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n2:pmcid: PMC0
bibo:doi: 10.1186%2F1741-7007-7-62
n7:contains: _:vb6088245 _:vb6088221 _:vb6088203 _:vb6088189 _:vb6088168

Subject Item: _:vb6088168
rdf:type: n7:Section
dc:title: results
n7:contains: _:vb6088172 _:vb6088173 _:vb6088174 _:vb6088175 _:vb6088169 _:vb6088170 _:vb6088171 _:vb6088180 _:vb6088181 _:vb6088182 _:vb6088183 _:vb6088176 _:vb6088177 _:vb6088178 _:vb6088179 _:vb6088188 _:vb6088184 _:vb6088185 _:vb6088186 _:vb6088187

Subject Item: _:vb6088169
rdf:type: n2:Context
rdf:value: in scrib mutant mosaic discs and blocking JNK signalling in scrib mutant clones by expressing a dominant-negative form of Drosophila JNK, Basket dominant negative (BskDN), dramatically increasing the scrib mutant clonal tissue size [>>2<<]. In agreement with these observations and other previously published reports [17,30], we confirmed that expressing BskDN in scrib mutant clones reduced apoptosis in the mutant tissue, although cell death was still observed in some wild
n2:mentions: n3:14592975

Subject Item: _:vb6088170
rdf:type: n2:Context
rdf:value: In agreement with these observations and other previously published reports [>>17<<,30], we confirmed that expressing BskDN in scrib mutant clones reduced apoptosis in the mutant tissue, although cell death was still observed in some wild type cells abutting the mutant clones (see Additional file 1, panels A-C).
n2:mentions: n3:16753569

Subject Item: _:vb6088171
rdf:type: n2:Context
rdf:value: In agreement with these observations and other previously published reports [17,>>30<<], we confirmed that expressing BskDN in scrib mutant clones reduced apoptosis in the mutant tissue, although cell death was still observed in some wild type cells abutting the mutant clones (see Additional file 1, panels A-C).
n2:mentions: n3:16150723

Subject Item: _:vb6088172
rdf:type: n2:Context
rdf:value: Furthermore, using a reporter of JNK signalling, the lacZ enhancer trap, misshapen (msn)-lacZ [>>31<<], we also confirmed that JNK signalling was ectopically activated within some scrib mutant cells, including those undergoing apoptosis, and that expressing BskDN in scrib mutant clones effectively prevented the ectopic expression of
n2:mentions: n3:15968584

Subject Item: _:vb6088173
rdf:type: n2:Context
rdf:value: Loss-of-function crb clones, using the null allele crb11A22 [>>25<<], exhibited no apparent defects in differentiation or cell morphology (see Additional file 2, panels A-B), although during pupal development cell morphology defects become apparent within the developing photoreceptor cells [32,33].
n2:mentions: n3:2344615

Subject Item: _:vb6088174
rdf:type: n2:Context
rdf:value: crb11A22 [25], exhibited no apparent defects in differentiation or cell morphology (see Additional file 2, panels A-B), although during pupal development cell morphology defects become apparent within the developing photoreceptor cells [>>32<<,33]. In contrast, third instar larval eye disc clones over-expressing a wild-type Crb transgene were small and mutant cells tended to be excluded from the epithelium with severely altered, more rounded, cell morphology.
n2:mentions: n3:11850624

Subject Item: _:vb6088175
rdf:type: n2:Context
rdf:value: [25], exhibited no apparent defects in differentiation or cell morphology (see Additional file 2, panels A-B), although during pupal development cell morphology defects become apparent within the developing photoreceptor cells [32,>>33<<]. In contrast, third instar larval eye disc clones over-expressing a wild-type Crb transgene were small and mutant cells tended to be excluded from the epithelium with severely altered, more rounded, cell morphology.
n2:mentions: n3:11850625

Subject Item: _:vb6088176
rdf:type: n2:Context
rdf:value: Similar overgrowth and polarity defects, but not JNK-dependent cell death, have been described when Crb was over-expressed in the wing disc epithelium [>>15<<].
n2:mentions: n3:16258546

Subject Item: _:vb6088177
rdf:type: n2:Context
rdf:value: Previously it has been shown that ectopic expression of aPKC in Drosophila can disrupt epithelial cell morphology and induce CycE expression [>>13<<], although this was not in a clonal context.
n2:mentions: n3:16116079

Subject Item: _:vb6088178
rdf:type: n2:Context
rdf:value: Therefore, in order to verify that the over-expression of aPKC could mimic the scrib mutant phenotype in the eye, we over-expressed wild-type aPKC incorporating a membrane-tethering CAAX motif (aPKCCAAXWT) in eye disc clones [>>14<<]. This produced a variable phenotype but generally led to only mild defects in tissue organization and very weak ectopic CycE expression (data not shown). In order to investigate the more extreme consequences of aPKC activation we
n2:mentions: n3:15302858

Subject Item: _:vb6088179
rdf:type: n2:Context
rdf:value: In order to investigate the more extreme consequences of aPKC activation we analysed clones of eye disc tissue ectopically expressing an activated version of aPKC lacking its N-terminal regulatory domain (aPKCΔN) [>>10<<]. This resulted in small eye disc clones, however, blocking JNK signalling in the aPKCΔN-expressing clones restored clone viability and most of the mutant tissue had aberrant morphology and was extruded basally to form large masses of
n2:mentions: n3:12629552

Subject Item: _:vb6088180
rdf:type: n2:Context
rdf:value: To determine if the scrib mutant defects could be due to deregulated aPKC activity we utilized a transgene expressing a kinase-dead, CAAX membrane-tethered, allele of aPKC (aPKCCAAXDN) [>>14<<]. The expression of aPKCCAAXDN in otherwise wild-type clones of tissue produced no discernible defects in cell morphology, proliferation or differentiation during larval stages of development (data not shown).
n2:mentions: n3:15302858

Subject Item: _:vb6088181
rdf:type: n2:Context
rdf:value: Such a phenomena is reminiscent of the non-cell autonomous compensatory cell proliferation that can be induced by dying cells within imaginal discs [reviewed in [>>34<<]]. Indeed, although the expression of aPKCCAAXDN in scrib mutant clones rescued most of the scrib mutant defects, the viability of the mutant tissue remained poor and the remnants of many apoptotic cells were evident. TUNEL detection
n2:mentions: n3:18774295

Subject Item: _:vb6088182
rdf:type: n2:Context
rdf:value: we have also observed that they cooperate with activated alleles of dRas1 (RasACT) RasACT or Notch (NACT) to repress pupal development and, throughout an extended 'giant larvae' phase of development, form massive and invasive tumours [>>2<<,3]. scrib- + RasACT tumour cells grow out basally from the eye disc, fail to differentiate (Figure 5A, B) and appear to invade the brain lobes along F-actin rich cables extending from between the eye/antennal disc to the brain, eventually
n2:mentions: n3:14592975

Subject Item: _:vb6088183
rdf:type: n2:Context
rdf:value: we have also observed that they cooperate with activated alleles of dRas1 (RasACT) RasACT or Notch (NACT) to repress pupal development and, throughout an extended 'giant larvae' phase of development, form massive and invasive tumours [2,>>3<<]. scrib- + RasACT tumour cells grow out basally from the eye disc, fail to differentiate (Figure 5A, B) and appear to invade the brain lobes along F-actin rich cables extending from between the eye/antennal disc to the brain, eventually
n2:mentions: n3:14551319

Subject Item: _:vb6088184
rdf:type: n2:Context
rdf:value: Consistent with previous reports [>>16<<,17], we found that blocking JNK signalling in scrib- + RasACT tumours, by co-expressing BskDN, restored pupation to the tumour-bearing larvae and repressed tumour invasion (see Additional file 4, panels F, G).
n2:mentions: n3:17082773

Subject Item: _:vb6088185
rdf:type: n2:Context
rdf:value: Consistent with previous reports [16,>>17<<], we found that blocking JNK signalling in scrib- + RasACT tumours, by co-expressing BskDN, restored pupation to the tumour-bearing larvae and repressed tumour invasion (see Additional file 4, panels F, G).
n2:mentions: n3:16753569

Subject Item: _:vb6088186
rdf:type: n2:Context
rdf:value: Proteins involved in both cell migration and invasion, including the matrix metalloproteinase, Mmp1 (data not shown) [>>16<<,18] and the integrin-associated scaffolding protein, Paxillin (Pax), were up-regulated in scrib mutant clones and at the invasive front of scrib- + RasACT tumours, in a JNK-dependent manner (see Additional file 5).
n2:mentions: n3:17082773

Subject Item: _:vb6088187
rdf:type: n2:Context
rdf:value: Proteins involved in both cell migration and invasion, including the matrix metalloproteinase, Mmp1 (data not shown) [16,>>18<<] and the integrin-associated scaffolding protein, Paxillin (Pax), were up-regulated in scrib mutant clones and at the invasive front of scrib- + RasACT tumours, in a JNK-dependent manner (see Additional file 5).
n2:mentions: n3:17301221

Subject Item: _:vb6088188
rdf:type: n2:Context
rdf:value: cell death, the failure of aPKCCAAXDN to block scrib- + RasACT tumourigenesis supports the hypothesis that JNK signalling alone is both necessary and sufficient in cooperation with RasACT to lead to neoplastic transformation [>>16<<].
n2:mentions: n3:17082773

Subject Item: _:vb6088189
rdf:type: n7:Section
dc:title: methods
n7:contains: _:vb6088190 _:vb6088191 _:vb6088192 _:vb6088193 _:vb6088194 _:vb6088195 _:vb6088196 _:vb6088197 _:vb6088198 _:vb6088199 _:vb6088200 _:vb6088201 _:vb6088202

Subject Item: _:vb6088190
rdf:type: n2:Context
rdf:value: All clonal analysis was carried out using MARCM (mosaic analysis with repressible cell marker) [>>19<<] with FRT82B and eyeless-FLP1 to induce clones and UAS-mCD8-GFP to visualize mutant tissue.
n2:mentions: n3:10197526

Subject Item: _:vb6088191
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [>>20<<]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned
n2:mentions: n3:11290291

Subject Item: _:vb6088192
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [>>21<<]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this
n2:mentions: n3:10471706

Subject Item: _:vb6088193
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [>>22<<]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:10688207

Subject Item: _:vb6088194
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [>>23<<]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:7925015

Subject Item: _:vb6088195
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [>>24<<]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:10022918

Subject Item: _:vb6088196
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [>>25<<]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:2344615

Subject Item: _:vb6088197
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [>>26<<]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:7606787

Subject Item: _:vb6088198
rdf:type: n2:Context
rdf:value: The following Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [>>10<<]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:12629552

Subject Item: _:vb6088199
rdf:type: n2:Context
rdf:value: Drosophila stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [>>14<<]; UAS-dRas1V12 [27]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:15302858

Subject Item: _:vb6088200
rdf:type: n2:Context
rdf:value: stocks were used: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [>>27<<]; UAS-Nintra [28]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:9389658

Subject Item: _:vb6088201
rdf:type: n2:Context
rdf:value: eyFLP1, UAS-mCD8-GFP;;Tub-GAL4 FRT82B Tub-GAL80 [20]; msn06946 [21]; scrib1 [22]; UAS-P35 [23]; UAS-bskDN [24]; crb11A22 [25]; UAS-crbwt2e [26]; UAS-DaPKCΔN [10]; UAS-DaPKCCAAXWT and UAS-DaPKCCAAXDN [14]; UAS-dRas1V12 [27]; UAS-Nintra [>>28<<]; UASp-scribFL19.2 (full length Scrib cDNA cloned into pUASP, this study).
n2:mentions: n3:9570768

Subject Item: _:vb6088202
rdf:type: n2:Context
rdf:value: mouse anti-Elav (Developmental Studies Hybridoma Bank) at 1 in 20, rat anti-Cyc E (Helen McNeill) at 1 in 400, rabbit anti-GFP (Invitrogen) at 1 in 1000, mouse anti-BrdU (Becton-Dickinson) at 1 in 50, rabbit anti-Paxillin at 1 in 400 [>>29<<]. Secondary antibodies were; anti-mouse/rat/rabbit Alexa647 (Invitrogen) at 1 in 400, anti-mouse/rat biotin (Jackson ImmunoResearch Laboratories) at 1 in 400 and streptavidin-conjugated fluorophores (Jackson ImmunoResearch Laboratories)
n2:mentions: n3:11520860

Subject Item: _:vb6088203
rdf:type: n7:Section
dc:title: discussion
n7:contains: _:vb6088204 _:vb6088205 _:vb6088206 _:vb6088207 _:vb6088216 _:vb6088217 _:vb6088218 _:vb6088219 _:vb6088220 _:vb6088208 _:vb6088209 _:vb6088210 _:vb6088211 _:vb6088212 _:vb6088213 _:vb6088214 _:vb6088215

Subject Item: _:vb6088204
rdf:type: n2:Context
rdf:value: This supports the epistatic relationship between scrib and crb described in the embryo, with the scrib mutant phenotype being dominant over the crb mutant phenotype, and suggesting that Crb acts upstream or independently of Scrib [>>8<<]. In contrast, the strong rescue of scrib mutant defects by expressing a dominant negative aPKC transgene suggests that aPKC either acts to inactivate Scrib and blocking aPKC restores Scrib activity as has been proposed for Lgl, or
n2:mentions: n3:12510194

Subject Item: _:vb6088205
rdf:type: n2:Context
rdf:value: Crb over-expression phenotypes can also be suppressed by aPKCCAAXDN co-expression, and aPKC can phosphorylate Crb to modulate its activity [>>14<<]. Similarly, aPKC can phosphorylate and inactivate Lgl, although Lgl also functions genetically upstream of aPKC in restraining the formation of neuroblastomas, by acting as a competitive substrate of aPKC and impeding aPKC's ability to
n2:mentions: n3:15302858

Subject Item: _:vb6088206
rdf:type: n2:Context
rdf:value: and inactivate Lgl, although Lgl also functions genetically upstream of aPKC in restraining the formation of neuroblastomas, by acting as a competitive substrate of aPKC and impeding aPKC's ability to phosphorylate and inactivate Numb [>>35<<]. In mammals, Scrib can also function upstream of aPKC via the correct localization of Cdc42 during cell migration [36,37].
n2:mentions: n3:18854163

Subject Item: _:vb6088207
rdf:type: n2:Context
rdf:value: In mammals, Scrib can also function upstream of aPKC via the correct localization of Cdc42 during cell migration [>>36<<,37]. However, in this context loss of Scrib appears to impair localized aPKC activity, suggesting that the relationship between Scrib and aPKC activity can vary in different contexts.
n2:mentions: n3:17081755

Subject Item: _:vb6088208
rdf:type: n2:Context
rdf:value: In mammals, Scrib can also function upstream of aPKC via the correct localization of Cdc42 during cell migration [36,>>37<<]. However, in this context loss of Scrib appears to impair localized aPKC activity, suggesting that the relationship between Scrib and aPKC activity can vary in different contexts.
n2:mentions: n3:17043654

Subject Item: _:vb6088209
rdf:type: n2:Context
rdf:value: avl mutant hyperplasia is also rescued by the expression of aPKCCAAXDN and this was suggested to reflect the ability of aPKCCAAXDN to reduce Crb activity, since Crb levels were elevated and mislocalized in both avl and Rab5 mutants [>>15<<]. Whether Crb or aPKC is the key to the formation of avl or Rab5 neoplasias, clearly an intimate relationship exists between the different neoplastic tumour suppressors and the polarity complex proteins.
n2:mentions: n3:16258546

Subject Item: _:vb6088210
rdf:type: n2:Context
rdf:value: Our studies confirm previous studies with respect to the key role for JNK in mediating the cooperative neoplastic overgrowth of scrib mutants with RasACT [>>16<<,17]. Oncogenic signals subvert a protective apoptotic JNK response, to an invasive neoplasia.
n2:mentions: n3:17082773

Subject Item: _:vb6088211
rdf:type: n2:Context
rdf:value: Our studies confirm previous studies with respect to the key role for JNK in mediating the cooperative neoplastic overgrowth of scrib mutants with RasACT [16,>>17<<]. Oncogenic signals subvert a protective apoptotic JNK response, to an invasive neoplasia.
n2:mentions: n3:16753569

Subject Item: _:vb6088212
rdf:type: n2:Context
rdf:value: Two identified JNK targets in scrib- + RasACT tumours are the matrix metalloproteinase protein, Mmp1 [our unpublished observations, [>>16<<,18]], and the integrin-associated scaffolding protein, Paxillin (this study).
n2:mentions: n3:17082773

Subject Item: _:vb6088213
rdf:type: n2:Context
rdf:value: Two identified JNK targets in scrib- + RasACT tumours are the matrix metalloproteinase protein, Mmp1 [our unpublished observations, [16,>>18<<]], and the integrin-associated scaffolding protein, Paxillin (this study).
n2:mentions: n3:17301221

Subject Item: _:vb6088214
rdf:type: n2:Context
rdf:value: of Timp (Tissue inhibitor of metalloproteases) restrained scrib- + RasACT tumour cells from fusing with and invading the brain lobes but did not abrogate tumour overgrowth or restore pupal development [our unpublished observations, [>>16<<,18]]. Both Mmp1 and Pax were induced by JNK signalling, independent of both scrib or Ras, since clones of cells expressing an activated allele of the Drosophila JNKK homologue, hemipterous (HepACT), also showed strong up-regulation of Pax
n2:mentions: n3:17082773

Subject Item: _:vb6088215
rdf:type: n2:Context
rdf:value: of Timp (Tissue inhibitor of metalloproteases) restrained scrib- + RasACT tumour cells from fusing with and invading the brain lobes but did not abrogate tumour overgrowth or restore pupal development [our unpublished observations, [16,>>18<<]]. Both Mmp1 and Pax were induced by JNK signalling, independent of both scrib or Ras, since clones of cells expressing an activated allele of the Drosophila JNKK homologue, hemipterous (HepACT), also showed strong up-regulation of Pax
n2:mentions: n3:17301221

Subject Item: _:vb6088216
rdf:type: n2:Context
rdf:value: However, it is also likely that Ras and N synergize with JNK to drive expression of novel target genes since, in scrib mutants kept alive with P35, JNK remains activated but this does not recapitulate the oncogenic effects of Ras or N [>>2<<]. One possible key to the ability of JNK to promote overgrowth in combination with RasACT is through blocking differentiation, since expressing BskDN in scrib- + RasACT tumours restored Ras-induced differentiation and thus restrained
n2:mentions: n3:14592975

Subject Item: _:vb6088217
rdf:type: n2:Context
rdf:value: The ability to repress pupation appears to be a property shared by all neoplastic overgrowth in Drosophila [>>38<<], although the contribution that JNK plays to this in other contexts is not yet known.
n2:mentions: n3:17947427

Subject Item: _:vb6088218
rdf:type: n2:Context
rdf:value: Many different cell polarity mutants apart from scrib share in the capacity to cooperate with RasACT in neoplastic transformation through JNK signalling, including genes that genetically act in opposition to scrib such as sdt [>>17<<] and crb (our unpublished observation). This is consistent with JNK being activated indirectly as a consequence of disturbed cell polarity/morphology and further suggests that JNK alone might be sufficient for cooperation.
n2:mentions: n3:16753569

Subject Item: _:vb6088219
rdf:type: n2:Context
rdf:value: Indeed, co-expression of Hep with RasACT has been shown to result in invasive neoplasia [>>16<<]. Our results are consistent with this since the cell morphology and proliferative defects of scrib mutant clones are rescued by aPKCCAAXDN, however, the mutant cells still undergo JNK-mediated apoptosis and can still cooperate with
n2:mentions: n3:17082773

Subject Item: _:vb6088220
rdf:type: n2:Context
rdf:value: In contrast, co-expressing HepACT with RasACT inefficiently results in neoplastic transformation, presumably because the levels of JNK signalling are too high and this restrains overgrowth or leads to cell death [>>16<<]. A loss of Scrib appears to contribute a level of JNK activity strong enough to result in either cell death or neoplastic transformation in cooperation with RasACT.
n2:mentions: n3:17082773

Subject Item: _:vb6088221
rdf:type: n7:Section
dc:title: background
n7:contains: _:vb6088222 _:vb6088223 _:vb6088240 _:vb6088241 _:vb6088242 _:vb6088243 _:vb6088244 _:vb6088224 _:vb6088225 _:vb6088226 _:vb6088227 _:vb6088228 _:vb6088229 _:vb6088230 _:vb6088231 _:vb6088232 _:vb6088233 _:vb6088234 _:vb6088235 _:vb6088236 _:vb6088237 _:vb6088238 _:vb6088239

Subject Item: _:vb6088222
rdf:type: n2:Context
rdf:value: giant larvae (Lgl), cooperatively establish and maintain apico-basal cell polarity and repress inappropriate cell proliferation and neoplasia (invasive overgrowth with a failure to differentiate) in both epithelial and neuronal tissues [>>1<<]. Furthermore, in a fly 'two-hit' model of tumourigenesis, the loss of any one of these three genes has also been shown to cooperate with oncogenic alleles of Ras resulting in neoplasia [2,3].
n2:mentions: n3:10884224

Subject Item: _:vb6088223
rdf:type: n2:Context
rdf:value: Furthermore, in a fly 'two-hit' model of tumourigenesis, the loss of any one of these three genes has also been shown to cooperate with oncogenic alleles of Ras resulting in neoplasia [>>2<<,3]. As the function of this group of proteins is conserved in humans (including human Scrib's ability to cooperate with oncogenes in promoting tumourigenesis [4,5]) a deeper understanding is needed of the way in which these genes function
n2:mentions: n3:14592975

Subject Item: _:vb6088224
rdf:type: n2:Context
rdf:value: Furthermore, in a fly 'two-hit' model of tumourigenesis, the loss of any one of these three genes has also been shown to cooperate with oncogenic alleles of Ras resulting in neoplasia [2,>>3<<]. As the function of this group of proteins is conserved in humans (including human Scrib's ability to cooperate with oncogenes in promoting tumourigenesis [4,5]) a deeper understanding is needed of the way in which these genes function
n2:mentions: n3:14551319

Subject Item: _:vb6088225
rdf:type: n2:Context
rdf:value: As the function of this group of proteins is conserved in humans (including human Scrib's ability to cooperate with oncogenes in promoting tumourigenesis [>>4<<,5]) a deeper understanding is needed of the way in which these genes function to repress neoplasia.
n2:mentions: n3:19041750

Subject Item: _:vb6088226
rdf:type: n2:Context
rdf:value: As the function of this group of proteins is conserved in humans (including human Scrib's ability to cooperate with oncogenes in promoting tumourigenesis [4,>>5<<]) a deeper understanding is needed of the way in which these genes function to repress neoplasia.
n2:mentions: n3:18641685

Subject Item: _:vb6088227
rdf:type: n2:Context
rdf:value: a powerful model organism, can be used to investigate these questions as pathways regulating cell proliferation, survival, differentiation and tumour cell invasion are all highly conserved between flies and humans [reviewed in [>>6<<]].
n2:mentions: n3:16034367

Subject Item: _:vb6088228
rdf:type: n2:Context
rdf:value: of aberrant cell polarity), ectopic expression of Cyclin E (CycE) and excessive cell proliferation, the mutant clones of tissue do not become overgrown because they are removed by Jun N-terminal kinase (JNK)-dependent apoptosis [>>2<<]. If, however, activated oncogenic alleles of either the small GTPase Ras (dRas1V12 or shortened to RasACT) or the receptor/transcription regulator Notch (Nintra or shortened to NACT) are specifically expressed within the mutant tissue,
n2:mentions: n3:14592975

Subject Item: _:vb6088229
rdf:type: n2:Context
rdf:value: to NACT) are specifically expressed within the mutant tissue, tumours are formed which become massively overgrown throughout an extended larval stage of development and which then invade the adjacent brain and ventral nerve cord [>>2<<,3].
n2:mentions: n3:14592975

Subject Item: _:vb6088230
rdf:type: n2:Context
rdf:value: to NACT) are specifically expressed within the mutant tissue, tumours are formed which become massively overgrown throughout an extended larval stage of development and which then invade the adjacent brain and ventral nerve cord [2,>>3<<].
n2:mentions: n3:14551319

Subject Item: _:vb6088231
rdf:type: n2:Context
rdf:value: Most of what is known of the way in which Scrib represses epithelial neoplasia in Drosophila has focused on how Scrib regulates cell polarity, particularly in the embryonic ectoderm [reviewed in [>>7<<]]. Genetic analysis suggests that Scrib, in cooperation with Dlg and Lgl, promotes basolateral membrane identity and functions antagonistically towards two other protein complexes, the Crumbs (Crb) complex and the Bazooka (Baz) complex,
n2:mentions: n3:19029932

Subject Item: _:vb6088232
rdf:type: n2:Context
rdf:value: with Dlg and Lgl, promotes basolateral membrane identity and functions antagonistically towards two other protein complexes, the Crumbs (Crb) complex and the Bazooka (Baz) complex, both functioning to promote apical cell identity [>>8<<,9]. The Crb complex, incorporating Crb, Stardust (Sdt) and Patj, is anchored apically through Crb's transmembrane domain.
n2:mentions: n3:12510194

Subject Item: _:vb6088233
rdf:type: n2:Context
rdf:value: with Dlg and Lgl, promotes basolateral membrane identity and functions antagonistically towards two other protein complexes, the Crumbs (Crb) complex and the Bazooka (Baz) complex, both functioning to promote apical cell identity [8,>>9<<]. The Crb complex, incorporating Crb, Stardust (Sdt) and Patj, is anchored apically through Crb's transmembrane domain.
n2:mentions: n3:12510193

Subject Item: _:vb6088234
rdf:type: n2:Context
rdf:value: act antagonistically towards one another is still incomplete, aPKC directly phosphorylates Lgl resulting in its inactivation and the binding of Lgl to aPKC has the potential to repress the ability of aPKC to phosphorylate other targets [>>10<<].
n2:mentions: n3:12629552

Subject Item: _:vb6088235
rdf:type: n2:Context
rdf:value: Studies have suggested that the proliferation and polarity functions of Scrib can be separated [>>11<<]. However, whether Scrib operates antagonistically to Crb and aPKC to repress proliferation is not known. In lgl mutants, tumour overgrowth can be rescued through reduced levels of aPKC [12], and aPKC over-expression is capable of
n2:mentions: n3:15611336

Subject Item: _:vb6088236
rdf:type: n2:Context
rdf:value: In lgl mutants, tumour overgrowth can be rescued through reduced levels of aPKC [>>12<<], and aPKC over-expression is capable of inducing CycE [13].
n2:mentions: n3:14657233

Subject Item: _:vb6088237
rdf:type: n2:Context
rdf:value: In lgl mutants, tumour overgrowth can be rescued through reduced levels of aPKC [12], and aPKC over-expression is capable of inducing CycE [>>13<<]. However, it is not known if aPKC functions upstream of Lgl, or if Lgl acts to restrain aPKC phosphorylation of alternative key targets that promote epithelial overgrowth. In fact, aPKC can activate Crb through phosphorylation [14] and
n2:mentions: n3:16116079

Subject Item: _:vb6088238
rdf:type: n2:Context
rdf:value: In fact, aPKC can activate Crb through phosphorylation [>>14<<] and Crb over-expression in the wing disc promotes epithelial neoplasia similar to loss of function mutants in scrib, dlg or lgl [15].
n2:mentions: n3:15302858

Subject Item: _:vb6088239
rdf:type: n2:Context
rdf:value: In fact, aPKC can activate Crb through phosphorylation [14] and Crb over-expression in the wing disc promotes epithelial neoplasia similar to loss of function mutants in scrib, dlg or lgl [>>15<<]. Thus, deregulated Crb activity could be primarily responsible for neoplastic overgrowth in scrib mutants as has been suggested for mutants in the syntaxin avalanche (avl) [15]. Deciphering the hierarch that operates amongst these key
n2:mentions: n3:16258546

Subject Item: _:vb6088240
rdf:type: n2:Context
rdf:value: Thus, deregulated Crb activity could be primarily responsible for neoplastic overgrowth in scrib mutants as has been suggested for mutants in the syntaxin avalanche (avl) [>>15<<]. Deciphering the hierarch that operates amongst these key polarity players in scrib mutant epithelial neoplasias is required.
n2:mentions: n3:16258546

Subject Item: _:vb6088241
rdf:type: n2:Context
rdf:value: of studies have shown how RasACT subverts the pro-apoptotic JNK signalling response in scrib mutants into a potent inducer of tumour overgrowth and invasion through the JNK-dependent expression of Matrix metalloproteinase 1 (Mmp1) [>>16<<-18]. However, whilst there is agreement on the key role of JNK in mediating cooperative overgrowth, these reports give conflicting conclusions about the role of Scrib.
n2:mentions: n3:17082773 n3:16753569 n3:17301221

Subject Item: _:vb6088242
rdf:type: n2:Context
rdf:value: It has been suggested that loss of Scrib contributes JNK-independent roles in promoting cooperation with RasACT [>>17<<], while others offer evidence that JNK is itself sufficient for the cooperation with RasACT [16] and, thus, cell polarity genes such as Scrib repress oncogene-mediated transformation merely by restraining JNK activation.
n2:mentions: n3:16753569

Subject Item: _:vb6088243
rdf:type: n2:Context
rdf:value: It has been suggested that loss of Scrib contributes JNK-independent roles in promoting cooperation with RasACT [17], while others offer evidence that JNK is itself sufficient for the cooperation with RasACT [>>16<<] and, thus, cell polarity genes such as Scrib repress oncogene-mediated transformation merely by restraining JNK activation.
n2:mentions: n3:17082773

Subject Item: _:vb6088244
rdf:type: n2:Context
rdf:value: As mammalian studies have recently demonstrated that human Scrib similarly restrains RasACT-mediated transformation [>>5<<], it is important to more fully understand how Drosophila Scrib exerts its tumour suppressor function.
n2:mentions: n3:18641685

Subject Item: _:vb6088245
rdf:type: n7:Section
dc:title: conclusion
n7:contains: _:vb6088246 _:vb6088247 _:vb6088248 _:vb6088249 _:vb6088250 _:vb6088251 _:vb6088252 _:vb6088253 _:vb6088254 _:vb6088255 _:vb6088256

Subject Item: _:vb6088246
rdf:type: n2:Context
rdf:value: Growing evidence links increased levels of aPKC with the development of human cancers [>>13<<,39] and accumulating data support a role for human Scrib in restraining carcinogenesis [reviewed in [7]].
n2:mentions: n3:16116079

Subject Item: _:vb6088247
rdf:type: n2:Context
rdf:value: Growing evidence links increased levels of aPKC with the development of human cancers [13,>>39<<] and accumulating data support a role for human Scrib in restraining carcinogenesis [reviewed in [7]].
n2:mentions: n3:15994303

Subject Item: _:vb6088248
rdf:type: n2:Context
rdf:value: Growing evidence links increased levels of aPKC with the development of human cancers [13,39] and accumulating data support a role for human Scrib in restraining carcinogenesis [reviewed in [>>7<<]]. Furthermore, the knockdown of human Scrib in MCF10A cells has recently been shown to cooperate with RasACT or Myc in promoting transformation.
n2:mentions: n3:19029932

Subject Item: _:vb6088249
rdf:type: n2:Context
rdf:value: In the case of RasACT expression with Scrib knockdown, cells grown in three-dimensional (3D) culture failed to form the normal polarized acini structures with a central luminal and, instead, adopted a highly invasive morphology [>>5<<]. Cooperation with RasACT was linked to the ability of Scrib knockdown to potentiate MAPK signalling [5], however, phospho-JNK levels were also increased.
n2:mentions: n3:18641685

Subject Item: _:vb6088250
rdf:type: n2:Context
rdf:value: Cooperation with RasACT was linked to the ability of Scrib knockdown to potentiate MAPK signalling [>>5<<], however, phospho-JNK levels were also increased.
n2:mentions: n3:18641685

Subject Item: _:vb6088251
rdf:type: n2:Context
rdf:value: JNK signalling is increasingly implicated in mammalian carcinogenesis [>>40<<-42], although, as in Drosophila, its role can be complex as it also promotes tumour regression through cell death in different contexts [43].
n2:mentions: n3:17440097 n3:17210697 n3:18593901

Subject Item: _:vb6088252
rdf:type: n2:Context
rdf:value: JNK signalling is increasingly implicated in mammalian carcinogenesis [40-42], although, as in Drosophila, its role can be complex as it also promotes tumour regression through cell death in different contexts [>>43<<]. In fact, MCF10A cells grown in 3D culture were also used to investigate Myc-induced transformation of human Scrib knockdown cells, and, in these experiments, luminal filling resulted from Scrib knockdown blocking Myc-induced
n2:mentions: n3:17303404

Subject Item: _:vb6088253
rdf:type: n2:Context
rdf:value: cells grown in 3D culture were also used to investigate Myc-induced transformation of human Scrib knockdown cells, and, in these experiments, luminal filling resulted from Scrib knockdown blocking Myc-induced JNK-dependent cell death [>>4<<]. While this is at odds with our Drosophila observations, that loss of Scrib promotes JNK-mediated cell death, JNK activation in scrib mutant clones was variable and possibly regulated through interactions with neighbouring wild type
n2:mentions: n3:19041750

Subject Item: _:vb6088254
rdf:type: n2:Context
rdf:value: promotes JNK-mediated cell death, JNK activation in scrib mutant clones was variable and possibly regulated through interactions with neighbouring wild type cells rather than through a cell autonomous up-regulation in JNK signalling [>>2<<,44]. Furthermore, other studies in flies have revealed that the Drosophila inhibitor of apoptosis 1 (Diap1) is upregulated in scrib mutants [45] and, thus, the loss of Scrib may potentially protect Drosophila cells from apoptosis in some
n2:mentions: n3:14592975

Subject Item: _:vb6088255
rdf:type: n2:Context
rdf:value: promotes JNK-mediated cell death, JNK activation in scrib mutant clones was variable and possibly regulated through interactions with neighbouring wild type cells rather than through a cell autonomous up-regulation in JNK signalling [2,>>44<<]. Furthermore, other studies in flies have revealed that the Drosophila inhibitor of apoptosis 1 (Diap1) is upregulated in scrib mutants [45] and, thus, the loss of Scrib may potentially protect Drosophila cells from apoptosis in some
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rdf:value: Furthermore, other studies in flies have revealed that the Drosophila inhibitor of apoptosis 1 (Diap1) is upregulated in scrib mutants [>>45<<] and, thus, the loss of Scrib may potentially protect Drosophila cells from apoptosis in some contexts.
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