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n2:pmcid: PMC0
bibo:doi: 10.1038%2Fsrep08654
n5:contains: _:vb38496877 _:vb38496855

Subject Item: _:vb38496855
rdf:type: n5:Section
dc:title: results
n5:contains: _:vb38496872 _:vb38496873 _:vb38496874 _:vb38496875 _:vb38496876 _:vb38496864 _:vb38496865 _:vb38496866 _:vb38496867 _:vb38496868 _:vb38496869 _:vb38496870 _:vb38496871 _:vb38496856 _:vb38496857 _:vb38496858 _:vb38496859 _:vb38496860 _:vb38496861 _:vb38496862 _:vb38496863

Subject Item: _:vb38496856
rdf:type: n2:Context
rdf:value: S1B). As previously reported>>20<<, we found that in normal human colonic crypts, Ki67 (MKI67) expression was restricted to the lower quarter of the crypt, and cytokeratin-20 (KRT20) expression was only found on the luminal surface, with a portion of the crypt expressing
n2:mentions: n3:18162766

Subject Item: _:vb38496857
rdf:type: n2:Context
rdf:value: 2A), consistent with reported isotopic ISH>>3<<10 and expression patterns seen in murine colonic crypts2.
n2:mentions: n3:22287596

Subject Item: _:vb38496858
rdf:type: n2:Context
rdf:value: 2A), consistent with reported isotopic ISH>>310<< and expression patterns seen in murine colonic crypts2.
n2:mentions: n3:23766371

Subject Item: _:vb38496859
rdf:type: n2:Context
rdf:value: 2A), consistent with reported isotopic ISH310 and expression patterns seen in murine colonic crypts>>2<<. We noted that normal small intestine also expresses LGR5 mRNA in a similarly small number of cells at the crypt base (Supplementary Fig. S2A and B).
n2:mentions: n3:17934449

Subject Item: _:vb38496860
rdf:type: n2:Context
rdf:value: HPPs are defined as small, non-dysplastic serrated lesions of the colon, which are generally thought to exhibit little or no malignant potential, although certain subtypes may be precursors to sessile serrated adenomas/polyps>>21<< (see Fig. 1). All of the HPPs described in this study were small lesions (<5 mm) and were validated independently by two expert pathologists as having no features suggestive of sessile serrated adenomas/polyps (SSA/Ps). As previously
n2:mentions: n3:22710576

Subject Item: _:vb38496861
rdf:type: n2:Context
rdf:value: As previously reported>>20<<, we found that all HPPs exhibited KRT20 expression that extends further down into the crypt than in normal crypts, and there was expansion of the Ki67+ population.
n2:mentions: n3:18162766

Subject Item: _:vb38496862
rdf:type: n2:Context
rdf:value: We chose to use CD44 as an additional stem cell marker to validate the LGR5 ISH, as it is one of the most widely used putative markers of stem cell populations in a broad range of normal and cancerous tissues>>22<<2324.
n2:mentions: n3:18980968

Subject Item: _:vb38496863
rdf:type: n2:Context
rdf:value: We chose to use CD44 as an additional stem cell marker to validate the LGR5 ISH, as it is one of the most widely used putative markers of stem cell populations in a broad range of normal and cancerous tissues>>2223<<24.
n2:mentions: n3:12629218

Subject Item: _:vb38496864
rdf:type: n2:Context
rdf:value: We chose to use CD44 as an additional stem cell marker to validate the LGR5 ISH, as it is one of the most widely used putative markers of stem cell populations in a broad range of normal and cancerous tissues>>222324<<.
n2:mentions: n3:19415765

Subject Item: _:vb38496865
rdf:type: n2:Context
rdf:value: We report that within subregions of adenomatous tissue, LGR5 is expressed in cells spread along the entire gland length and at a relatively uniform level (Fig. 3A, Fig. 4A), an expression pattern that has been previously documented>>10<<. Immunohistochemical analysis showed that the glands expressing high LGR5 were often lacking KRT20 expression suggesting the cells were indeed in an undifferentiated state, and furthermore the expression of the stem cell marker CD44 was
n2:mentions: n3:23766371

Subject Item: _:vb38496866
rdf:type: n2:Context
rdf:value: This ‘patchy’ expression of LGR5 has previously been reported in colonic adenomas in studies using LGR5-targeting antibodies>>14<<15. Interestingly, we noted the presence of structures in which expansion of the LGR5 compartment was restricted along the length of half of the adenomatous gland (Fig. 4C), suggestive of dysregulation of the stem cell niche. It is
n2:mentions: n3:19030762

Subject Item: _:vb38496867
rdf:type: n2:Context
rdf:value: This ‘patchy’ expression of LGR5 has previously been reported in colonic adenomas in studies using LGR5-targeting antibodies>>1415<<. Interestingly, we noted the presence of structures in which expansion of the LGR5 compartment was restricted along the length of half of the adenomatous gland (Fig. 4C), suggestive of dysregulation of the stem cell niche. It is possible
n2:mentions: n3:20195621

Subject Item: _:vb38496868
rdf:type: n2:Context
rdf:value: Although the majority of colorectal cancers arise via the conventional adenoma-carcinoma pathway, approximately 10–20% of colorectal carcinomas are thought to arise from serrated lesions>>25<<. These tumors follow a different pathway of progression from adenoma to carcinoma (Fig.
n2:mentions: n3:23431044

Subject Item: _:vb38496869
rdf:type: n2:Context
rdf:value: 1), and have a distinct genetic and epigenetic profile to their non-serrated counterparts>>26<<, and it has been proposed that serrated lesions can be identified by differential expression of certain markers27.
n2:mentions: n3:20869746

Subject Item: _:vb38496870
rdf:type: n2:Context
rdf:value: 1), and have a distinct genetic and epigenetic profile to their non-serrated counterparts26, and it has been proposed that serrated lesions can be identified by differential expression of certain markers>>27<<. We therefore sought to determine if the disruption of the stem cell niche was a feature of the crypts of sessile serrated adenomas/polyps (SSA/Ps) and traditional serrated adenomas (TSAs).
n2:mentions: n3:23570326

Subject Item: _:vb38496871
rdf:type: n2:Context
rdf:value: S3), consistent with a report suggesting that the putative stem cell marker CD133 is more highly expressed in SSA/Ps than HPPs>>28<<. The expression of CD44 was more extensive than normal or hyperplastic crypts, although still localized to the crypt base (Fig. 3B). SSA/Ps frequently displayed expansion of the Ki67+ population compared to normal crypts, and KRT20
n2:mentions: n3:23436931

Subject Item: _:vb38496872
rdf:type: n2:Context
rdf:value: Ectopic crypts are a defining feature of TSAs in which small crypts grow laterally from the side of the original crypt>>20<<. We identified regions containing ectopic crypts in the TSAs that were analyzed for LGR5 expression, and present novel evidence to show that these crypts contain a small population of LGR5-expressing cells at the base.
n2:mentions: n3:18162766

Subject Item: _:vb38496873
rdf:type: n2:Context
rdf:value: It has been reported that human colorectal cancers can contain an LGR5-expressing stem cell niche at the base of tumor glands resembling crypts>>4<<5. We therefore analyzed LGR5 ISH and Ki67, KRT20 and CD44 IHC in 23 adenocarcinomas of all stages (pT1:
n2:mentions: n3:22637696

Subject Item: _:vb38496874
rdf:type: n2:Context
rdf:value: It has been reported that human colorectal cancers can contain an LGR5-expressing stem cell niche at the base of tumor glands resembling crypts>>45<<. We therefore analyzed LGR5 ISH and Ki67, KRT20 and CD44 IHC in 23 adenocarcinomas of all stages (pT1:
n2:mentions: n3:21419747

Subject Item: _:vb38496875
rdf:type: n2:Context
rdf:value: This is in apparent contrast to a previous report>>29<< in which low Ki67 was described at the invasive front, however such comparisons are confounded by inconsistency of staining protocols and inter-observer heterogeneity.
n2:mentions: n3:11696421

Subject Item: _:vb38496876
rdf:type: n2:Context
rdf:value: The expression of LGR5 has been reported to be correlated with invasion and metastasis of gastric cancer>>30<<. We therefore sought to examine LGR5 expression within invasive cell populations in our samples, and noted that LGR5 expression was generally high in populations of invading cells (Fig.
n2:mentions: n3:24594994

Subject Item: _:vb38496877
rdf:type: n5:Section
dc:title: discussion
n5:contains: _:vb38496880 _:vb38496881 _:vb38496882 _:vb38496883 _:vb38496884 _:vb38496885 _:vb38496886 _:vb38496887 _:vb38496888 _:vb38496889 _:vb38496890 _:vb38496891 _:vb38496892 _:vb38496878 _:vb38496879

Subject Item: _:vb38496878
rdf:type: n2:Context
rdf:value: A previously published report has suggested that the organization of the cell hierarchy within murine adenomas closely resembles that of the normal crypt, with Lgr5 expression remaining at the crypt base>>7<<. However there is also evidence to suggest that in the human colon, stem cell architecture is lost during the development of adenomas, and that expression of the colonic stem cell marker LGR5 is widespread throughout adenomatous glands10.
n2:mentions: n3:22855427

Subject Item: _:vb38496879
rdf:type: n2:Context
rdf:value: However there is also evidence to suggest that in the human colon, stem cell architecture is lost during the development of adenomas, and that expression of the colonic stem cell marker LGR5 is widespread throughout adenomatous glands>>10<<. This report clarifies these discrepancies by providing new evidence that while conventional adenomas have a disrupted stem cell architecture as determined by frequent and widespread non-basal expression of LGR5, the stem cell
n2:mentions: n3:23766371

Subject Item: _:vb38496880
rdf:type: n2:Context
rdf:value: These structures extend laterally from crypts of a TSA, and little is known about the mechanism of their formation, although in the murine colon it has been proposed to be due to a repression of bone morphogenetic protein (BMP) signaling>>31<<. The observation that ectopic crypts express basal LGR5 suggests that their growth is indeed sustained by cell division within a ‘mini stem cell niche’.
n2:mentions: n3:15017003

Subject Item: _:vb38496881
rdf:type: n2:Context
rdf:value: This data is consistent with previous reports suggesting that mouse>>7<< and human10 adenomatous glands display a dramatic increase in the number of LGR5-positive cells.
n2:mentions: n3:22855427

Subject Item: _:vb38496882
rdf:type: n2:Context
rdf:value: This data is consistent with previous reports suggesting that mouse7 and human>>10<< adenomatous glands display a dramatic increase in the number of LGR5-positive cells.
n2:mentions: n3:23766371

Subject Item: _:vb38496883
rdf:type: n2:Context
rdf:value: We note that in the murine colon, only a small percentage of the total Lgr5-expressing cells actually function as stem cells at any one time>>8<<, though they are all endowed with stem cell potential9.
n2:mentions: n3:24035355

Subject Item: _:vb38496884
rdf:type: n2:Context
rdf:value: We note that in the murine colon, only a small percentage of the total Lgr5-expressing cells actually function as stem cells at any one time8, though they are all endowed with stem cell potential>>9<<. In addition, we have recently shown that normal human colonic crypts are sustained by a small number of ‘functional’ stem cells (similar to that of murine crypts), however adenomatous crypts display an elevated number of functional stem
n2:mentions: n3:24531760

Subject Item: _:vb38496885
rdf:type: n2:Context
rdf:value: we have recently shown that normal human colonic crypts are sustained by a small number of ‘functional’ stem cells (similar to that of murine crypts), however adenomatous crypts display an elevated number of functional stem cells>>6<<. Thus, applying this insight to conventional adenomas, the large number of LGR5+ cells suggests a large pool of cells with stem cell potential, but not necessarily a large number of functioning stem cells.
n2:mentions: n3:25127143

Subject Item: _:vb38496886
rdf:type: n2:Context
rdf:value: LGR5 is reported to positively regulate invasion and metastasis of gastric cancer via a matrix metalloproteinase 2 (MMP2)-dependent mechanism>>30<<, however in vitro and mouse xenograft models of colorectal cancer have shown that LGR5 knockdown increases invasion32.
n2:mentions: n3:24594994

Subject Item: _:vb38496887
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rdf:value: regulate invasion and metastasis of gastric cancer via a matrix metalloproteinase 2 (MMP2)-dependent mechanism30, however in vitro and mouse xenograft models of colorectal cancer have shown that LGR5 knockdown increases invasion>>32<<. Our analysis revealed that LGR5-positive cells are substantially enriched in invasive cell populations, suggesting that LGR5 can support invasion rather than inhibit it.
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rdf:value: that occurs upon invasion is also seen in lineage-labelled murine stem cells, whereby measurements of clone size and proliferative potential in invasive squamous cell carcinoma showed geometric expansion of the stem cell population>>33<<.
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rdf:value: Interestingly, a previous report suggests that in established human colon carcinomas, LGR5 expression remains basal and is exclusive from KRT20 expression>>5<<. We were able to examine LGR5 expression in 18 cases of adenocarcinoma, and consistent with the findings of Merlos-Suarez and colleagues, we report that in the majority of established carcinomas with a glandular structure,
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rdf:value: These are significant observations, as there is little currently known about the histogenesis of serrated adenomas, although the origin of conventional adenomas has been intensively investigated>>34<<35. Our findings may reflect an important difference in their origin and mode of progression. Moreover, we note that significant shortcomings in the routine diagnosis of serrated colorectal polyps have been reported36, and due to its
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rdf:value: These are significant observations, as there is little currently known about the histogenesis of serrated adenomas, although the origin of conventional adenomas has been intensively investigated>>3435<<. Our findings may reflect an important difference in their origin and mode of progression. Moreover, we note that significant shortcomings in the routine diagnosis of serrated colorectal polyps have been reported36, and due to its
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rdf:value: Moreover, we note that significant shortcomings in the routine diagnosis of serrated colorectal polyps have been reported>>36<<, and due to its consistent basal localization in serrated lesions, the examination of LGR5 distribution might play a role in improving discrimination of these enigmatic lesions.
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