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1 0 1 0 1 0 1 ] /Size [ 64 ] >> stream H,GDQϽs~MMͻii$I#!DDDDDDh-E-EZ'D AQ/eY1UI[&m,EdI_HؤVmzlzM߆Cuvq\F=F<=G`D0q*ȜLL]d/rWx+)I!!ǐSD9b%h)f͏ϘWL;a'We"x endstream endobj 135 0 obj << /BitsPerSample 8 /Decode [ 0 1 0 1 0 1 0 1 ] /Domain [ 0 1 ] /Encode [ 0 63 ] /Filter /FlateDecode /FunctionType 0 /Length 18 /Order 1 /Range [ 0 1 0 1 0 1 0 1 ] /Size [ 64 ] >> stream H/`91@/B endstream endobj 136 0 obj << /AntiAlias false /ColorSpace 249 0 R /Coords [ 0 0 0 0 0 1 ] /Domain [ 0 1 ] /Extend [ true true ] /Function 137 0 R /ShadingType 3 >> endobj 137 0 obj << /Bounds [ ] /Domain [ 0 1 ] /Encode [ 0 1 ] /FunctionType 3 /Functions [ 138 0 R ] >> endobj 138 0 obj << /BitsPerSample 8 /Decode [ 0 1 0 1 0 1 0 1 ] /Domain [ 0 1 ] /Encode [ 0 63 ] /Filter /FlateDecode /FunctionType 0 /Length 222 /Order 1 /Range [ 0 1 0 1 0 1 0 1 ] /Size [ 64 ] >> stream H k+aߛܷlÜl6l!"D"B_sE~$ P ('d@IJ(M e!ʧ?-TUBP3/.BݒPhXq4:eѺhvw=y:Tc?UΔseBTkFUFȽ}0ؓl_W#f$ߍԇiL|o#cLٿ4> endobj 140 0 obj << /Bounds [ ] /Domain [ 0 1 ] 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These cells establish a barrier between sometimes hostile external environments and the internal milieu. However, mucosae are also responsible for nutrient absorption and waste secretion, which require a selectively permeable barrier. These functions place the mucosal epithelium at the centre of interactions between the mucosal immune system and luminal contents, including dietary antigens and microbial products. Recent advances have uncovered mechanisms by which the intestinal mucosal barrier is regulated in response to physiological and immunological stimuli. Here I discuss these discoveries along with evidence that this regulation shapes mucosal immune responses in the gut and, when dysfunctional, may contribute to disease.) >> endobj 290 0 obj << /D (nri_2653_nov09.indd:.15786) /S /GoTo >> endobj 291 0 obj << /A 292 0 R /Parent 288 0 R /Prev 293 0 R /Title (\376\377Conclusions) >> endobj 292 0 obj << /D (nri_2653_nov09.indd:.15853) /S /GoTo >> endobj 293 0 obj << /A 294 0 R /Next 291 0 R /Parent 288 0 R /Prev 295 0 R /Title (\376\377Tight junctions integrate mucosal homeostasis) >> endobj 294 0 obj << /D (nri_2653_nov09.indd:.15847) /S /GoTo >> endobj 295 0 obj << /A 296 0 R /Next 293 0 R /Parent 288 0 R /Prev 297 0 R /Title (\376\377Figure 3 | The epithelium and tight junction as integrators of mucosal homeostasis. Minor barrier defects allow bacterial products and dietary antigens to cross the epithelium and enter the lamina propria. This can lead to disease or homeostasis. If the foreign materials are taken up by antigen-presenting cells \(APCs\), such as dendritic cells, that direct the differentiation of T helper 1 \(Th1\) or Th2 cells, disease can develop. In this process, APCs and Th1 cells can release tumour necrosis factor \(TNF\) and interferon-\263 \(IFN\263\), which signal to epithelial cells to increase flux across the tight junction leak pathway, thereby allowing further leakage of bacterial products and dietary antigens from the lumen into the lamina propria and amplifying the cycle of inflammation. This may, ultimately, culminate in established disease. Alternatively, interleukin-13 \(IL 13\) released by Th2 cells increases flux across small cation-selective pores, potentially contributing to ongoing disease. Conversely, homeostasis may dominate if APCs promote regulatory T\240\(Treg\) cell differentiation, which can be enhanced by epithelial cell-derived transforming growth factor-\262 \(TGF\262\) and retinoic acid. The Treg cells display latency-associated peptide \(LAP\) on their surfaces and may secrete IL 10\240and TGF\262 to prevent disease. MLCK, myosin light chain kinase; TLR, Toll-like receptor; TCR, T cell receptor.) >> endobj 296 0 obj << /D (nri_2653_nov09.indd:.15841) /S /GoTo >> endobj 297 0 obj << /A 298 0 R /Next 295 0 R /Parent 288 0 R /Prev 299 0 R /Title (\376\377Barrier function and immunity) >> endobj 298 0 obj << /D (nri_2653_nov09.indd:.15835) /S /GoTo >> endobj 299 0 obj << /A 300 0 R /Next 297 0 R /Parent 288 0 R /Prev 301 0 R /Title (\376\377Table 1 | Barrier defects associated with intestinal disease) >> endobj 300 0 obj << /D (nri_2653_nov09.indd:.15829) /S /GoTo >> endobj 301 0 obj << /A 302 0 R /Next 299 0 R /Parent 288 0 R /Prev 303 0 R /Title (\376\377Figure 2 | Differential effects of cytokines on tight junction structure and function. a | Occludin is normally concentrated at the tight junction \(arrow\) in jejunal villus epithelium. The perijunctional actomyosin ring \(red\) and nuclei \(blue\) are shown for reference. b | Exogenous tumour necrosis factor increases myosin light chain kinase activity, which causes perijunctional myosin II regulatory light chain phosphorylation and triggers occludin \(green\) endocytosis \(arrow\). This increases flux across the tight junction leak pathway and enhances paracellular permeability to large solutes. c | Claudin 2 expression \(green\) is limited to crypt epithelial cells; it is not expressed by epithelial cells at the mucosal surface in the normal colon. F actin \(red\) and nuclei \(blue\) are shown for reference. d | Interleukin-13 can stimulate claudin 2 expression \(green\) in surface epithelial cells \(arrows\). This increases flux across small tight junction pores, thereby enhancing paracellular cation permeability.) >> endobj 302 0 obj << /D (nri_2653_nov09.indd:.15823) /S /GoTo >> endobj 303 0 obj << /A 304 0 R /Next 301 0 R /Parent 288 0 R /Prev 305 0 R /Title (\376\377Barrier regulation by immune stimuli ) >> endobj 304 0 obj << /D (nri_2653_nov09.indd:.15817) /S /GoTo >> endobj 305 0 obj << /A 306 0 R /Next 303 0 R /Parent 288 0 R /Prev 307 0 R /Title (\376\377Box 1 | Coordination of transcellular and paracellular transport) >> endobj 306 0 obj << /D (nri_2653_nov09.indd:.15810) /S /GoTo >> endobj 307 0 obj << /A 308 0 R /Next 305 0 R /Parent 288 0 R /Prev 309 0 R /Title (\376\377Interdependence of transport routes) >> endobj 308 0 obj << /D (nri_2653_nov09.indd:.15804) /S /GoTo >> endobj 309 0 obj << /A 310 0 R /Next 307 0 R /Parent 288 0 R /Prev 311 0 R /Title (\376\377Anatomy of mucosal barriers) >> endobj 310 0 obj << /D (nri_2653_nov09.indd:.15798) /S /GoTo >> endobj 311 0 obj << /A 312 0 R /Next 309 0 R /Parent 288 0 R /Prev 289 0 R /Title (\376\377Figure 1 | Anatomy of the mucosal barrier. a | The human intestinal mucosa is composed of a simple layer of columnar epithelial cells, as well as the underlying lamina propria and muscular mucosa. Goblet cells, which synthesize and release mucin, as well as other differentiated epithelial cell types, are present. The unstirred layer, which cannot be seen histologically, is located immediately above the epithelial cells. The tight junction, a component of the apical junctional complex, seals the paracellular space between epithelial cells. Intraepithelial lymphocytes are located above the basement membrane, but are subjacent to the tight junction. The lamina propria is located beneath the basement membrane and contains immune cells, including macrophages, dendritic cells, plasma cells, lamina propria lymphocytes and, in some cases, neutrophils. b | An electron micrograph and corresponding line drawing of the junctional complex of an intestinal epithelial cell. Just below the base of the microvilli, the plasma membranes of adjacent cells seem to fuse at the tight junction, where claudins, zonula occludens 1 \(ZO1\), occludin and F actin interact. E cadherin, \261-catenin 1, \262-catenin, catenin \2641 \(also known as p120 catenin; not shown\) and F actin interact to form the adherens junction. Myosin light chain kinase \(MLCK\) is associated with the perijunctional actomyosin ring. 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Turner Nature Reviews Immunology 9, 799 (2009). doi:10.1038/nri2653 Nature Publishing Group © 2009 Nature Publishing Group Intestinal mucosal barrier function in health and disease Adobe PDF Library 8.0 © 2009 Nature Publishing Group 10.1038/nri2653 1474-1741 809 1474-1733 11 Nature Reviews Immunology permissions@nature.com 799 9 2009-11-01 http://dx.doi.org/10.1038/nri2653 2009-10-15T10:11:38+05:30 Adobe InDesign CS3 (5.0.4) Nature Reviews Immunology 9, 799 (2009). doi:10.1038/nri2653 2009-10-20T18:12:38+05:30 2009-10-20T18:12:38+05:30 doi:10.1038/nri2653 True uuid:da15695e-9a4c-466b-8fc6-3f80ac5a28ff uuid:1cf49700-ea0f-42d6-8bf6-b42de54529b5 endstream endobj 325 0 obj << /Nums [ 0 315 0 R ] >> endobj 326 0 obj << /Dests 313 0 R /EmbeddedFiles 314 0 R >> endobj 327 0 obj << /CreationDate (D:20091015101138+05'30') /Subject (Nature Reviews Immunology 9, 799 \(2009\). doi:10.1038/nri2653) /Author (Jerrold R. 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