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Research Review Fibrocartilage

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Research Review Fibrocartilage J. Anat. (1990), 171, pp. 1-15 1 Printed in Great Britain Research Review Fibrocartilage M. BENJAMIN AND E. J. EVANS Department of Anatomy, University of Wales College of Cardiff, PO Box 900, Cardif CF1 3 YF, Wales Fibrocartilage has long been a neglected tissue that is too often viewed as a poor relation of hyaline cartilage. It failed to achieve the status of a tissue with the early histologists, but it is beginning to come of age, for modem techniques are revealing some exciting secrets about fibrocartilage in knee joint menisci and intervertebral discs in particular. Yet there has never been any general review on fibrocartilage, and workers concerned with the tissue in one organ rarely consider it in another. Consequently, we lack any global picture that would encourage the spread of interest in the tissue and the effective exchange of ideas. Our review deals largely with the white fibrocartilage of standard texts and for reasons of space excludes yellow elastic cartilage. We have concentrated on fibrocartilage as a tissue rather than fibrocartilages as organs. HISTORICAL CONSIDERATIONS The most important work on cartilage in the older literature is that of Schaffer (1930). His monograph is a thorough, comparative account of cartilage and related tissues throughout the animal kingdom. The reader interested in fibrocartilage must also study Schaffer's account of chondroid tissue, for some tissues that would now be regarded as fibrocartilage were viewed by Schaffer as hyaline-cell chondroid tissue. He had a narrow vision of 'true' cartilage and called tissues where the cells were not shrunken in lacunae, 'chondroid'. GENERAL ASPECTS OF STRUCTURE Fibrocartilage is a transitional tissue that lacks a perichondrium and has structural and functional properties intermediate between those of dense fibrous connective tissue and hyaline cartilage. Fibrocartilage merges with the hyaline cartilage of the radius in the triangular fibrocartilage complex of the wrist (Benjamin, Evans & Pemberton, 1990) and with dense fibrous connective tissue in ligaments and tendons (Benjamin, Evans & Copp, 1986; Woo et al. 1988). In view of the gradual transition between fibrocartilage and dense fibrous connective tissue, it is worth noting that Masson's trichrome stains collagen under tension red and collagen under compression green (Flint, Lyons, Meaney & Williams, 1975). As fibrocartilage resists compression, it stains predominantly green. 2 M. BENJAMIN AND E. J. EVANS Cells The cells of fibrocartilage may be irregularly arranged or lie in longitudinal rows. Except for the occasional cell that is more typical of loose connective tissues (e.g. mast cells), the majority ofcells look like chondrocytes or fibroblasts, but it is often difficult to know which to call them. Furthermore, Somer & Somer (1983) argue that parts of the knee joint menisci where chondroitin sulphate is absent from around the cells should be regarded as chondroid tissue. Edwards & Chrisman (1979) refer to both chondrocytes and fibroblasts in the fibrocartilage that appears in healing articular cartilage. Cooper & Misol (1970) distinguish typical chondrocytes from typical fibroblasts in ligament attachments, but comment that "the cells gradually change structural characteristics" from one cell type to the other. Ghadially (1983) considers that most meniscal cells that look like fibroblasts are really flattened chondrocytes. At the electron microscope level, these cells have numerous short processes and are surrounded by a territorial matrix. The flattened meniscal cells are thus similar to those in the superficial zone of articular cartilage. As a general rule, the more chondrocytic cells are found in the centre of fibrocartilage, and the more fibroblastic at its periphery. Although Badi (1972) recognises only chondrocytes in the fibrocartilage at the attachment of the rat patellar ligament, he distinguishes two subclasses of tissue according to the size of the cells. Small-celled fibrocartilage can calcify without cell hypertrophy and remains throughout life. It is then the Type II chondroid bone of Beresford (1981). Large-celled fibrocartilage develops from the small and disappears at the end of skeletal growth. The ultrastructural features of fibrocartilaginous chondrocytes described by Cooper & Misol (1970), Merrilees & Flint (1980), Buckwalter (1982), Ghadially (1983) and by Okuda, Gorski & Amadio (1987) suggest that the fine structure of the cells is similar to that of the cells in hyaline cartilage. Eyre et al. (1988) distinguish between notochordal cells, fibroblasts and chondrocytes in intervertebral disc. His descriptions of the chondrocytes are in broad agreement with those of Ghadially (1983) in menisci, but he makes no mention of chondrocytic features of the fibroblasts in the outer part of the annulus fibrosus. Fibres The collagen fibres can be irregularly arranged or form striking patterns. Circumferential hoops are conspicuous in knee joint menisci (Arnoczky et al. 1988) and collagen fibres running in the long axis ofa ligament or tendon are typical ofmany attachment zones (Woo et al. 1988). However, in the fibrocartilaginous regions of tendons that pass around bony pulleys, the collagen fibres have a basket-weave appearance and some bundles run at right angles to the long axis of the tendon (Merrilees & Flint, 1980). The fibres in the inner, fibrocartilaginous part of the annulus fibrosus of intervertebral discs do not form such obvious lamellae as those in the outer, more fibrous parts (Buckwalter, 1982; Eyre et al. 1988). Seven different types ofcollagen have now been documented in the annulus fibrosus of the intervertebral disc (Eyre, 1988) and at least four in the menisci of the knee joint (Arnoczky et al. 1988). Type I collagen is the most abundant and accounts for about 90 % of the total meniscal collagen and 80% of the collagen of the annulus fibrosus. There is a reciprocal gradient in the distribution of Types I and II collagen in the intervertebral disc. Type II is absent from the most peripheral part of the disc, but accounts for 80% of collagen in the nucleus pulposus, where the proportion of Type Fibrocartilage 3 I collagen has fallen to low levels. Type II collagen accounts for 1-2 % of total meniscal collagens. Penile fibrocartilage of the rat is also predominantly Type I collagen, though Type II is present around the cells (Murakami, 1987). The abundant presence of Type I collagen (tensile in function) and the relative paucity of Type II collagen (characteristic of tissues subject to pressure) is often taken to be a key biochemical feature that distinguishes fibro- from hyaline cartilage (Arnoczky et al. 1988). Other collagens are present in fibrocartilage in small quantities. Types V and VI have been identified in both menisci and intervertebral discs, but Types IX and XI have only been found in discs (Eyre, 1988; Arnoczky et al. 1988). In addition, Type M collagen occurs in fibrocartilage that forms on the surface of osteophytes in degenerating femoral articular cartilage (Nemeth-Csoka & MeszaLros, 1983). The minor collagens may be important in anchoring chondrocytes to their matrix (Melrose & Ghosh, 1988) and in allowing collagen fibres to interact with one another and with proteoglycans (Eyre, 1988). Small numbers of elastic fibres and/or elastic system fibres are present in the fibrocartilage of intervertebral discs (Buckwalter, Cooper & Maynard, 1976; Cotta- Pereira, Del-Caro & Montes, 1984), menisci (Peters & Smillie, 1972; Ghadially, 1983; Arnoczky et al. 1988) and tendon or ligament attachment zones (Cooper & Misol, 1970). Amorphous matrix The major constituents of the amorphous matrix are proteoglycans. Our knowledge of them is based mainly on studies of knee joint menisci (Arnoczky et al. 1988) and intervertebral discs (Bogduk & Twomey, 1987; Eyre et al. 1988). It is only here that sufficient quantities ofthe tissue can be obtained for analysis. There is less proteoglycan in fibro- than in hyaline cartilage, but more than in pure fibrous tissue (Gillard, Reilly, Bell-Booth & Flint, 1979; Koob & Vogel, 1987). The mean amount of glycosamino- glycans in knee joint menisci is 10-12 % of that in hyaline articular cartilage (Arnoczky et al. 1988). The proteoglycans of fibrocartilage differ biochemically from those of hyaline cartilage (McNicol & Roughley, 1980; Roughley, McNicol, Santer & Buckwalter, 1981; Arnoczky et al. 1988; Eyre et al. 1988). They have been identified histochemically in fibrocartilaginous finger menisci where they are prominent in the territorial matrix of the chondrocytes (Fisher, Elliott, Cooke & Forrest, 1985). Little is known of the other non-collagenous proteins (NCP) in fibrocartilage. However, it is likely that the microenvironment of the cartilage cells is highly structured and that NCP (link proteins, calcium-binding proteins and matrix glycoproteins) are important for cell-matrix interrelationships (Melrose & Ghosh, 1988). The attachment of a chondrocyte to its matrix via specific cell membrane receptors and intercalated transmembrane glycoproteins may allow the cell to respond to any changes in its matrix and to co-ordinate growth and repair (Huang, 1977). It is thus interesting to note that degraded link proteins have been found in human intervertebral disc (Eyre et al. 1988). Blood supply Fibrocartilage is generally poorly vascularised. The lack of blood vessels is a striking feature of fibrocartilage in intervertebral discs (Humzah & Soames, 1988), knee joint menisci (Arnoczky et al. 1988), the articular
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