The interplay between intrinsic and extrinsic Wnt signaling in controlling intestinal transformation

Albuquerque, 2002, The ‘just-right’ signaling model: APC somatic mutations are selected based on a specific level of activation of the beta-catenin signaling cascade, Hum. Mol. Genet., 10.1093/hmg/11.13.1549 Albuquerque, 2011, Colorectal cancers choosing sides, Biochim. Biophys. Acta - Rev. Cancer, 10.1016/j.bbcan.2011.07.005 Anastas, 2012, WNT signalling pathways as therapeutic targets in cancer, Nat. Rev. Cancer Bänziger, 2006, Wntless, a conserved membrane protein dedicated to the secretion of wnt proteins from signaling cells, Cell, 10.1016/j.cell.2006.02.049 Barker, 2007, Identification of stem cells in small intestine and colon by marker gene Lgr5, Nature, 449, 1003, 10.1038/nature06196 Barker, 2009, Crypt stem cells as the cells-of-origin of intestinal cancer, Nature, 457, 608, 10.1038/nature07602 Batts, 2006, Bmp signaling is required for intestinal growth and morphogenesis, Dev. Dynam., 10.1002/dvdy.20741 Behrens, 1998, Functional interaction of an axin homolog, conductin, with β-catenin, APC, and GSK3β, Science (80-. ), 10.1126/science.280.5363.596 Bovolenta, 2008, Beyond Wnt inhibition: new functions of secreted Frizzled-related proteins in development and disease, J. Cell Sci., 10.1242/jcs.026096 Castellone, 2005, Prostaglandin E2 promotes colon cancer cell growth through a Gs-axin-beta-catenin signaling axis, Science, 10.1126/science.1116221 Chulada, 2000, Genetic disruption of Ptgs-1, as well as of Ptgs-2, reduces intestinal tumorigenesis in Min mice, Cancer Res., 60, 4705 Crabtree, 2003, Refining the relation between ‘first hits’ and ‘second hits’ at the APC locus: the ‘loose fit’ model and evidence for differences in somatic mutation spectra among patients, Oncogene, 10.1038/sj.onc.1206471 Davis, 2015, Aberrant epithelial GREM1 expression initiates colonic tumorigenesis from cells outside the stem cell niche, Nat. Med., 21, 62, 10.1038/nm.3750 de Lau, 2012, The R-spondin protein family, Genome Biol., 10.1186/gb-2012-13-3-242 Degirmenci, 2018, GLI1-expressing mesenchymal cells form the essential Wnt-secreting niche for colon stem cells, Nature, 10.1038/s41586-018-0190-3 Delmas, 2011, Wif1 is a frequent target for epigenetic silencing in squamous cell carcinoma of the cervix, Carcinogenesis, 10.1093/carcin/bgr193 Dijksterhuis, 2014, WNT/Frizzled signalling: receptor-ligand selectivity with focus on FZD-G protein signalling and its physiological relevance: IUPHAR Review 3, Br. J. Pharmacol., 10.1111/bph.12364 Dow, 2015, Apc restoration promotes cellular differentiation and reestablishes crypt homeostasis in colorectal cancer, Cell, 161, 1539, 10.1016/j.cell.2015.05.033 Durand, 2012, Functional intestinal stem cells after Paneth cell ablation induced by the loss of transcription factor Math1 (Atoh1), Proc. Natl. Acad. Sci., 10.1073/pnas.1201652109 Farin, 2012, Redundant sources of Wnt regulate intestinal stem cells and promote formation of paneth cells, Gastroenterology, 10.1053/j.gastro.2012.08.031 Fearon, 1990, For colorectal tumorigenesis, Cell, 61, 759, 10.1016/0092-8674(90)90186-I Fevr, 2007, Wnt/beta-catenin is essential for intestinal homeostasis and maintenance of intestinal stem cells, Mol. Cell Biol., 10.1128/MCB.01034-07 Ficari, 2000, APC gene mutations and colorectal adenomatosis in familial adenomatous polyposis, Br. J. Cancer, 82, 348, 10.1054/bjoc.1999.0925 Flanagan, 2015, Frizzled7 functions as a Wnt receptor in intestinal epithelial Lgr5+stem cells, Stem Cell Rep., 10.1016/j.stemcr.2015.03.003 Fodde, 2002, The APC gene in colorectal cancer, Eur. J. Cancer, 10.1016/S0959-8049(02)00040-0 Fodde, 2001, APC, signal transduction and genetic instability in colorectal cancer, Nat. Rev. Cancer, 10.1038/35094067 Gregorieff, 2005, Expression pattern of Wnt signaling components in the adult intestine, Gastroenterology, 10.1016/j.gastro.2005.06.007 Greicius, 2018, PDGFRα + pericryptal stromal cells are the critical source of Wnts and RSPO3 for murine intestinal stem cells in vivo, Proc. Natl. Acad. Sci., 10.1073/pnas.1713510115 Gross, 2012, Active Wnt proteins are secreted on exosomes, Nat. Cell Biol., 10.1038/ncb2574 Grün, 2015, Single-cell messenger RNA sequencing reveals rare intestinal cell types, Nature, 10.1038/nature14966 Hao, 2012, ZNRF3 promotes Wnt receptor turnover in an R-spondin-sensitive manner, Nature, 10.1038/nature11019 Haramis, 2004, De novo crypt formation and juvenile polyposis on BMP inhibition in mouse intestine, Science, 303, 1684, 10.1126/science.1093587 Herring, 2018, Unsupervised trajectory analysis of single-cell RNA-seq and imaging data reveals alternative tuft cell origins in the gut, Cell Syst., 10.1016/j.cels.2017.10.012 Hilkens, 2017, RSPO3 expands intestinal stem cell and niche compartments and drives tumorigenesis, Gut, 10.1136/gutjnl-2016-311606 Horiguchi, 2017, ANGPTL2 expression in the intestinal stem cell niche controls epithelial regeneration and homeostasis, EMBO J., 10.15252/embj.201695690 Howe, 1998, Mutations in the SMAD4/DPC4 gene in juvenile polyposis, Science (80-. ), 10.1126/science.280.5366.1086 Howe, 2001, Germline mutations of the gene encoding bone morphogenetic protein receptor 1A in juvenile polyposis, Nat. Genet., 10.1038/88919 Huels, 2018, Wnt ligands influence tumour initiation by controlling the number of intestinal stem cells, Nat. Commun., 10.1038/s41467-018-03426-2 Jadhav, 2017, Dynamic reorganization of chromatin accessibility signatures during dedifferentiation of secretory precursors into Lgr5+ intestinal stem cells, Cell Stem Cell, 21, 65, 10.1016/j.stem.2017.05.001 Jaeger, 2012, Hereditary mixed polyposis syndrome is caused by a 40-kb upstream duplication that leads to increased and ectopic expression of the BMP antagonist GREM1, Nat. Genet., 44, 699, 10.1038/ng.2263 Junttila, 2013, Influence of tumour micro-environment heterogeneity on therapeutic response, Nature, 501, 346, 10.1038/nature12626 Kabiri, 2014 Kakugawa, 2015, Notum deacylates Wnt proteins to suppress signalling activity, Nature, 10.1038/nature14259 Kalluri, 2006, Fibroblasts in cancer, Nat. Rev. Cancer, 6, 392, 10.1038/nrc1877 Karagiannis, 2012, Cancer-associated fibroblasts drive the progression of metastasis through both paracrine and mechanical pressure on cancer tissue, Mol. Cancer Res., 10.1158/1541-7786.MCR-12-0307 Kim, 2005, Mitogenic influence of human R-spondin1 on the intestinal epithelium, Science (80-. ), 10.1126/science.1112521 Kim, 2006, R-spondin proteins: a novel link to β-catenin activation, Cell Cycle, 10.4161/cc.5.1.2305 Kim, 2008, R-Spondin family members regulate the Wnt pathway by a common mechanism, Mol. Biol. Cell, 10.1091/mbc.e08-02-0187 Knudsen, 2003, Attenuated familial adenomatous polyposis (AFAP). A review of the literature, Fam. Cancer, 10.1023/A:1023286520725 Kohler, 2008, Functional definition of the mutation cluster region of adenomatous polyposis coli in colorectal tumours, Hum. Mol. Genet., 10.1093/hmg/ddn095 Korkut, 2009, Trans-synaptic transmission of vesicular Wnt signals through Evi/Wntless, Cell, 10.1016/j.cell.2009.07.051 Kosinski, 2007, Gene expression patterns of human colon tops and basal crypts and BMP antagonists as intestinal stem cell niche factors, Proc. Natl. Acad. Sci. U. S. A., 104, 15418, 10.1073/pnas.0707210104 Krausova, 2014, Wnt signaling in adult intestinal stem cells and cancer, Cell. Signal., 10.1016/j.cellsig.2013.11.032 Kuhnert, 2004, Essential requirement for Wnt signaling in proliferation of adult small intestine and colon revealed by adenoviral expression of Dickkopf-1, Proc. Natl. Acad. Sci., 10.1073/pnas.2536800100 Kwong, 2009, APC and its modifiers in colon cancer, Adv. Exp. Med. Biol., 10.1007/978-1-4419-1145-2_8 Lamprecht, 2016, Small intestinal cancer: why the rarity?, Trends Cancer, 10.1016/j.trecan.2016.06.006 Leedham, 2013, A basal gradient of Wnt and stem-cell number influences regional tumour distribution in human and mouse intestinal tracts, Gut, 10.1136/gutjnl-2011-301601 Lenos, 2018, Stem cell functionality is microenvironmentally defined during tumour expansion and therapy response in colon cancer, Nat. Cell Biol., 20, 1, 10.1038/s41556-018-0179-z Lopez-Garcia, 2010, Intestinal stem cell replacement follows a pattern of neutral drift, Science, 330, 822, 10.1126/science.1196236 Lotti, 2013, Chemotherapy activates cancer-associated fibroblasts to maintain colorectal cancer-initiating cells by IL-17A, J. Exp. Med., 10.1084/jem.20131195 Madar, 2013, ‘Cancer associated fibroblasts’--more than meets the eye, Trends Mol. Med., 19, 447, 10.1016/j.molmed.2013.05.004 Maywald, 2015, IL-33 activates tumor stroma to promote intestinal polyposis, Proc. Natl. Acad. Sci., 10.1073/pnas.1422445112 Meran, 2017, Intestinal stem cell niche: the extracellular matrix and cellular components, Stem Cell. Int., 10.1155/2017/7970385 Merlos-Suárez, 2011, The intestinal stem cell signature identifies colorectal cancer stem cells and predicts disease relapse, Cell Stem Cell, 8, 511, 10.1016/j.stem.2011.02.020 Neumann, 2014, Gut commensal bacteria and regional Wnt gene expression in the proximal versus distal colon, Am. J. Pathol., 10.1016/j.ajpath.2013.11.029 Niehrs, 2006, Function and biological roles of the Dickkopf family of Wnt modulators, Oncogene, 10.1038/sj.onc.1210054 Nik, 2013, Foxf2 in intestinal fibroblasts reduces numbers of Lgr5+stem cells and adenoma formation by inhibiting wnt signaling, Gastroenterology, 10.1053/j.gastro.2013.01.045 Nile, 2018, A selective peptide inhibitor of Frizzled 7 receptors disrupts intestinal stem cells article, Nat. Chem. Biol. Nusse, 2017, Wnt/β-Catenin signaling, disease, and emerging therapeutic modalities, Cell, 10.1016/j.cell.2017.05.016 Ootani, 2009, Sustained in vitro intestinal epithelial culture within a Wnt-dependent stem cell niche, Nat. Med., 10.1038/nm.1951 Ormestad, 2006 Oshima, 1996, Suppression of intestinal polyposis in Apc delta716 knockout mice by inhibition of cyclooxygenase 2 (COX-2), Cell, 87, 803, 10.1016/S0092-8674(00)81988-1 Powell, 1992, APC mutations occur early during colorectal tumorigenesis, Nature, 10.1038/359235a0 Pugh, 1994, Patients with adenomatous polyps and carcinomas have increased colonic mucosal prostaglandin E2, Gut, 10.1136/gut.35.5.675 Qi, 2017, BMP restricts stemness of intestinal Lgr5 + stem cells by directly suppressing their signature genes, Nat. Commun. Reynolds, 2014, Canonical Wnt signals combined with suppressed TGFβ/BMP pathways promote renewal of the native human colonic epithelium, Gut, 10.1136/gutjnl-2012-304067 Rudin, 2012, Comprehensive genomic analysis identifies SOX2 as a frequently amplified gene in small-cell lung cancer, Nat. Genet., 44, 1111, 10.1038/ng.2405 San Roman, 2014, Wnt secretion from epithelial cells and subepithelial myofibroblasts is not required in the mouse intestinal stem cell niche in vivo, Stem Cell Rep., 10.1016/j.stemcr.2013.12.012 Sato, 2007, Frequent epigenetic inactivation of DICKKOPF family genes in human gastrointestinal tumors, Carcinogenesis, 10.1093/carcin/bgm178 Sato, 2011, Paneth cells constitute the niche for Lgr5 stem cells in intestinal crypts, Nature, 10.1038/nature09637 Segditsas, 2006, Colorectal cancer and genetic alterations in the Wnt pathway, Oncogene, 10.1038/sj.onc.1210059 Seshagiri, 2012, Recurrent R-spondin fusions in colon cancer, Nature, 10.1038/nature11282 Snippert, 2010, Intestinal crypt homeostasis results from neutral competition between symmetrically dividing Lgr5 stem cells, Cell, 143, 134, 10.1016/j.cell.2010.09.016 Snippert, 2014, Biased competition between Lgr5 intestinal stem cells driven by oncogenic mutation induces clonal expansion, EMBO Rep., 15, 62, 10.1002/embr.201337799 Sonoshita, 2001, Acceleration of intestinal polyposis through prostaglandin receptor EP2 in ApcΔ716knockout mice, Nat. Med., 10.1038/nm0901-1048 Stamos, 2013, The β-catenin destruction complex, Cold Spring Harb. Perspect. Biol., 10.1101/cshperspect.a007898 Stankiewicz, 2009, Genomic and genic deletions of the FOX gene cluster on 16q24.1 and inactivating mutations of FOXF1 cause alveolar capillary dysplasia and other malformations, Am. J. Hum. Genet., 10.1016/j.ajhg.2009.05.005 Stzepourginski, 2017, CD34 + mesenchymal cells are a major component of the intestinal stem cells niche at homeostasis and after injury, Proc. Natl. Acad. Sci., 10.1073/pnas.1620059114 Su, 1993, Association of the APC tumor suppressor protein with catenins, Science (80-. ), 10.1126/science.8259519 Szenker-Ravi, 2018, RSPO2 inhibition of RNF43 and ZNRF3 governs limb development independently of LGR4/5/6, Nature, 10.1038/s41586-018-0118-y Takada, 2006, Monounsaturated fatty acid modification of wnt protein: its role in wnt secretion, Dev. Cell, 10.1016/j.devcel.2006.10.003 Tao, 2017, Cancer associated fibroblasts: an essential role in the tumor microenvironment, Oncol. Lett., 10.3892/ol.2017.6497 Tata, 2013, Dedifferentiation of committed epithelial cells into stem cells in vivo, Nature, 503, 218, 10.1038/nature12777 Tetteh, 2016, Replacement of lost Lgr5-positive stem cells through plasticity of their enterocyte-lineage daughters, Cell Stem Cell, 10.1016/j.stem.2016.01.001 Valenta, 2016, Wnt ligands secreted by subepithelial mesenchymal cells are essential for the survival of intestinal stem cells and gut homeostasis, Cell Rep., 10.1016/j.celrep.2016.03.088 van Es, 2012, A critical role for the wnt effector Tcf4 in adult intestinal homeostatic self-renewal, Mol. Cell Biol., 10.1128/MCB.06288-11 van Neerven, 2016, Bidirectional interconversion of stem and non-stem cancer cell populations: a reassessment of theoretical models for tumor heterogeneity, Mol. Cell. Oncol., 3 Vermeulen, 2014, Stem cell dynamics in homeostasis and cancer of the intestine, Nat. Rev. Cancer, 14, 468, 10.1038/nrc3744 Vermeulen, 2013, Defining stem cell dynamics in models of intestinal tumor initiation, Science, 342, 995, 10.1126/science.1243148 Vincan, 2008, The upstream components of the Wnt signalling pathway in the dynamic EMT and MET associated with colorectal cancer progression, Clin. Exp. Metastasis, 10.1007/s10585-008-9156-4 Wang, 2003, Forkhead transcription factor Foxf2 (LUN)-deficient mice exhibit abnormal development of secondary palate, Dev. Biol., 10.1016/S0012-1606(03)00176-3 Wang, 2013, Structural basis for R-spondin recognition by LGR4/5/6 receptors, Genes Dev., 10.1101/gad.219360.113 Warner, 2015, Engineering high-potency R-spondin adult stem cell growth factors s, Mol. Pharmacol. Mol Pharmacol. Will, 2010, Location in the large bowel influences the APC mutations observed in FAP adenomas, Fam. Cancer, 10.1007/s10689-010-9332-y Willert, 2003, Wnt proteins are lipid-modified and can act as stem cell growth factors, Nature, 10.1038/nature01611 Yan, 2017, Non-equivalence of Wnt and R-spondin ligands during Lgr5 + intestinal stem-cell self-renewal, Nature, 10.1038/nature22313 Yan, 2017, Intestinal enteroendocrine lineage cells possess homeostatic and injury-inducible stem cell activity, Cell Stem Cell, 21, 78, 10.1016/j.stem.2017.06.014 Young, 2013, What are the best routes to effectively model human colorectal cancer?, Mol. Oncol., 10.1016/j.molonc.2013.02.006