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Cartilage
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Cartilage is a resilient and smooth type of connective tissue. It is a semi-transparent and non-porous
type of tissue. It is usually covered by a tough and fibrous membrane called perichondrium. In
tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage,[1] and is a
structural component of many body parts including the rib cage, the neck and the bronchial tubes,
and the intervertebral discs. In other taxa, such as chondrichthyans, but also in cyclostomes, it may
constitute a much greater proportion of the skeleton.[2] It is not as hard and rigid as bone, but it is
much stiffer and much less flexible than muscle. The matrix of cartilage is made up of
glycosaminoglycans, proteoglycans, collagen fibers and, sometimes, elastin. It usually grows quicker
than bone.

Because of its rigidity, cartilage often serves

Cartilage
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 or

nerves, hence it is insensitive. Some
fibrocartilage such as the meniscus of the
knee does however have blood supply in part.
Nutrition is supplied to the chondrocytes by
Light micrograph of undecalcified hyaline cartilage
diffusion. The compression of the articular
showing chondrocytes and organelles, lacunae and
cartilage or flexion of the elastic cartilage
matrix.
generates fluid flow, which assists the diffusion
of nutrients to the chondrocytes. Compared to Identifiers
other connective tissues, cartilage has a very
slow turnover of its extracellular matrix and is MeSH D002356

documented to repair at only a very slow rate TA98 A02.0.00.005

relative to other tissues.
TA2 381

Contents Anatomical terminology

[edit on Wikidata]
Structure
Development

Articular cartilage

Function
Mechanical properties

Frictional properties
There are three different types of cartilage: elastic
Repair (A), hyaline (B), and fibrous (C). In elastic
cartilage, the cells are closer together creating
Clinical significance
less intercellular space. Elastic cartilage is found
Disease in the external ear flaps and in parts of the larynx.
Hyaline cartilage has fewer cells than elastic
Imaging
cartilage; there is more intercellular space. Hyaline
Other animals cartilage is found in the nose, ears, trachea, parts
of the larynx, and smaller respiratory tubes.
Cartilaginous fish
Fibrous cartilage has the fewest cells so it has the
Invertebrate cartilage most intercellular space. Fibrous cartilage is found
in the spine and the menisci.
Arthropods

Mollusks

Sabellid polychaetes

Plants and fungi

References

Further reading

External links

Structure

Development
Main article: Chondrogenesis

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. In all vertebrates, cartilage is the main skeletal
tissue in early ontogenetic stages;[3][4] in osteichthyans, many cartilaginous elements subsequently
ossify through endochondral and perichondral ossification.[5]

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.[6] It has been identified that non-coding RNAs (e.g. miRNAs and long non-
coding RNAs) as the most important epigenetic modulators can affect the chondrogenesis. This also
justifies the non-coding RNAs' contribution in various cartilage-dependent pathological conditions
such as arthritis, and so on.[7]

Articular cartilage

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.[8] Cartilage growth thus refers to the matrix deposition, Section from mouse joint showing cartilage

but can also refer to both the growth and remodeling of the (purple)

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.

Function

Mechanical properties

The mechanical properties of articular cartilage in load-bearing joints such as the 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.[9]

Frictional properties

Lubricin, a glycoprotein abundant in cartilage and synovial fluid, plays a major role in bio-lubrication
and wear protection of cartilage.[10]

Repair

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. Over the last years,
surgeons and scientists have elaborated a series of cartilage repair procedures that help to postpone
the need for joint replacement. A tear of the meniscus of the knee cartilage can often be surgically
trimmed to reduce problems.

Biological engineering techniques are being developed to generate new cartilage, using a cellular
"scaffolding" material and cultured cells to grow artificial cartilage.[11] Extensive researches have been
conducted on freeze-thawed PVA hydrogels as a base material for such a purpose.[12] These gels have
exhibited great promises in terms of biocompatibility, wear resistance, shock absorption, friction
coefficient, flexibility, and lubrication, and thus are considered superior to polyethylene-based
cartilages. A two-year implantation of the PVA hydrogels as artificial meniscus in rabbits showed that
the gels remain intact without degradation, fracture, or loss of properties.[12]

Clinical significance

Disease
Main article: Chondropathy

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
Human skeleton
considered the result of "wear and tear" rather than a true disease. It is treated by
with articular
arthroplasty, the replacement of the joint by a synthetic joint often made of a stainless cartilage shown in
steel alloy (cobalt chromoly) and ultra-high-molecular-weight polyethylene blue

(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.[13] In osteoarthritis, increased expression of inflammatory cytokines and chemokines cause
aberrant changes in differentiated chondrocytes function which leads to an excess of chondrocyte
catabolic activity, mediated by factors including matrix metalloproteinases (MMPs) and aggrecanases.[14]

Traumatic rupture or detachment: The cartilage in the knee is frequently damaged but 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 asphyxiation 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.

Imaging

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.[15]

Other animals

Cartilaginous fish

Cartilaginous fish (chondrichthyes) or sharks, rays

and chimaeras have a skeleton composed entirely of
cartilage.

Invertebrate cartilage

Cartilage tissue can also be found among some Histological image of hyaline cartilage stained with
arthropods such as horseshoe crabs, some mollusks haematoxylin and eosin, under polarized light

such as marine snails and cephalopods, and some

annelids like sabellid polychaetes.

Arthropods

The most studied cartilage in arthropods is the branchial cartilage of Limulus polyphemus. It is a
vesicular cell-rich cartilage due to the large, spherical and vacuolated chondrocytes with no
homologies in other arthropods. Other type of cartilage found in Limulus polyphemus is the
endosternite cartilage, a fibrous-hyaline cartilage with chondrocytes of typical morphology in a fibrous
component, much more fibrous than vertebrate hyaline cartilage, with mucopolysaccharides
immunoreactive against chondroitin sulfate antibodies. There are homologous tissues to the
endosternite cartilage in other arthropods.[16] The embryos of Limulus polyphemus express ColA and
hyaluronan in the gill cartilage and the endosternite, which indicates that these tissues are fibrillar-
collagen-based cartilage. The endosternite cartilage forms close to Hh-expressing ventral nerve cords
and expresses ColA and SoxE, a Sox9 analog. This is also seen in gill cartilage tissue.[17]

Mollusks

In cephalopods, the models used for the studies of cartilage are Octopus vulgaris and Sepia officinalis.
The cephalopod cranial cartilage is the invertebrate cartilage that shows more resemblance to the
vertebrate hyaline cartilage. The growth is thought to take place throughout the movement of cells
from the periphery to the center. The chondrocytes present different morphologies related to their
position in the tissue.[16] The embryos of Sepia officinalis express ColAa, ColAb, and hyaluronan in the
cranial cartilages and other regions of chondrogenesis. This implies that the cartilage is fibrillar-
collagen-based. The Sepia officinalis embryo expresses hh, whose presence causes ColAa and ColAb
expression and is also able to maintain proliferating cells undiferentiated. It has been observed that
this species presents the expression SoxD and SoxE, analogs of the vertebrate Sox5/6 and Sox9, in
the developing cartilage. The cartilage growth pattern is the same as in vertebrate cartilage.[17]

In gastropods, the interest lies in the odontophore, a cartilaginous structure that supports the radula.
The most studied species regarding this particular tissue is Busycotypus canaliculatus. The
odontophore is a vesicular cell rich cartilage, consisting of vacuolated cells containing myoglobin,
surrounded by a low amount of extra cellular matrix containing collagen. The odontophore contains
muscle cells along with the chondrocytes in the case of Lymnaea and other mollusks that graze
vegetation.[16]

Sabellid polychaetes

The Sabellid polychaetes, or feather duster worms, have cartilage tissue with cellular and matrix
specialization supporting their tentacles. They present two distinct extracellular matrix regions. These
regions are an acellular fibrous region with a high collagen content, called cartilage-like matrix, and
collagen lacking a highly cellularized core, called osteoid-like matrix. The cartilage-like matrix
surrounds the osteoid-like matrix. The amount of the acellular fibrous region is variable. The model
organisms used in the study of cartilage in sabellid polychaetes are Potamilla sp and Myxicola
infundibulum.[16]

Plants and fungi

Vascular plants, particularly seeds, and the stems of some mushrooms, are sometimes called
"cartilaginous", although they contain no cartilage.[18]

References

1. ^ Sophia Fox, AJ; Bedi, A; Rodeo, SA (November 2009). "The basic science of articular cartilage: structure,
composition, and function" . Sports Health. 1 (6): 461–8. doi:10.1177/1941738109350438 .
PMC 3445147 . PMID 23015907 .

2. ^ de Buffrénil, Vivian; de Ricqlès, Armand J; Zylberberg, Louise; Padian, Kevin; Laurin, Michel; Quilhac,
Alexandra (2021). Vertebrate skeletal histology and paleohistology (Firstiton ed.). Boca Raton, FL: CRC Press.
pp. xii + 825. ISBN 978-1351189576.

3. ^ Buffrénil, Vivian de; Quilhac, Alexandra (2021). "An Overview of the Embryonic Development of the Bony
Skeleton" . Vertebrate Skeletal Histology and Paleohistology. CRC Press: 29–38. doi:10.1201/9781351189590-
2 . ISBN 9781351189590. S2CID 236422314 .

4. ^ Quilhac, Alexandra (2021). "An Overview of Cartilage Histology" . Vertebrate Skeletal Histology and
Paleohistology. CRC Press: 123–146. doi:10.1201/9781351189590-7 . ISBN 9781351189590.
S2CID 236413810 .

5. ^ Cervantes-Diaz, Fret; Contreras, Pedro; Marcellini, Sylvain (March 2017). "Evolutionary origin of
endochondral ossification: the transdifferentiation hypothesis". Development Genes and Evolution. 227 (2):
121–127. doi:10.1007/s00427-016-0567-y . PMID 27909803 . S2CID 21024809 .

6. ^ Asanbaeva A, Masuda K, Thonar EJ, Klisch SM, Sah RL (January 2008). "Cartilage growth and remodeling:
modulation of balance between proteoglycan and collagen network in vitro with beta-aminopropionitrile" .
Osteoarthritis and Cartilage. 16 (1): 1–11. doi:10.1016/j.joca.2007.05.019 . PMID 17631390 .

7. ^ Razmara E, Bitaraf A, Yousefi H, Nguyen TH, Garshasbi M, Cho WC, Babashah S (September 2019). "Non-
Coding RNAs in Cartilage Development: An Updated Review" . International Journal of Molecular Sciences. 20
(18): 4475. doi:10.3390/ijms20184475 . PMC 6769748 . PMID 31514268 .

8. ^ Asanbaeva A, Tam J, Schumacher BL, Klisch SM, Masuda K, Sah RL (June 2008). "Articular cartilage tensile
integrity: modulation by matrix depletion is maturation-dependent" . Archives of Biochemistry and
Biophysics. 474 (1): 175–82. doi:10.1016/j.abb.2008.03.012 . PMC 2440786 . PMID 18394422 .

9. ^ Hayes WC, Mockros LF (October 1971). "Viscoelastic properties of human articular cartilage" (PDF).
Journal of Applied Physiology. 31 (4): 562–8. doi:10.1152/jappl.1971.31.4.562 . PMID 5111002 .

10. ^ Rhee DK, Marcelino J, Baker M, Gong Y, Smits P, Lefebvre V, et al. (March 2005). "The secreted glycoprotein
lubricin protects cartilage surfaces and inhibits synovial cell overgrowth" . The Journal of Clinical
Investigation. 115 (3): 622–31. doi:10.1172/JCI22263 . PMC 548698 . PMID 15719068 .

11. ^ International Cartilage Repair Society ICRS

12. ^ a b
Adelnia, Hossein; Ensandoost, Reza; Shebbrin Moonshi, Shehzahdi; Gavgani, Jaber Nasrollah; Vasafi,
Emad Izadi; Ta, Hang Thu (2022-02-05). "Freeze/thawed polyvinyl alcohol hydrogels: Present, past and
future" . European Polymer Journal. 164: 110974. doi:10.1016/j.eurpolymj.2021.110974 .
hdl:10072/417476 . ISSN 0014-3057 . S2CID 245576810 .

13. ^ "Supplements for osteoarthritis 'do not work' " . BBC News. 16 September 2010.

14. ^ Ansari, Mohammad Y.; Ahmad, Nashrah; Haqqi, Tariq M. (2018-09-05). "Butein Activates Autophagy
Through AMPK/TSC2/ULK1/mTOR Pathway to Inhibit IL-6 Expression in IL-1β Stimulated Human
Chondrocytes" . Cellular Physiology and Biochemistry. 49 (3): 932–946. doi:10.1159/000493225 .
ISSN 1015-8987 . PMID 30184535 . S2CID 52166938 .

15. ^ Osteoarthritis Archived 2011-07-07 at the Wayback Machine. Osteoarthritis.about.com. Retrieved on

2015-10-26.

16. ^ a b c d
Cole AG, Hall BK (2004). "The nature and significance of invertebrate cartilages revisited: distribution
and histology of cartilage and cartilage-like tissues within the Metazoa". Zoology. 107 (4): 261–73.
doi:10.1016/j.zool.2004.05.001 . PMID 16351944 .

17. ^ a b
Tarazona OA, Slota LA, Lopez DH, Zhang G, Cohn MJ (May 2016). "The genetic program for cartilage
development has deep homology within Bilateria". Nature. 533 (7601): 86–9. Bibcode:2016Natur.533...86T .
doi:10.1038/nature17398 . PMID 27111511 . S2CID 3932905 .

18. ^ Eflora – Glossary . University of Sydney (2010-06-16). Retrieved on 2015-10-26.

Further reading

Keller-Peck C (2008). Vertebrate Histology, ZOOL 400. Boise State University.

External links

Cartilage.org , International Cartilage Regeneration & Joint Preservation Society

KUMC.edu Archived 2011-04-08 at the Wayback Machine, Cartilage tutorial, University of Kansas
Medical Center

Bartleby.com , text from Gray's anatomy

MadSci.org , I've heard 'Ears and nose do not ever stop growing.' Is this false?

CartilageHealth.com , Information on Articular Cartilage Injury Prevention, Repair and Rehabilitation

About.com Archived 2011-07-07 at the Wayback Machine, Osteoarthritis

Cartilage types

Different cartilages on TheFreeDictionary

Cartilage photomicrographs

Last edited on 19 February 2024, at 05:17

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