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n2:pmcid: PMC0
bibo:doi: 10.1083%2Fjcb.200811030
n5:contains: _:vb5405873 _:vb5405888 _:vb5405895

Subject Item: _:vb5405873
rdf:type: n5:Section
dc:title: introduction
n5:contains: _:vb5405874 _:vb5405875 _:vb5405876 _:vb5405877 _:vb5405878 _:vb5405879 _:vb5405880 _:vb5405881 _:vb5405882 _:vb5405883 _:vb5405884 _:vb5405885 _:vb5405886 _:vb5405887

Subject Item: _:vb5405874
rdf:type: n2:Context
rdf:value: Although several molecules have been identified as regulators of epithelial apico-basal polarity (Shin et al., >>2006<<), precise mechanisms to regulate the apical domain organization in the cortical progenitor cells remain unknown.
n2:mentions: n3:16771626

Subject Item: _:vb5405875
rdf:type: n2:Context
rdf:value: In Drosophila melanogaster, there are two Fats, Fat and Fat-like (Tanoue and Takeichi, >>2005<<), which are different in their functions.
n2:mentions: n3:15923647

Subject Item: _:vb5405876
rdf:type: n2:Context
rdf:value: Fat is well known as a regulator of cell proliferation and planar cell polarity (PCP; Bryant et al., >>1988<<; Mahoney et al., 1991; Yang et al., 2002; Ma et al., 2003; Casal et al., 2006).
n2:mentions: n3:3417051

Subject Item: _:vb5405877
rdf:type: n2:Context
rdf:value: Fat is well known as a regulator of cell proliferation and planar cell polarity (PCP; Bryant et al., 1988; Mahoney et al., >>1991<<; Yang et al., 2002; Ma et al., 2003; Casal et al., 2006).
n2:mentions: n3:1959133

Subject Item: _:vb5405878
rdf:type: n2:Context
rdf:value: Fat is well known as a regulator of cell proliferation and planar cell polarity (PCP; Bryant et al., 1988; Mahoney et al., 1991; Yang et al., >>2002<<; Ma et al., 2003; Casal et al., 2006).
n2:mentions: n3:11893338

Subject Item: _:vb5405879
rdf:type: n2:Context
rdf:value: Fat is well known as a regulator of cell proliferation and planar cell polarity (PCP; Bryant et al., 1988; Mahoney et al., 1991; Yang et al., 2002; Ma et al., >>2003<<; Casal et al., 2006).
n2:mentions: n3:12540853

Subject Item: _:vb5405880
rdf:type: n2:Context
rdf:value: Fat is well known as a regulator of cell proliferation and planar cell polarity (PCP; Bryant et al., 1988; Mahoney et al., 1991; Yang et al., 2002; Ma et al., 2003; Casal et al., >>2006<<). In the PCP regulation, Fat interacts with Dachsous, another huge cadherin, through the extracellular region of each. dachsous mutants show several morphogenetic defects (Clark et al., 1995), and both fat and dachsous mutants commonly
n2:mentions: n3:17075008

Subject Item: _:vb5405881
rdf:type: n2:Context
rdf:value: dachsous mutants show several morphogenetic defects (Clark et al., >>1995<<), and both fat and dachsous mutants commonly exhibit defects in PCP (Yang et al., 2002; Ma et al., 2003).
n2:mentions: n3:7601355

Subject Item: _:vb5405882
rdf:type: n2:Context
rdf:value: dachsous mutants show several morphogenetic defects (Clark et al., 1995), and both fat and dachsous mutants commonly exhibit defects in PCP (Yang et al., >>2002<<; Ma et al., 2003).
n2:mentions: n3:11893338

Subject Item: _:vb5405883
rdf:type: n2:Context
rdf:value: dachsous mutants show several morphogenetic defects (Clark et al., 1995), and both fat and dachsous mutants commonly exhibit defects in PCP (Yang et al., 2002; Ma et al., >>2003<<).
n2:mentions: n3:12540853

Subject Item: _:vb5405884
rdf:type: n2:Context
rdf:value: In mammals, four types of Fats have been identified: Fat1, Fat2, Fat3, and Fat4 (Rock et al., >>2005<<; Tanoue and Takeichi, 2005). Among them, Fat4 is thought to be the vertebrate counterpart of Drosophila Fat based on their sequence similarity.
n2:mentions: n3:16059920

Subject Item: _:vb5405885
rdf:type: n2:Context
rdf:value: In mammals, four types of Fats have been identified: Fat1, Fat2, Fat3, and Fat4 (Rock et al., 2005; Tanoue and Takeichi, >>2005<<). Among them, Fat4 is thought to be the vertebrate counterpart of Drosophila Fat based on their sequence similarity.
n2:mentions: n3:15923647

Subject Item: _:vb5405886
rdf:type: n2:Context
rdf:value: phenotypes of Fat4-null mice were reported; they exhibit complicated morphological abnormalities, including abnormal orientation of inner ear hair cells, cystic dilation of kidney tubules, and defects in organ shapes (Saburi et al., >>2008<<). However, the molecular and cellular functions of Fat4 still remain to be analyzed.
n2:mentions: n3:18604206

Subject Item: _:vb5405887
rdf:type: n2:Context
rdf:value: At the mRNA level, the former is broadly expressed in mouse embryos, whereas the latter exhibits a very restricted expression (Rock et al., >>2005<<). Their functions also remain to be explored.
n2:mentions: n3:16059920

Subject Item: _:vb5405888
rdf:type: n5:Section
dc:title: materials and methods
n5:contains: _:vb5405892 _:vb5405893 _:vb5405894 _:vb5405889 _:vb5405890 _:vb5405891

Subject Item: _:vb5405889
rdf:type: n2:Context
rdf:value: Cells were cultured as described previously (Tanoue and Takeichi, >>2004<<). For isolating stable cDNA transfectants, 400 µg/ml G418 (Invitrogen) or 250 µg/ml hygromycin B (Invitrogen) were used.
n2:mentions: n3:15148305

Subject Item: _:vb5405890
rdf:type: n2:Context
rdf:value: Immunostaining was performed as described previously (Tanoue and Takeichi, >>2004<<). Cell aggregation assays were performed as described previously (Nagafuchi et al., 1987).
n2:mentions: n3:15148305

Subject Item: _:vb5405891
rdf:type: n2:Context
rdf:value: Cell aggregation assays were performed as described previously (Nagafuchi et al., >>1987<<).
n2:mentions: n3:3498123

Subject Item: _:vb5405892
rdf:type: n2:Context
rdf:value: N terminus–truncated forms of Fat4 and Dachsous1 were constructed by using pCA-Sig-IRES vector (Tanoue and Takeichi, >>2004<<). For production of a GST-fused cytoplasmic region and deletion mutants of Fat4 in Escherichia coli, the pGEX-2T vector (GE Healthcare) was used.
n2:mentions: n3:15148305

Subject Item: _:vb5405893
rdf:type: n2:Context
rdf:value: Detailed procedures were described previously (Saito, >>2006<<). The following Stealth siRNAs were used for the RNAi experiments:
n2:mentions: n3:17406448

Subject Item: _:vb5405894
rdf:type: n2:Context
rdf:value: Detailed procedures were described previously (Tanoue and Takeichi, >>2004<<).
n2:mentions: n3:15148305

Subject Item: _:vb5405895
rdf:type: n5:Section
dc:title: results and discussion
n5:contains: _:vb5405912 _:vb5405913 _:vb5405914 _:vb5405904 _:vb5405905 _:vb5405906 _:vb5405907 _:vb5405908 _:vb5405909 _:vb5405910 _:vb5405911 _:vb5405896 _:vb5405897 _:vb5405898 _:vb5405899 _:vb5405900 _:vb5405901 _:vb5405902 _:vb5405903

Subject Item: _:vb5405896
rdf:type: n2:Context
rdf:value: The discontinuous distribution of Fat and Dachsous was also observed for the Drosophila homologues (Ma et al., >>2003<<), suggesting that this unique localization pattern reflects certain conserved nature of these molecules.
n2:mentions: n3:12540853

Subject Item: _:vb5405897
rdf:type: n2:Context
rdf:value: In Drosophila, in the absence of fat, the level of Dachsous protein is down-regulated; conversely, without dachsous, the Fat protein level is up-regulated (Ma et al., >>2003<<). To test whether such a relationship also exists for the mammalian homologues, we depleted these molecules in the embryonic cortices by electroporating siRNAs targeting their transcripts.
n2:mentions: n3:12540853

Subject Item: _:vb5405898
rdf:type: n2:Context
rdf:value: MUPP1 is a paralogue of Patj, which regulates apico-basal polarity and apical membrane organization in cultured cells as well as in Drosophila (Bachmann et al., >>2001<<; Shin et al., 2005; Richard et al., 2006; Sugihara-Mizuno et al., 2007).
n2:mentions: n3:11740560

Subject Item: _:vb5405899
rdf:type: n2:Context
rdf:value: MUPP1 is a paralogue of Patj, which regulates apico-basal polarity and apical membrane organization in cultured cells as well as in Drosophila (Bachmann et al., 2001; Shin et al., >>2005<<; Richard et al., 2006; Sugihara-Mizuno et al., 2007).
n2:mentions: n3:15738264

Subject Item: _:vb5405900
rdf:type: n2:Context
rdf:value: MUPP1 is a paralogue of Patj, which regulates apico-basal polarity and apical membrane organization in cultured cells as well as in Drosophila (Bachmann et al., 2001; Shin et al., 2005; Richard et al., >>2006<<; Sugihara-Mizuno et al., 2007).
n2:mentions: n3:16245332

Subject Item: _:vb5405901
rdf:type: n2:Context
rdf:value: MUPP1 is a paralogue of Patj, which regulates apico-basal polarity and apical membrane organization in cultured cells as well as in Drosophila (Bachmann et al., 2001; Shin et al., 2005; Richard et al., 2006; Sugihara-Mizuno et al., >>2007<<). The interaction between Fat4 and MUPP1 was confirmed by detecting MUPP1 from the pulled-down materials with anti-MUPP1 antibodies (Fig.
n2:mentions: n3:17397395

Subject Item: _:vb5405902
rdf:type: n2:Context
rdf:value: As previous studies showed that MUPP1 forms a complex with Pals1, a vertebrate counterpart of Drosophila Stardust (van de Pavert et al., >>2004<<; Sugihara-Mizuno et al., 2007), we examined whether this interaction also occurred in the cortex, and as a result, we indeed detected Pals1 in the MUPP1 immunoprecipitates prepared from cortical lysates (Fig.
n2:mentions: n3:15316081

Subject Item: _:vb5405903
rdf:type: n2:Context
rdf:value: As previous studies showed that MUPP1 forms a complex with Pals1, a vertebrate counterpart of Drosophila Stardust (van de Pavert et al., 2004; Sugihara-Mizuno et al., >>2007<<), we examined whether this interaction also occurred in the cortex, and as a result, we indeed detected Pals1 in the MUPP1 immunoprecipitates prepared from cortical lysates (Fig.
n2:mentions: n3:17397395

Subject Item: _:vb5405904
rdf:type: n2:Context
rdf:value: Because the Stardust complex has been reported to regulate the apical membrane organization in Drosophila (Pellikka et al., >>2002<<; Hong et al., 2003; Richard et al., 2006), we asked whether Fat4 and Dachsous1 had similar functions.
n2:mentions: n3:11850625

Subject Item: _:vb5405905
rdf:type: n2:Context
rdf:value: Because the Stardust complex has been reported to regulate the apical membrane organization in Drosophila (Pellikka et al., 2002; Hong et al., >>2003<<; Richard et al., 2006), we asked whether Fat4 and Dachsous1 had similar functions.
n2:mentions: n3:14569003

Subject Item: _:vb5405906
rdf:type: n2:Context
rdf:value: Because the Stardust complex has been reported to regulate the apical membrane organization in Drosophila (Pellikka et al., 2002; Hong et al., 2003; Richard et al., >>2006<<), we asked whether Fat4 and Dachsous1 had similar functions. We prepared ultrathin sections for electron microscopy of E15.5 cortices, which had been electroporated with siRNAs targeting Fat4, Dachsous1, or Pals1 (Fig.
n2:mentions: n3:16245332

Subject Item: _:vb5405907
rdf:type: n2:Context
rdf:value: 5 A, red), as observed by others (Ho et al., >>2000<<). This gap appeared to contain amorphous electron-dense materials, but the cytoplasmic sides exhibited no specific structures.
n2:mentions: n3:10753311

Subject Item: _:vb5405908
rdf:type: n2:Context
rdf:value: We did not examine the effect of MUPP1 removal because of the presence of its paralogue, Patj, in the cortex (Srinivasan et al., >>2008<<).
n2:mentions: n3:18403412

Subject Item: _:vb5405909
rdf:type: n2:Context
rdf:value: The AJ is known as the major cell junctional structure observed in the apical portions of neural progenitor cells (Ho et al., >>2000<<; Lien et al., 2006; Kadowaki et al., 2007).
n2:mentions: n3:10753311

Subject Item: _:vb5405910
rdf:type: n2:Context
rdf:value: The AJ is known as the major cell junctional structure observed in the apical portions of neural progenitor cells (Ho et al., 2000; Lien et al., >>2006<<; Kadowaki et al., 2007).
n2:mentions: n3:16543460

Subject Item: _:vb5405911
rdf:type: n2:Context
rdf:value: The AJ is known as the major cell junctional structure observed in the apical portions of neural progenitor cells (Ho et al., 2000; Lien et al., 2006; Kadowaki et al., >>2007<<). We found that Fat4 and Dachsous1 were located more apical to the AJ and that the plasma membranes at the corresponding region showed a simple apposition, which we defined as the subapical membrane apposition. Such apically extended
n2:mentions: n3:17222817

Subject Item: _:vb5405912
rdf:type: n2:Context
rdf:value: We found that Fat4 and Dachsous1 could interact in a heterophilic fashion and regulated the protein level of the partner in a way similar to that found in Drosophila (Ma et al., >>2003<<). Then, the question arises as to how the Fat4–Dachsous1 system regulates the apical membrane architecture.
n2:mentions: n3:12540853

Subject Item: _:vb5405913
rdf:type: n2:Context
rdf:value: Stardust, the Drosophila counterpart of Pals1, and Crumbs are well known to regulate the apical membrane organization in Drosophila retinal cells (Pellikka et al., >>2002<<). It is attractive to hypothesize that Fat may cooperate with these apical regulators and in turn takes part in the apical membrane organization.
n2:mentions: n3:11850625

Subject Item: _:vb5405914
rdf:type: n2:Context
rdf:value: Because Crumbs also forms a complex with MUPP1/PatJ and Pals1 (van de Pavert et al., >>2004<<), it is important to determine how the Fat and Crumbs systems share these binding partners for their potential cooperation in future studies.
n2:mentions: n3:15316081

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