Cartilage is a resilient and smooth elastic tissue, rubber-like padding that covers and protects the ends of long bones at the joints, and is a structural component of the rib cage, the ear, the nose, the bronchial tubes, the intervertebral discs, and many other body components. It is not as hard and rigid as bone, but it is much stiffer and much less flexible than muscle.
Because of its rigidity, cartilage often serves the purpose of holding tubes open in the body. Examples include the rings of the trachea, such as the cricoid cartilage and carina.
Cartilage is composed of specialized cells called chondrocytes that produce a large amount of collagenous extracellular matrix, abundant ground substance that is rich in proteoglycan and elastin fibers. Cartilage is classified in three types, elastic cartilage, hyaline cartilage and fibrocartilage, which differ in relative amounts of collagen and proteoglycan.
Cartilage does not contain blood vessels (it is avascular) or nerves (it is aneural). Nutrition is supplied to the chondrocytes by diffusion. The compression of the articular cartilage or flexion of the elastic cartilage generates fluid flow, which assists diffusion of nutrients to the chondrocytes. Compared to other connective tissues, cartilage has a very slow turnover of its extracellular matrix and does not repair.
There are three different types of cartilage; elastic (A), hyaline (B), and fibrous (C). In elastic cartilage the cells are closer together creating less intercellular space. Elastic cartilage is found in the external ear flaps and in parts of the larynx. Hyaline cartilage has fewer cells than elastic cartilage, there is more intercellular space. Hyaline cartilage is found in the nose, ears, trachea, parts of the larynx, and smaller respiratory tubes. Fibrous cartilage has the least amount of cells so it has the most amount of intercellular space. Fibrous cartilage is found in the spine and the menisci.
In embryogenesis, the skeletal system is derived from the mesoderm germ layer. Chondrification (also known as chondrogenesis) is the process by which cartilage is formed from condensed mesenchyme tissue, which differentiates into chondroblasts and begins secreting the molecules (aggrecan and collagen type II) that form the extracellular matrix.
Following the initial chondrification that occurs during embryogenesis, cartilage growth consists mostly of the maturing of immature cartilage to a more mature state. The division of cells within cartilage occurs very slowly, and thus growth in cartilage is usually not based on an increase in size or mass of the cartilage itself.
The articular cartilage function is dependent on the molecular composition of the extracellular matrix (ECM). The ECM consists mainly of proteoglycan and collagens. The main proteoglycan in cartilage is aggrecan, which, as its name suggests, forms large aggregates with hyaluronan. These aggregates are negatively charged and hold water in the tissue. The collagen, mostly collagen type II, constrains the proteoglycans. The ECM responds to tensile and compressive forces that are experienced by the cartilage. Cartilage growth thus refers to the matrix deposition, but can also refer to both the growth and remodeling of the extracellular matrix. Due to the great stress on the patellofemoral joint during resisted knee extension, the articular cartilage of the patella is among the thickest in the human body.
The mechanical properties of articular cartilage in load bearing joints such as knee and hip have been studied extensively at macro, micro and nano-scales. These mechanical properties include the response of cartilage in frictional, compressive, shear and tensile loading. Cartilage is resilient and displays viscoelastic properties.
Lubricin, a glycoprotein abundant in cartilage and synovial fluid, plays a major role in bio-lubrication and wear protection of cartilage.
Cartilage has limited repair capabilities: Because chondrocytes are bound in lacunae, they cannot migrate to damaged areas. Therefore, cartilage damage is difficult to heal. Also, because hyaline cartilage does not have a blood supply, the deposition of new matrix is slow. Damaged hyaline cartilage is usually replaced by fibrocartilage scar tissue. Over the last years, surgeons and scientists have elaborated a series of cartilage repair procedures that help to postpone the need for joint replacement.
Bioengineering techniques are being developed to generate new cartilage, using a cellular "scaffolding" material and cultured cells to grow artificial cartilage.
Several diseases can affect cartilage. Chondrodystrophies are a group of diseases, characterized by the disturbance of growth and subsequent ossification of cartilage. Some common diseases that affect the cartilage are listed below.
- Osteoarthritis: Osteoarthritis is a disease of the whole joint, however one of the most affected tissues is the articular cartilage. The cartilage covering bones (articular cartilage--a subset of hyaline cartilage) is thinned, eventually completely wearing away, resulting in a "bone against bone" within the joint, leading to reduced motion, and pain. Osteoarthritis affects the joints exposed to high stress and is therefore considered the result of "wear and tear" rather than a true disease. It is treated by arthroplasty, the replacement of the joint by a synthetic joint often made of a stainless steel alloy (cobalt chromoly) and ultra high molecular weight polyethylene (UHMWPE). Chondroitin sulfate or glucosamine sulfate supplements, have been claimed to reduce the symptoms of osteoarthritis but there is little good evidence to support this claim.
- Traumatic rupture or detachment: The cartilage in the knee is frequently damaged, and can be partially repaired through knee cartilage replacement therapy. Often when athletes talk of damaged "cartilage" in their knee, they are referring to a damaged meniscus ( a fibrocartilage structure) and not the articular cartilage.
- Achondroplasia: Reduced proliferation of chondrocytes in the epiphyseal plate of long bones during infancy and childhood, resulting in dwarfism.
- Costochondritis: Inflammation of cartilage in the ribs, causing chest pain.
- Spinal disc herniation : Asymmetrical compression of an intervertebral disc ruptures the sac-like disc, causing a herniation of its soft content. The hernia often compresses the adjacent nerves and causes back pain.
- Relapsing polychondritis: a destruction, probably autoimmune, of cartilage, especially of the nose and ears, causing disfiguration. Death occurs by suffocation as the larynx loses its rigidity and collapses.
Tumors made up of cartilage tissue, either benign or malignant, can occur. They usually appear in bone, rarely in pre-existing cartilage. The benign tumors are called chondroma, the malignant ones chondrosarcoma. Tumors arising from other tissues may also produce a cartilage-like matrix, the best known being pleomorphic adenoma of the salivary glands.
The matrix of cartilage acts as a barrier, preventing the entry of lymphocytes or diffusion of immunoglobulins. This property allows for the transplantation of cartilage from one individual to another without fear of tissue rejection.
Cartilage does not absorb x-rays under normal In vivo conditions, but a dye can be injected into the synovial membrane that will cause the x-rays to be absorbed by the dye. The resulting void on the radiographic film between the bone and meniscus represents the cartilage. For In vitro x-ray scans, the outer soft tissue is most likely removed, so the cartilage and air boundary are enough to contrast the presence of cartilage due to the refraction of the x-ray.
Cartilaginous fish (chondrichthyes) like sharks, rays and skates have a skeleton composed entirely of cartilage.
Cartilage tissue can also be found among invertebrates such as horseshoe crabs, marine snails, and cephalopods.
Plants and fungi
- Vascular plants, particularly seeds, and the stems of some mushrooms, may be referred to as being "cartilaginous," although they contain no cartilage.
- ^ Asanbaeva A, Masuda K, Thonar EJ, Klisch SM, Sah RL (2008). "Cartilage growth and remodeling: Modulation of balance between proteoglycan and collagen network in vitro with ?-aminopropionitrile1". Osteoarthritis and Cartilage. 16 (1): 1-11. doi:10.1016/j.joca.2007.05.019. PMID 17631390.
- ^ Asanbaeva A, Tam J, Schumacher BL, Klisch SM, Masuda K, Sah RL (2008). "Articular cartilage tensile integrity: Modulation by matrix depletion is maturation-dependent". Archives of Biochemistry and Biophysics. 474 (1): 175-182. doi:10.1016/j.abb.2008.03.012. PMC 2440786 . PMID 18394422.
- ^ Hayes WC, Mockros LF (1971). "Viscoelastic properties of human articular cartilage" (PDF). J Appl Physiol. 31 (4): 562-8. PMID 5111002.
- ^ Rhee DK, Marcelino J, Baker M, Gong Y, Smits P, Lefebvre V, Jay GD, Stewart M, Wang H, Warman ML, Carpten JD (2005). "The secreted glycoprotein lubricin protects cartilage surfaces and inhibits synovial cell overgrowth". J Clin Invest. 115 (3): 622-31. doi:10.1172/JCI22263. PMC 548698 . PMID 15719068.
- ^ International Cartilage Repair Society ICRS
- ^ "Supplements for osteoarthritis 'do not work'". BBC News. 16 September 2010.
- ^ Osteoarthritis. Osteoarthritis.about.com. Retrieved on 2015-10-26.
- ^ Eflora - Glossary. University of Sydney (2010-06-16). Retrieved on 2015-10-26.
- General references