dinosaurs, ossified axial muscle tendons form a latticework along the neural and haemal spines on the tail, presumably for support. Fibrous connective tissue The term "connective tissue" (in German, Bindegewebe ) was introduced in 1830 by Johannes Peter Müller . The tissue was already recognized as a distinct class in the 18th century. Connective tissue can be broadly classified into connective tissue proper, and special connective tissue. Connective tissue proper includes loose connective tissue, and dense connective tissue. Loose and dense connective tissue are distinguished by

2378-488: The mechanical properties. While the collagen fibrils allow tendons to resist tensile stress, the proteoglycans allow them to resist compressive stress. These molecules are very hydrophilic, meaning that they can absorb a large amount of water and therefore have a high swelling ratio. Since they are noncovalently bound to the fibrils, they may reversibly associate and disassociate so that the bridges between fibrils can be broken and reformed. This process may be involved in allowing

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2436-427: The mineralized fibrocartilage near the interface with the bone. Collagen fibres coalesce into macroaggregates . After secretion from the cell, cleaved by procollagen N- and C- proteases , the tropocollagen molecules spontaneously assemble into insoluble fibrils. A collagen molecule is about 300 nm long and 1–2 nm wide, and the diameter of the fibrils that are formed can range from 50–500 nm. In tendons,

2494-565: The organisation and structure of this matrix, the different mechanical properties required by different tendons can be achieved. Energy storing tendons have been shown to utilise significant amounts of sliding between fascicles to enable the high strain characteristics they require, whilst positional tendons rely more heavily on sliding between collagen fibres and fibrils. However, recent data suggests that energy storing tendons may also contain fascicles which are twisted, or helical, in nature - an arrangement that would be highly beneficial for providing

2552-475: The past two decades, much research has focused on the elastic properties of some tendons and their ability to function as springs. Not all tendons are required to perform the same functional role, with some predominantly positioning limbs, such as the fingers when writing (positional tendons) and others acting as springs to make locomotion more efficient (energy storing tendons). Energy storing tendons can store and recover energy at high efficiency. For example, during

2610-728: The potential for another rupture to occur. In response to repeated mechanical loading or injury, cytokines may be released by tenocytes and can induce the release of MMPs, causing degradation of the ECM and leading to recurring injury and chronic tendinopathies. A variety of other molecules are involved in tendon repair and regeneration. There are five growth factors that have been shown to be significantly upregulated and active during tendon healing: insulin-like growth factor 1 (IGF-I), platelet-derived growth factor (PDGF), vascular endothelial growth factor (VEGF), basic fibroblast growth factor (bFGF), and transforming growth factor beta (TGF-β). These growth factors all have different roles during

2668-483: The processes meet. Blood vessels may be visualized within the endotendon running parallel to collagen fibres, with occasional branching transverse anastomoses . The internal tendon bulk is thought to contain no nerve fibres, but the epitenon and paratenon contain nerve endings, while Golgi tendon organs are present at the myotendinous junction between tendon and muscle. Tendon length varies in all major groups and from person to person. Tendon length is, in practice,

2726-511: The ratio of ground substance to fibrous tissue. Loose connective tissue has much more ground substance and a relative lack of fibrous tissue, while the reverse is true of dense connective tissue. Loose connective tissue includes reticular connective tissue , and adipose tissue . Dense connective tissue also known as fibrous tissue is subdivided into dense regular and dense irregular connective tissue . Dense regular connective tissue, found in structures such as tendons and ligaments ,

2784-465: The response of tenocytes to mechanical force that enable them to alter their gene expression, protein synthesis, and cell phenotype, and eventually cause changes in tendon structure. A major factor is mechanical deformation of the extracellular matrix , which can affect the actin cytoskeleton and therefore affect cell shape, motility, and function. Mechanical forces can be transmitted by focal adhesion sites, integrins , and cell-cell junctions. Changes in

2842-412: The site of injury, and the levels of GAG and water are high. After about six weeks, the remodeling stage begins. The first part of this stage is consolidation, which lasts from about six to ten weeks after the injury. During this time, the synthesis of collagen and GAGs is decreased, and the cellularity is also decreased as the tissue becomes more fibrous as a result of increased production of collagen I and

2900-537: The site where they actively engage in movement, passing through regions where space is premium, like the carpal tunnel . There are about 4000 tendons in the human body, of which 55 are listed here: Naming convention for the table: Traditionally, tendons have been considered to be a mechanism by which muscles connect to bone as well as muscles itself, functioning to transmit forces. This connection allows tendons to passively modulate forces during locomotion, providing additional stability with no active work. However, over

2958-407: The spring-like behaviour required in these tendons. Tendons are viscoelastic structures, which means they exhibit both elastic and viscous behaviour. When stretched, tendons exhibit typical "soft tissue" behavior. The force-extension, or stress-strain curve starts with a very low stiffness region, as the crimp structure straightens and the collagen fibres align suggesting negative Poisson's ratio in

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3016-463: The steppe nomads of Eurasia, and Native Americans. The first stone throwing artillery also used the elastic properties of sinew. Sinew makes for an excellent cordage material for three reasons: It is extremely strong, it contains natural glues, and it shrinks as it dries, doing away with the need for knots. Tendon (in particular, beef tendon) is used as a food in some Asian cuisines (often served at yum cha or dim sum restaurants). One popular dish

3074-412: The strain of the collagen fibrils alone have been shown to be much lower than the total elongation and strain of the entire tendon under the same amount of stress, demonstrating that the proteoglycan-rich matrix must also undergo deformation, and stiffening of the matrix occurs at high strain rates. This deformation of the non-collagenous matrix occurs at all levels of the tendon hierarchy, and by modulating

3132-431: The tendon extracellular matrix (ECM), and their classification has been difficult because their symptoms and histopathology often are similar. Types of tendinopathy include: Tendinopathies may be caused by several intrinsic factors including age, body weight, and nutrition. The extrinsic factors are often related to sports and include excessive forces or loading, poor training techniques, and environmental conditions. It

3190-451: The theory that movement and activity assist in tendon healing, it has been shown that immobilization of the tendons after injury often has a negative effect on healing. In rabbits, collagen fascicles that are immobilized have shown decreased tensile strength, and immobilization also results in lower amounts of water, proteoglycans, and collagen crosslinks in the tendons. Several mechanotransduction mechanisms have been proposed as reasons for

3248-612: The total protein content of the mammalian body. Connective tissue has a wide variety of functions that depend on the types of cells and the different classes of fibers involved. Loose and dense irregular connective tissue , formed mainly by fibroblasts and collagen fibers , have an important role in providing a medium for oxygen and nutrients to diffuse from capillaries to cells, and carbon dioxide and waste substances to diffuse from cells back into circulation. They also allow organs to resist stretching and tearing forces. Dense regular connective tissue , which forms organized structures,

3306-455: The triple helix and results in the development of crimps. The crimps in the collagen fibrils allow the tendons to have some flexibility as well as a low compressive stiffness. In addition, because the tendon is a multi-stranded structure made up of many partially independent fibrils and fascicles, it does not behave as a single rod, and this property also contributes to its flexibility. The proteoglycan components of tendons also are important to

3364-408: Was believed that tendons could not undergo matrix turnover and that tenocytes were not capable of repair. However, it has since been shown that, throughout the lifetime of a person, tenocytes in the tendon actively synthesize matrix components as well as enzymes such as matrix metalloproteinases (MMPs) can degrade the matrix. Tendons are capable of healing and recovering from injuries in a process that

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