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Recently, TLRs have been directly linked to collagen synthesis or breakdown by
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TABLE 4-3 Types and Function of Select MMPs GROUP Collagenases ENZYME MMP-1 (Collagenase-1) MMP-8 (Collagenase-2) MMP-13 (Collagenase-3) MMP-2 (Gelatinase-A) ECM SUBSTRATE Collagen I, II, III, VII, X Collagen I, II, III Collagen I, II, III, IV, X Gelatin I Collagen IV, V, VII, X Fibronectin Elastin Gelatin I, V Collagen IV, V Fibronectin Elastin Proteoglycans Fibronectin Laminin Gelatin I, III, IV, V Fibronectin Gelatin I, III, IV, V Fibronectin Gelatin Laminin Collagen IV OTHER SELECT SUBSTRATES Pro-TNF, IL-1 , MMP-2, MMP-9 MMP-9 IL-1 , MMP-1, MMP-9, MMP-13
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MMP-1, MMP-8 IGF binding protein
mediating the expression of various metalloproteinases Matrix metalloproteinases (MMPs) are a group of enzymes responsible for the breakdown of collagen and can be classified into four subfamilies: (1) Collagenases, (2) gelatinases, (3) stromelysins, and (4) membrane-type MMPs (Table 4-3 for a summary of the functions of the first three types) Initial breakdown of collagen depends on members of the collagenase family that are capable of cleaving native triple helical collagen After the initial cleavage of collagen, the resultant fragments are further degraded by gelatinases and stromelysins43 The expression of MMPs is tightly regulated and regulation of the extracellular matrix involves a balance between synthesis of structural components and MMPs MMPs are expressed primarily by fibroblasts, but also by macrophages and keratinocytes and the expression of MMPs is modulated by cytokines For example, MMP-1 production from fibroblasts is stimulated by IL-1, IL-6, TNF- , and TGF- 44 46 Moreover, other cytokines such as IL-4 inhibit MMP expression and are chemoattractant for fibroblasts, favoring collagen and fibronectin synthesis and matrix preservation47 In addition to regulation at the transcriptional level, MMP activity is regulated by tissue inhibitors of metal-
loproteinase (TIMP) TIMPs, low molecular weight glycoproteins, are synthesized mainly in fibroblasts and macrophages,48 and inhibit MMP activity by forming heat-stable 1:1 stoichiometric complexes The expression of TIMPs is also regulated by cytokines and growth factors For example, TIMP-1 is induced by IL-1, IL-6, and EGF 45,49 Although both MMPs and TIMPs can be induced by similar stimuli, the expression can be regulated in both a coordinated and reciprocal manner The critical balance between MMP and TIMP expression determines the balance between matrix degradation and matrix preservation During periods of extracellular matrix homeostasis, the expression of MMP and TIMP is tightly coordinated providing for appropriate remodeling without excessive tissue breakdown However, if the amount of MMP expression is increased relative to TIMP expression, excessive matrix degradation is thought to occur The role of MMPs in photoaging has been well documented Both UVA and UVB radiation induce AP-1, a transcription factor important for the expression of MMP-1, 3, and 950 It is then hypothesized that these MMPs are involved in collagen breakdown, and subsequent imperfect repair yields molecular scar-
ring23 Cumulative UV exposure and the additive effect of molecular remodeling results in visible photoaging, characterized by wrinkles and decreased skin tone Histologically, photoaged skin reveals disorganized dermal collagen fibrils and increased elastin In addition to their ability to induce cytokines and chemokines, TLRs have been implicated in the induction of MMPs Several preliminary studies have shown that microbial agents are capable of inducing MMP expression through a TLR-dependent pathway For example, in Lyme disease the causative agent Borrelia burgdorferi is capable of inducing MMP-9 through a TLR2-dependent mechanism51 Moreover, mycobacterial cell wall components are also thought to increase MMP-9 through TLR252 More recently, CpG oligodeoxynucleotide, the ligand for TLR9, exhibited the capacity to induce MMP-9 expression in macrophages via a TLR9/NF- B-dependent signaling pathway53 Not all TLR pathways behave equally with regard to MMP regulation Imiquimod, a TLR7 and 8 ligand, downregulates production of MMP-9 while simultaneously upregulating TIMP expression54 Clinical evidence to support the role of 5% imiquimod cream in the reversal of photoaging and actinic damage was recently described by Kligman and colleagues The daily application of imiquimod cream for 5 days each week for 4 weeks resulted in a decrease in wrinkles, dyspigmentation, and hyperkeratotic pores Histologically, reversal of epidermal atypia and atrophy were observed in posttreatment biopsies55 In this regard, imiquimod appears to have potential as a novel therapy for reversal of photoaging as well as the prevention of cutaneous neoplasms The exact role imiquimod exerts in regulating the expression of MMP in vivo has not been determined and further studies are warranted in this area
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