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n3:pmcid: PMC0
bibo:doi: 10.1002%2Femmm.200900039
n7:contains: _:vb6757854 _:vb6757838 _:vb6757887 _:vb6757811

Subject Item: _:vb6757811
rdf:type: n7:Section
dc:title: introduction
n7:contains: _:vb6757816 _:vb6757817 _:vb6757818 _:vb6757819 _:vb6757820 _:vb6757821 _:vb6757822 _:vb6757823 _:vb6757812 _:vb6757813 _:vb6757814 _:vb6757815 _:vb6757832 _:vb6757833 _:vb6757834 _:vb6757835 _:vb6757836 _:vb6757837 _:vb6757824 _:vb6757825 _:vb6757826 _:vb6757827 _:vb6757828 _:vb6757829 _:vb6757830 _:vb6757831

Subject Item: _:vb6757812
rdf:type: n3:Context
rdf:value: Recurrence and metastatic spread have been proposed to depend on cancer stem cells, which express CD133 (AC133; Ieta et al, >>2008<<; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, 2009).
n3:mentions: n2:17932721

Subject Item: _:vb6757813
rdf:type: n3:Context
rdf:value: Recurrence and metastatic spread have been proposed to depend on cancer stem cells, which express CD133 (AC133; Ieta et al, 2008; O'Brien et al, >>2007<<; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, 2009).
n3:mentions: n2:17122772

Subject Item: _:vb6757814
rdf:type: n3:Context
rdf:value: Recurrence and metastatic spread have been proposed to depend on cancer stem cells, which express CD133 (AC133; Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, >>2007<<; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, 2009).
n3:mentions: n2:17122771

Subject Item: _:vb6757815
rdf:type: n3:Context
rdf:value: Recurrence and metastatic spread have been proposed to depend on cancer stem cells, which express CD133 (AC133; Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, >>2007<<; Vermeulen et al, 2008; Zhu et al, 2009).
n3:mentions: n2:18371377

Subject Item: _:vb6757816
rdf:type: n3:Context
rdf:value: Recurrence and metastatic spread have been proposed to depend on cancer stem cells, which express CD133 (AC133; Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, >>2008<<; Zhu et al, 2009).
n3:mentions: n2:18765800

Subject Item: _:vb6757817
rdf:type: n3:Context
rdf:value: Recurrence and metastatic spread have been proposed to depend on cancer stem cells, which express CD133 (AC133; Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, >>2009<<).
n3:mentions: n2:19092805

Subject Item: _:vb6757818
rdf:type: n3:Context
rdf:value: Whereas CCs are thought to derive from constitutive activation of WNT signalling by mutation of APC of βCAT (e.g. Morin et al, >>1997<<), the involvement of HH-GLI signalling in CC is not clear, even though it is critical for many types of human cancers (reviewed in Ruiz i Altaba et al, 2007).
n3:mentions: n2:9065402

Subject Item: _:vb6757819
rdf:type: n3:Context
rdf:value: activation of WNT signalling by mutation of APC of βCAT (e.g. Morin et al, 1997), the involvement of HH-GLI signalling in CC is not clear, even though it is critical for many types of human cancers (reviewed in Ruiz i Altaba et al, >>2007<<). Secreted HH ligands block the function of PATCHED1 (PTCH1) in the responding cell, which normally inhibits SMOOTHENED (SMOH).
n3:mentions: n2:17845852

Subject Item: _:vb6757820
rdf:type: n3:Context
rdf:value: Hh has been suggested to regulate the differentiation of normal intestinal villi (van den Brink et al, >>2004<<), and in mice it affects mostly the surrounding mesenchyme (Alinger et al, 2009; Madison et al, 2005).
n3:mentions: n2:14770182

Subject Item: _:vb6757821
rdf:type: n3:Context
rdf:value: Hh has been suggested to regulate the differentiation of normal intestinal villi (van den Brink et al, 2004), and in mice it affects mostly the surrounding mesenchyme (Alinger et al, >>2009<<; Madison et al, 2005).
n3:mentions: n2:19280222

Subject Item: _:vb6757822
rdf:type: n3:Context
rdf:value: Hh has been suggested to regulate the differentiation of normal intestinal villi (van den Brink et al, 2004), and in mice it affects mostly the surrounding mesenchyme (Alinger et al, 2009; Madison et al, >>2005<<). Moreover, while limited in vitro cell line and expression data suggest a controversial role in epithelial cells (Akiyoshi et al, 2006; Douard et al, 2006; Monzo et al, 2006; Oniscu et al, 2004; Qualtrough et al, 2004), this has been
n3:mentions: n2:15590741

Subject Item: _:vb6757823
rdf:type: n3:Context
rdf:value: Moreover, while limited in vitro cell line and expression data suggest a controversial role in epithelial cells (Akiyoshi et al, >>2006<<; Douard et al, 2006; Monzo et al, 2006; Oniscu et al, 2004; Qualtrough et al, 2004), this has been either not observed (Berman et al, 2003) or denied (Chatel et al, 2007; van den Brink et al, 2004).
n3:mentions: n2:16299030

Subject Item: _:vb6757824
rdf:type: n3:Context
rdf:value: Moreover, while limited in vitro cell line and expression data suggest a controversial role in epithelial cells (Akiyoshi et al, 2006; Douard et al, >>2006<<; Monzo et al, 2006; Oniscu et al, 2004; Qualtrough et al, 2004), this has been either not observed (Berman et al, 2003) or denied (Chatel et al, 2007; van den Brink et al, 2004).
n3:mentions: n2:16701100

Subject Item: _:vb6757825
rdf:type: n3:Context
rdf:value: Moreover, while limited in vitro cell line and expression data suggest a controversial role in epithelial cells (Akiyoshi et al, 2006; Douard et al, 2006; Monzo et al, 2006; Oniscu et al, >>2004<<; Qualtrough et al, 2004), this has been either not observed (Berman et al, 2003) or denied (Chatel et al, 2007; van den Brink et al, 2004).
n3:mentions: n2:15258993

Subject Item: _:vb6757826
rdf:type: n3:Context
rdf:value: cell line and expression data suggest a controversial role in epithelial cells (Akiyoshi et al, 2006; Douard et al, 2006; Monzo et al, 2006; Oniscu et al, 2004; Qualtrough et al, 2004), this has been either not observed (Berman et al, >>2003<<) or denied (Chatel et al, 2007; van den Brink et al, 2004). Moreover, recent studies indicate that in mice Hh may act in an exclusive paracrine fashion in pancreatic and gastrointestinal (GI) tumours (Tian et al, 2009; Yauch et al, 2008).
n3:mentions: n2:14520411

Subject Item: _:vb6757827
rdf:type: n3:Context
rdf:value: in epithelial cells (Akiyoshi et al, 2006; Douard et al, 2006; Monzo et al, 2006; Oniscu et al, 2004; Qualtrough et al, 2004), this has been either not observed (Berman et al, 2003) or denied (Chatel et al, 2007; van den Brink et al, >>2004<<). Moreover, recent studies indicate that in mice Hh may act in an exclusive paracrine fashion in pancreatic and gastrointestinal (GI) tumours (Tian et al, 2009; Yauch et al, 2008).
n3:mentions: n2:14770182

Subject Item: _:vb6757828
rdf:type: n3:Context
rdf:value: Moreover, recent studies indicate that in mice Hh may act in an exclusive paracrine fashion in pancreatic and gastrointestinal (GI) tumours (Tian et al, >>2009<<; Yauch et al, 2008).
n3:mentions: n2:19246386

Subject Item: _:vb6757829
rdf:type: n3:Context
rdf:value: Moreover, recent studies indicate that in mice Hh may act in an exclusive paracrine fashion in pancreatic and gastrointestinal (GI) tumours (Tian et al, 2009; Yauch et al, >>2008<<). This idea has been extended to human CCs without direct data (Yauch et al, 2008; discussed in Ruiz i Altaba, 2008) and it contradicts the presence of active signalling in the tumour cells of human melanomas, prostate and basal cell
n3:mentions: n2:18754008

Subject Item: _:vb6757830
rdf:type: n3:Context
rdf:value: This idea has been extended to human CCs without direct data (Yauch et al, >>2008<<; discussed in Ruiz i Altaba, 2008) and it contradicts the presence of active signalling in the tumour cells of human melanomas, prostate and basal cell carcinomas and other human tumours (Dahmane et al, 1997; Sanchez et al, 2004; Stecca
n3:mentions: n2:18754008

Subject Item: _:vb6757831
rdf:type: n3:Context
rdf:value: direct data (Yauch et al, 2008; discussed in Ruiz i Altaba, 2008) and it contradicts the presence of active signalling in the tumour cells of human melanomas, prostate and basal cell carcinomas and other human tumours (Dahmane et al, >>1997<<; Sanchez et al, 2004; Stecca et al, 2007; see Ruiz i Altaba, 2008).
n3:mentions: n2:9349822

Subject Item: _:vb6757832
rdf:type: n3:Context
rdf:value: al, 2008; discussed in Ruiz i Altaba, 2008) and it contradicts the presence of active signalling in the tumour cells of human melanomas, prostate and basal cell carcinomas and other human tumours (Dahmane et al, 1997; Sanchez et al, >>2004<<; Stecca et al, 2007; see Ruiz i Altaba, 2008).
n3:mentions: n2:15314219

Subject Item: _:vb6757833
rdf:type: n3:Context
rdf:value: in Ruiz i Altaba, 2008) and it contradicts the presence of active signalling in the tumour cells of human melanomas, prostate and basal cell carcinomas and other human tumours (Dahmane et al, 1997; Sanchez et al, 2004; Stecca et al, >>2007<<; see Ruiz i Altaba, 2008).
n3:mentions: n2:17392427

Subject Item: _:vb6757834
rdf:type: n3:Context
rdf:value: A role for HH-GLI in human CC has also not been highlighted by genomic approaches (Wood et al, >>2007<<)—although this was also the case of HH-GLI-dependent glioblastomas (Parsons et al, 2008; see also Clement et al, 2007)—and the few mutations in SMOH found in GI tumours are not activating (Guleng et al, 2006).
n3:mentions: n2:17932254

Subject Item: _:vb6757835
rdf:type: n3:Context
rdf:value: A role for HH-GLI in human CC has also not been highlighted by genomic approaches (Wood et al, 2007)—although this was also the case of HH-GLI-dependent glioblastomas (Parsons et al, >>2008<<; see also Clement et al, 2007)—and the few mutations in SMOH found in GI tumours are not activating (Guleng et al, 2006).
n3:mentions: n2:18772396

Subject Item: _:vb6757836
rdf:type: n3:Context
rdf:value: A role for HH-GLI in human CC has also not been highlighted by genomic approaches (Wood et al, 2007)—although this was also the case of HH-GLI-dependent glioblastomas (Parsons et al, 2008; see also Clement et al, >>2007<<)—and the few mutations in SMOH found in GI tumours are not activating (Guleng et al, 2006).
n3:mentions: n2:17196391

Subject Item: _:vb6757837
rdf:type: n3:Context
rdf:value: approaches (Wood et al, 2007)—although this was also the case of HH-GLI-dependent glioblastomas (Parsons et al, 2008; see also Clement et al, 2007)—and the few mutations in SMOH found in GI tumours are not activating (Guleng et al, >>2006<<). Some of the potential problems with the studies mentioned above in terms of resolving an action of HH-GLI in human CCs include the use of only one or few established human cell lines and the lack of single-cell resolution data on the
n3:mentions: n2:17287906

Subject Item: _:vb6757838
rdf:type: n7:Section
dc:title: materials and methods
n7:contains: _:vb6757852 _:vb6757853 _:vb6757848 _:vb6757849 _:vb6757850 _:vb6757851 _:vb6757844 _:vb6757845 _:vb6757846 _:vb6757847 _:vb6757840 _:vb6757841 _:vb6757842 _:vb6757843 _:vb6757839

Subject Item: _:vb6757839
rdf:type: n3:Context
rdf:value: In situ hybridizations with frozen sections were performed with digoxygenin-labelled antisense RNA probes for GLI1 and PTCH1 as described previously (Dahmane et al, >>2001<<; Stecca et al, 2007).
n3:mentions: n2:11748155

Subject Item: _:vb6757840
rdf:type: n3:Context
rdf:value: In situ hybridizations with frozen sections were performed with digoxygenin-labelled antisense RNA probes for GLI1 and PTCH1 as described previously (Dahmane et al, 2001; Stecca et al, >>2007<<). Immunohistotochemistry of frozen sections used mouse anti-CEA (DAKO) mouse anti-Ki67 (Cell Signalling), mouse anti-SHH (University of Iowa Hybridoma Bank), mouse anti-PanCytokeratin (Sigma), anti-GLI1 monoclonal and polyclonal (Cell
n3:mentions: n2:17392427

Subject Item: _:vb6757841
rdf:type: n3:Context
rdf:value: were pretreated by bleaching with hydrogen peroxide and boiling for detection of βCAT with horseradish peroxidase-coupled secondary antibodies and development with diaminobenzidine as described previously (van de Wetering et al, >>2002<<).
n3:mentions: n2:12408868

Subject Item: _:vb6757842
rdf:type: n3:Context
rdf:value: Full-length GLI1, GLI3 and GLI3R plasmids were described previously (Ruiz i Altaba, >>1999<<). 106 of the total or CD133+ selected cell populations of HT29, Ls174T, CC14 and mCC11 were mixed with 5 µg of total DNA and transfected with a Nucleofector (Amaxa) according to the manufacturer's instructions.
n3:mentions: n2:10375510

Subject Item: _:vb6757843
rdf:type: n3:Context
rdf:value: The sequences for the siRNAs were 5′–3′: First set (Clement et al, >>2007<<; Sanchez et al, 2004; Stecca et al, 2007) – GLI1, AACUCCACAGGCAUACAGGAU; GLI2, AAGAUCUGGACAGGGAUGACU; GLI3, AAUGAGGAUGAAAGUCCUGGA.
n3:mentions: n2:17196391

Subject Item: _:vb6757844
rdf:type: n3:Context
rdf:value: The sequences for the siRNAs were 5′–3′: First set (Clement et al, 2007; Sanchez et al, >>2004<<; Stecca et al, 2007) – GLI1, AACUCCACAGGCAUACAGGAU; GLI2, AAGAUCUGGACAGGGAUGACU; GLI3, AAUGAGGAUGAAAGUCCUGGA.
n3:mentions: n2:15314219

Subject Item: _:vb6757845
rdf:type: n3:Context
rdf:value: The sequences for the siRNAs were 5′–3′: First set (Clement et al, 2007; Sanchez et al, 2004; Stecca et al, >>2007<<) – GLI1, AACUCCACAGGCAUACAGGAU; GLI2, AAGAUCUGGACAGGGAUGACU; GLI3, AAUGAGGAUGAAAGUCCUGGA.
n3:mentions: n2:17392427

Subject Item: _:vb6757846
rdf:type: n3:Context
rdf:value: LV-shSMOH was as in Clement et al (>>2007<<). LV-shPTCH1 was made in the same LV-CTH vector as LV-shSMOH with the target sequence 5′-GCACTATGCTCCTTTCCTC-3′.
n3:mentions: n2:17196391

Subject Item: _:vb6757847
rdf:type: n3:Context
rdf:value: CD133+ CC cells (O'Brien et al, >>2007<<; Ricci-Vitiani et al, 2007) were sorted from human resections, mouse xenografts of cultured cells using magnetic cell sorting (MACS) as described by the manufacturer (MiltenyiBiotech).
n3:mentions: n2:17122772

Subject Item: _:vb6757848
rdf:type: n3:Context
rdf:value: CD133+ CC cells (O'Brien et al, 2007; Ricci-Vitiani et al, >>2007<<) were sorted from human resections, mouse xenografts of cultured cells using magnetic cell sorting (MACS) as described by the manufacturer (MiltenyiBiotech).
n3:mentions: n2:17122771

Subject Item: _:vb6757849
rdf:type: n3:Context
rdf:value: For metastatic growth (Stecca et al, >>2007<<), 1 × 106 HT29-LacZ and CC14-LacZ infected with different lentivectors were injected into the tail vein (i.
n3:mentions: n2:17392427

Subject Item: _:vb6757850
rdf:type: n3:Context
rdf:value: Primers were as described in Dahmane et al (>>2001<<), Sanchez et al (2004), Stecca et al (2007) and Clement et al (2007).
n3:mentions: n2:11748155

Subject Item: _:vb6757851
rdf:type: n3:Context
rdf:value: Primers were as described in Dahmane et al (2001), Sanchez et al (>>2004<<), Stecca et al (2007) and Clement et al (2007).
n3:mentions: n2:15314219

Subject Item: _:vb6757852
rdf:type: n3:Context
rdf:value: Primers were as described in Dahmane et al (2001), Sanchez et al (2004), Stecca et al (>>2007<<) and Clement et al (2007).
n3:mentions: n2:17392427

Subject Item: _:vb6757853
rdf:type: n3:Context
rdf:value: Primers were as described in Dahmane et al (2001), Sanchez et al (2004), Stecca et al (2007) and Clement et al (>>2007<<). Additional primer sets were 5′ to 3′: KLF4-Forward – AGAAGGATCTCGGCCAATTT; and KLF4-R, GTGGAGAAAGATGGGAGCAG; VIMENTIN-F, TGCCCTTAAAGGAACCAATG and VIMENTIN-R: TCCAGCAGCTTCCTGTAGGT; IHH -F: CACCCCCAATTACAATCCAG and IHH-R:
n3:mentions: n2:17196391

Subject Item: _:vb6757854
rdf:type: n7:Section
dc:title: results
n7:contains: _:vb6757855 _:vb6757884 _:vb6757885 _:vb6757886 _:vb6757880 _:vb6757881 _:vb6757882 _:vb6757883 _:vb6757876 _:vb6757877 _:vb6757878 _:vb6757879 _:vb6757872 _:vb6757873 _:vb6757874 _:vb6757875 _:vb6757868 _:vb6757869 _:vb6757870 _:vb6757871 _:vb6757864 _:vb6757865 _:vb6757866 _:vb6757867 _:vb6757860 _:vb6757861 _:vb6757862 _:vb6757863 _:vb6757856 _:vb6757857 _:vb6757858 _:vb6757859

Subject Item: _:vb6757855
rdf:type: n3:Context
rdf:value: HT29 (APC/BRAF/p53 mutant) and Caco2 (APC/p53 mutant) (Table S1 of Supporting Information), showed consistent expression of GLI1, PTCH1 and SHH as well as other HH-GLI pathway components, extending previous findings (Bian et al, >>2007<<; Monzo et al, 2006; Oniscu et al, 2004).
n3:mentions: n2:17461467

Subject Item: _:vb6757856
rdf:type: n3:Context
rdf:value: (APC/p53 mutant) (Table S1 of Supporting Information), showed consistent expression of GLI1, PTCH1 and SHH as well as other HH-GLI pathway components, extending previous findings (Bian et al, 2007; Monzo et al, 2006; Oniscu et al, >>2004<<). Expression was detected in epithelial tumour cells of CCs in situ in the colon (Fig 1A) and of metastatic CCs in the liver (Fig 1C), that were positive for carcinoembryonic antigen (CEA), cytokeratins and βCAT (Fig S1 and Table S1 of
n3:mentions: n2:15258993

Subject Item: _:vb6757857
rdf:type: n3:Context
rdf:value: GLI1 is undetectable by immunocytochemisty in normal development given its very low levels, detectable levels of GLI1 protein in transfected cells or in cancer cells can be found in the nucleus and/or cytoplasm (e.g. Dahmane et al, >>1997<<; Ruiz i Altaba, 1999; Stecca et al, 2007). Here we found GLI1 labelling mostly in the cytoplasm (Fig 1B,D).
n3:mentions: n2:9349822

Subject Item: _:vb6757858
rdf:type: n3:Context
rdf:value: by immunocytochemisty in normal development given its very low levels, detectable levels of GLI1 protein in transfected cells or in cancer cells can be found in the nucleus and/or cytoplasm (e.g. Dahmane et al, 1997; Ruiz i Altaba, >>1999<<; Stecca et al, 2007). Here we found GLI1 labelling mostly in the cytoplasm (Fig 1B,D).
n3:mentions: n2:10375510

Subject Item: _:vb6757859
rdf:type: n3:Context
rdf:value: in normal development given its very low levels, detectable levels of GLI1 protein in transfected cells or in cancer cells can be found in the nucleus and/or cytoplasm (e.g. Dahmane et al, 1997; Ruiz i Altaba, 1999; Stecca et al, >>2007<<). Here we found GLI1 labelling mostly in the cytoplasm (Fig 1B,D).
n3:mentions: n2:17392427

Subject Item: _:vb6757860
rdf:type: n3:Context
rdf:value: To measure gene expression levels in different CC cell populations and since CC stem cells express CD133 (AC133; O'Brien et al, >>2007<<; Ricci-Vitiani et al, 2007), these were selected from fresh non-metastatic (TNM stages 1,2) and metastatic (TNM3,4) CCs as well as liver metastases (Fig 2A and Fig S2 of Supporting Information).
n3:mentions: n2:17122772

Subject Item: _:vb6757861
rdf:type: n3:Context
rdf:value: To measure gene expression levels in different CC cell populations and since CC stem cells express CD133 (AC133; O'Brien et al, 2007; Ricci-Vitiani et al, >>2007<<), these were selected from fresh non-metastatic (TNM stages 1,2) and metastatic (TNM3,4) CCs as well as liver metastases (Fig 2A and Fig S2 of Supporting Information).
n3:mentions: n2:17122771

Subject Item: _:vb6757862
rdf:type: n3:Context
rdf:value: An HH-GLI activity signature (GLI1, PTCH1, GLI2, SHH and HIP) was detected in all CCs (Fig 2A,B and Fig S2 of Supporting Information), consistent with increased SHH expression (Fig 2B; Douard et al, >>2006<<; Oniscu et al, 2004). This signature was not enriched in CD133+ cells of normal colon or liver and it was only slightly enriched in non-metastatic tumours (TNM1,2) (Fig 2A,B).
n3:mentions: n2:16701100

Subject Item: _:vb6757863
rdf:type: n3:Context
rdf:value: An HH-GLI activity signature (GLI1, PTCH1, GLI2, SHH and HIP) was detected in all CCs (Fig 2A,B and Fig S2 of Supporting Information), consistent with increased SHH expression (Fig 2B; Douard et al, 2006; Oniscu et al, >>2004<<). This signature was not enriched in CD133+ cells of normal colon or liver and it was only slightly enriched in non-metastatic tumours (TNM1,2) (Fig 2A,B).
n3:mentions: n2:15258993

Subject Item: _:vb6757864
rdf:type: n3:Context
rdf:value: GLI1 itself and SNAIL1, also a GLI target (Li et al, >>2006<<) involved in epithelial-to-mesenchymal transition (EMT; Batlle et al, 2000; Cano et al, 2000), similarly showed an increased CD133+/CD133− average in liver metastases versus TNM3,4 CCs (6 in liver metastases vs.
n3:mentions: n2:16158046

Subject Item: _:vb6757865
rdf:type: n3:Context
rdf:value: GLI1 itself and SNAIL1, also a GLI target (Li et al, 2006) involved in epithelial-to-mesenchymal transition (EMT; Batlle et al, >>2000<<; Cano et al, 2000), similarly showed an increased CD133+/CD133− average in liver metastases versus TNM3,4 CCs (6 in liver metastases vs.
n3:mentions: n2:10655587

Subject Item: _:vb6757866
rdf:type: n3:Context
rdf:value: GLI1 itself and SNAIL1, also a GLI target (Li et al, 2006) involved in epithelial-to-mesenchymal transition (EMT; Batlle et al, 2000; Cano et al, >>2000<<), similarly showed an increased CD133+/CD133− average in liver metastases versus TNM3,4 CCs (6 in liver metastases vs.
n3:mentions: n2:10655586

Subject Item: _:vb6757867
rdf:type: n3:Context
rdf:value: First, to knock-down HH-GLI signalling at the level of GLI function we used siRNAs specific for GLI1, GLI2 or GLI3 with ±80–85% targeting efficiencies (Clement et al, >>2007<<; Sanchez et al, 2004; Stecca et al, 2007) (Fig 3B and Fig S3 of Supporting Information).
n3:mentions: n2:17196391

Subject Item: _:vb6757868
rdf:type: n3:Context
rdf:value: First, to knock-down HH-GLI signalling at the level of GLI function we used siRNAs specific for GLI1, GLI2 or GLI3 with ±80–85% targeting efficiencies (Clement et al, 2007; Sanchez et al, >>2004<<; Stecca et al, 2007) (Fig 3B and Fig S3 of Supporting Information).
n3:mentions: n2:15314219

Subject Item: _:vb6757869
rdf:type: n3:Context
rdf:value: First, to knock-down HH-GLI signalling at the level of GLI function we used siRNAs specific for GLI1, GLI2 or GLI3 with ±80–85% targeting efficiencies (Clement et al, 2007; Sanchez et al, 2004; Stecca et al, >>2007<<) (Fig 3B and Fig S3 of Supporting Information).
n3:mentions: n2:17392427

Subject Item: _:vb6757870
rdf:type: n3:Context
rdf:value: An independent second set of siRNAs (Clement et al, >>2007<<; Sanchez et al, 2004; Stecca et al, 2007) gave the same results (Fig 3B and Fig S3 of Supporting Information).
n3:mentions: n2:17196391

Subject Item: _:vb6757871
rdf:type: n3:Context
rdf:value: An independent second set of siRNAs (Clement et al, 2007; Sanchez et al, >>2004<<; Stecca et al, 2007) gave the same results (Fig 3B and Fig S3 of Supporting Information).
n3:mentions: n2:15314219

Subject Item: _:vb6757872
rdf:type: n3:Context
rdf:value: An independent second set of siRNAs (Clement et al, 2007; Sanchez et al, 2004; Stecca et al, >>2007<<) gave the same results (Fig 3B and Fig S3 of Supporting Information).
n3:mentions: n2:17392427

Subject Item: _:vb6757873
rdf:type: n3:Context
rdf:value: Expression of an shRNA to SMOOTHENED (shSMOH; Clement et al, >>2007<<) with 85–90% kd targeting efficiency induced a 3.6-fold decrease in BrdU incorporation (p < 0.001) 4–5 days after transduction; and a 26-fold increase in cleaved Caspase-3 labelling (p < 0.001) (Fig 3C).
n3:mentions: n2:17196391

Subject Item: _:vb6757874
rdf:type: n3:Context
rdf:value: As an additional test for the requirement of HH-GLI function, we also inhibited SMOH pharmacologically by treatment with cyclopamine, a plant alkaloid (Keeler, >>1970<<, 1978) that inhibits SMOH (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba,
n3:mentions: n2:4986632

Subject Item: _:vb6757875
rdf:type: n3:Context
rdf:value: As an additional test for the requirement of HH-GLI function, we also inhibited SMOH pharmacologically by treatment with cyclopamine, a plant alkaloid (Keeler, 1970, >>1978<<) that inhibits SMOH (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, 2005;
n3:mentions: n2:723484

Subject Item: _:vb6757876
rdf:type: n3:Context
rdf:value: As an additional test for the requirement of HH-GLI function, we also inhibited SMOH pharmacologically by treatment with cyclopamine, a plant alkaloid (Keeler, 1970, 1978) that inhibits SMOH (Cooper et al, >>1998<<; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, 2005; Stecca et al, 2007).
n3:mentions: n2:9616123

Subject Item: _:vb6757877
rdf:type: n3:Context
rdf:value: As an additional test for the requirement of HH-GLI function, we also inhibited SMOH pharmacologically by treatment with cyclopamine, a plant alkaloid (Keeler, 1970, 1978) that inhibits SMOH (Cooper et al, 1998; Incardona et al, >>1998<<) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, 2005; Stecca et al, 2007).
n3:mentions: n2:9716521

Subject Item: _:vb6757878
rdf:type: n3:Context
rdf:value: by treatment with cyclopamine, a plant alkaloid (Keeler, 1970, 1978) that inhibits SMOH (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, >>2007<<; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, 2005; Stecca et al, 2007).
n3:mentions: n2:17196391

Subject Item: _:vb6757879
rdf:type: n3:Context
rdf:value: with cyclopamine, a plant alkaloid (Keeler, 1970, 1978) that inhibits SMOH (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, >>2001<<; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, 2005; Stecca et al, 2007).
n3:mentions: n2:11748155

Subject Item: _:vb6757880
rdf:type: n3:Context
rdf:value: alkaloid (Keeler, 1970, 1978) that inhibits SMOH (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, >>2004<<; Sanchez & Ruiz i Altaba, 2005; Stecca et al, 2007).
n3:mentions: n2:15314219

Subject Item: _:vb6757881
rdf:type: n3:Context
rdf:value: that inhibits SMOH (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, >>2005<<; Stecca et al, 2007). Cyclopamine treatment produced 1.9-fold (p < 0.01) and 3.5-fold (p < 0.01) less BrdU incorporation on average at 5 and 10 µM as compared with the control compound tomatidine (10 µM) (Fig 3D).
n3:mentions: n2:15652709

Subject Item: _:vb6757882
rdf:type: n3:Context
rdf:value: (Cooper et al, 1998; Incardona et al, 1998) and that we found to be a specific inhibitor of HH-GLI at effective doses (e.g. Clement et al, 2007; Dahmane et al, 2001; Sanchez et al, 2004; Sanchez & Ruiz i Altaba, 2005; Stecca et al, >>2007<<). Cyclopamine treatment produced 1.9-fold (p < 0.01) and 3.5-fold (p < 0.01) less BrdU incorporation on average at 5 and 10 µM as compared with the control compound tomatidine (10 µM) (Fig 3D).
n3:mentions: n2:17392427

Subject Item: _:vb6757883
rdf:type: n3:Context
rdf:value: These can then be easily visualized by XGal staining yielding blue metastatic cells (Stecca et al, >>2007<<).
n3:mentions: n2:17392427

Subject Item: _:vb6757884
rdf:type: n3:Context
rdf:value: The increase of metastatic growth of cells with high HH-GLI activity could be due to a growth advantage and/or to an induction of EMT (Li et al, >>2006<<). To test the possibility that GLI1 directly drives EMT advanced CC cells, GLI1 was expressed at higher levels in CC14 cells, which derive from an advanced tumour in the colon (Table S1 of Supporting Information).
n3:mentions: n2:16158046

Subject Item: _:vb6757885
rdf:type: n3:Context
rdf:value: on whole tumour growth in vitro and in xenografts additionally indicates an action of HH-GLI on the tumour bulk, possibly paralleling the requirement of HH-GLI in both neural stem cells and derived precursors (Palma & Ruiz i Altaba, >>2004<<; Stecca & Ruiz i Altaba, 2009).
n3:mentions: n2:14681189

Subject Item: _:vb6757886
rdf:type: n3:Context
rdf:value: and in xenografts additionally indicates an action of HH-GLI on the tumour bulk, possibly paralleling the requirement of HH-GLI in both neural stem cells and derived precursors (Palma & Ruiz i Altaba, 2004; Stecca & Ruiz i Altaba, >>2009<<).
n3:mentions: n2:19214186

Subject Item: _:vb6757887
rdf:type: n7:Section
dc:title: discussion
n7:contains: _:vb6757904 _:vb6757905 _:vb6757906 _:vb6757907 _:vb6757908 _:vb6757909 _:vb6757910 _:vb6757911 _:vb6757896 _:vb6757897 _:vb6757898 _:vb6757899 _:vb6757900 _:vb6757901 _:vb6757902 _:vb6757903 _:vb6757888 _:vb6757889 _:vb6757890 _:vb6757891 _:vb6757892 _:vb6757893 _:vb6757894 _:vb6757895

Subject Item: _:vb6757888
rdf:type: n3:Context
rdf:value: A possible role of HH-GLI in human CC had been proposed based on limited in vitro data (e.g. Douard et al, >>2006<<; Qualtrough et al, 2004). This, however, has been questioned and denied (e.g. Berman et al, 2003; Chatel et al, 2007; van den Brink et al, 2004).
n3:mentions: n2:16701100

Subject Item: _:vb6757889
rdf:type: n3:Context
rdf:value: This, however, has been questioned and denied (e.g. Berman et al, >>2003<<; Chatel et al, 2007; van den Brink et al, 2004).
n3:mentions: n2:14520411

Subject Item: _:vb6757890
rdf:type: n3:Context
rdf:value: This, however, has been questioned and denied (e.g. Berman et al, 2003; Chatel et al, 2007; van den Brink et al, >>2004<<). Our extensive analyses with cell lines and primary tumours utilizing multiple in vitro and in vivo assays resolves the existing controversy in favor of an essential role of HH-GLI in human sporadic CCs.
n3:mentions: n2:14770182

Subject Item: _:vb6757891
rdf:type: n3:Context
rdf:value: We have previously mapped the site of action of HH-GLI signalling in several human cancers by in situ RNA hybridization to the epithelial tumour cells, as these express GLI1 and PTCH1 (Clement et al, >>2007<<; Dahmane et al, 2001; Stecca et al, 2007).
n3:mentions: n2:17196391

Subject Item: _:vb6757892
rdf:type: n3:Context
rdf:value: We have previously mapped the site of action of HH-GLI signalling in several human cancers by in situ RNA hybridization to the epithelial tumour cells, as these express GLI1 and PTCH1 (Clement et al, 2007; Dahmane et al, >>2001<<; Stecca et al, 2007).
n3:mentions: n2:11748155

Subject Item: _:vb6757893
rdf:type: n3:Context
rdf:value: previously mapped the site of action of HH-GLI signalling in several human cancers by in situ RNA hybridization to the epithelial tumour cells, as these express GLI1 and PTCH1 (Clement et al, 2007; Dahmane et al, 2001; Stecca et al, >>2007<<). However, this conclusion has recently been challenged for colon and other cancers, with suggestions of purely paracrine signalling—from SHH-expressing epithelium to GLI1-expressing responding stroma—largely based on extrapolation of
n3:mentions: n2:17392427

Subject Item: _:vb6757894
rdf:type: n3:Context
rdf:value: for colon and other cancers, with suggestions of purely paracrine signalling—from SHH-expressing epithelium to GLI1-expressing responding stroma—largely based on extrapolation of data from mice to the human situation (Tian et al, >>2009<<; Yauch et al, 2008; discussed in Ruiz i Altaba, 2008).
n3:mentions: n2:19246386

Subject Item: _:vb6757895
rdf:type: n3:Context
rdf:value: other cancers, with suggestions of purely paracrine signalling—from SHH-expressing epithelium to GLI1-expressing responding stroma—largely based on extrapolation of data from mice to the human situation (Tian et al, 2009; Yauch et al, >>2008<<; discussed in Ruiz i Altaba, 2008).
n3:mentions: n2:18754008

Subject Item: _:vb6757896
rdf:type: n3:Context
rdf:value: For instance, in rodents there is convincing data that signalling occurs from epithelium to mesenchyme in the prostate and colon (e.g. Madison et al, >>2005<<; Pu et al, 2004) but from epithelium to epithelium in humans (Sanchez et al, 2004; this work).
n3:mentions: n2:15590741

Subject Item: _:vb6757897
rdf:type: n3:Context
rdf:value: For instance, in rodents there is convincing data that signalling occurs from epithelium to mesenchyme in the prostate and colon (e.g. Madison et al, 2005; Pu et al, >>2004<<) but from epithelium to epithelium in humans (Sanchez et al, 2004; this work).
n3:mentions: n2:15328011

Subject Item: _:vb6757898
rdf:type: n3:Context
rdf:value: For instance, in rodents there is convincing data that signalling occurs from epithelium to mesenchyme in the prostate and colon (e.g. Madison et al, 2005; Pu et al, 2004) but from epithelium to epithelium in humans (Sanchez et al, >>2004<<; this work).
n3:mentions: n2:15314219

Subject Item: _:vb6757899
rdf:type: n3:Context
rdf:value: In CC, as in stomach (Berman et al, >>2003<<) and prostate (Sanchez et al, 2004) cancers, ligand-driven autocrine signalling in SHH+/GLI1+ epithelial cells may be active.
n3:mentions: n2:14520411

Subject Item: _:vb6757900
rdf:type: n3:Context
rdf:value: In CC, as in stomach (Berman et al, 2003) and prostate (Sanchez et al, >>2004<<) cancers, ligand-driven autocrine signalling in SHH+/GLI1+ epithelial cells may be active.
n3:mentions: n2:15314219

Subject Item: _:vb6757901
rdf:type: n3:Context
rdf:value: The rare GLI1-dependent, SMOH-independent cases we observe (see also Sanchez et al, >>2004<<) could have alterations in components downstream of SMOH.
n3:mentions: n2:15314219

Subject Item: _:vb6757902
rdf:type: n3:Context
rdf:value: Finally, the promotion of EMT by HH-GLI1 activity further supports the role of GLI1 as a central nexus regulating cancer cell behavior (Ruiz i Altaba et al, >>2007<<), including the promotion of metastases.
n3:mentions: n2:17845852

Subject Item: _:vb6757903
rdf:type: n3:Context
rdf:value: Several studies have established the human CD133+ (AC133+) CC fraction as a population of cells highly enriched in CC cancer stem cells (Ieta et al, >>2008<<; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, 2009).
n3:mentions: n2:17932721

Subject Item: _:vb6757904
rdf:type: n3:Context
rdf:value: Several studies have established the human CD133+ (AC133+) CC fraction as a population of cells highly enriched in CC cancer stem cells (Ieta et al, 2008; O'Brien et al, >>2007<<; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, 2009).
n3:mentions: n2:17122772

Subject Item: _:vb6757905
rdf:type: n3:Context
rdf:value: Several studies have established the human CD133+ (AC133+) CC fraction as a population of cells highly enriched in CC cancer stem cells (Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, >>2007<<; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, 2009). Moreover, CD133+ (Prominin1+) mouse intestinal cells are stem cells that can induce tumourigenesis (Snippert et al, 2009; Zhu et al, 2009).
n3:mentions: n2:17122771

Subject Item: _:vb6757906
rdf:type: n3:Context
rdf:value: Several studies have established the human CD133+ (AC133+) CC fraction as a population of cells highly enriched in CC cancer stem cells (Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, >>2007<<; Vermeulen et al, 2008; Zhu et al, 2009). Moreover, CD133+ (Prominin1+) mouse intestinal cells are stem cells that can induce tumourigenesis (Snippert et al, 2009; Zhu et al, 2009).
n3:mentions: n2:18371377

Subject Item: _:vb6757907
rdf:type: n3:Context
rdf:value: studies have established the human CD133+ (AC133+) CC fraction as a population of cells highly enriched in CC cancer stem cells (Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, >>2008<<; Zhu et al, 2009). Moreover, CD133+ (Prominin1+) mouse intestinal cells are stem cells that can induce tumourigenesis (Snippert et al, 2009; Zhu et al, 2009).
n3:mentions: n2:18765800

Subject Item: _:vb6757908
rdf:type: n3:Context
rdf:value: the human CD133+ (AC133+) CC fraction as a population of cells highly enriched in CC cancer stem cells (Ieta et al, 2008; O'Brien et al, 2007; Ricci-Vitiani et al, 2007; Todaro et al, 2007; Vermeulen et al, 2008; Zhu et al, >>2009<<). Moreover, CD133+ (Prominin1+) mouse intestinal cells are stem cells that can induce tumourigenesis (Snippert et al, 2009; Zhu et al, 2009).
n3:mentions: n2:19092805

Subject Item: _:vb6757909
rdf:type: n3:Context
rdf:value: Moreover, CD133+ (Prominin1+) mouse intestinal cells are stem cells that can induce tumourigenesis (Snippert et al, >>2009<<; Zhu et al, 2009).
n3:mentions: n2:19324043

Subject Item: _:vb6757910
rdf:type: n3:Context
rdf:value: Moreover, CD133+ (Prominin1+) mouse intestinal cells are stem cells that can induce tumourigenesis (Snippert et al, 2009; Zhu et al, >>2009<<). Our results support these findings and not the suggestion that all epithelial cells of local human CCs are CD133+ and that CD133+ cells are not tumourigenic (Shmelkov et al, 2008). We find only ±9–18% CD133+ cells in local CCs, that
n3:mentions: n2:19092805

Subject Item: _:vb6757911
rdf:type: n3:Context
rdf:value: Our results support these findings and not the suggestion that all epithelial cells of local human CCs are CD133+ and that CD133+ cells are not tumourigenic (Shmelkov et al, >>2008<<). We find only ±9–18% CD133+ cells in local CCs, that CD133+ cells are tumourigenic and that their population size is modulated by HH-GLI activity in vivo.
n3:mentions: n2:18497886

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n3:RelevantScore: 21
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rdf:type: n3:RelevantBibliographicResource
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n3:RelevantScore: 16
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n3:RelevantScore: 15
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