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n4:pmcid
PMC0
bibo:doi
10.1186%2F1471-213X-11-57
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_:vb12702139 _:vb12702188 _:vb12702181 _:vb12702212 _:vb12702242
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dc:title
results
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We have previously reported that scrib mutant clones of tissue in the eye disc exhibit ectopic CycE expression and excessive cell proliferation, although this is restrained through JNK-dependent apoptosis [>>39<<]. If JNK signaling is blocked in scrib mutant clones by expressing a dominant negative version of Drosophila JNK, bsk (bskDN), apoptosis is prevented and mutant cells are observed to ectopically proliferate posterior to the morphogenetic
n4:mentions
n2:14592975
Subject Item
_:vb12702141
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clones by expressing a dominant negative version of Drosophila JNK, bsk (bskDN), apoptosis is prevented and mutant cells are observed to ectopically proliferate posterior to the morphogenetic furrow (MF) resulting in clonal overgrowth [>>38<<]. The ectopic cell proliferation in scrib mutant clones expressing bskDN is similar to mutants in the Hippo pathway, and therefore to determine if the Hippo pathway is impaired by the loss of scrib, we examined known targets of
n4:mentions
n2:19778415
Subject Item
_:vb12702142
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Targets examined included protein levels of DIAP1 [>>40<<], and expression of the enhancer traps for four-jointed (fj-lacZ) and ex (ex-lacZ), both of which are known to function as readouts of impaired Hippo signaling [17,41].
n4:mentions
n2:12202036
Subject Item
_:vb12702143
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Targets examined included protein levels of DIAP1 [40], and expression of the enhancer traps for four-jointed (fj-lacZ) and ex (ex-lacZ), both of which are known to function as readouts of impaired Hippo signaling [>>17<<,41].
n4:mentions
n2:16341207
Subject Item
_:vb12702144
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Targets examined included protein levels of DIAP1 [40], and expression of the enhancer traps for four-jointed (fj-lacZ) and ex (ex-lacZ), both of which are known to function as readouts of impaired Hippo signaling [17,>>41<<].
n4:mentions
n2:16980976
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As we had previously shown that scrib mutant cells continue to ectopically proliferate in the eye disc even when JNK signaling is blocked [>>38<<], it seemed likely that this impairment to Hippo signaling would not depend upon JNK activation.
n4:mentions
n2:19778415
Subject Item
_:vb12702146
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However, a previous report has highlighted the role that JNK signaling can play in promoting Hippo pathway impairment [>>37<<], and therefore we also examined the expression of the reporters in scrib mutant clones in which JNK signaling was blocked by the expression of bskDN.
n4:mentions
n2:21145886
Subject Item
_:vb12702147
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The bskDN transgene is highly effective at blocking JNK signaling since it completely abrogates both the ectopic expression of the JNK pathway reporter, msn-lacZ, and the JNK pathway target, Paxillin, in scrib mutant clones [>>38<<]. However, to confirm that JNK signaling was not required for Hippo pathway impairment, we also knocked down bsk expression in scrib mutant cells using a bskRNAi transgene.
n4:mentions
n2:19778415
Subject Item
_:vb12702148
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Removing Yki function can rescue Hippo pathway mutant overgrowth, however, Yki is also required for normal cell proliferation in the eye disc [>>7<<]. In contrast, Sd is largely dispensable for normal eye disc growth and proliferation and specifically mediates Hippo pathway mutant overgrowth [4,5]. Therefore, to determine if the ectopic cell proliferation and altered cell morphology
n4:mentions
n2:16096061
Subject Item
_:vb12702149
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In contrast, Sd is largely dispensable for normal eye disc growth and proliferation and specifically mediates Hippo pathway mutant overgrowth [>>4<<,5]. Therefore, to determine if the ectopic cell proliferation and altered cell morphology of scrib mutant cells were due to loss of Hippo pathway signaling we utilized RNAi-mediated knockdown of sd function in scrib mutant eye disc clones
n4:mentions
n2:18258486
Subject Item
_:vb12702150
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In contrast, Sd is largely dispensable for normal eye disc growth and proliferation and specifically mediates Hippo pathway mutant overgrowth [4,>>5<<]. Therefore, to determine if the ectopic cell proliferation and altered cell morphology of scrib mutant cells were due to loss of Hippo pathway signaling we utilized RNAi-mediated knockdown of sd function in scrib mutant eye disc clones
n4:mentions
n2:18258485
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are apically localized within the pseudo-stratified columnar epithelium (Figure 3A), however, in scrib mutant clones, cells are often extruded basally and Elav-positive nuclei are aberrantly localized basally within the epithelium [>>38<<]. In scrib mutants (data not shown), or in scrib mutants protected from cell death by the expression of bskDN, knockdown of sd function with sdRNAi failed to restore normal cell morphology to the mutant tissue (Figure 3B, C).
n4:mentions
n2:19778415
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Loss of lgl in the wing disc also impairs Hippo signaling, and this has been shown to depend upon JNK signaling [>>37<<]. Indeed, as in the eye disc [38,39], loss of scrib in the wing disc similarly led to the ectopic expression of the JNK pathway reporter msn-lacZ (Figure 4C).
n4:mentions
n2:21145886
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Indeed, as in the eye disc [>>38<<,39], loss of scrib in the wing disc similarly led to the ectopic expression of the JNK pathway reporter msn-lacZ (Figure 4C).
n4:mentions
n2:19778415
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_:vb12702154
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Indeed, as in the eye disc [38,>>39<<], loss of scrib in the wing disc similarly led to the ectopic expression of the JNK pathway reporter msn-lacZ (Figure 4C).
n4:mentions
n2:14592975
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scrib mutants was dependent upon impaired Hippo signaling we could not remove Sd function, as was done in the eye disc, since Sd is required for normal wing disc growth through association with the wing determination factor Vestigial [>>42<<,43]. Therefore, we utilized a yki null allele, ykiB5, to halve the gene dosage of yki in a scrib1/scrib3 mutant background.
n4:mentions
n2:9869643
Subject Item
_:vb12702156
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mutants was dependent upon impaired Hippo signaling we could not remove Sd function, as was done in the eye disc, since Sd is required for normal wing disc growth through association with the wing determination factor Vestigial [42,>>43<<]. Therefore, we utilized a yki null allele, ykiB5, to halve the gene dosage of yki in a scrib1/scrib3 mutant background.
n4:mentions
n2:9869635
Subject Item
_:vb12702157
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Consistent with previous reports [>>35<<], although giant larvae were still formed throughout an extended larval phase of development, wing disc overgrowth was dramatically reduced, resulting in significantly smaller wing discs with disorganized morphology (Figure 4F).
n4:mentions
n2:20362447
Subject Item
_:vb12702158
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Loss of scrib also promotes tumorigenesis in cooperation with oncogenic Ras signaling [reviewed in [>>28<<]]. In a "two hit" Drosophila tumorigenesis model we have previously shown that although scrib mutant eye clones die via JNK-mediated apoptosis, if RasACT or its downstream effector, Rafgof, is expressed in the mutant clones, cell death is
n4:mentions
n2:19029932
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_:vb12702159
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mutant eye clones die via JNK-mediated apoptosis, if RasACT or its downstream effector, Rafgof, is expressed in the mutant clones, cell death is prevented and massive and invasive tumors develop throughout an extended larval stage [>>39<<]. To determine if downregulation of Hippo pathway signaling is also an important mediator of these overgrowths we examined the expression of the Hippo pathway reporters, ex-lacZ and fj-lacZ, in scrib- + Rafgof tumors.
n4:mentions
n2:14592975
Subject Item
_:vb12702160
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To further confirm the importance of impaired Hippo signaling in scrib- + Rafgof tumorigenesis, we also halved the gene dosage of yki in the tumor background, and knocked down yki function within the tumor using a ykiRNAi transgene [>>5<<]. Consistent with the effects of sdRNAi, both methods of limiting yki function were unable to restore pupation or prevent tumor overgrowth throughout an extended larval phase of development, however, they significantly reduced tumor size
n4:mentions
n2:18258485
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n4:Context
rdf:value
Despite the reduction in tumor overgrowth, however, knockdown of yki was unable to prevent tumor cells from adopting an invasive morphology and appearing to move between the brain lobes, as has been previously described [>>38<<], indicating that the tumor cells retained invasive capabilities (see additional file 6).
n4:mentions
n2:19778415
Subject Item
_:vb12702162
rdf:type
n4:Context
rdf:value
in scrib mutant clones could both be rescued by the expression of a membrane-tethered, kinase-dead (dominant negative) form of aPKC (aPKCCAAXDN), although this was not sufficient to rescue the mutant cells from JNK-dependent cell death [>>38<<]. Therefore to determine if the scrib mutant defects in Hippo pathway activity were aPKC-dependent, we examined ex-lacZ and fj-lacZ in scrib mutant clones expressing aPKCCAAXDN, that were also protected from apoptosis by the coexpression
n4:mentions
n2:19778415
Subject Item
_:vb12702163
rdf:type
n4:Context
rdf:value
Consistent with the scrib mutant defects in Hippo signaling being aPKC-dependent, ectopic activation of aPKC has been shown to be sufficient to impair Hippo signaling [>>35<<,37]. This has been linked to a capacity for aPKC to activate JNK [37], however, we have previously shown that although the expression of a truncated activated allele of aPKC (aPKCΔN) in eye disc clones induces JNK-dependent cell death, if
n4:mentions
n2:20362447
Subject Item
_:vb12702164
rdf:type
n4:Context
rdf:value
Consistent with the scrib mutant defects in Hippo signaling being aPKC-dependent, ectopic activation of aPKC has been shown to be sufficient to impair Hippo signaling [35,>>37<<]. This has been linked to a capacity for aPKC to activate JNK [37], however, we have previously shown that although the expression of a truncated activated allele of aPKC (aPKCΔN) in eye disc clones induces JNK-dependent cell death, if
n4:mentions
n2:21145886
Subject Item
_:vb12702165
rdf:type
n4:Context
rdf:value
This has been linked to a capacity for aPKC to activate JNK [>>37<<], however, we have previously shown that although the expression of a truncated activated allele of aPKC (aPKCΔN) in eye disc clones induces JNK-dependent cell death, if JNK signaling is blocked, ectopic cell proliferation still ensues
n4:mentions
n2:21145886
Subject Item
_:vb12702166
rdf:type
n4:Context
rdf:value
however, we have previously shown that although the expression of a truncated activated allele of aPKC (aPKCΔN) in eye disc clones induces JNK-dependent cell death, if JNK signaling is blocked, ectopic cell proliferation still ensues [>>38<<]. Indeed, the Hippo pathway reporters fj-lacZ and ex-lacZ were ectopically expressed in aPKCΔN and bskDN coexpressing eye disc clones (Figure 7A-D).
n4:mentions
n2:19778415
Subject Item
_:vb12702167
rdf:type
n4:Context
rdf:value
aPKC-dependent impairment to Hippo signaling, we next wanted to determine if aPKC signaling was also important for promoting tissue overgrowth in core Hippo pathway mutants such as wts that are known to upregulate aPKC protein levels [>>30<<,31]. However, although wtsRNAi-expressing clones ectopically express CycE and DIAP1 and overgrow (Figure 8A, B), similar to scrib mutant clones, coexpression of aPKCCAAXDN in wtsRNAi clones neither prevented the ectopic expression of CycE
n4:mentions
n2:19531584
Subject Item
_:vb12702168
rdf:type
n4:Context
rdf:value
impairment to Hippo signaling, we next wanted to determine if aPKC signaling was also important for promoting tissue overgrowth in core Hippo pathway mutants such as wts that are known to upregulate aPKC protein levels [30,>>31<<]. However, although wtsRNAi-expressing clones ectopically express CycE and DIAP1 and overgrow (Figure 8A, B), similar to scrib mutant clones, coexpression of aPKCCAAXDN in wtsRNAi clones neither prevented the ectopic expression of CycE or
n4:mentions
n2:19531586
Subject Item
_:vb12702169
rdf:type
n4:Context
rdf:value
Upstream regulation of Hippo signaling occurs through at least two transmembrane proteins, Ft and Crb [reviewed in [>>44<<]]. Crb acts through Ex [32-34], and Ft acts through both Ex [14,15,19] and the unconventional myosin Dachs (D) [12]. The overexpression of ex can promote ectopic Hippo signaling, and when expressed in otherwise wild type clones can
n4:mentions
n2:20619814
Subject Item
_:vb12702170
rdf:type
n4:Context
rdf:value
Crb acts through Ex [>>32<<-34], and Ft acts through both Ex [14,15,19] and the unconventional myosin Dachs (D) [12].
n4:mentions
n2:20798049 n2:20362445 n2:20498073
Subject Item
_:vb12702171
rdf:type
n4:Context
rdf:value
Crb acts through Ex [32-34], and Ft acts through both Ex [>>14<<,15,19] and the unconventional myosin Dachs (D) [12].
n4:mentions
n2:16996265
Subject Item
_:vb12702172
rdf:type
n4:Context
rdf:value
Crb acts through Ex [32-34], and Ft acts through both Ex [14,>>15<<,19] and the unconventional myosin Dachs (D) [12].
n4:mentions
n2:16996266
Subject Item
_:vb12702173
rdf:type
n4:Context
rdf:value
Crb acts through Ex [32-34], and Ft acts through both Ex [14,15,>>19<<] and the unconventional myosin Dachs (D) [12].
n4:mentions
n2:17045801
Subject Item
_:vb12702174
rdf:type
n4:Context
rdf:value
Crb acts through Ex [32-34], and Ft acts through both Ex [14,15,19] and the unconventional myosin Dachs (D) [>>12<<]. The overexpression of ex can promote ectopic Hippo signaling, and when expressed in otherwise wild type clones can restrain clonal growth in a Hpo and Wts-dependent manner [13,17,41]. Significantly, the overexpression of ex in scrib
n4:mentions
n2:16735478
Subject Item
_:vb12702175
rdf:type
n4:Context
rdf:value
The overexpression of ex can promote ectopic Hippo signaling, and when expressed in otherwise wild type clones can restrain clonal growth in a Hpo and Wts-dependent manner [>>13<<,17,41]. Significantly, the overexpression of ex in scrib mutant eye disc clones also restrained clonal growth, and this was the case even if JNK-mediated apoptosis of the clonal tissue was prevented (Figure 9A-F). Indeed, scrib clones
n4:mentions
n2:17359963
Subject Item
_:vb12702176
rdf:type
n4:Context
rdf:value
The overexpression of ex can promote ectopic Hippo signaling, and when expressed in otherwise wild type clones can restrain clonal growth in a Hpo and Wts-dependent manner [13,>>17<<,41]. Significantly, the overexpression of ex in scrib mutant eye disc clones also restrained clonal growth, and this was the case even if JNK-mediated apoptosis of the clonal tissue was prevented (Figure 9A-F). Indeed, scrib clones
n4:mentions
n2:16341207
Subject Item
_:vb12702177
rdf:type
n4:Context
rdf:value
The overexpression of ex can promote ectopic Hippo signaling, and when expressed in otherwise wild type clones can restrain clonal growth in a Hpo and Wts-dependent manner [13,17,>>41<<]. Significantly, the overexpression of ex in scrib mutant eye disc clones also restrained clonal growth, and this was the case even if JNK-mediated apoptosis of the clonal tissue was prevented (Figure 9A-F). Indeed, scrib clones
n4:mentions
n2:16980976
Subject Item
_:vb12702178
rdf:type
n4:Context
rdf:value
Ft can act in parallel to Ex, and indeed the overexpression of ex in ft mutant clones is unable to restrain ft mutant tissue overgrowth [>>16<<]. We therefore next determined if impaired Ft signaling could be responsible for downregulating the Hippo pathway in scrib mutants.
n4:mentions
n2:18077345
Subject Item
_:vb12702179
rdf:type
n4:Context
rdf:value
To do this we generated scrib clones in a d mutant animal background, since Ft acts through inhibiting D and loss of d fully abrogates ft mutant overgrowth [>>12<<] (see Figure 10).
n4:mentions
n2:16735478
Subject Item
_:vb12702180
rdf:type
n4:Context
rdf:value
In the wing disc, loss of scrib is also associated with JNK activation, and although JNK signaling has been shown to be sufficient to downregulate the Hippo pathway in the wing disc [>>37<<], Hippo pathway signaling remains impaired in scrib mutants even when JNK is blocked.
n4:mentions
n2:21145886
Subject Item
_:vb12702181
rdf:type
n5:Section
dc:title
conclusions
n5:contains
_:vb12702182 _:vb12702183 _:vb12702184 _:vb12702185 _:vb12702186 _:vb12702187
Subject Item
_:vb12702182
rdf:type
n4:Context
rdf:value
Loss of Hippo pathway signaling or ectopic activation of the Yki or Sd homologues, (YAP and TEAD family proteins respectively), is emerging as a powerful oncogenic force [reviewed in [>>21<<]]. Similarly, the mammalian Scrib module is increasingly implicated in tumorigenesis [reviewed in [28]], and mammalian Scrib can restrain tissue transformation by oncogenic Ras [61], and Myc [62]. Mammalian Scrib also functions within
n4:mentions
n2:18955139
Subject Item
_:vb12702183
rdf:type
n4:Context
rdf:value
Similarly, the mammalian Scrib module is increasingly implicated in tumorigenesis [reviewed in [>>28<<]], and mammalian Scrib can restrain tissue transformation by oncogenic Ras [61], and Myc [62].
n4:mentions
n2:19029932
Subject Item
_:vb12702184
rdf:type
n4:Context
rdf:value
Similarly, the mammalian Scrib module is increasingly implicated in tumorigenesis [reviewed in [28]], and mammalian Scrib can restrain tissue transformation by oncogenic Ras [>>61<<], and Myc [62].
n4:mentions
n2:18641685
Subject Item
_:vb12702185
rdf:type
n4:Context
rdf:value
Similarly, the mammalian Scrib module is increasingly implicated in tumorigenesis [reviewed in [28]], and mammalian Scrib can restrain tissue transformation by oncogenic Ras [61], and Myc [>>62<<]. Mammalian Scrib also functions within planar cell polarity pathways [63], and although links with the Hippo pathway have not yet been described, the connection is likely to be conserved since studies in the zebrafish indicate that
n4:mentions
n2:19041750
Subject Item
_:vb12702186
rdf:type
n4:Context
rdf:value
Mammalian Scrib also functions within planar cell polarity pathways [>>63<<], and although links with the Hippo pathway have not yet been described, the connection is likely to be conserved since studies in the zebrafish indicate that zScrib binds to zFat1 and also promotes Hippo pathway activation [44].
n4:mentions
n2:12724779
Subject Item
_:vb12702187
rdf:type
n4:Context
rdf:value
[63], and although links with the Hippo pathway have not yet been described, the connection is likely to be conserved since studies in the zebrafish indicate that zScrib binds to zFat1 and also promotes Hippo pathway activation [>>44<<]. The uniting of these two powerful tumor suppressor pathways clearly has important implications for human carcinogenesis.
n4:mentions
n2:20619814
Subject Item
_:vb12702188
rdf:type
n5:Section
dc:title
methods
n5:contains
_:vb12702196 _:vb12702197 _:vb12702198 _:vb12702199 _:vb12702192 _:vb12702193 _:vb12702194 _:vb12702195 _:vb12702204 _:vb12702205 _:vb12702206 _:vb12702207 _:vb12702200 _:vb12702201 _:vb12702202 _:vb12702203 _:vb12702189 _:vb12702190 _:vb12702191 _:vb12702208 _:vb12702209 _:vb12702210 _:vb12702211
Subject Item
_:vb12702189
rdf:type
n4:Context
rdf:value
The following Drosophila stocks were used: ey-FLP1, UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [>>64<<]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except
n4:mentions
n2:11290291
Subject Item
_:vb12702190
rdf:type
n4:Context
rdf:value
The following Drosophila stocks were used: ey-FLP1, UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [64]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [>>26<<]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71];
n4:mentions
n2:12629552
Subject Item
_:vb12702191
rdf:type
n4:Context
rdf:value
The following Drosophila stocks were used: ey-FLP1, UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [64]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [>>66<<]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A
n4:mentions
n2:8946916
Subject Item
_:vb12702192
rdf:type
n4:Context
rdf:value
The following Drosophila stocks were used: ey-FLP1, UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [64]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [>>67<<]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof
n4:mentions
n2:10022918
Subject Item
_:vb12702193
rdf:type
n4:Context
rdf:value
were used: ey-FLP1, UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [64]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [>>68<<]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74];
n4:mentions
n2:11566858
Subject Item
_:vb12702194
rdf:type
n4:Context
rdf:value
ey-FLP1, UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [64]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [>>12<<]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12
n4:mentions
n2:16735478
Subject Item
_:vb12702195
rdf:type
n4:Context
rdf:value
UAS-mCD8-GFP;;Tub-GAL4, FRT82B, Tub-GAL80 [64]; y, w, hs-FLP; FRT82B, Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [>>69<<]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75];
n4:mentions
n2:8269855
Subject Item
_:vb12702196
rdf:type
n4:Context
rdf:value
Ubi-GFP; en-GAL4; UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [>>69<<]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC
n4:mentions
n2:8269855
Subject Item
_:vb12702197
rdf:type
n4:Context
rdf:value
UAS-DaPKCΔN [26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [>>70<<]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424);
n4:mentions
n2:9144921
Subject Item
_:vb12702198
rdf:type
n4:Context
rdf:value
[26]; UAS-DaPKCCAAXDN [65]; UAS-aPKCRNAi (VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [>>71<<]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG
n4:mentions
n2:3417051
Subject Item
_:vb12702199
rdf:type
n4:Context
rdf:value
(VDRC #2907); bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [>>72<<]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG
n4:mentions
n2:8606003
Subject Item
_:vb12702200
rdf:type
n4:Context
rdf:value
bsk2 [66]; UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [>>26<<]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1);
n4:mentions
n2:12629552
Subject Item
_:vb12702201
rdf:type
n4:Context
rdf:value
UAS-bskDN [67]; UAS-bskRNAi (NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [>>26<<]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7];
n4:mentions
n2:12629552
Subject Item
_:vb12702202
rdf:type
n4:Context
rdf:value
(NIG #5680R-1); d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [>>73<<]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:7926754
Subject Item
_:vb12702203
rdf:type
n4:Context
rdf:value
d1 [68]; dGC13 [12]; exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [>>74<<]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:1934068
Subject Item
_:vb12702204
rdf:type
n4:Context
rdf:value
exe1 [69]; ex697 (ex-lacZ in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [>>75<<]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:9389658
Subject Item
_:vb12702205
rdf:type
n4:Context
rdf:value
in all figures except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [>>38<<]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:19778415
Subject Item
_:vb12702206
rdf:type
n4:Context
rdf:value
except Figure 9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [>>76<<]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:10688207
Subject Item
_:vb12702207
rdf:type
n4:Context
rdf:value
9J which is exe1) [69]; UAS-ex [70]; ftfd [71]; UAS-ftRNAi (VDRC #9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [>>77<<]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:15611336
Subject Item
_:vb12702208
rdf:type
n4:Context
rdf:value
#9396); fj-lacZ [72]; UAS-lgl5.1 [26]; UAS-lgl3A [26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [>>78<<]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [5].
n4:mentions
n2:7743921
Subject Item
_:vb12702209
rdf:type
n4:Context
rdf:value
[26]; UAS-phlgof (UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [>>7<<]; UAS-ykiRNAi [5].
n4:mentions
n2:16096061
Subject Item
_:vb12702210
rdf:type
n4:Context
rdf:value
(UAS-Rafgof) [73]; Ras85De1b [74]; UAS-dRas1V12 [75]; UAS-scrib19.2 [38]; FRT82B, scrib1 [76]; scrib3 [77]; UAS-scribRNAi (VDRC #27424); UAS-sdRNAi (NIG #8544R-2); wtsX1 [78]; UAS-wtsRNAi (NIG #12072R-1); ykiB5 [7]; UAS-ykiRNAi [>>5<<].
n4:mentions
n2:18258485
Subject Item
_:vb12702211
rdf:type
n4:Context
rdf:value
Clonal analysis utilized either MARCM (mosaic analysis with repressible cell marker) [>>79<<] with FRT82B and ey-FLP1 to induce clones and mCD8-GFP expression to mark mutant tissue, or for negatively marked scrib1 clones, hs-FLP with FRT82B, Ubi-GFP.
n4:mentions
n2:10197526
Subject Item
_:vb12702212
rdf:type
n5:Section
dc:title
discussion
n5:contains
_:vb12702240 _:vb12702241 _:vb12702228 _:vb12702229 _:vb12702230 _:vb12702231 _:vb12702224 _:vb12702225 _:vb12702226 _:vb12702227 _:vb12702236 _:vb12702237 _:vb12702238 _:vb12702239 _:vb12702232 _:vb12702233 _:vb12702234 _:vb12702235 _:vb12702213 _:vb12702214 _:vb12702215 _:vb12702220 _:vb12702221 _:vb12702222 _:vb12702223 _:vb12702216 _:vb12702217 _:vb12702218 _:vb12702219
Subject Item
_:vb12702213
rdf:type
n4:Context
rdf:value
Indeed, in zebrafish Scrib binds to, and functionally cooperates with, Ft [>>45<<]; and as this interaction could be conserved in Drosophila [45] very direct points of intersection between Scrib and the upstream regulators of Hippo signaling can be envisaged.
n4:mentions
n2:19439659
Subject Item
_:vb12702214
rdf:type
n4:Context
rdf:value
Indeed, in zebrafish Scrib binds to, and functionally cooperates with, Ft [45]; and as this interaction could be conserved in Drosophila [>>45<<] very direct points of intersection between Scrib and the upstream regulators of Hippo signaling can be envisaged.
n4:mentions
n2:19439659
Subject Item
_:vb12702215
rdf:type
n4:Context
rdf:value
However, this interpretation is complicated by Ex's capacity to directly bind to, and sequester Yki activity [>>46<<]. In fact, even ex overexpression was not sufficient to block ectopic CycE expression in scrib mutant clones, and epistasis experiments confirm that the deregulated Hippo signaling in scrib mutants is at least partially epistatic to both
n4:mentions
n2:19289086
Subject Item
_:vb12702216
rdf:type
n4:Context
rdf:value
Indeed, a number of recent reports indicating that increased levels of F-actin are sufficient to inhibit Hippo pathway activity [>>47<<,48] are also suggestive, since the aPKC-dependent loss of apico-basal cell polarity in scrib mutants is often associated with F-actin accumulations (data not shown).
n4:mentions
n2:21556047
Subject Item
_:vb12702217
rdf:type
n4:Context
rdf:value
Indeed, a number of recent reports indicating that increased levels of F-actin are sufficient to inhibit Hippo pathway activity [47,>>48<<] are also suggestive, since the aPKC-dependent loss of apico-basal cell polarity in scrib mutants is often associated with F-actin accumulations (data not shown).
n4:mentions
n2:21525075
Subject Item
_:vb12702218
rdf:type
n4:Context
rdf:value
however, lgl mutant eye disc clones are not associated with the severe alterations in cell morphology characteristic of scrib mutant cells, yet the impairment to Hippo signaling in lgl mutant clones is also dependent upon aPKC activity [>>35<<]. Furthermore, Ex and Ft localization were unaffected in the absence of lgl, whilst both Hpo and Ras-associated domain family protein (RASSF) were co-mislocalized, consistent with the Hippo pathway being impaired downstream of Ft and Ex
n4:mentions
n2:20362447
Subject Item
_:vb12702219
rdf:type
n4:Context
rdf:value
Furthermore, Ex and Ft localization were unaffected in the absence of lgl, whilst both Hpo and Ras-associated domain family protein (RASSF) were co-mislocalized, consistent with the Hippo pathway being impaired downstream of Ft and Ex [>>35<<]. Whether a common aPKC-dependent mechanism of Hippo pathway inhibition is operative in both lgl and scrib mutant eye discs will require further investigation.
n4:mentions
n2:20362447
Subject Item
_:vb12702220
rdf:type
n4:Context
rdf:value
A recent report indicates that the impaired Hippo pathway signaling in lgl mutant wing discs is dependent upon JNK signaling, and that ectopic aPKC signaling in the wing disc also acts through JNK to promote Yki activity [>>37<<]. In the eye disc, however, we show that Hippo pathway impairment in scrib mutants, as well as upon ectopic activation of aPKC signaling, cannot be rescued by blocking JNK, and this is also likely to be the case for lgl mutants [49].
n4:mentions
n2:21145886
Subject Item
_:vb12702221
rdf:type
n4:Context
rdf:value
In the eye disc, however, we show that Hippo pathway impairment in scrib mutants, as well as upon ectopic activation of aPKC signaling, cannot be rescued by blocking JNK, and this is also likely to be the case for lgl mutants [>>49<<]. Furthermore, in scrib mutant wing discs, although we demonstrate that JNK signaling is ectopically activated upon scrib knockdown, we also show that Hippo pathway impairment in the wing occurs, at least in part, even when JNK signaling
n4:mentions
n2:20724829
Subject Item
_:vb12702222
rdf:type
n4:Context
rdf:value
In lgl mutants, the cell polarity defects in the wing disc are also JNK-dependent [>>37<<], and while we did not analyze cell polarity markers upon scrib knockdown in the wing, one possibility is that loss of scrib is associated with JNK-independent cell polarity defects that impact upon the Hippo pathway in a similar manner
n4:mentions
n2:21145886
Subject Item
_:vb12702223
rdf:type
n4:Context
rdf:value
Certainly loss of scrib is notable for eliciting much stronger cell morphology defects in the eye disc than loss of lgl [>>39<<,50]. As the cell morphology defects in scrib mutant eye disc clones are aPKC-dependent [38], it will be important to determine what role aPKC signaling plays in the scrib mutant wing disc phenotypes.
n4:mentions
n2:14592975
Subject Item
_:vb12702224
rdf:type
n4:Context
rdf:value
Certainly loss of scrib is notable for eliciting much stronger cell morphology defects in the eye disc than loss of lgl [39,>>50<<]. As the cell morphology defects in scrib mutant eye disc clones are aPKC-dependent [38], it will be important to determine what role aPKC signaling plays in the scrib mutant wing disc phenotypes.
n4:mentions
n2:17870065
Subject Item
_:vb12702225
rdf:type
n4:Context
rdf:value
As the cell morphology defects in scrib mutant eye disc clones are aPKC-dependent [>>38<<], it will be important to determine what role aPKC signaling plays in the scrib mutant wing disc phenotypes.
n4:mentions
n2:19778415
Subject Item
_:vb12702226
rdf:type
n4:Context
rdf:value
Indeed, the role of JNK in neoplasia is proving to be complex, since in some contexts it functions as an oncogene to promote neoplasia, whilst in different contexts it acts as a tumor suppressor through the induction of apoptosis [>>38<<,39,51-53]. Its effects upon Hippo pathway regulation may therefore also be context dependent.
n4:mentions
n2:19778415
Subject Item
_:vb12702227
rdf:type
n4:Context
rdf:value
Indeed, the role of JNK in neoplasia is proving to be complex, since in some contexts it functions as an oncogene to promote neoplasia, whilst in different contexts it acts as a tumor suppressor through the induction of apoptosis [38,>>39<<,51-53]. Its effects upon Hippo pathway regulation may therefore also be context dependent.
n4:mentions
n2:14592975
Subject Item
_:vb12702228
rdf:type
n4:Context
rdf:value
Indeed, the role of JNK in neoplasia is proving to be complex, since in some contexts it functions as an oncogene to promote neoplasia, whilst in different contexts it acts as a tumor suppressor through the induction of apoptosis [38,39,>>51<<-53]. Its effects upon Hippo pathway regulation may therefore also be context dependent.
n4:mentions
n2:17082773 n2:16753569 n2:21368274
Subject Item
_:vb12702229
rdf:type
n4:Context
rdf:value
effects, with increased expression in wild type cells surrounding mutant clones, were also sometimes observed (data not shown), consistent with the involvement of the Hippo pathway in regenerative proliferation around dying tissue [>>54<<]. Much of this variability appeared to be reduced when JNK signaling was blocked in the mutant clones, and whilst the rescue of some non-cell autonomous effects would be consistent with the role that JNK can play in promoting non-cell
n4:mentions
n2:21111727
Subject Item
_:vb12702230
rdf:type
n4:Context
rdf:value
signaling was blocked in the mutant clones, and whilst the rescue of some non-cell autonomous effects would be consistent with the role that JNK can play in promoting non-cell autonomous compensatory proliferation through Yki activity [>>37<<], it also remains possible that JNK may act in opposite ways to downregulate DIAP1 or other reporters in mutant cells destined to die.
n4:mentions
n2:21145886
Subject Item
_:vb12702231
rdf:type
n4:Context
rdf:value
Loss of wts is capable of eliciting neoplastic overgrowth [>>55<<], and Hippo pathway mutants induce apical hypertrophy with increased levels of apical cell determinants such as Crb, aPKC [30,31], and F-actin [48], although, interestingly, despite the potential for increased Crb, aPKC and F-actin to
n4:mentions
n2:17947427
Subject Item
_:vb12702232
rdf:type
n4:Context
rdf:value
Loss of wts is capable of eliciting neoplastic overgrowth [55], and Hippo pathway mutants induce apical hypertrophy with increased levels of apical cell determinants such as Crb, aPKC [>>30<<,31], and F-actin [48], although, interestingly, despite the potential for increased Crb, aPKC and F-actin to promote tissue hyperplasia, the overgrowth in Hippo pathway mutants is independent of the apical hypertrophy [30,31].
n4:mentions
n2:19531584
Subject Item
_:vb12702233
rdf:type
n4:Context
rdf:value
Loss of wts is capable of eliciting neoplastic overgrowth [55], and Hippo pathway mutants induce apical hypertrophy with increased levels of apical cell determinants such as Crb, aPKC [30,>>31<<], and F-actin [48], although, interestingly, despite the potential for increased Crb, aPKC and F-actin to promote tissue hyperplasia, the overgrowth in Hippo pathway mutants is independent of the apical hypertrophy [30,31].
n4:mentions
n2:19531586
Subject Item
_:vb12702234
rdf:type
n4:Context
rdf:value
Loss of wts is capable of eliciting neoplastic overgrowth [55], and Hippo pathway mutants induce apical hypertrophy with increased levels of apical cell determinants such as Crb, aPKC [30,31], and F-actin [>>48<<], although, interestingly, despite the potential for increased Crb, aPKC and F-actin to promote tissue hyperplasia, the overgrowth in Hippo pathway mutants is independent of the apical hypertrophy [30,31].
n4:mentions
n2:21525075
Subject Item
_:vb12702235
rdf:type
n4:Context
rdf:value
as Crb, aPKC [30,31], and F-actin [48], although, interestingly, despite the potential for increased Crb, aPKC and F-actin to promote tissue hyperplasia, the overgrowth in Hippo pathway mutants is independent of the apical hypertrophy [>>30<<,31]. This is consistent with our own work demonstrating that wts and ft mutant tissue overgrowth was independent of aPKC signaling.
n4:mentions
n2:19531584
Subject Item
_:vb12702236
rdf:type
n4:Context
rdf:value
Crb, aPKC [30,31], and F-actin [48], although, interestingly, despite the potential for increased Crb, aPKC and F-actin to promote tissue hyperplasia, the overgrowth in Hippo pathway mutants is independent of the apical hypertrophy [30,>>31<<]. This is consistent with our own work demonstrating that wts and ft mutant tissue overgrowth was independent of aPKC signaling.
n4:mentions
n2:19531586
Subject Item
_:vb12702237
rdf:type
n4:Context
rdf:value
cell polarity pathways is not confined to traditional apico-basal epithelial polarity regulators such as Scrib and aPKC, since the Hippo pathway components ft, fj and ds also participate in planar cell polarity pathways [reviewed in [>>56<<]], as do scrib [57], lgl [58] and aPKC [59,60].
n4:mentions
n2:17230199
Subject Item
_:vb12702238
rdf:type
n4:Context
rdf:value
is not confined to traditional apico-basal epithelial polarity regulators such as Scrib and aPKC, since the Hippo pathway components ft, fj and ds also participate in planar cell polarity pathways [reviewed in [56]], as do scrib [>>57<<], lgl [58] and aPKC [59,60].
n4:mentions
n2:19563796
Subject Item
_:vb12702239
rdf:type
n4:Context
rdf:value
not confined to traditional apico-basal epithelial polarity regulators such as Scrib and aPKC, since the Hippo pathway components ft, fj and ds also participate in planar cell polarity pathways [reviewed in [56]], as do scrib [57], lgl [>>58<<] and aPKC [59,60].
n4:mentions
n2:16251968
Subject Item
_:vb12702240
rdf:type
n4:Context
rdf:value
traditional apico-basal epithelial polarity regulators such as Scrib and aPKC, since the Hippo pathway components ft, fj and ds also participate in planar cell polarity pathways [reviewed in [56]], as do scrib [57], lgl [58] and aPKC [>>59<<,60]. The emerging picture is therefore one of a complex network of interactions whereby multiple components regulating cellular architecture are employed by cells to read their position within a morphogenetic field and respond with
n4:mentions
n2:15907474
Subject Item
_:vb12702241
rdf:type
n4:Context
rdf:value
apico-basal epithelial polarity regulators such as Scrib and aPKC, since the Hippo pathway components ft, fj and ds also participate in planar cell polarity pathways [reviewed in [56]], as do scrib [57], lgl [58] and aPKC [59,>>60<<]. The emerging picture is therefore one of a complex network of interactions whereby multiple components regulating cellular architecture are employed by cells to read their position within a morphogenetic field and respond with
n4:mentions
n2:17488624
Subject Item
_:vb12702242
rdf:type
n5:Section
dc:title
background
n5:contains
_:vb12702252 _:vb12702253 _:vb12702254 _:vb12702255 _:vb12702248 _:vb12702249 _:vb12702250 _:vb12702251 _:vb12702244 _:vb12702245 _:vb12702246 _:vb12702247 _:vb12702243 _:vb12702264 _:vb12702265 _:vb12702260 _:vb12702261 _:vb12702262 _:vb12702263 _:vb12702256 _:vb12702257 _:vb12702258 _:vb12702259
Subject Item
_:vb12702243
rdf:type
n4:Context
rdf:value
Drosophila has long been recognized as an important model organism for elucidating oncogenic and tumor suppressor pathways [reviewed in [>>1<<]]. Traditionally two distinct classes of tumor suppressor mutants have been described, the loss of which cause either hyperplastic or neoplastic overgrowth [reviewed in [2]].
n4:mentions
n2:16034367
Subject Item
_:vb12702244
rdf:type
n4:Context
rdf:value
Traditionally two distinct classes of tumor suppressor mutants have been described, the loss of which cause either hyperplastic or neoplastic overgrowth [reviewed in [>>2<<]]. Hyperplastic overgrowth is characterized by excessive cell proliferation that is eventually restrained by terminal differentiation, while neoplastic overgrowth exhibits impaired differentiation, defects in cell polarity and the
n4:mentions
n2:16872256
Subject Item
_:vb12702245
rdf:type
n4:Context
rdf:value
Over recent years, a large number of hyperplastic tumor suppressor mutants have been united into a single pathway, the Hippo pathway [reviewed in [>>3<<]]. Core components of the Hippo pathway include the serine-threonine kinases Hippo (Hpo) and Warts (Wts), and their adaptor proteins, Salvador (Sav) and Mob-As-Tumor-Suppressor (Mats).
n4:mentions
n2:17318211
Subject Item
_:vb12702246
rdf:type
n4:Context
rdf:value
the nucleus where it binds to its DNA binding partner, Scalloped (Sd), and promotes expression of proteins involved in cell proliferation (Cyclin E; CycE), cell growth (Myc) and cell survival (Drosophila Inhibitor of Apoptosis 1; DIAP1) [>>4<<-11]. It is the dual role of the Hippo pathway in regulating both cell proliferation and survival functions that makes its loss such a potent driver of tissue overgrowth.
n4:mentions
n2:18258485 n2:18258486 n2:16096061 n2:18313299 n2:18256197 n2:17889654 n2:20951343
Subject Item
_:vb12702247
rdf:type
n4:Context
rdf:value
The pathway is regulated through input from upstream components including Merlin and Expanded (Ex), and the transmembrane proteins Fat (Ft) and Dachsous (Ds) [>>12<<-19]. It is proposed that the primary function of the Hippo pathway is to incorporate positional cues within an epithelial field to dictate the ultimate size of organ development [20]. The pathway is highly conserved and also functions to
n4:mentions
n2:17258190 n2:18077345 n2:16341207 n2:16996265 n2:16996266 n2:17045801 n2:17359963 n2:16735478
Subject Item
_:vb12702248
rdf:type
n4:Context
rdf:value
It is proposed that the primary function of the Hippo pathway is to incorporate positional cues within an epithelial field to dictate the ultimate size of organ development [>>20<<]. The pathway is highly conserved and also functions to restrain organ size in mammals. Furthermore, increasing evidence links Hippo pathway deregulation to tumorigenesis [reviewed in [21]].
n4:mentions
n2:18694569
Subject Item
_:vb12702249
rdf:type
n4:Context
rdf:value
Furthermore, increasing evidence links Hippo pathway deregulation to tumorigenesis [reviewed in [>>21<<]].
n4:mentions
n2:18955139
Subject Item
_:vb12702250
rdf:type
n4:Context
rdf:value
genes that regulate cell polarity, scrib, discs large (dlg) and lethal giant larvae (lgl), as well as mutants within the endocytic pathway including avalanche (avl), tumor suppressor protein 101 (TSG101) and Rab5 [reviewed in [>>22<<]]. The loss of apico-basal cell polarity and overgrowth phenotypes of a number of these mutants, including scrib, lgl, avl and TSG101 are dependent upon atypical protein kinase C (aPKC) activity, since mutant phenotypes can be rescued by
n4:mentions
n2:19560990
Subject Item
_:vb12702251
rdf:type
n4:Context
rdf:value
phenotypes of a number of these mutants, including scrib, lgl, avl and TSG101 are dependent upon atypical protein kinase C (aPKC) activity, since mutant phenotypes can be rescued by reducing atypical protein kinase C function [>>23<<-25]. Direct and mutual antagonism between the junctional tumor suppressors and aPKC has been demonstrated by the ability of aPKC to associate with and phosphorylate Lgl, thereby releasing Lgl from the cell cortex and thus potentially
n4:mentions
n2:14657233 n2:19855819 n2:16258546
Subject Item
_:vb12702252
rdf:type
n4:Context
rdf:value
antagonism between the junctional tumor suppressors and aPKC has been demonstrated by the ability of aPKC to associate with and phosphorylate Lgl, thereby releasing Lgl from the cell cortex and thus potentially inhibiting Lgl function [>>26<<], and the ability of Lgl to inhibit aPKC-dependent phosphorylation of other key targets [27].
n4:mentions
n2:12629552
Subject Item
_:vb12702253
rdf:type
n4:Context
rdf:value
of aPKC to associate with and phosphorylate Lgl, thereby releasing Lgl from the cell cortex and thus potentially inhibiting Lgl function [26], and the ability of Lgl to inhibit aPKC-dependent phosphorylation of other key targets [>>27<<]. Like the Hippo pathway, mammalian homologues of the Drosophila neoplastic tumor suppressors, as well as aPKC, are increasingly implicated as important players in human cancers [reviewed in [28]].
n4:mentions
n2:18854163
Subject Item
_:vb12702254
rdf:type
n4:Context
rdf:value
Like the Hippo pathway, mammalian homologues of the Drosophila neoplastic tumor suppressors, as well as aPKC, are increasingly implicated as important players in human cancers [reviewed in [>>28<<]].
n4:mentions
n2:19029932
Subject Item
_:vb12702255
rdf:type
n4:Context
rdf:value
Indeed, wts mutants were originally identified based upon mutant cell morphology [>>29<<], and this is now known to be a phenotype associated with other Hippo pathway mutants and due to Yki-dependent upregulation of the apical cell polarity determinant Crumbs (Crb) and apical hypertrophy [30,31].
n4:mentions
n2:7698644
Subject Item
_:vb12702256
rdf:type
n4:Context
rdf:value
upon mutant cell morphology [29], and this is now known to be a phenotype associated with other Hippo pathway mutants and due to Yki-dependent upregulation of the apical cell polarity determinant Crumbs (Crb) and apical hypertrophy [>>30<<,31]. Crb itself acts to regulate Hippo signaling by binding to Ex [32], and either excessive Crb activity or loss of Crb results in deregulation of Ex and an impairment to Hippo signaling resulting in tissue overgrowth [32-35].
n4:mentions
n2:19531584
Subject Item
_:vb12702257
rdf:type
n4:Context
rdf:value
upon mutant cell morphology [29], and this is now known to be a phenotype associated with other Hippo pathway mutants and due to Yki-dependent upregulation of the apical cell polarity determinant Crumbs (Crb) and apical hypertrophy [30,>>31<<]. Crb itself acts to regulate Hippo signaling by binding to Ex [32], and either excessive Crb activity or loss of Crb results in deregulation of Ex and an impairment to Hippo signaling resulting in tissue overgrowth [32-35].
n4:mentions
n2:19531586
Subject Item
_:vb12702258
rdf:type
n4:Context
rdf:value
Crb itself acts to regulate Hippo signaling by binding to Ex [>>32<<], and either excessive Crb activity or loss of Crb results in deregulation of Ex and an impairment to Hippo signaling resulting in tissue overgrowth [32-35].
n4:mentions
n2:20498073
Subject Item
_:vb12702259
rdf:type
n4:Context
rdf:value
Crb itself acts to regulate Hippo signaling by binding to Ex [32], and either excessive Crb activity or loss of Crb results in deregulation of Ex and an impairment to Hippo signaling resulting in tissue overgrowth [>>32<<-35]. The neoplastic tumor suppressors scrib, dlg and lgl also interact with the Hippo pathway. dlg, lgl or scrib mutant follicle cells surrounding the female ovary have elevated levels of Yki targets Cyclin E and DIAP1, and exhibit strong
n4:mentions
n2:20362447 n2:20362445 n2:20498073 n2:20798049
Subject Item
_:vb12702260
rdf:type
n4:Context
rdf:value
dlg, lgl or scrib mutant follicle cells surrounding the female ovary have elevated levels of Yki targets Cyclin E and DIAP1, and exhibit strong genetic interactions with wts [>>36<<]. Furthermore, loss of lgl has been shown to impair Hippo signaling in the eye disc in an aPKC signaling-dependent manner [35]. Indeed in the wing disc both the loss of lgl or ectopic aPKC activity promotes Yki activity through an
n4:mentions
n2:18430928
Subject Item
_:vb12702261
rdf:type
n4:Context
rdf:value
Furthermore, loss of lgl has been shown to impair Hippo signaling in the eye disc in an aPKC signaling-dependent manner [>>35<<]. Indeed in the wing disc both the loss of lgl or ectopic aPKC activity promotes Yki activity through an upregulation of Jun N-terminal kinase (JNK) signaling [37]. Whether scrib regulates the Hippo pathway in the eye or wing disc is not
n4:mentions
n2:20362447
Subject Item
_:vb12702262
rdf:type
n4:Context
rdf:value
Indeed in the wing disc both the loss of lgl or ectopic aPKC activity promotes Yki activity through an upregulation of Jun N-terminal kinase (JNK) signaling [>>37<<]. Whether scrib regulates the Hippo pathway in the eye or wing disc is not yet clear. In contrast to lgl, reduced levels of Scrib in the eye disc to levels where apico-basal cell polarity is only mildly affected does not impair Hippo
n4:mentions
n2:21145886
Subject Item
_:vb12702263
rdf:type
n4:Context
rdf:value
In contrast to lgl, reduced levels of Scrib in the eye disc to levels where apico-basal cell polarity is only mildly affected does not impair Hippo signaling [>>35<<], although prior studies with null alleles have demonstrated both aPKC-dependent proliferation as well as ectopic JNK signaling in scrib mutant eye disc clones [38].
n4:mentions
n2:20362447
Subject Item
_:vb12702264
rdf:type
n4:Context
rdf:value
cell polarity is only mildly affected does not impair Hippo signaling [35], although prior studies with null alleles have demonstrated both aPKC-dependent proliferation as well as ectopic JNK signaling in scrib mutant eye disc clones [>>38<<]. Furthermore, whilst homozygous scrib mutant wing disc overgrowth is reduced in response to limiting Yki levels [35], it has not been determined if Hippo signaling is impaired in this tissue and whether the genetic interaction with yki
n4:mentions
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Furthermore, whilst homozygous scrib mutant wing disc overgrowth is reduced in response to limiting Yki levels [>>35<<], it has not been determined if Hippo signaling is impaired in this tissue and whether the genetic interaction with yki reflects a general sensitivity of scrib mutant tissue to limiting levels of survival/proliferation functions.
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