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n3:pmcid: PMC0
bibo:doi: 10.1083%2Fjcb.200704152
n5:contains: _:vb2614398 _:vb2614376 _:vb2614365 _:vb2614342

Subject Item: _:vb2614342
rdf:type: n5:Section
dc:title: introduction
n5:contains: _:vb2614364 _:vb2614360 _:vb2614361 _:vb2614362 _:vb2614363 _:vb2614356 _:vb2614357 _:vb2614358 _:vb2614359 _:vb2614352 _:vb2614353 _:vb2614354 _:vb2614355 _:vb2614348 _:vb2614349 _:vb2614350 _:vb2614351 _:vb2614344 _:vb2614345 _:vb2614346 _:vb2614347 _:vb2614343

Subject Item: _:vb2614343
rdf:type: n3:Context
rdf:value: stem/progenitor cells found at the bottom third of Lieberkühn crypts throughout the length of the intestine and in the Paneth cells of the small intestine, as well as in human tumors of the intestinal epithelium (Blache et al., >>2004<<). Sox9 was first identified as a key regulator of cartilage and male gonad development.
n3:mentions: n2:15240568

Subject Item: _:vb2614344
rdf:type: n3:Context
rdf:value: for the campomelic dysplasia syndrome, a skeletal dysmorphology syndrome characterized by skeletal malformation of endochondral bones and by male-to-female sex reversal in the majority of genotypically XY individuals (Foster et al., >>1994<<; Wagner et al., 1994).
n3:mentions: n2:7990924

Subject Item: _:vb2614345
rdf:type: n3:Context
rdf:value: dysplasia syndrome, a skeletal dysmorphology syndrome characterized by skeletal malformation of endochondral bones and by male-to-female sex reversal in the majority of genotypically XY individuals (Foster et al., 1994; Wagner et al., >>1994<<). Sox9 has also been implicated in the development of cranial neural crest derivatives (Spokony et al., 2002), in the neural stem cell switch from neurogenesis to gliogenesis (Stolt et al., 2003) and in heart (Akiyama et al., 2004a), hair
n3:mentions: n2:8001137

Subject Item: _:vb2614346
rdf:type: n3:Context
rdf:value: Sox9 has also been implicated in the development of cranial neural crest derivatives (Spokony et al., >>2002<<), in the neural stem cell switch from neurogenesis to gliogenesis (Stolt et al., 2003) and in heart (Akiyama et al., 2004a), hair (Vidal et al., 2005), and pancreas (Seymour et al., 2007) development.
n3:mentions: n2:11807034

Subject Item: _:vb2614347
rdf:type: n3:Context
rdf:value: Sox9 has also been implicated in the development of cranial neural crest derivatives (Spokony et al., 2002), in the neural stem cell switch from neurogenesis to gliogenesis (Stolt et al., >>2003<<) and in heart (Akiyama et al., 2004a), hair (Vidal et al., 2005), and pancreas (Seymour et al., 2007) development.
n3:mentions: n2:12842915

Subject Item: _:vb2614348
rdf:type: n3:Context
rdf:value: Sox9 has also been implicated in the development of cranial neural crest derivatives (Spokony et al., 2002), in the neural stem cell switch from neurogenesis to gliogenesis (Stolt et al., 2003) and in heart (Akiyama et al., 2004a), hair (Vidal et al., 2005), and pancreas (Seymour et al., 2007) development.
n3:mentions: n2:15096597

Subject Item: _:vb2614349
rdf:type: n3:Context
rdf:value: implicated in the development of cranial neural crest derivatives (Spokony et al., 2002), in the neural stem cell switch from neurogenesis to gliogenesis (Stolt et al., 2003) and in heart (Akiyama et al., 2004a), hair (Vidal et al., >>2005<<), and pancreas (Seymour et al., 2007) development. In each of these tissues, Sox9 expression is restricted to specific cell types, suggesting a complex transcriptional regulation.
n3:mentions: n2:16085486

Subject Item: _:vb2614350
rdf:type: n3:Context
rdf:value: neural crest derivatives (Spokony et al., 2002), in the neural stem cell switch from neurogenesis to gliogenesis (Stolt et al., 2003) and in heart (Akiyama et al., 2004a), hair (Vidal et al., 2005), and pancreas (Seymour et al., >>2007<<) development. In each of these tissues, Sox9 expression is restricted to specific cell types, suggesting a complex transcriptional regulation.
n3:mentions: n2:17267606

Subject Item: _:vb2614351
rdf:type: n3:Context
rdf:value: In addition, the currently identified Sox9 target genes, for instance, in the cartilage and in the gonad, display tissue-specific expression (Ng et al., >>1997<<; de Santa Barbara et al., 1998), indicating that Sox9 may regulate distinct sets of genes in the different tissues in which it is expressed.
n3:mentions: n2:9119111

Subject Item: _:vb2614352
rdf:type: n3:Context
rdf:value: In addition, the currently identified Sox9 target genes, for instance, in the cartilage and in the gonad, display tissue-specific expression (Ng et al., 1997; de Santa Barbara et al., >>1998<<), indicating that Sox9 may regulate distinct sets of genes in the different tissues in which it is expressed.
n3:mentions: n2:9774680

Subject Item: _:vb2614353
rdf:type: n3:Context
rdf:value: In the intestinal epithelium, the function of Sox9 remains unresolved, although in vitro studies suggested a role in the control of cell differentiation (Blache et al., >>2004<<). In vitro and in vivo data indicate that Sox9 is a transcriptional target of Wnt signaling.
n3:mentions: n2:15240568

Subject Item: _:vb2614354
rdf:type: n3:Context
rdf:value: For instance, Sox9 expression is abrogated in Tcf4-null embryos, and it is strongly expressed in colorectal carcinoma cell lines containing activating mutations in components of the Wnt pathway (Blache et al., >>2004<<).
n3:mentions: n2:15240568

Subject Item: _:vb2614355
rdf:type: n3:Context
rdf:value: In particular, deletion of the gene encoding Tcf4, another HMG-box transcription factor (Korinek et al., >>1998<<), or overexpression of the inhibitor Dickkopf (Pinto et al., 2003; Kuhnert et al., 2004), resulted in a loss of the proliferative compartment and in impaired differentiation of secretory cell lineages.
n3:mentions: n2:9697701

Subject Item: _:vb2614356
rdf:type: n3:Context
rdf:value: In particular, deletion of the gene encoding Tcf4, another HMG-box transcription factor (Korinek et al., 1998), or overexpression of the inhibitor Dickkopf (Pinto et al., >>2003<<; Kuhnert et al., 2004), resulted in a loss of the proliferative compartment and in impaired differentiation of secretory cell lineages.
n3:mentions: n2:12865297

Subject Item: _:vb2614357
rdf:type: n3:Context
rdf:value: In particular, deletion of the gene encoding Tcf4, another HMG-box transcription factor (Korinek et al., 1998), or overexpression of the inhibitor Dickkopf (Pinto et al., 2003; Kuhnert et al., >>2004<<), resulted in a loss of the proliferative compartment and in impaired differentiation of secretory cell lineages.
n3:mentions: n2:14695885

Subject Item: _:vb2614358
rdf:type: n3:Context
rdf:value: Conversely, mutation of the gene encoding Apc, a negative regulator of the pathway, resulted in crypt expansion, abrogation of cell migration, and amplification of the Paneth cell population (Sansom et al., >>2004<<; Andreu et al., 2005).
n3:mentions: n2:15198980

Subject Item: _:vb2614359
rdf:type: n3:Context
rdf:value: Conversely, mutation of the gene encoding Apc, a negative regulator of the pathway, resulted in crypt expansion, abrogation of cell migration, and amplification of the Paneth cell population (Sansom et al., 2004; Andreu et al., >>2005<<). In addition, deletion of the Wnt receptor Frizzled-5 revealed an essential role of the Wnt–Frizzled-5 pathway in the maturation of Paneth cells (van Es et al., 2005).
n3:mentions: n2:15716339

Subject Item: _:vb2614360
rdf:type: n3:Context
rdf:value: In addition, deletion of the Wnt receptor Frizzled-5 revealed an essential role of the Wnt–Frizzled-5 pathway in the maturation of Paneth cells (van Es et al., >>2005<<). The sorting process of epithelial cells along the crypt–villus axis also depends on the Wnt pathway, via a modulation of Ephrin–Eph receptor interactions (Batlle et al., 2002). The Wnt signaling pathway can thus induce diverse cellular
n3:mentions: n2:15778706

Subject Item: _:vb2614361
rdf:type: n3:Context
rdf:value: The sorting process of epithelial cells along the crypt–villus axis also depends on the Wnt pathway, via a modulation of Ephrin–Eph receptor interactions (Batlle et al., >>2002<<). The Wnt signaling pathway can thus induce diverse cellular responses in the intestinal epithelium. In addition to these physiological functions, the Wnt pathway is centrally implicated in cancer, as mutations in components of this
n3:mentions: n2:12408869

Subject Item: _:vb2614362
rdf:type: n3:Context
rdf:value: In addition to these physiological functions, the Wnt pathway is centrally implicated in cancer, as mutations in components of this pathway have been identified in the majority of human colorectal carcinoma (Morin et al., >>1997<<). Such mutations mimic activation of the pathway by Wnt ligands (i.e., stabilization of β-catenin) and result in constitutive transcriptional activity of the β-catenin–Tcf4 complex and in aberrant expression of its target genes (Korinek
n3:mentions: n2:9065402

Subject Item: _:vb2614363
rdf:type: n3:Context
rdf:value: mimic activation of the pathway by Wnt ligands (i.e., stabilization of β-catenin) and result in constitutive transcriptional activity of the β-catenin–Tcf4 complex and in aberrant expression of its target genes (Korinek et al., >>1997<<). Despite the central importance of this pathway in the physiopathology of the intestinal epithelium, little is known about the molecular mechanisms involved in restricting this wide spectrum of potential functions to elicit a specific
n3:mentions: n2:9065401

Subject Item: _:vb2614364
rdf:type: n3:Context
rdf:value: The fact that Sox9 is transcriptionally regulated by the β-catenin–Tcf4 complex (Blache et al., >>2004<<), together with the particular expression of Sox9 in the compartment of the intestinal epithelium that contains Wnt-stimulated cells, suggests distinct functions in proliferating stem/progenitor cells and in the postmitotic Paneth cells
n3:mentions: n2:15240568

Subject Item: _:vb2614365
rdf:type: n5:Section
dc:title: materials and methods
n5:contains: _:vb2614366 _:vb2614367 _:vb2614368 _:vb2614369 _:vb2614370 _:vb2614371 _:vb2614372 _:vb2614373 _:vb2614374 _:vb2614375

Subject Item: _:vb2614366
rdf:type: n3:Context
rdf:value: The vil-Cre strain (el Marjou et al., >>2004<<) was in a nearly pure C57BL/6 background (at least 14 backcrosses to this background).
n3:mentions: n2:15282745

Subject Item: _:vb2614367
rdf:type: n3:Context
rdf:value: To generate Sox9+/flox mice (Kist et al., >>2002<<), exons 2 and 3 of the Sox9 gene were flanked by LoxP sequences.
n3:mentions: n2:11857796

Subject Item: _:vb2614368
rdf:type: n3:Context
rdf:value: The HT29.16E-Sox9 and HT29.16EΔCSox9 cell lines were described previously (Blache et al., >>2004<<; Jay et al., 2005).
n3:mentions: n2:15240568

Subject Item: _:vb2614369
rdf:type: n3:Context
rdf:value: The HT29.16E-Sox9 and HT29.16EΔCSox9 cell lines were described previously (Blache et al., 2004; Jay et al., >>2005<<).
n3:mentions: n2:15781631

Subject Item: _:vb2614370
rdf:type: n3:Context
rdf:value: Full-length human Sox9 and C-terminally truncated (dominant-negative) human Sox9 constructs were described previously (Südbeck et al., >>1996<<), as were the pTOP-FLASH and pFOP-FLASH reporter constructs (Morin et al., 1997).
n3:mentions: n2:8640233

Subject Item: _:vb2614371
rdf:type: n3:Context
rdf:value: Full-length human Sox9 and C-terminally truncated (dominant-negative) human Sox9 constructs were described previously (Südbeck et al., 1996), as were the pTOP-FLASH and pFOP-FLASH reporter constructs (Morin et al., >>1997<<). The Sox9-VP16 construct (Kamachi et al., 1999) was a gift from H. Kondoh (Institute for Molecular and Cell Biology, Osaka University, Osaka, Japan).
n3:mentions: n2:9065402

Subject Item: _:vb2614372
rdf:type: n3:Context
rdf:value: The Sox9-VP16 construct (Kamachi et al., >>1999<<) was a gift from H. Kondoh (Institute for Molecular and Cell Biology, Osaka University, Osaka, Japan).
n3:mentions: n2:9858536

Subject Item: _:vb2614373
rdf:type: n3:Context
rdf:value: Immunohistochemistry was performed essentially as described previously (Blache et al., >>2004<<). Sections of human intestinal metaplasia were provided by F. Bibeau (CRLC Val d'Aurelle Lamarque, Montpellier, France).
n3:mentions: n2:15240568

Subject Item: _:vb2614374
rdf:type: n3:Context
rdf:value: Protein lysates and immunoblotting were performed as described previously (Hollande et al., >>2003<<). Proteins were visualized using ECL Plus (GE Healthcare), and the bands were quantified by densitometry using ImageJ 1.32J (NIH).
n3:mentions: n2:12615962

Subject Item: _:vb2614375
rdf:type: n3:Context
rdf:value: Okano (Keio University, Tokyo, Japan), and the anti-Sox9 was described previously (de Santa Barbara et al., >>1998<<). β-Actin A5441 (Western blot; 1:5,000) was purchased from Sigma-Aldrich, anti-Ki-67 (1:200) and anti-lysozyme (1:1,000) were purchased from DakoCytomation, anti-PCNA (1:100) and anti–cyclin D1 (1:100) were purchased from Neomarker,
n3:mentions: n2:9774680

Subject Item: _:vb2614376
rdf:type: n5:Section
dc:title: results
n5:contains: _:vb2614380 _:vb2614381 _:vb2614382 _:vb2614383 _:vb2614377 _:vb2614378 _:vb2614379 _:vb2614388 _:vb2614389 _:vb2614390 _:vb2614391 _:vb2614384 _:vb2614385 _:vb2614386 _:vb2614387 _:vb2614396 _:vb2614397 _:vb2614392 _:vb2614393 _:vb2614394 _:vb2614395

Subject Item: _:vb2614377
rdf:type: n3:Context
rdf:value: the function of Sox9 in the turnover of the adult intestinal epithelium, Villin-Cre (vil-Cre) mice, in which the Cre recombinase is expressed specifically in the intestinal epithelium from 10.5 d postcoitum onward (el Marjou et al., >>2004<<) were crossed with Sox9flox/flox (Kist et al., 2002) mice, which have both Sox9 alleles flanked by loxP sequences.
n3:mentions: n2:15282745

Subject Item: _:vb2614378
rdf:type: n3:Context
rdf:value: intestinal epithelium, Villin-Cre (vil-Cre) mice, in which the Cre recombinase is expressed specifically in the intestinal epithelium from 10.5 d postcoitum onward (el Marjou et al., 2004) were crossed with Sox9flox/flox (Kist et al., >>2002<<) mice, which have both Sox9 alleles flanked by loxP sequences. This generated Sox9flox/flox-vil-Cre mice, with an intestinal epithelium lacking Sox9 protein, indicating effective vil-Cre–mediated recombination of the Sox9flox allele (Fig.
n3:mentions: n2:11857796

Subject Item: _:vb2614379
rdf:type: n3:Context
rdf:value: Among these, mucus-producing goblet cells represent the largest population and are responsible for epithelium protection and lubrication (Velcich et al., >>2002<<). Most of the other cells are enterocytes, with few interspersed enteroendocrine cells. Alcian blue and Muc2 stainings showed that the goblet cell population was strongly decreased in the colon of Sox9-deficient animals (Fig. 3, a–d). No
n3:mentions: n2:11872843

Subject Item: _:vb2614380
rdf:type: n3:Context
rdf:value: These cells are involved in regulating the interactions between epithelial cells and the indigenous microorganism population, which, in turn, is essential to elaborate the microvasculature underlying the epithelium (Wilson et al., >>1999<<; Stappenbeck et al., 2002).
n3:mentions: n2:10506557

Subject Item: _:vb2614381
rdf:type: n3:Context
rdf:value: in regulating the interactions between epithelial cells and the indigenous microorganism population, which, in turn, is essential to elaborate the microvasculature underlying the epithelium (Wilson et al., 1999; Stappenbeck et al., >>2002<<). Remarkably, morphological identification coupled with staining of small intestinal sections from Sox9flox/flox-vil-Cre mice for lysozyme, an early marker of Paneth cell differentiation, revealed that almost all the crypts were
n3:mentions: n2:12432102

Subject Item: _:vb2614382
rdf:type: n3:Context
rdf:value: Indeed, the number of cells positively stained with Musashi-1, a putative marker of stem cells in the nervous system and the intestinal epithelium (Sakakibara et al., >>1996<<; Potten et al., 2003), was increased in Sox9-deficient mice (Fig.
n3:mentions: n2:8660864

Subject Item: _:vb2614383
rdf:type: n3:Context
rdf:value: Indeed, the number of cells positively stained with Musashi-1, a putative marker of stem cells in the nervous system and the intestinal epithelium (Sakakibara et al., 1996; Potten et al., >>2003<<), was increased in Sox9-deficient mice (Fig.
n3:mentions: n2:12558601

Subject Item: _:vb2614384
rdf:type: n3:Context
rdf:value: In some pathological situations, such as intestinal metaplasia, ectopic Paneth cells can also be found in the esophagus (Barrett's esophagus) or in the stomach (Schreiber et al., >>1978<<). Thus, if Sox9 is required for the differentiation of the Paneth cell lineage, it should be expressed in Paneth cells found in such aberrant structures. To test this, we analyzed biopsy sections from a patient with Barrett's esophagus.
n3:mentions: n2:648823

Subject Item: _:vb2614385
rdf:type: n3:Context
rdf:value: (c and d) Analysis of the transcriptional activity of the β-catenin–Tcf4 complex, using a luciferase reporter system (Morin et al., >>1997<<). Endogenous transcriptional activity of the β-catenin–Tcf4 complex. Transient overexpression of Sox9 inhibits the β-catenin–Tcf4 activity, whereas overexpressing ΔCSox9 increases it (c). Similarily, induction of Sox9 overexpression in
n3:mentions: n2:9065402

Subject Item: _:vb2614386
rdf:type: n3:Context
rdf:value: We and others have shown that the level of Sox9 expression regulates the transcriptional activity of the β-catenin–Tcf4 complex in cultured HEK293 cells (Akiyama et al., 2004b; Blache et al., 2004). Physical interaction between Sox9 and β-catenin has been reported, resulting in a competition between Sox9 and Tcf4 for binding to β-catenin.
n3:mentions: n2:15132997

Subject Item: _:vb2614387
rdf:type: n3:Context
rdf:value: We and others have shown that the level of Sox9 expression regulates the transcriptional activity of the β-catenin–Tcf4 complex in cultured HEK293 cells (Akiyama et al., 2004b; Blache et al., >>2004<<). Physical interaction between Sox9 and β-catenin has been reported, resulting in a competition between Sox9 and Tcf4 for binding to β-catenin.
n3:mentions: n2:15240568

Subject Item: _:vb2614388
rdf:type: n3:Context
rdf:value: Formation of the Sox9–β-catenin complex results in degradation of the two proteins (Akiyama et al., 2004b). We thus hypothesized that the absence of Sox9 in crypt cells of Sox9flox/flox-vil-Cre mice, where Wnt signaling is physiologically active, might result in increased availability of the nuclear pool of β-catenin for binding to Tcf4. To
n3:mentions: n2:15132997

Subject Item: _:vb2614389
rdf:type: n3:Context
rdf:value: analyzed the transcriptional activity of the β-catenin–Tcf4 complex after manipulation of Sox9 expression in colon carcinoma cell lines containing a constitutive activation of the β-catenin–Tcf transcriptional complex (Korinek et al., >>1997<<). The basal β-catenin–Tcf4 activity present in DLD-1 cells was efficiently inhibited by Sox9 and increased after overexpression of a dominant-negative version of Sox9 (Fig.
n3:mentions: n2:9065401

Subject Item: _:vb2614390
rdf:type: n3:Context
rdf:value: expression of key β-catenin–Tcf target genes by Sox9, but the molecular mechanism underlying this regulation remained unclear, as the ΔC-Sox9 construct does not contain the domain thought to interact with β-catenin (Akiyama et al., 2004b).
n3:mentions: n2:15132997

Subject Item: _:vb2614391
rdf:type: n3:Context
rdf:value: The previously reported physical interaction between Sox9 and β-catenin had been detected after overexpression of tagged Sox9 and β-catenin in COS cells (Akiyama et al., 2004b). We reasoned that if this interaction was important to modulate Wnt signaling in colon carcinoma cells, then the endogenous complex should be readily detectable in these cells.
n3:mentions: n2:15132997

Subject Item: _:vb2614392
rdf:type: n3:Context
rdf:value: 8 a) was originally identified in a campomelic dysplasia patient and abolishes the DNA binding properties of Sox9 (Meyer et al., >>1997<<). We introduced this mutation in both the Sox9 and ΔCSox9 constructs. The resulting products did not have any transcriptional activity using Sox-luciferase reporters (Fig. 8 b) and failed to modify β-catenin–Tcf activity (Fig. 8 c). This
n3:mentions: n2:9002675

Subject Item: _:vb2614393
rdf:type: n3:Context
rdf:value: To test this, we used a Sox9 construct in which the C-terminal domain of Sox9, involved in transactivation, and in the interaction with β-catenin (Akiyama et al., 2004b), is removed and replaced by the unrelated VP16 transactivating domain (Kamachi et al., 1999; Fig.
n3:mentions: n2:15132997

Subject Item: _:vb2614394
rdf:type: n3:Context
rdf:value: a Sox9 construct in which the C-terminal domain of Sox9, involved in transactivation, and in the interaction with β-catenin (Akiyama et al., 2004b), is removed and replaced by the unrelated VP16 transactivating domain (Kamachi et al., >>1999<<; Fig. 8 a). Transient transfection of this construct in colon carcinoma cells showed that the Sox9-VP16 chimeric protein potently activates transcription of a Sox-luciferase reporter gene (Fig.
n3:mentions: n2:9858536

Subject Item: _:vb2614395
rdf:type: n3:Context
rdf:value: We then aimed to identify potential Wnt pathway inhibitors, such as the inhibitor of β-catenin and Tcf (ICAT; Tago et al., >>2000<<) and Groucho-related (Grg/TLE) corepressors (Cavallo et al., 1998; Roose et al., 1998), which might be transcriptionally regulated by Sox9.
n3:mentions: n2:10898789

Subject Item: _:vb2614396
rdf:type: n3:Context
rdf:value: We then aimed to identify potential Wnt pathway inhibitors, such as the inhibitor of β-catenin and Tcf (ICAT; Tago et al., 2000) and Groucho-related (Grg/TLE) corepressors (Cavallo et al., >>1998<<; Roose et al., 1998), which might be transcriptionally regulated by Sox9. In vitro, the induction of Sox9 expression in HT29Cl.
n3:mentions: n2:9783586

Subject Item: _:vb2614397
rdf:type: n3:Context
rdf:value: We then aimed to identify potential Wnt pathway inhibitors, such as the inhibitor of β-catenin and Tcf (ICAT; Tago et al., 2000) and Groucho-related (Grg/TLE) corepressors (Cavallo et al., 1998; Roose et al., >>1998<<), which might be transcriptionally regulated by Sox9. In vitro, the induction of Sox9 expression in HT29Cl.
n3:mentions: n2:9783587

Subject Item: _:vb2614398
rdf:type: n5:Section
dc:title: discussion
n5:contains: _:vb2614399 _:vb2614428 _:vb2614424 _:vb2614425 _:vb2614426 _:vb2614427 _:vb2614420 _:vb2614421 _:vb2614422 _:vb2614423 _:vb2614416 _:vb2614417 _:vb2614418 _:vb2614419 _:vb2614412 _:vb2614413 _:vb2614414 _:vb2614415 _:vb2614408 _:vb2614409 _:vb2614410 _:vb2614411 _:vb2614404 _:vb2614405 _:vb2614406 _:vb2614407 _:vb2614400 _:vb2614401 _:vb2614402 _:vb2614403

Subject Item: _:vb2614399
rdf:type: n3:Context
rdf:value: Interference with the Wnt pathway results in depletion of the Paneth cell lineage (Pinto et al., >>2003<<) and, conversely, loss of the key negative regulator of the pathway, Apc, results in an expansion of the Paneth cell lineage (Sansom et al., 2004; Andreu et al., 2005).
n3:mentions: n2:12865297

Subject Item: _:vb2614400
rdf:type: n3:Context
rdf:value: with the Wnt pathway results in depletion of the Paneth cell lineage (Pinto et al., 2003) and, conversely, loss of the key negative regulator of the pathway, Apc, results in an expansion of the Paneth cell lineage (Sansom et al., >>2004<<; Andreu et al., 2005). In addition, the terminal maturation of Paneth cells requires Wnt signaling through the Frizzled-5 receptor (van Es et al., 2005).
n3:mentions: n2:15198980

Subject Item: _:vb2614401
rdf:type: n3:Context
rdf:value: results in depletion of the Paneth cell lineage (Pinto et al., 2003) and, conversely, loss of the key negative regulator of the pathway, Apc, results in an expansion of the Paneth cell lineage (Sansom et al., 2004; Andreu et al., >>2005<<). In addition, the terminal maturation of Paneth cells requires Wnt signaling through the Frizzled-5 receptor (van Es et al., 2005).
n3:mentions: n2:15716339

Subject Item: _:vb2614402
rdf:type: n3:Context
rdf:value: In addition, the terminal maturation of Paneth cells requires Wnt signaling through the Frizzled-5 receptor (van Es et al., >>2005<<). As we previously showed that Sox9 is a transcriptional target of Wnt signaling, the Sox9 requirement for the differentiation of the Paneth cell lineage is likely to reflect its role to specify this specific aspect of the Wnt pathway. In
n3:mentions: n2:15778706

Subject Item: _:vb2614403
rdf:type: n3:Context
rdf:value: an earlier stage than the Wnt–Frizzled-5 pathway, during the course of Paneth cell differentiation, as this pathway was previously shown to be involved in the terminal differentiation of already committed Paneth cells (van Es et al., >>2005<<). Although less likely, an alternative in which Sox9 and Wnt signaling, as two distinct pathways, would control Paneth cell differentiation cannot be formally excluded.
n3:mentions: n2:15778706

Subject Item: _:vb2614404
rdf:type: n3:Context
rdf:value: In addition, no major alteration of microorganism population along the crypt–villus axis was detected (Garabedian et al., >>1997<<). On the other hand, a role of Paneth cells in regulating the villus angiogenesis (Stappenbeck et al., 2002) and in clearing bacterial infections (Wilson et al., 1999) have been described, although no major impact was reported at the
n3:mentions: n2:9295317

Subject Item: _:vb2614405
rdf:type: n3:Context
rdf:value: On the other hand, a role of Paneth cells in regulating the villus angiogenesis (Stappenbeck et al., >>2002<<) and in clearing bacterial infections (Wilson et al., 1999) have been described, although no major impact was reported at the physiopathological level.
n3:mentions: n2:12432102

Subject Item: _:vb2614406
rdf:type: n3:Context
rdf:value: On the other hand, a role of Paneth cells in regulating the villus angiogenesis (Stappenbeck et al., 2002) and in clearing bacterial infections (Wilson et al., >>1999<<) have been described, although no major impact was reported at the physiopathological level.
n3:mentions: n2:10506557

Subject Item: _:vb2614407
rdf:type: n3:Context
rdf:value: relevant given the key role of mucus in protecting the epithelium against the luminal content, well illustrated, for instance, by the colon epithelium tumorigenesis resulting from a deletion of the Muc2 gene (Velcich et al., >>2002<<). This function of Sox9 in regulating the differentiation of goblet cells might also play a part in the cellular response to Wnt signals, as altered goblet cell differentiation was also found in mice in which the Wnt pathway has been
n3:mentions: n2:11872843

Subject Item: _:vb2614408
rdf:type: n3:Context
rdf:value: in regulating the differentiation of goblet cells might also play a part in the cellular response to Wnt signals, as altered goblet cell differentiation was also found in mice in which the Wnt pathway has been impaired (Pinto et al., >>2003<<). That Sox9 deficiency results in the complete absence of Paneth cells but leads only to a reduction of the goblet cell lineage might indicate different Sox9 requirements for the differentiation of these two lineages, and suggests that
n3:mentions: n2:12865297

Subject Item: _:vb2614409
rdf:type: n3:Context
rdf:value: In a previous report, we showed that overexpression of Sox9 in the LS174T colon carcinoma cell line repressed the Muc2 gene (Blache et al., >>2004<<). This apparently contrasts with the present finding that the absence of Sox9 in the intestinal epithelium causes an obvious reduction of the number of goblet cells in vivo.
n3:mentions: n2:15240568

Subject Item: _:vb2614410
rdf:type: n3:Context
rdf:value: a role in the differentiation of the goblet cell lineage, and we hypothesize that this role occurs primarily at the level of cell fate choice, similar to what was shown previously in the developing nervous system (Spokony et al., >>2002<<). In addition, as suggested by in vitro data and the respective expression patterns of Sox9 and Muc2, down-regulation of Sox9 expression might be necessary to allow terminal maturation of committed goblet cells, although this hypothesis
n3:mentions: n2:11807034

Subject Item: _:vb2614411
rdf:type: n3:Context
rdf:value: Although previous studies indicated that Sox9 could inhibit the activity of the β-catenin–Tcf4 complex in HEK293 cells transiently transfected with artificial luciferase reporters (Akiyama et al., 2004b; Blache et al., 2004), the observed regulation of the β-catenin–Tcf4 complex transcriptional activity on endogenous target gene promoters, in cultured colon carcinoma cells known to have a constitutive β-catenin–Tcf4 activity, was
n3:mentions: n2:15132997

Subject Item: _:vb2614412
rdf:type: n3:Context
rdf:value: Although previous studies indicated that Sox9 could inhibit the activity of the β-catenin–Tcf4 complex in HEK293 cells transiently transfected with artificial luciferase reporters (Akiyama et al., 2004b; Blache et al., >>2004<<), the observed regulation of the β-catenin–Tcf4 complex transcriptional activity on endogenous target gene promoters, in cultured colon carcinoma cells known to have a constitutive β-catenin–Tcf4 activity, was somewhat unexpected.
n3:mentions: n2:15240568

Subject Item: _:vb2614413
rdf:type: n3:Context
rdf:value: In agreement with this, a down-regulation of the β-catenin–Tcf4 complex has been reported as cultured colon carcinoma cells reach confluence (Mariadason et al., >>2001<<). Thus, in low- confluence culture conditions, the endogenous level of β-catenin–Tcf4 complex likely saturates the endogenous target gene promoters, even in the presence of exogenous Sox9. In contrast, the transfected luciferase Tcf
n3:mentions: n2:11309309

Subject Item: _:vb2614414
rdf:type: n3:Context
rdf:value: This also explains the apparent discrepancy between the results presented here and in our previous report, in which we showed that c-Myc was not regulated by Sox9 in low-confluence cell culture conditions (Blache et al., >>2004<<). Thus, even in cells in which the Wnt pathway is constitutively active, Sox9 may play an important retroinhibition role.
n3:mentions: n2:15240568

Subject Item: _:vb2614415
rdf:type: n3:Context
rdf:value: cells obtained in this study with the Sox9flox/flox control mice (46–56% with a 2-h BrdU pulse) is consistent with previous analyses showing a 3H labeling index of 29–33% after a 1-h treatment with 3H-thymidine (Cheng and Bjerknes, >>1982<<). The 16–32% increase in BrdU-positive cells in Sox9flox/flox-vil-Cre mice is therefore likely to significantly impact epithelial homeostasis, given the rate of renewal of the intestinal epithelium.
n3:mentions: n2:7114498

Subject Item: _:vb2614416
rdf:type: n3:Context
rdf:value: A role of Sox9 in regulating cell proliferation has already been proposed in chondrocytic cell lines, in which Sox9 promoted differentiation (Panda et al., >>2001<<), as well as in breast and prostate tumor cell lines (Afonja et al., 2002; Drivdahl et al., 2004).
n3:mentions: n2:11514554

Subject Item: _:vb2614417
rdf:type: n3:Context
rdf:value: A role of Sox9 in regulating cell proliferation has already been proposed in chondrocytic cell lines, in which Sox9 promoted differentiation (Panda et al., 2001), as well as in breast and prostate tumor cell lines (Afonja et al., >>2002<<; Drivdahl et al., 2004).
n3:mentions: n2:12420222

Subject Item: _:vb2614418
rdf:type: n3:Context
rdf:value: regulating cell proliferation has already been proposed in chondrocytic cell lines, in which Sox9 promoted differentiation (Panda et al., 2001), as well as in breast and prostate tumor cell lines (Afonja et al., 2002; Drivdahl et al., >>2004<<). Here, however, Sox9 is specifically expressed, in a normal physiological situation, in the proliferative compartment of the intestinal epithelium, and activity of the protooncogenic Wnt pathway is required for its expression (Blache et
n3:mentions: n2:15077158

Subject Item: _:vb2614419
rdf:type: n3:Context
rdf:value: however, Sox9 is specifically expressed, in a normal physiological situation, in the proliferative compartment of the intestinal epithelium, and activity of the protooncogenic Wnt pathway is required for its expression (Blache et al., >>2004<<). Thus, Sox9 might function as a gatekeeper in intestinal crypts to prevent overactivity of the Wnt-dependent transcriptional program.
n3:mentions: n2:15240568

Subject Item: _:vb2614420
rdf:type: n3:Context
rdf:value: Wnt signaling is known to be important for the maintenance of the proliferating compartment (Korinek et al., >>1998<<; Pinto et al., 2003; Kuhnert et al., 2004), as well as for the differentiation and maturation of Paneth cells (Pinto et al., 2003; Andreu et al., 2005; van Es et al., 2005).
n3:mentions: n2:9697701

Subject Item: _:vb2614421
rdf:type: n3:Context
rdf:value: Wnt signaling is known to be important for the maintenance of the proliferating compartment (Korinek et al., 1998; Pinto et al., >>2003<<; Kuhnert et al., 2004), as well as for the differentiation and maturation of Paneth cells (Pinto et al., 2003; Andreu et al., 2005; van Es et al., 2005).
n3:mentions: n2:12865297

Subject Item: _:vb2614422
rdf:type: n3:Context
rdf:value: Wnt signaling is known to be important for the maintenance of the proliferating compartment (Korinek et al., 1998; Pinto et al., 2003; Kuhnert et al., >>2004<<), as well as for the differentiation and maturation of Paneth cells (Pinto et al., 2003; Andreu et al., 2005; van Es et al., 2005).
n3:mentions: n2:14695885

Subject Item: _:vb2614423
rdf:type: n3:Context
rdf:value: signaling is known to be important for the maintenance of the proliferating compartment (Korinek et al., 1998; Pinto et al., 2003; Kuhnert et al., 2004), as well as for the differentiation and maturation of Paneth cells (Pinto et al., >>2003<<; Andreu et al., 2005; van Es et al., 2005).
n3:mentions: n2:12865297

Subject Item: _:vb2614424
rdf:type: n3:Context
rdf:value: be important for the maintenance of the proliferating compartment (Korinek et al., 1998; Pinto et al., 2003; Kuhnert et al., 2004), as well as for the differentiation and maturation of Paneth cells (Pinto et al., 2003; Andreu et al., >>2005<<; van Es et al., 2005).
n3:mentions: n2:15716339

Subject Item: _:vb2614425
rdf:type: n3:Context
rdf:value: maintenance of the proliferating compartment (Korinek et al., 1998; Pinto et al., 2003; Kuhnert et al., 2004), as well as for the differentiation and maturation of Paneth cells (Pinto et al., 2003; Andreu et al., 2005; van Es et al., >>2005<<). Here, we found that c-Myc, a central effector of the Wnt pathway, as well as cyclin D1, was expressed in the proliferative compartment of the small intestinal epithelium but not, or at a much lower level, in the Paneth cell compartment
n3:mentions: n2:15778706

Subject Item: _:vb2614426
rdf:type: n3:Context
rdf:value: This was unexpected given that Paneth cells have been reported to express the highest levels of nuclear β-catenin (van Es et al., >>2005<<). This suggests that distinct genetic programs are set, downstream to the Wnt pathway, in the stem/progenitor versus Paneth cell compartments, and this was reflected by a recent transcriptomic study (Van der Flier et al., 2007). The
n3:mentions: n2:15778706

Subject Item: _:vb2614427
rdf:type: n3:Context
rdf:value: This suggests that distinct genetic programs are set, downstream to the Wnt pathway, in the stem/progenitor versus Paneth cell compartments, and this was reflected by a recent transcriptomic study (Van der Flier et al., >>2007<<). The absence of Sox9 results in a single proliferative stem/progenitor compartment, in which all the cells express c-Myc and cyclin D1. This suggests that Sox9 is required to specify the Paneth cell compartment, in which the presence of
n3:mentions: n2:17320548

Subject Item: _:vb2614428
rdf:type: n3:Context
rdf:value: to control colon epithelium morphology, a role that had not been previously reported for the Wnt pathway, although architectural degeneration was described after Dickkopf-1 overexpression in the colon epithelium (Kuhnert et al., >>2004<<).
n3:mentions: n2:14695885

Subject Item: _:vb358712453
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 41
n3:hasRelevantPaperId: n2:17681175

Subject Item: _:vb358712454
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 32
n3:hasRelevantPaperId: n2:17934449

Subject Item: _:vb358712455
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 28
n3:hasRelevantPaperId: n2:15240568

Subject Item: _:vb358712456
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 21
n3:hasRelevantPaperId: n2:21113151

Subject Item: _:vb358712457
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 20
n3:hasRelevantPaperId: n2:15778706

Subject Item: _:vb358712458
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 18
n3:hasRelevantPaperId: n2:19329995

Subject Item: _:vb358712459
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 17
n3:hasRelevantPaperId: n2:19019766

Subject Item: _:vb358712460
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 17
n3:hasRelevantPaperId: n2:12865297

Subject Item: _:vb358712461
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 15
n3:hasRelevantPaperId: n2:11739954

Subject Item: _:vb358712462
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 15
n3:hasRelevantPaperId: n2:12408869

Subject Item: _:vb358712463
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 14
n3:hasRelevantPaperId: n2:23870119

Subject Item: _:vb358712464
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 14
n3:hasRelevantPaperId: n2:22246670

Subject Item: _:vb358712465
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 14
n3:hasRelevantPaperId: n2:15959515

Subject Item: _:vb358712466
rdf:type: n3:RelevantBibliographicResource
n3:RelevantScore: 13
n3:hasRelevantPaperId: n2:15282745

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