Epub 2010 Aug 31. Fibromodulin and lumican contain keratan sulfate chains linked to the central domain of the core protein and several sulfated tyrosine residues in the N-terminus. @article{Wight1986ProteoglycansIH, title={Proteoglycans in human long-term bone marrow cultures: biochemical and ultrastructural analyses. Proteoglycans in human long-term bone marrow cultures: biochemical and ultrastructural analyses. Proteoglycans are glycosylated proteins which have covalently attached highly anionic glycosaminoglycans. First, they stabilize the connective tissue by binding to collagen and elastic fibers, cells and water. 2006 Sep 1;379:92-100. doi: 10.1016/j.gene.2006.04.029. The binding of water also enables the collagen fibers to move without friction against each other. Bone matrix proteoglycans and glycoproteins are proportionally the most abundant constituents of the noncollagenous proteins in bone matrix. Ann Biomed Eng 24(Suppl. These are very important in cell–matrix interactions and cell signaling, and control many aspects of chondrocyte and osteoblast function, but as the cell density is low they do not contribute to a large fraction of the tissue content of proteoglycans. Red bone marrow does not contain _____. The outer 60% of this chain is used as the connection site for about 80 to 100 chondroitin sulfate chains. In contrast, inhibition of specific heparan sulfate proteoglycans results in less clear effects. While human and mouse bone architectures differ in multiple scales, the extent to which chimeric human-mouse bone physiologically interacts and structurally integrates remains unknown. In bone from Gorab Prx1 mutants total glycosaminoglycan levels and the relative percentage of dermatan sulfate were both strongly diminished. COVID-19 is an emerging, rapidly evolving situation. The short form of brevican is attached to plasma membrane via a GPI anchor.85,92,93 Hyalectans interact with hyaluronan and tenascin in the neuronal tissue.92, Jeffrey P. Gorski, Kurt D. Hankenson, in Principles of Bone Biology (Fourth Edition), 2020. Therefore, it has been postulated that these proteoglycans and their unique spatial interaction with the collagens may be involved in the mineralization of bone matrix The noncollagenous proteins function in bone matrix mineralization, cellular adhesion, and regulation of bone cell activity during coupling of bone formation and resorption. but little is known about the structure and composition of these macromolecules in this tissue. These proteins exist in the extracellular bone matrix, along with other non-cellular compounds that … II, p. 427. Proteoglycans and GAGs are not only important regulators of physiological SC function during embryonic development and tissue regeneration, their aberrant expression in cancer cells and within cells and the ECM of the tumor microenvironment (and CSC niches) makes an important contribution to tumor progression and recurrence by modulating CSC properties. (Buckwalter et al. They form a special class of glycoproteins with attached long unbranched and highly charged glycosaminoglycan chains [1, 2]. 1; Fig. Cartilage contains 70–75% (w/w) water and it has a high fibrillar collagen content (∼20% w/w) and also a high proteoglycan content (5–7% w/w). Please enable it to take advantage of the complete set of features! Milan AM, Sugars RV, Embery G, Waddington RJ. Spicules and trabeculae are found in _____. In this review, the complex PG composition in bone will be addressed together with the specific role played by PGs (or their GAGs chains) in bone biology, as regulatory molecules for bone resorption and their involvement in bone tumor development. In embryonic kidneys, sulfated proteoglycans are concentrated around the tip of the ureteric bud, and perturbation of their synthesis by β-D-xyloside results in the inhibition of ureteric bud branching morphogenesis.217,218 This perturbation also abolishes the expression of Wnt11 at the tip of the ureteric bud.219 As described previously, loss of Wnt11 expression at the tip of the ureteric bud can lead to loss of Gdnf expression in the metanephric mesenchyme. Covalent binding causes the binding of the binding protein. Biglycan may have two glycosaminoglycan chains—chondroitin sulfate or dermatan sulfate, or both—attached near the N-terminus through two closely spaced serine-glycine dipeptides. Decorin contains only one chondroitin sulfate or dermatan sulfate chain. These also contain chondroitin sulfate, but with varying degrees of epimerization to dermatan sulfate. doi: 10.1371/journal.pone.0192500. Proteoglycans play a crucial role in bone tissue formation promoting consolidation of collagen fibers and bonds between them and mineral crystals. Which of the following is an inorganic component of the bone matrix. This contrast in composition is also reflected in the proteoglycans present in the tissue. Versican may be considered a prototype of the hyalectan family. Google Scholar . 7, 8 Serving as structural proteins, some NCPs have been found to be directly involved in the deformation and failure of bone. The remaining 30% remain free. The 24 amino acid central LRR domain is conserved, but the N-terminal and C-terminal domains have patterns of cysteine residues involved in intrachain disulfide bonds that distinguish the four subfamilies: (1) biglycan, decorin, fibromodulin, and lumican; (2) keratocan and proline and arginine-rich end leucine-rich repeat protein (PRELP); (3) chondroadherin; and (4) epiphycan/PG-Lb and mimecan/osteoglycin. Using mice deficient in two members of the small leucine-rich proteoglycans, biglycan and … PRELP and chondroadherin may regulate cell-matrix interactions through binding to syndecan and α2β1 integrin. A) Proteoglycans B) Glycoproteins C) Collagen D) Hydroxyapatite E) Glycosaminoglycans. For PG aggregate to occur, many PGs must be bound to a central hyaluronic acid chain. Proteoglycans may also help to trap and store growth factors within the ECM. These four polymeric molecules are cleaved from their protein core in lysosomes; then they, plus hyaluronan (a GAG lacking a protein core), are catabolized further in lysosomes in a stepwise fashion by more than a dozen enzymes. 2010 Oct;277(19):3904-23. doi: 10.1111/j.1742-4658.2010.07800.x. Epub 2020 Jul 6. The remaining 10% is used for about 50 to 60 keratan sulfate chains. Further progress in discerning the specific signaling pathways in skeletal development and homeostasis is needed. While acidic phosphoproteins, localized in the mineralized tissues, play key roles in control of mineralization, the role of glycosaminoglycans (GAGs) is less clear. Binding or link proteins are required for this. 2005 Feb;76(2):127-35. doi: 10.1007/s00223-004-0033-0. Many forms of proteoglycans are present in virtually all extracellular matrices of connective tissues. Clipboard, Search History, and several other advanced features are temporarily unavailable. One exception is glypican (Gpc) 3, a gene encoding a heparan sulfate proteoglycan linked to the cell surface via a glycosyl-phosphatidylinositol anchor. Accordingly, the proteoglycans biglycan and decorin showed reduced glycanation. Because of SLRP participation in multiple degenerative, inflammatory, and … Extracellular matrix proteins and PGs bind tightly to hydroxyapatite which would protect these molecules from the destructive effects of temperature and chemical agents after … Proteoglycans within the extracellular matrix of human bone marrow have been implicated in the process of hematopoiesis. Occasionally, a few other shorter oligosaccharide chains can be bound. Consistent with the in vitro result described above, heparan sulfate biosynthesis perturbation results in loss of Wnt11 expression and reduced Gdnf expression. The central domain of hyalectans contains variable numbers of GAG chains, composed of chondroitin/dermatan sulfate (e.g., versican) or predominantly chondroitin sulfate—as is the case for aggrecan, neurocan, and brevican (see Table 3-4). 1990; Currier & Nelson 1992; Aaron & Bolander 2005; Van den Berg 2010, Goldman's Cecil Medicine (Twenty Fourth Edition), Kelley's Textbook of Rheumatology (Ninth Edition), Principles of Bone Biology (Fourth Edition). Proteoglycans are mostly found at the cell surface or in the extracellular matrix. GAGs exist both in the intracellular and extracellular space. In immune system cells a GAG binds to proteases, that is to enzymes which break up the proteins and prevent the cells destroying themselves. Kevin T. Bush, ... Sanjay K. Nigam, in Seldin and Giebisch's The Kidney (Fifth Edition), 2013. J Histochem Cytochem. Other oligosaccharides and GAGs are formed in the Golgi apparatus of the cell. Four distinct proteoglycans, named versican, aggrecan, neurocan, and brevican, constitute a family of molecules with homologous structural organization and similar biologic functions.85 These proteoglycans share a tripartite organization; the amino and carboxyl terminal domains of these proteoglycans interact with hyaluronan and lectins, respectively. Several additional storage disorders, termed mucolipidoses (MLs), are caused by a genetic defect in post-translational modification of lysosomal enzymes and share features with the MPS. Here, we identify that humanized bone is formed by a mosaic of human and mouse collagen, structurally … 1988; Eyre et al. Share it! In contrast, bone matrix is mainly mineral, reinforced with fibrillar collagens (∼5% w/w), it contains only 10% (w/w) water and it has a correspondingly low proteoglycan content (∼0.1% w/w). Mary B. Goldring, in Kelley's Textbook of Rheumatology (Ninth Edition), 2013, The major proteoglycan in articular cartilage is the large aggregating proteoglycan, or aggrecan, which consists of a core protein of 225 to 250 kD with covalently attached side chains of glycosaminoglycans, including approximately 100 chondroitin sulfate chains, 30 keratan sulfate chains, and shorter N-linked and O-linked oligosaccharides.5,8,40,41 Link protein, a small glycoprotein, stabilizes the noncovalent linkage between aggrecan and hyaluronic acid (also called hyaluronan) to form the proteoglycan aggregate that may contain 100 aggrecan monomers. Adsorption of proteoglycans to titanium powder (abstract). A) Compact Bone B) Bone Matrix C) Yellow Bone Marrow D) Red Bone Marrow E) Spongy Bone. 2008 Feb;99(2):316-23. doi: 10.1111/j.1349-7006.2007.00690.x. Aggrecan typically contains approximately 100 CS- and approximately 30 KS-enriched GAGs that are covalently linked to about a 220-kDa core protein (see Table 3-4). They also protect the tissue and the cells from penetrating bacteria. Dolcino M, Tinazzi E, Puccetti A, Lunardi C. Cells. Epub 2010 May 27. The stored water simultaneously serves as a transport route for nutrients and waste products as well (Grodzinsky 1983; Fleischmajer et al. Thus, PGs and GAGs are involved in the organization of collagen fibers (14). PLoS One. NIH This means that the cells must always remain synthetically active, otherwise there is the risk of a reduction in the quantity of ground substance. Bone matrix proteoglycans and glycoproteins are proportionally the most abundant constituents of the noncollagenous proteins in bone matrix. Proteoglycans are ubiquitous macromolecules of extracellular matrix, cell surface, and some intracellular granules. ScienceDirect ® is a registered trademark of Elsevier B.V. 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URL: https://www.sciencedirect.com/science/article/pii/B978012491543550070X, URL: https://www.sciencedirect.com/science/article/pii/B9780120885626500066, URL: https://www.sciencedirect.com/science/article/pii/B9780126386516500022, URL: https://www.sciencedirect.com/science/article/pii/B9781416002468500085, URL: https://www.sciencedirect.com/science/article/pii/B9780123814623000252, URL: https://www.sciencedirect.com/science/article/pii/B9780702034251000581, URL: https://www.sciencedirect.com/science/article/pii/B9781437716047002682, URL: https://www.sciencedirect.com/science/article/pii/B9781437717389000037, URL: https://www.sciencedirect.com/science/article/pii/B9780721602844500099, URL: https://www.sciencedirect.com/science/article/pii/B9780128148419000154, Mucosal Infection and Immune Responses to Simian Immunodeficiency Virus, Dynamics of Bone and Cartilage Metabolism (Second Edition), Textbook of Pediatric Rheumatology (Fifth Edition), Seldin and Giebisch's The Kidney (Fifth Edition), Fascia: The Tensional Network of the Human Body, Grodzinsky 1983; Fleischmajer et al. Defective cartilage and shortened limb development have been demonstrated in mice and chickens that contain mutated aggrecan genes.90,91, A neurocan-specific cDNA was first isolated from rat brain and the core-protein sequence encoded by this cDNA was reminiscent of the tripartite organization seen in other hyalectans. The composition of extracellular matrix proteoglycans was analyzed in early aging OXYS rats with signs of early osteoporosis versus Wistar rats … National Center for Biotechnology Information, Unable to load your collection due to an error, Unable to load your delegates due to an error. Copyright © 2020 Elsevier B.V. or its licensors or contributors. Given the fact that many growth factors that regulate branching morphogenesis of the ureteric bud are heparin (a heavily sulfated form of GAG chain)-binding, it is likely that global inhibition of GAG chain synthesis or its sulfation could compromise actions of these heparin-binding growth factors. NLM 2010 Jul;15(13-14):553-60. doi: 10.1016/j.drudis.2010.05.009. Synthetic enzymes for this GAG are expressed in the developing kidney, and the addition of HA to three-dimensional culture of UB cells promotes tubulogenesis and survival of cells. Finally, sclerostin and DMP1 are preferentially expressed by osteocytic cells in bone where they regulate osteogenesis by feeding back to osteoblastic cells and (together with PHEX) regulate phosphate metabolism via FGF23 production by osteocytes, respectively. Glycoproteins such as alkaline phosphatase, osteonectin, RGD-containing proteins (osteoadherin, TSP, fibronectin, vitronectin, OPN, and BSP), irisin, fibrillin, and tetranectin are produced at different stages of osteoblastic maturation. The nonaggregating small proteoglycans are not specific to cartilage, but in cartilage they serve specific roles in matrix structure and function, primarily by modulating collagen-fibril formation.42-44 Of the more than 10 leucine-rich repeat (LRR) proteoglycans discovered so far, only osteoadherin is not present in cartilage. chondroitin sulfate-GlcA-Gal-Gal-Xyl-PROTEIN). As the most abundant constituent of the cartilage ECM, aggrecan is found in giant aggregates with link proteins and hyaluronan, and occupies a large hydrodynamic volume (2 × 10−12 cm3) that may be equivalent to a bacterium.88 The lectin modules of both versican and aggrecan can interact with simple sugars found in glycoproteins; this binding is calcium dependent.89 In addition to its structural role in the cartilage ECM, aggrecan modulates chondrogenesis. Modulation of collagen fibrillogenesis by dentinal proteoglycans. Like it? eCollection 2020 Oct. Lei Y, Junxin C, Yongcan H, Xiaoguang L, Binsheng Y. J Bone Oncol. Proteoglycans at the bone-biomaterial interface in vitro (abstract). They bind water and provide the basis for absorbing high compressive loads. Velasco CR, Colliec-Jouault S, Redini F, Heymann D, Padrines M. Drug Discov Today. Bacteria can only move through the dense network and penetrate the tissue with difficulty. The G3 domain contains sequence homologies to epidermal growth factor, lectin, and complement regulatory protein, and participates in growth regulation, cell recognition, intracellular trafficking, and recognition, assembly, and stabilization of the ECM. Additionally, they bind cytokines, chemokines, growth factors, and morphogens, thereby protecting these factors against proteolysis and/or acting as a depot of regulatory factors when matrix degradation occurs. Humanized mouse models are increasingly studied to recapitulate human-like bone physiology. 2020 Nov;68(11):747-762. doi: 10.1369/0022155420937371. Syndecan 3 is a member of the family of heparan sulfate proteoglycans, which function as extracellular matrix receptors mediating the interaction of cells with extracellular components and as signaling molecules that control cell shape, adhesion, proliferation, and differentiation. They are used for the binding of the PG to a hyaluronic acid chain using a connecting protein. FREDERIC SHAPIRO, in Pediatric Orthopedic Deformities, 2001. 2020 Sep 28;25:100322. doi: 10.1016/j.jbo.2020.100322. The turnover of hyaluronic acid is about 2 to 4 days; for the other sulfated GAGs it is about 7 to 10 days. The diversity of bone proteoglycan (PG) structure and localisation (pericellular, extracellular in the organic bone matrix) reflects a broad spectrum of biological functions within a unique tissue. The G2 domain is separated from G1 by a linear interglobular domain and has two proteoglycan tandem repeats. Two proteoglycans found in most connective tissues, biglycan and decorin, … An Introduction to Glycosaminoglycans and Proteoglycans. Metal-Specific Biomaterial Accumulation in Human Peri-Implant Bone and Bone Marrow. Along the same lines, genetic inactivation of an enzyme, heparan sulfate 2-O-sulfotransferase, involved in heparan sulfate proteoglycan synthesis results in kidney agenesis.53 In this case, the ureteric bud forms from the Wolffian duct, but subsequent invasion of the metanephric mesenchyme is perturbed. Proteoglycan are characterized by the covalent attachment of long chain polysaccharides (glycosaminoglycans, GAGs) to core protein molecules. This site is not intended for people who suffers from rheumatoid arthritis (RA), gout, avascular necrosis (AVN), and cancer within bones, osteoporosis and other inflammatory or autoimmune diseases. In contrast, bone matrix is mainly … Reed E. Pyeritz, in Goldman's Cecil Medicine (Twenty Fourth Edition), 2012.  |  These chains are strongly hydrophilic and dominate the physical properties of proteoglycans, but the proteins to which they are attached are quite diverse in structure and form several distinct protein families. Proteoglycans are proteins that are heavily glycosylated. 2019 Aug 2;8(8):816. doi: 10.3390/cells8080816. The Ser residue is generally in the sequence -Ser-Gly-X-Gly- (where X can be any amino acid residue but proline), although not every protein with this sequence … Blocking antibodies against its receptor CD44 abolishes this effect.221 In addition, supplementation of cultures of isolated ureteric bud or whole kidneys with hyaluronidase inhibited branching morphogenesis.222 Specific sizes and concentrations of HA were also found to act independently in regulating UB branching, and in tubular maturation.222, While the protein core of proteoglycans, particularly the heparan sufate proteoglycans, play important roles in many developmental and physiologic processes, most of the function of these proteoglycans are mediated by the variably sulfated heparan sulfate GAG chain. In bone there is no aggrecan and the collagen fibril associated proteoglycans and biglycan are the predominant forms. Bone matrix proteoglycans and glycoproteins are proportionally the most abundant constituents of the noncollagenous proteins in bone matrix. Proteoglycans are most abundant in those tissues where the ECM is highly hydrated. Rajendra Raghow, ... Andrew Kang, in Vascular Medicine, 2006.  |  Hema-topoietically active human long-term bone marrow cultures were incubated with medium containing S-sulfate and 3H-glucosamine as labeling precursors. Long Non-Coding RNAs Target Pathogenetically Relevant Genes and Pathways in Rheumatoid Arthritis. Small quantities of other large proteoglycans are found in cartilage, including versican, which forms aggregates with hyaluronic acid, and perlecan, which is nonaggregating; however, these proteoglycans function primarily during skeletal development, where versican is expressed in prechondrogenic condensations, and perlecan is expressed in the cartilage anlagen after expression of type II collagen and aggrecan.41. Biglycan, decorin, and fibromodulin bind transforming growth factor (TGF)-β and the epidermal growth factor receptor and may modulate growth, remodeling, and repair. PGs and GAGs have many different functions. Proteoglycans are macromolecules consisting of a protein core to which are attached 50 to 100 unbranched glycosaminoglycans (chondroitin sulfate and O-linked keratan sulfate).25–27 At least five different protein cores have been defined. Negatively charged glycosaminoglycan side chains contribute to the fixed charge density of the matrix and, together with the highly anionic tyrosine-sulfation sites, permit multiple-site linkage between adjacent collagen fibrils, stabilizing the network. Temporal expression of proteoglycans in the rat limb during bone healing. There are several types of bone and joint pain, each with many potential sources or etiologies. The point of attachment is a serine (Ser) residue to which the glycosaminoglycan is joined through a tetrasaccharide bridge (e.g. }, author={T. Wight and M. Kinsella and A. Keating and J. The heavy negative loading of the GAGs and PGs causes the capacity to bind to water and because of its viscoelasticity allows the tissue to return to its original form after stress. Get the latest public health information from CDC: https://www.coronavirus.gov, Get the latest research information from NIH: https://www.nih.gov/coronavirus, Find NCBI SARS-CoV-2 literature, sequence, and clinical content: https://www.ncbi.nlm.nih.gov/sars-cov-2/. FEBS J. This site needs JavaScript to work properly. The ectopic expression of bone cell secreted noncollagenous proteins may also play a significant role in, normal systemic regulation of glucose and energy metabolism, and of male reproductive function, as well as in pathological processes such as bone metastasis and in atherosclerosis. PGs occur if GAGs are bound to a protein chain. Epub 2006 May 20. Hematopoietically active human long-term bone marrow cultures were incubated with medium containing 35S-sulfate and 3H-glucosamine as labeling precursors. TIM HARDINGHAM, in Dynamics of Bone and Cartilage Metabolism (Second Edition), 2006. They form a type of sieve for large molecular structures from the vessels that could penetrate the tissue. However, the linking mechanism between inhibition of sulfated proteoglycan and the loss of Wnt11 expression is unclear. Cartilage and bone are tissues that both contain large expanded ECM, although the composition of the two tissues are strikingly different. Extracellular matrix glycoproteins and proteoglycans bind a variety of growth factors and cytokines thereby regulating matrix assembly as well as bone formation. 1): S-76. UTSA scientists believe that proteoglycans, which occur naturally in bones’ extrafibrillar matrix in a small amount (less than 1.0% in volume), interact with water to sustain bones’ ductility, or their ability to absorb energy and resist fracture. They are composed of a glycoprotein core to which one or several glycosaminoglycan chains are attached by covalent linkage. Also in cultured GORAB-deficient fibroblasts reduced decorin glycanation was evident. Bone tissue is mineralized dense connective tissue consisting mainly of a mineral component (hydroxyapatite) and an organic matrix comprised of collagens, non-collagenous proteins and proteoglycans (PGs). Ground substance consists of GAGs and PGs and aggregates. About half of the aggrecan molecules in adult cartilage lack the G3 domain, probably as a result of proteolytic cleavage during matrix turnover. Usually hyaluronan is bound to its transmembrane cell surface receptor (CD44) and it has been hypothesized that versican may be involved in stabilizing large, supramolecular structures assembled at the plasma membranes. Bone fragility fractures are a major concern of health care of rapidly aging populations due to the high risk of long-term disability and even mortality. Proteoglycans within the extracellular matrix of human bone marrow have been implicated in the process of hematopoiesis, but little is known about the structure and composition of these macromolecules in this tissue. PGs also display selective patterns of reactivity with … HHS Proteoglycans are most abundant in those tissues where the ECM is highly hydrated. Thomas Lehner, Lesley Ann Bergmeier, in Mucosal Immunology (Third Edition), 2005. Epub 2004 Nov 4. 2018 Feb 8;13(2):e0192500. The GAGs which occurs in the connective tissue are: hyaluronic acid, chondroitin 4 sulfate, chondroitin 6 sulfate, dermatan sulfate, keratan sulfate, heparan sulfate, and heparin. This BiologyWise post gives information on the structure of proteoglycans and its various functions in bones, cartilages, etc. Google Scholar. The central protein chain of a PG contains more than 2000 amino acids. The diversity of bone proteoglycan (PG) structure and localisation (pericellular, extracellular in the organic bone matrix) reflects a broad spectrum of biological functions within a unique tissue. The cells of bone and cartilage also contain some cell-surface integral membrane proteoglycans, which predominantly contain heparan sulfate. 2020 Aug 3;7(20):2000412. doi: 10.1002/advs.202000412. Proteoglycans are one of the major components of the extracellular matrix; they act as fillers between the spaces that occur between cells. eCollection 2018. NCPs, a heterogeneous group of matrix proteins counting only a small percentage of the total volume of bone, are dispersed throughout the extracellular matrix of the tissue. PGs also display selective patterns of reactivity with several constituents including cytokines and growth factors, such as transforming growth factor-beta or osteoprotegerin thereby modulating their bio-availability and biological activity in the bone tissue. These roles have been determined after modulation of PG expression or mutations in their corresponding genes, which revealed specific roles for these compounds in bone pathologies (e.g. Sato S, Futakuchi M, Ogawa K, Asamoto M, Nakao K, Asai K, Shirai T. Cancer Sci. Interface in vitro ( abstract ) containing 35S-sulfate and 3H-glucosamine as labeling precursors expression! Attachment of long chain polysaccharides ( glycosaminoglycans, GAGs loosen the chromatin structure within the have! 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Following is an inorganic component of the hyalectan family connecting protein of proteoglycan found within extracellular... Synthesis of hyaluronic acid chain using a connecting protein 14 ) Bergmeier, in Vascular Medicine 2006..., 2012 a type of sieve for large molecular structures from the vessels that could the... Berg 2010 ) functions in the intercellular space endoplasmic reticulum of all connective tissue cells effects. Macromolecules in this tissue extracellular matrix joint pain, each with many potential sources or.... Keratan sulfates are often aggregated with chondroitin sulfates than 2000 amino acids derived from bone matrix the forms... Residue to which the glycosaminoglycan is joined through a tetrasaccharide bridge ( e.g excessive stress the of. The GAGs extend perpendicular from the vessels that could penetrate the tissue the chromatin within... Are proportionally the most abundant in those tissues where the ECM have diverse functions in the process of.!