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10.1038%2Fonc.2011.131
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introduction
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LIN28B is a homolog of LIN28 (also called LIN28A) (Guo et al., >>2006<<), which induces pluripotency in somatic cells when expressed in concert with KLF4, SOX2, and NANOG(Yu et al., 2007).
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LIN28B is a homolog of LIN28 (also called LIN28A) (Guo et al., 2006), which induces pluripotency in somatic cells when expressed in concert with KLF4, SOX2, and NANOG(Yu et al., >>2007<<). A high degree of homology exists between LIN28B, LIN28 and the heterochronic gene lin-28 in C. elegans (Moss and Tang, 2003).
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A high degree of homology exists between LIN28B, LIN28 and the heterochronic gene lin-28 in C. elegans (Moss and Tang, >>2003<<). Human LIN28 and LIN28B each contain a cold-shock domain and CCHC zinc fingers that confer RNA binding ability(Moss and Tang, 2003). The ability to bind RNA is critical to both Lin28 and Lin28b, as inhibition of let-7 microRNA biogenesis
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Human LIN28 and LIN28B each contain a cold-shock domain and CCHC zinc fingers that confer RNA binding ability(Moss and Tang, >>2003<<). The ability to bind RNA is critical to both Lin28 and Lin28b, as inhibition of let-7 microRNA biogenesis is a cardinal feature of their functions.
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The let-7 family of microRNAs comprises isoforms with highly conserved sequences that exhibit functional redundancy (Pasquinelli et al., >>2000<<; Zhao et al., 2010).
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that of most microRNAs in that the various isoforms are transcribed initially by RNA polymerase II as pri-microRNAs, and processed by Drosha and DGCR8 into pre-microRNAs that are subsequently exported from the nucleus (Gregory et al., >>2004<<; Han et al., 2004; Lee et al., 2003; Lee et al., 2002). The hairpin loops of pre-microRNAs are cleaved by dicer in the cytoplasm to yield microRNA:
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in that the various isoforms are transcribed initially by RNA polymerase II as pri-microRNAs, and processed by Drosha and DGCR8 into pre-microRNAs that are subsequently exported from the nucleus (Gregory et al., 2004; Han et al., >>2004<<; Lee et al., 2003; Lee et al., 2002). The hairpin loops of pre-microRNAs are cleaved by dicer in the cytoplasm to yield microRNA:
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isoforms are transcribed initially by RNA polymerase II as pri-microRNAs, and processed by Drosha and DGCR8 into pre-microRNAs that are subsequently exported from the nucleus (Gregory et al., 2004; Han et al., 2004; Lee et al., >>2003<<; Lee et al., 2002). The hairpin loops of pre-microRNAs are cleaved by dicer in the cytoplasm to yield microRNA:
n2:mentions
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transcribed initially by RNA polymerase II as pri-microRNAs, and processed by Drosha and DGCR8 into pre-microRNAs that are subsequently exported from the nucleus (Gregory et al., 2004; Han et al., 2004; Lee et al., 2003; Lee et al., >>2002<<). The hairpin loops of pre-microRNAs are cleaved by dicer in the cytoplasm to yield microRNA:
n2:mentions
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The hairpin loops of pre-microRNAs are cleaved by dicer in the cytoplasm to yield microRNA:microRNA duplexes that are disassociated to release mature let-7 (Lee et al., >>2003<<; Lee et al., 2002).
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The hairpin loops of pre-microRNAs are cleaved by dicer in the cytoplasm to yield microRNA:microRNA duplexes that are disassociated to release mature let-7 (Lee et al., 2003; Lee et al., >>2002<<). MicroRNAs are incorporated into the RNA induced silencing complex (RISC) and bind the 3′ UTR of target transcripts to provide post-transcriptional gene regulation by mRNA sequestration or cleavage (Esquela-Kerscher and Slack, 2006).
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MicroRNAs are incorporated into the RNA induced silencing complex (RISC) and bind the 3′ UTR of target transcripts to provide post-transcriptional gene regulation by mRNA sequestration or cleavage (Esquela-Kerscher and Slack, >>2006<<).
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Several established let-7 mRNA targets have tumor promoting properties, including the canonical target HMGA2 (Lee and Dutta, >>2007<<; Mayr et al., 2007; Park et al., 2007) and the classic oncogenes KRAS and c-MYC (Akao et al., 2006; Johnson et al., 2007; Johnson et al., 2005).
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Several established let-7 mRNA targets have tumor promoting properties, including the canonical target HMGA2 (Lee and Dutta, 2007; Mayr et al., >>2007<<; Park et al., 2007) and the classic oncogenes KRAS and c-MYC (Akao et al., 2006; Johnson et al., 2007; Johnson et al., 2005).
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Several established let-7 mRNA targets have tumor promoting properties, including the canonical target HMGA2 (Lee and Dutta, 2007; Mayr et al., 2007; Park et al., >>2007<<) and the classic oncogenes KRAS and c-MYC (Akao et al., 2006; Johnson et al., 2007; Johnson et al., 2005).
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Several established let-7 mRNA targets have tumor promoting properties, including the canonical target HMGA2 (Lee and Dutta, 2007; Mayr et al., 2007; Park et al., 2007) and the classic oncogenes KRAS and c-MYC (Akao et al., >>2006<<; Johnson et al., 2007; Johnson et al., 2005).
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let-7 mRNA targets have tumor promoting properties, including the canonical target HMGA2 (Lee and Dutta, 2007; Mayr et al., 2007; Park et al., 2007) and the classic oncogenes KRAS and c-MYC (Akao et al., 2006; Johnson et al., >>2007<<; Johnson et al., 2005). In addition, let-7 microRNAs have been described as tumor suppressors and are implicated as prognostic factors in a variety of divergent cancers (Akao et al., 2006; Shell et al., 2007; Takamizawa et al., 2004).
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have tumor promoting properties, including the canonical target HMGA2 (Lee and Dutta, 2007; Mayr et al., 2007; Park et al., 2007) and the classic oncogenes KRAS and c-MYC (Akao et al., 2006; Johnson et al., 2007; Johnson et al., >>2005<<). In addition, let-7 microRNAs have been described as tumor suppressors and are implicated as prognostic factors in a variety of divergent cancers (Akao et al., 2006; Shell et al., 2007; Takamizawa et al., 2004).
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In addition, let-7 microRNAs have been described as tumor suppressors and are implicated as prognostic factors in a variety of divergent cancers (Akao et al., >>2006<<; Shell et al., 2007; Takamizawa et al., 2004).
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In addition, let-7 microRNAs have been described as tumor suppressors and are implicated as prognostic factors in a variety of divergent cancers (Akao et al., 2006; Shell et al., >>2007<<; Takamizawa et al., 2004).
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In addition, let-7 microRNAs have been described as tumor suppressors and are implicated as prognostic factors in a variety of divergent cancers (Akao et al., 2006; Shell et al., 2007; Takamizawa et al., >>2004<<). Importantly, Lin28 and Lin28b may relieve let-7 target suppression by binding to immature let-7 molecules and blocking further processing (Hagan et al., 2009; Heo et al., 2008; Heo et al., 2009).
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Importantly, Lin28 and Lin28b may relieve let-7 target suppression by binding to immature let-7 molecules and blocking further processing (Hagan et al., >>2009<<; Heo et al., 2008; Heo et al., 2009).
n2:mentions
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Importantly, Lin28 and Lin28b may relieve let-7 target suppression by binding to immature let-7 molecules and blocking further processing (Hagan et al., 2009; Heo et al., >>2008<<; Heo et al., 2009).
n2:mentions
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Importantly, Lin28 and Lin28b may relieve let-7 target suppression by binding to immature let-7 molecules and blocking further processing (Hagan et al., 2009; Heo et al., 2008; Heo et al., >>2009<<).
n2:mentions
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The regulator of let-7 biogenesis LIN28B, also a let-7 target (Boyerinas et al., >>2008<<), is specifically implicated in this process because it is transactivated by c-myc (Chang et al., 2009).
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The regulator of let-7 biogenesis LIN28B, also a let-7 target (Boyerinas et al., 2008), is specifically implicated in this process because it is transactivated by c-myc (Chang et al., >>2009<<). Nearly 70% of colorectal tumors harbor elevated levels of c-myc (Erisman et al., 1985); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006;
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Nearly 70% of colorectal tumors harbor elevated levels of c-myc (Erisman et al., >>1985<<); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006; He et al., 1998; Powell et al., 1992; Rubinfeld et al., 1993; Sikora et al., 1987;
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Nearly 70% of colorectal tumors harbor elevated levels of c-myc (Erisman et al., 1985); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, >>2006<<; He et al., 1998; Powell et al., 1992; Rubinfeld et al., 1993; Sikora et al., 1987; Stewart et al., 1986).
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colorectal tumors harbor elevated levels of c-myc (Erisman et al., 1985); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006; He et al., >>1998<<; Powell et al., 1992; Rubinfeld et al., 1993; Sikora et al., 1987; Stewart et al., 1986).
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elevated levels of c-myc (Erisman et al., 1985); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006; He et al., 1998; Powell et al., >>1992<<; Rubinfeld et al., 1993; Sikora et al., 1987; Stewart et al., 1986).
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c-myc (Erisman et al., 1985); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006; He et al., 1998; Powell et al., 1992; Rubinfeld et al., >>1993<<; Sikora et al., 1987; Stewart et al., 1986).
n2:mentions
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1985); up-regulation occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006; He et al., 1998; Powell et al., 1992; Rubinfeld et al., 1993; Sikora et al., >>1987<<; Stewart et al., 1986).
n2:mentions
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occurs in the early stages of colon carcinoma as a consequence of Wnt pathway deregulation and β-catenin stabilization (Clevers, 2006; He et al., 1998; Powell et al., 1992; Rubinfeld et al., 1993; Sikora et al., 1987; Stewart et al., >>1986<<).
n2:mentions
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materials and methods
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Intestinal crypts were isolated from adult BL6 females as previously described by Flint, et al, >>1991<<. RNA was isolated using TRIzol (Invitrogen, Carsblad, CA) and 1 μg total RNA use for reverse transcriptase reactions with superscript III (Invitrogen, Carlsblad, CA) and oligo dT.
n2:mentions
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; Keene et al., >>2006<<) supplemented with protease, phosphatase and RNase inhibitors; lysates were cleared by centrifugation and frozen at −80°C overnight.
n2:mentions
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; Keene et al., >>2006<<) and blocked in 5% BSA, 200 μg/ml yeast tRNA for 1 hour at 4°C.
n2:mentions
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results
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As expected, we found increased expression of known let-7 targets, including IGF2BP1, CDC34, and HMGA2 (Boyerinas et al., >>2008<<; Johnson et al., 2007; Lee and Dutta, 2007) (Figure 6a).
n2:mentions
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As expected, we found increased expression of known let-7 targets, including IGF2BP1, CDC34, and HMGA2 (Boyerinas et al., 2008; Johnson et al., >>2007<<; Lee and Dutta, 2007) (Figure 6a).
n2:mentions
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As expected, we found increased expression of known let-7 targets, including IGF2BP1, CDC34, and HMGA2 (Boyerinas et al., 2008; Johnson et al., 2007; Lee and Dutta, >>2007<<) (Figure 6a).
n2:mentions
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A similar phenomenon has been described for the Lin28b homolog Lin28, which binds to and stabilizes IGF2 mRNA during differentiation of myoblasts (Polesskaya et al., >>2007<<). In order to determine whether Lin28b associates with LGR5 and PROM1 mRNA transcripts, we first examined RNA-protein interactions of Lin28b in colon cancer cell lines through RNA immunoprecipitation followed by quantitative RT-qPCR. IGF2
n2:mentions
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discussion
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canonical Wnt signaling, it is possible that LIN28B is up-regulated in colon tumors as a consequence of APC mutation (or other changes that deregulate Wnt signaling), which occurs in the vast majority of colon tumors (Pino and Chung, >>2010<<). Alternatively, up-regulation of LIN28B in colon tumors may occur as a result of increased mRNA stabilization.
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recent evidence demonstrates that Lin28 also does not function exclusively through inhibition of let-7 processing, but blocks gliogenesis in favor of neurogenesis in undifferentiated cells by stabilizing IGF2 mRNA (Balzer et al., >>2010<<). Considering the high degree of homology between Lin28 and Lin28b, and the RNA-binding activity inherent to both, it is possible that both Lin28 and Lin28b modulate expression of a number of genes in addition to IGF2, LGR5, and PROM1,
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One might consider introducing point mutations into the cold-shock domain of Lin28b; this approach has been utilized previously for evaluating let-7 independent functions of Lin28 (Balzer et al., >>2010<<). However, cold-shock domain mutations may disrupt all RNA-binding activities of LIN28B, for example, LGR5 and PROM1 binding, thereby precluding assessment of specific let-7 independent functions. Thus, determining the specific domains
n2:mentions
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Within the intestine, expression of the cell surface protein PROM1 is restricted to the crypts and adjacent epithelial cells (Snippert et al., >>2009<<), while expression of the orphan receptor LGR5 occurs exclusively in cycling columnar cells within the crypt base (Barker et al., 2007).
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surface protein PROM1 is restricted to the crypts and adjacent epithelial cells (Snippert et al., 2009), while expression of the orphan receptor LGR5 occurs exclusively in cycling columnar cells within the crypt base (Barker et al., >>2007<<). Since co-expression of LGR5 and PROM1 marks intestinal and colonic epithelial stem cells, up-regulation of these factors by Lin28b suggests a possible role for Lin28b in establishment and/or maintenance of intestinal stem cells.
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Interestingly, adenomas may arise in the colon from PROM1+ crypt cells (Zhu et al., >>2009<<), and overexpression of LGR5 in colorectal adenocarcinomas correlates with late-stage tumorigenesis, invasion, and metastasis (Uchida et al., 2010).
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Interestingly, adenomas may arise in the colon from PROM1+ crypt cells (Zhu et al., 2009), and overexpression of LGR5 in colorectal adenocarcinomas correlates with late-stage tumorigenesis, invasion, and metastasis (Uchida et al., >>2010<<). We have demonstrated that constitutive LIN28B expression promotes both tumorigenesis as well as induction of LGR5 and PROM1 in colonic epithelial cells.
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