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_:vb591261832 _:vb591261833 _:vb591261834 _:vb591261835 _:vb591261836 _:vb591261837 _:vb591261838 _:vb591261839 _:vb591261824 _:vb591261825 _:vb591261826 _:vb591261827 _:vb591261828 _:vb591261829 _:vb591261830 _:vb591261831
n2:pmcid: PMC0
bibo:doi: 10.1371%2Fjournal.pgen.1005595
n9:contains: _:vb45456512 _:vb45456539 _:vb45456489 _:vb45456464

Subject Item: _:vb45456464
rdf:type: n9:Section
dc:title: introduction
n9:contains: _:vb45456484 _:vb45456485 _:vb45456486 _:vb45456487 _:vb45456480 _:vb45456481 _:vb45456482 _:vb45456483 _:vb45456488 _:vb45456468 _:vb45456469 _:vb45456470 _:vb45456471 _:vb45456465 _:vb45456466 _:vb45456467 _:vb45456476 _:vb45456477 _:vb45456478 _:vb45456479 _:vb45456472 _:vb45456473 _:vb45456474 _:vb45456475

Subject Item: _:vb45456465
rdf:type: n2:Context
rdf:value: The production of reactive oxygen species (ROS) by various redox metabolic reactions, which has generally been considered to be deleterious, is now emerging as an active participant in cell signaling events [>>1<<,2]. ROS are byproducts of aerobic metabolism that include superoxide O2 -, peroxide H2O2 and hydroxyl radicals OH·. ROS, and in particular H2O2 are required for inflammatory cell recruitment [3,4]. Amphibian and zebrafish injuries produce
n2:mentions: n3:21746850

Subject Item: _:vb45456466
rdf:type: n2:Context
rdf:value: The production of reactive oxygen species (ROS) by various redox metabolic reactions, which has generally been considered to be deleterious, is now emerging as an active participant in cell signaling events [1,>>2<<]. ROS are byproducts of aerobic metabolism that include superoxide O2 -, peroxide H2O2 and hydroxyl radicals OH·. ROS, and in particular H2O2 are required for inflammatory cell recruitment [3,4]. Amphibian and zebrafish injuries produce
n2:mentions: n3:25371358

Subject Item: _:vb45456467
rdf:type: n2:Context
rdf:value: ROS, and in particular H2O2 are required for inflammatory cell recruitment [>>3<<,4]. Amphibian and zebrafish injuries produce the ROS necessary to promote proliferation and regeneration [5–8]. In mammalian cells, ROS are known to act as second messengers to activate diverse redox-sensitive signaling transduction
n2:mentions: n3:20188558

Subject Item: _:vb45456468
rdf:type: n2:Context
rdf:value: ROS, and in particular H2O2 are required for inflammatory cell recruitment [3,>>4<<]. Amphibian and zebrafish injuries produce the ROS necessary to promote proliferation and regeneration [5–8]. In mammalian cells, ROS are known to act as second messengers to activate diverse redox-sensitive signaling transduction
n2:mentions: n3:19494811

Subject Item: _:vb45456469
rdf:type: n2:Context
rdf:value: Amphibian and zebrafish injuries produce the ROS necessary to promote proliferation and regeneration [>>5<<–8]. In mammalian cells, ROS are known to act as second messengers to activate diverse redox-sensitive signaling transduction cascades, including the stress-activated MAP kinases p38 and the Jun-N Terminal kinase (JNK) [9–11]. ROS-mediated
n2:mentions: n3:24413150 n3:23314862 n3:23803955 n3:17559804

Subject Item: _:vb45456470
rdf:type: n2:Context
rdf:value: In mammalian cells, ROS are known to act as second messengers to activate diverse redox-sensitive signaling transduction cascades, including the stress-activated MAP kinases p38 and the Jun-N Terminal kinase (JNK) [>>9<<–11]. ROS-mediated p38 activation occurs during the inflammatory response in rats [12] and during the loss of self-renewal and differentiation in glioma-initiating cells [13]. It has also been found that p38 and JNK are differentially
n2:mentions: n3:16987031 n3:11773609 n3:21228261

Subject Item: _:vb45456471
rdf:type: n2:Context
rdf:value: ROS-mediated p38 activation occurs during the inflammatory response in rats [12] and during the loss of self-renewal and differentiation in glioma-initiating cells [>>13<<]. It has also been found that p38 and JNK are differentially required during repair. In endothelial cells, TNF-α stimulates repair through the positive action of JNK and negative regulation of p38 [14], whereas in corneal repair, p38, and
n2:mentions: n3:24185179

Subject Item: _:vb45456472
rdf:type: n2:Context
rdf:value: In endothelial cells, TNF-α stimulates repair through the positive action of JNK and negative regulation of p38 [>>14<<], whereas in corneal repair, p38, and not JNK, is required for epithelial migration [15].
n2:mentions: n3:23258237

Subject Item: _:vb45456473
rdf:type: n2:Context
rdf:value: In endothelial cells, TNF-α stimulates repair through the positive action of JNK and negative regulation of p38 [14], whereas in corneal repair, p38, and not JNK, is required for epithelial migration [>>15<<]. In Drosophila both MAPK have been associated with stress responses [16]. Drosophila p38 pathway responds to different environmental stimuli and stressors [17,18]. Moreover, increasing ROS beyond basal level triggers precocious
n2:mentions: n3:12663671

Subject Item: _:vb45456474
rdf:type: n2:Context
rdf:value: In Drosophila both MAPK have been associated with stress responses [>>16<<]. Drosophila p38 pathway responds to different environmental stimuli and stressors [17,18].
n2:mentions: n3:22266315

Subject Item: _:vb45456475
rdf:type: n2:Context
rdf:value: Drosophila p38 pathway responds to different environmental stimuli and stressors [>>17<<,18]. Moreover, increasing ROS beyond basal level triggers precocious differentiation of Drosophila hematopoietic progenitors through JNK signaling [19].
n2:mentions: n3:21829386

Subject Item: _:vb45456476
rdf:type: n2:Context
rdf:value: Drosophila p38 pathway responds to different environmental stimuli and stressors [17,>>18<<]. Moreover, increasing ROS beyond basal level triggers precocious differentiation of Drosophila hematopoietic progenitors through JNK signaling [19].
n2:mentions: n3:15514678

Subject Item: _:vb45456477
rdf:type: n2:Context
rdf:value: Moreover, increasing ROS beyond basal level triggers precocious differentiation of Drosophila hematopoietic progenitors through JNK signaling [>>19<<].
n2:mentions: n3:19727075

Subject Item: _:vb45456478
rdf:type: n2:Context
rdf:value: The JNK signaling pathway has emerged as an early response to cell death and physical damage and appears to play a critical role in compensatory proliferation, regeneration and wound healing [>>20<<–28]. Moreover, upon apoptotic stimulus p53 and JNK are activated by the caspase Dronc and function upstream of pro-apoptotic genes, creating an amplifying loop that ensures cell death [29–33].
n2:mentions: n3:19531351 n3:15469838 n3:15968584 n3:20215351 n3:18956337 n3:19048077 n3:24497843 n3:16281037 n3:15766749

Subject Item: _:vb45456479
rdf:type: n2:Context
rdf:value: Moreover, upon apoptotic stimulus p53 and JNK are activated by the caspase Dronc and function upstream of pro-apoptotic genes, creating an amplifying loop that ensures cell death [>>29<<–33]. One of the early known responses to cell death is the transcriptional activation of the phosphatase puckered (puc), a downstream effector of the JNK pathway and a powerful negative regulator of the same pathway. Interestingly puc has
n2:mentions: n3:21886179 n3:16980627 n3:16920621 n3:15496444 n3:19244279

Subject Item: _:vb45456480
rdf:type: n2:Context
rdf:value: Interestingly puc has been found in surviving cells of nearby tissue after cell death [>>23<<,27] and after physical injury [22,34].
n2:mentions: n3:20215351

Subject Item: _:vb45456481
rdf:type: n2:Context
rdf:value: Interestingly puc has been found in surviving cells of nearby tissue after cell death [23,>>27<<] and after physical injury [22,34].
n2:mentions: n3:24497843

Subject Item: _:vb45456482
rdf:type: n2:Context
rdf:value: Interestingly puc has been found in surviving cells of nearby tissue after cell death [23,27] and after physical injury [>>22<<,34]. JNK activation of the cytokines unpaired (upd), a family of cytokines linked to the human interleukin-6, is necessary for hyperproliferation in Drosophila tumors and for wound healing [34–36]. Thus, we hypothesize here that the
n2:mentions: n3:18956337

Subject Item: _:vb45456483
rdf:type: n2:Context
rdf:value: Interestingly puc has been found in surviving cells of nearby tissue after cell death [23,27] and after physical injury [22,>>34<<]. JNK activation of the cytokines unpaired (upd), a family of cytokines linked to the human interleukin-6, is necessary for hyperproliferation in Drosophila tumors and for wound healing [34–36]. Thus, we hypothesize here that the
n2:mentions: n3:20072127

Subject Item: _:vb45456484
rdf:type: n2:Context
rdf:value: JNK activation of the cytokines unpaired (upd), a family of cytokines linked to the human interleukin-6, is necessary for hyperproliferation in Drosophila tumors and for wound healing [>>34<<–36]. Thus, we hypothesize here that the activation of JNK, which is amplified in dying cells, is in some way propagated to nearby surviving tissue where beneficial low levels of JNK promote upd expression.
n2:mentions: n3:20072127 n3:19563763 n3:25647511

Subject Item: _:vb45456485
rdf:type: n2:Context
rdf:value: Apoptotic cells have been observed in the early regeneration of different animals and are thought to provide signals that regulate wound healing and regeneration [>>37<<–39]. As apoptosis has been associated with oxidative stress and cytokines act as a functional link between oxidative stress and compensatory proliferation in mammals [40], we decided to investigate whether ROS occur upstream from the
n2:mentions: n3:22078876 n3:22391035 n3:24512708

Subject Item: _:vb45456486
rdf:type: n2:Context
rdf:value: As apoptosis has been associated with oxidative stress and cytokines act as a functional link between oxidative stress and compensatory proliferation in mammals [>>40<<], we decided to investigate whether ROS occur upstream from the stress-activated protein kinases p38 and JNK and cytokines during tissue repair.
n2:mentions: n3:22253262

Subject Item: _:vb45456487
rdf:type: n2:Context
rdf:value: We took advantage of the regeneration capacity of Drosophila imaginal disc epithelium (reviewed in [>>41<<,42]) to address these questions.
n2:mentions: n3:22934642

Subject Item: _:vb45456488
rdf:type: n2:Context
rdf:value: We took advantage of the regeneration capacity of Drosophila imaginal disc epithelium (reviewed in [41,>>42<<]) to address these questions.
n2:mentions: n3:20127699

Subject Item: _:vb45456489
rdf:type: n9:Section
dc:title: materials and methods
n9:contains: _:vb45456490 _:vb45456491 _:vb45456492 _:vb45456493 _:vb45456494 _:vb45456495 _:vb45456496 _:vb45456497 _:vb45456498 _:vb45456499 _:vb45456500 _:vb45456501 _:vb45456502 _:vb45456503 _:vb45456504 _:vb45456505 _:vb45456506 _:vb45456507 _:vb45456508 _:vb45456509 _:vb45456510 _:vb45456511

Subject Item: _:vb45456490
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [>>78<<], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2:
n2:mentions: n3:8287481

Subject Item: _:vb45456491
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [>>80<<], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2:
n2:mentions: n3:9846179

Subject Item: _:vb45456492
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [>>81<<], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2:
n2:mentions: n3:17035294

Subject Item: _:vb45456493
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [>>82<<] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2:
n2:mentions: n3:16950871

Subject Item: _:vb45456494
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [>>83<<], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2::GFP [43], TRE-DsRed.
n2:mentions: n3:15079076

Subject Item: _:vb45456495
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [>>47<<], dATF2 PB [49], p38a 1, [17], LexO-rCD2::GFP [43], TRE-DsRed.T4 [45] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [84], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:19917724

Subject Item: _:vb45456496
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [>>49<<], p38a 1, [17], LexO-rCD2::GFP [43], TRE-DsRed.T4 [45] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [84], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:21703449

Subject Item: _:vb45456497
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [>>17<<], LexO-rCD2::GFP [43], TRE-DsRed.T4 [45] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [84], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:21829386

Subject Item: _:vb45456498
rdf:type: n2:Context
rdf:value: The Drosophila melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2::GFP [>>43<<], TRE-DsRed.T4 [45] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [84], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:20805468

Subject Item: _:vb45456499
rdf:type: n2:Context
rdf:value: melanogaster strains used were ptc-Gal4 [78], tubGal80 TS [79], UAS-rpr [80], ci-Gal4 [81], nub-Gal4 [82] sal-Gal4 and sal E/Pv -Gal4 [83], p38b d27, lic d13 [47], dATF2 PB [49], p38a 1, [17], LexO-rCD2::GFP [43], TRE-DsRed.T4 [>>45<<] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [84], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:22509270

Subject Item: _:vb45456500
rdf:type: n2:Context
rdf:value: T4 [45] as AP1 reporter, puc-lacZ [>>44<<], puc E69-A -Gal4 [84], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:9472024

Subject Item: _:vb45456501
rdf:type: n2:Context
rdf:value: T4 [45] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [>>84<<], UAS-upd [85], upd-Gal4 (from D.
n2:mentions: n3:15363413

Subject Item: _:vb45456502
rdf:type: n2:Context
rdf:value: T4 [45] as AP1 reporter, puc-lacZ [44], puc E69-A -Gal4 [84], UAS-upd [>>85<<], upd-Gal4 (from D.
n2:mentions: n3:7796812

Subject Item: _:vb45456503
rdf:type: n2:Context
rdf:value: Harrison), 10XSTAT92E-GFP [>>86<<], en-Gal4, UAS-GFP, UAS-myrtomato, UAS-Sod.A (sod1), UAS-Cat.
n2:mentions: n3:17008134

Subject Item: _:vb45456504
rdf:type: n2:Context
rdf:value: A, UAS-dome DN, hop2, hop 27, stat92e 06346 (Bloomington Stock center), stat92e 397 [>>87<<], and hep r75 [88].
n2:mentions: n3:8608595

Subject Item: _:vb45456505
rdf:type: n2:Context
rdf:value: A, UAS-dome DN, hop2, hop 27, stat92e 06346 (Bloomington Stock center), stat92e 397 [87], and hep r75 [>>88<<]. Transgenic Drosophila shRNAi lines were obtained from the Vienna Drosophila RNAi Center (VDRC). Canton S was used as the wild type control.
n2:mentions: n3:8521475

Subject Item: _:vb45456506
rdf:type: n2:Context
rdf:value: The sal E/Pv -LHG construct was created cutting the wing specific enhancer of spalt, sal E/Pv [>>83<<] from pC4LacZ-Spalt PE EcoRI/BamHI and cloning this fragment into the plasmid attB-LHG containing a Gal80-suppressible form of LexA transcriptional activator (LHG) [43].
n2:mentions: n3:15079076

Subject Item: _:vb45456507
rdf:type: n2:Context
rdf:value: was created cutting the wing specific enhancer of spalt, sal E/Pv [83] from pC4LacZ-Spalt PE EcoRI/BamHI and cloning this fragment into the plasmid attB-LHG containing a Gal80-suppressible form of LexA transcriptional activator (LHG) [>>43<<]. LHG contains both the binding domain of LexA and the activator domain of Gal4, which is recognized by the inhibitor Gal80 TS.
n2:mentions: n3:20805468

Subject Item: _:vb45456508
rdf:type: n2:Context
rdf:value: The LexO-rpr strain was obtained subcloning the pro-apoptotic gene reaper (rpr) from pOT2-rpr (IP02529) EcoRI/XhoI in the pLOTattB plasmid [>>89<<] carrying the lexA operator LexO.
n2:mentions: n3:16582903

Subject Item: _:vb45456509
rdf:type: n2:Context
rdf:value: Cell death was genetically induced as previously described [>>23<<,90]. We used two different drivers to induce cell death. The first, ptc-Gal4 which is expressed in a narrow stripe in the center of the disc.
n2:mentions: n3:20215351

Subject Item: _:vb45456510
rdf:type: n2:Context
rdf:value: Cell death was genetically induced as previously described [23,>>90<<]. We used two different drivers to induce cell death. The first, ptc-Gal4 which is expressed in a narrow stripe in the center of the disc.
n2:mentions: n3:23903186

Subject Item: _:vb45456511
rdf:type: n2:Context
rdf:value: The second, sal E/Pv -Gal4 strain, which consists of sal wing enhancer with expression confined to the wing [>>83<<] has been used in this work to score adult wing parameters.
n2:mentions: n3:15079076

Subject Item: _:vb45456512
rdf:type: n9:Section
dc:title: results
n9:contains: _:vb45456516 _:vb45456517 _:vb45456518 _:vb45456519 _:vb45456513 _:vb45456514 _:vb45456515 _:vb45456524 _:vb45456525 _:vb45456526 _:vb45456527 _:vb45456520 _:vb45456521 _:vb45456522 _:vb45456523 _:vb45456532 _:vb45456533 _:vb45456534 _:vb45456535 _:vb45456528 _:vb45456529 _:vb45456530 _:vb45456531 _:vb45456536 _:vb45456537 _:vb45456538

Subject Item: _:vb45456513
rdf:type: n2:Context
rdf:value: gene reaper (rpr) under the control of a UAS (henceforth ptc>rpr); the Gal4/UAS system was controlled by the temperature-sensitive form of Gal80 (Gal80 TS), which inhibits Gal4 and enables examination of regeneration after cell death [>>23<<,24]. As previously described, ptc>rpr discs show a stripe of apoptotic cells that eventually extrudes basally and is replaced apically by living cells (Fig 1D) [23].
n2:mentions: n3:20215351

Subject Item: _:vb45456514
rdf:type: n2:Context
rdf:value: reaper (rpr) under the control of a UAS (henceforth ptc>rpr); the Gal4/UAS system was controlled by the temperature-sensitive form of Gal80 (Gal80 TS), which inhibits Gal4 and enables examination of regeneration after cell death [23,>>24<<]. As previously described, ptc>rpr discs show a stripe of apoptotic cells that eventually extrudes basally and is replaced apically by living cells (Fig 1D) [23].
n2:mentions: n3:19531351

Subject Item: _:vb45456515
rdf:type: n2:Context
rdf:value: As previously described, ptc>rpr discs show a stripe of apoptotic cells that eventually extrudes basally and is replaced apically by living cells (Fig 1D) [>>23<<]. CellROX Green was strongly incorporated into the ptc>rpr apoptotic cells (TO-PRO-3 positive) (Fig 1E). Strikingly, living cells adjacent to the apoptotic zone also showed ROS, albeit at much lower levels than in dead cells (Fig 1E and
n2:mentions: n3:20215351

Subject Item: _:vb45456516
rdf:type: n2:Context
rdf:value: We generated a sal E/Pv -LHG transgene, which includes a Gal80 suppressible form of LexA [>>43<<], to conditionally express lexO-rpr in the sal E/Pv domain.
n2:mentions: n3:20805468

Subject Item: _:vb45456517
rdf:type: n2:Context
rdf:value: We used two different reporters to monitor JNK activity: puc-lacZ, which marks puc-expressing cells [>>44<<], and the TRE-DsRed.
n2:mentions: n3:9472024

Subject Item: _:vb45456518
rdf:type: n2:Context
rdf:value: T4 reporter, which monitors the JNK substrate AP1 transcription factor (hereafter TRE-red reporter) (Fig 3A and S3A Fig) [>>45<<]. In ptc>rpr discs, we found high levels of TRE-red reporter in the basal apoptotic zone and, to a lesser extent, in the apical living cells (Fig 3A and 3B). In contrast, puc-lacZ positive cells were found in the apical zone, as described
n2:mentions: n3:22509270

Subject Item: _:vb45456519
rdf:type: n2:Context
rdf:value: In contrast, puc-lacZ positive cells were found in the apical zone, as described previously [>>23<<], and rarely in the apoptotic zone.
n2:mentions: n3:20215351

Subject Item: _:vb45456520
rdf:type: n2:Context
rdf:value: As NAC is an excellent source of sulfhydryl SH- groups and efficiently promotes scavenging of free radicals [>>46<<], it was the most suitable antioxidant to determine the relationship between ROS and JNK in stressed imaginal discs.
n2:mentions: n3:9577247

Subject Item: _:vb45456521
rdf:type: n2:Context
rdf:value: To test NAC effects on JNK, we used the TRE-red reporter because is more rapidly and extensively expressed than puc-lacZ (Figs 3A and S3B) and because its activity is blocked in JNK mutants [>>45<<] or after chemical JNK inhibitors (S3C Fig).
n2:mentions: n3:22509270

Subject Item: _:vb45456522
rdf:type: n2:Context
rdf:value: Another potential response to ROS increase is the activation of the p38 signaling cascade [>>10<<,17,18]. Active p-38 signaling can be monitored using anti-phosphorylated p38 (P-p38).
n2:mentions: n3:16987031

Subject Item: _:vb45456523
rdf:type: n2:Context
rdf:value: Another potential response to ROS increase is the activation of the p38 signaling cascade [10,>>17<<,18]. Active p-38 signaling can be monitored using anti-phosphorylated p38 (P-p38).
n2:mentions: n3:21829386

Subject Item: _:vb45456524
rdf:type: n2:Context
rdf:value: Another potential response to ROS increase is the activation of the p38 signaling cascade [10,17,>>18<<]. Active p-38 signaling can be monitored using anti-phosphorylated p38 (P-p38). We found that discs fixed and incubated with anti-P-p38 a few minutes after physical injury showed P-p38 staining around the wound.
n2:mentions: n3:15514678

Subject Item: _:vb45456525
rdf:type: n2:Context
rdf:value: As most of the alleles are lethal or semilethal in homozygosis [>>47<<], we tested them in heterozygosis.
n2:mentions: n3:19917724

Subject Item: _:vb45456526
rdf:type: n2:Context
rdf:value: Drosophila p38 signaling is activated by MKK3/licorne (lic)-mediated phosphorylation [>>48<<]. Heterozygous lic d13 showed all wing sectors albeit wings were smaller than controls. However, double heterozygotes for lic d13 and p38b d27 were unable to regenerate some wing sectors. We also tested Atf2 PB, a hypomorphic allele of
n2:mentions: n3:10364162

Subject Item: _:vb45456527
rdf:type: n2:Context
rdf:value: We also tested Atf2 PB, a hypomorphic allele of the ATF2 transcription factor downstream of p38 [>>49<<], either in homozygosis or in double heterozygous combinations (lic d13 Atf2 PB or p38b d27 Atf2 PB).
n2:mentions: n3:21703449

Subject Item: _:vb45456528
rdf:type: n2:Context
rdf:value: the pathway using the imidazole drug SB202190, a specific cell permeable p38 MAP kinase inhibitor that has been reported to do not interfere JNK or ERK kinases and is known to prevent phosphorylation of Atf2 in Drosophila S2 cells [>>50<<]. We first tested the specificity of the SB202190 on P-p38 in rpr-ablated discs and found significant differences between individuals fed with the drug and controls.
n2:mentions: n3:9753474

Subject Item: _:vb45456529
rdf:type: n2:Context
rdf:value: The evidence that JNK is active in the living tissue located near damaged zones arises from the expression of puc and TRE-red reporters (Fig 3A and 3B), and also because inhibition of JNK results in defects in repair [>>21<<,23,25,26,51]. Moreover, JNK activation promotes upd expression in different contexts [28,52].
n2:mentions: n3:15766749

Subject Item: _:vb45456530
rdf:type: n2:Context
rdf:value: The evidence that JNK is active in the living tissue located near damaged zones arises from the expression of puc and TRE-red reporters (Fig 3A and 3B), and also because inhibition of JNK results in defects in repair [21,>>23<<,25,26,51]. Moreover, JNK activation promotes upd expression in different contexts [28,52].
n2:mentions: n3:20215351

Subject Item: _:vb45456531
rdf:type: n2:Context
rdf:value: The evidence that JNK is active in the living tissue located near damaged zones arises from the expression of puc and TRE-red reporters (Fig 3A and 3B), and also because inhibition of JNK results in defects in repair [21,23,>>25<<,26,51]. Moreover, JNK activation promotes upd expression in different contexts [28,52].
n2:mentions: n3:15968584

Subject Item: _:vb45456532
rdf:type: n2:Context
rdf:value: The evidence that JNK is active in the living tissue located near damaged zones arises from the expression of puc and TRE-red reporters (Fig 3A and 3B), and also because inhibition of JNK results in defects in repair [21,23,25,>>26<<,51]. Moreover, JNK activation promotes upd expression in different contexts [28,52].
n2:mentions: n3:16281037

Subject Item: _:vb45456533
rdf:type: n2:Context
rdf:value: The evidence that JNK is active in the living tissue located near damaged zones arises from the expression of puc and TRE-red reporters (Fig 3A and 3B), and also because inhibition of JNK results in defects in repair [21,23,25,26,>>51<<]. Moreover, JNK activation promotes upd expression in different contexts [28,52].
n2:mentions: n3:15269788

Subject Item: _:vb45456534
rdf:type: n2:Context
rdf:value: Moreover, JNK activation promotes upd expression in different contexts [>>28<<,52]. We wondered whether those low non-deleterious JNK levels are capable of triggering tissue repair through upd expression.
n2:mentions: n3:19048077

Subject Item: _:vb45456535
rdf:type: n2:Context
rdf:value: Moreover, JNK activation promotes upd expression in different contexts [28,>>52<<]. We wondered whether those low non-deleterious JNK levels are capable of triggering tissue repair through upd expression.
n2:mentions: n3:25719210

Subject Item: _:vb45456536
rdf:type: n2:Context
rdf:value: cytokines are ligands that associate with the receptor domeless (dome) to stimulate the kinase activity of the receptor associated protein kinase hopscotch (hop), which in turn phosphorylates dimers of the transcription factor STAT92E [>>53<<]. We found upd and upd3 expression near the wound after both physical injury and cell death (Fig 7A, 7B, 7F and 7G).
n2:mentions: n3:24685611

Subject Item: _:vb45456537
rdf:type: n2:Context
rdf:value: This injury-induced upd expression was blocked in JNKK hep r75 mutants (Fig 7C and 7D) and by JNK Inhibitor IX (S3C Fig), which is consistent with previous observations [>>28<<,35].
n2:mentions: n3:19048077

Subject Item: _:vb45456538
rdf:type: n2:Context
rdf:value: This injury-induced upd expression was blocked in JNKK hep r75 mutants (Fig 7C and 7D) and by JNK Inhibitor IX (S3C Fig), which is consistent with previous observations [28,>>35<<].
n2:mentions: n3:25647511

Subject Item: _:vb45456539
rdf:type: n9:Section
dc:title: discussion
n9:contains: _:vb45456540 _:vb45456541 _:vb45456542 _:vb45456543 _:vb45456552 _:vb45456553 _:vb45456554 _:vb45456555 _:vb45456556 _:vb45456557 _:vb45456558 _:vb45456559 _:vb45456544 _:vb45456545 _:vb45456546 _:vb45456547 _:vb45456548 _:vb45456549 _:vb45456550 _:vb45456551 _:vb45456568 _:vb45456569 _:vb45456570 _:vb45456571 _:vb45456572 _:vb45456573 _:vb45456574 _:vb45456560 _:vb45456561 _:vb45456562 _:vb45456563 _:vb45456564 _:vb45456565 _:vb45456566 _:vb45456567

Subject Item: _:vb45456540
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [>>44<<,54], healing and initiation of regenerative growth [21,23–26,51,55,56].
n2:mentions: n3:9472024

Subject Item: _:vb45456541
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [44,>>54<<], healing and initiation of regenerative growth [21,23–26,51,55,56].
n2:mentions: n3:11311164

Subject Item: _:vb45456542
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [44,54], healing and initiation of regenerative growth [>>21<<,23–26,51,55,56].
n2:mentions: n3:15766749

Subject Item: _:vb45456543
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [44,54], healing and initiation of regenerative growth [21,>>23<<–26,51,55,56].
n2:mentions: n3:19531351 n3:15968584 n3:20215351 n3:16281037

Subject Item: _:vb45456544
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [44,54], healing and initiation of regenerative growth [21,23–26,>>51<<,55,56]. Thus, this early responsive module is crucial to maintain tissue in a healthy condition, trigger tissue repair and restore homeostasis.
n2:mentions: n3:15269788

Subject Item: _:vb45456545
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [44,54], healing and initiation of regenerative growth [21,23–26,51,>>55<<,56]. Thus, this early responsive module is crucial to maintain tissue in a healthy condition, trigger tissue repair and restore homeostasis.
n2:mentions: n3:20813879

Subject Item: _:vb45456546
rdf:type: n2:Context
rdf:value: Non-lethal levels of JNK may have multiple functions, among them cytoskeleton organization [44,54], healing and initiation of regenerative growth [21,23–26,51,55,>>56<<]. Thus, this early responsive module is crucial to maintain tissue in a healthy condition, trigger tissue repair and restore homeostasis.
n2:mentions: n3:11784101

Subject Item: _:vb45456547
rdf:type: n2:Context
rdf:value: In an apoptotic context, Rpr dimerizes and, through direct binding, brings the Drosophila inhibitor of apoptosis protein-1 (DIAP1) to mitochondria, concomitantly promoting DIAP1 auto-ubiquitination and destruction [>>57<<,58]. Rpr action on the mitochondria results in alteration of cytochrome C driven by caspases [59] and in mitochondrial disruption [60].
n2:mentions: n3:17998202

Subject Item: _:vb45456548
rdf:type: n2:Context
rdf:value: In an apoptotic context, Rpr dimerizes and, through direct binding, brings the Drosophila inhibitor of apoptosis protein-1 (DIAP1) to mitochondria, concomitantly promoting DIAP1 auto-ubiquitination and destruction [57,>>58<<]. Rpr action on the mitochondria results in alteration of cytochrome C driven by caspases [59] and in mitochondrial disruption [60].
n2:mentions: n3:20837774

Subject Item: _:vb45456549
rdf:type: n2:Context
rdf:value: Rpr action on the mitochondria results in alteration of cytochrome C driven by caspases [>>59<<] and in mitochondrial disruption [60].
n2:mentions: n3:10037791

Subject Item: _:vb45456550
rdf:type: n2:Context
rdf:value: Rpr action on the mitochondria results in alteration of cytochrome C driven by caspases [59] and in mitochondrial disruption [>>60<<]. The ROS dyes used here detect a wide range of ROS, and therefore we cannot discriminate between membrane oxidases or mitochondrial origin. However, since Rpr acts on mitochondria, mitochondrial alterations could cause the burst of ROS
n2:mentions: n3:17488629

Subject Item: _:vb45456551
rdf:type: n2:Context
rdf:value: It has been proposed that ROS can mediate the activation of JNK [>>61<<] by quenching the MAP kinase phosphatases [62].
n2:mentions: n3:8663189

Subject Item: _:vb45456552
rdf:type: n2:Context
rdf:value: It has been proposed that ROS can mediate the activation of JNK [61] by quenching the MAP kinase phosphatases [>>62<<]. Conversely, low levels of ROS detected in nearby surviving tissue correlate with low non-deleterious levels of JNK and activation of MAP kinase phosphatases. Thus, puc MAP kinase phosphatase could protect the living cells close to the
n2:mentions: n3:24603536

Subject Item: _:vb45456553
rdf:type: n2:Context
rdf:value: Additionally, the caspase Dronc, which acts downstream from Rpr, has functions beyond apoptosis [>>63<<]. Dronc is involved in the activation of JNK and p53, which activate the pro-apoptotic genes, creating an amplification loop that ensures apoptosis [27,29–31,64].
n2:mentions: n3:15268856

Subject Item: _:vb45456554
rdf:type: n2:Context
rdf:value: Dronc is involved in the activation of JNK and p53, which activate the pro-apoptotic genes, creating an amplification loop that ensures apoptosis [>>27<<,29–31,64].
n2:mentions: n3:24497843

Subject Item: _:vb45456555
rdf:type: n2:Context
rdf:value: Dronc is involved in the activation of JNK and p53, which activate the pro-apoptotic genes, creating an amplification loop that ensures apoptosis [27,>>29<<–31,64]. The JNK/p53 driven apoptosis stimulates proliferation of the nearby tissues [29–31,65]. Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and
n2:mentions: n3:21886179 n3:16980627 n3:16920621

Subject Item: _:vb45456556
rdf:type: n2:Context
rdf:value: Dronc is involved in the activation of JNK and p53, which activate the pro-apoptotic genes, creating an amplification loop that ensures apoptosis [27,29–31,>>64<<]. The JNK/p53 driven apoptosis stimulates proliferation of the nearby tissues [29–31,65]. Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and
n2:mentions: n3:22898807

Subject Item: _:vb45456557
rdf:type: n2:Context
rdf:value: The JNK/p53 driven apoptosis stimulates proliferation of the nearby tissues [>>29<<–31,65]. Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [33,66,67][31,68]. Alternatively, we show here that ROS operate as
n2:mentions: n3:21886179 n3:16980627 n3:16920621

Subject Item: _:vb45456558
rdf:type: n2:Context
rdf:value: The JNK/p53 driven apoptosis stimulates proliferation of the nearby tissues [29–31,>>65<<]. Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [33,66,67][31,68]. Alternatively, we show here that ROS operate as signals
n2:mentions: n3:22036477

Subject Item: _:vb45456559
rdf:type: n2:Context
rdf:value: Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [>>33<<,66,67][31,68].
n2:mentions: n3:19244279

Subject Item: _:vb45456560
rdf:type: n2:Context
rdf:value: Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [33,>>66<<,67][31,68].
n2:mentions: n3:15469838

Subject Item: _:vb45456561
rdf:type: n2:Context
rdf:value: Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [33,66,>>67<<][31,68]. Alternatively, we show here that ROS operate as signals responding to insults (apoptosis, mechanical stress) that turn on the homeostatic machinery to compensate the epithelial damage. This fits with a scenario in which ROS are
n2:mentions: n3:18331718

Subject Item: _:vb45456562
rdf:type: n2:Context
rdf:value: Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [33,66,67][>>31<<,68]. Alternatively, we show here that ROS operate as signals responding to insults (apoptosis, mechanical stress) that turn on the homeostatic machinery to compensate the epithelial damage. This fits with a scenario in which ROS are able
n2:mentions: n3:16920621

Subject Item: _:vb45456563
rdf:type: n2:Context
rdf:value: Although still unclear, it has been proposed that apoptotic cells can release the products of mitogenic genes such as wingless (wg) and decapentaplegic (dpp) [33,66,67][31,>>68<<]. Alternatively, we show here that ROS operate as signals responding to insults (apoptosis, mechanical stress) that turn on the homeostatic machinery to compensate the epithelial damage. This fits with a scenario in which ROS are able to
n2:mentions: n3:23633961

Subject Item: _:vb45456564
rdf:type: n2:Context
rdf:value: Indeed, ROS have been proven to cross cell membranes, to spread through gap junctions [>>69<<–71] and to enter into the cell through specific membrane aquaporin channels [71,72].
n2:mentions: n3:16566894 n3:22911831 n3:21388278

Subject Item: _:vb45456565
rdf:type: n2:Context
rdf:value: Indeed, ROS have been proven to cross cell membranes, to spread through gap junctions [69–71] and to enter into the cell through specific membrane aquaporin channels [>>71<<,72]. Therefore, ROS behave as an efficient paracrine signal that ultimately will result in Upd activation.
n2:mentions: n3:16566894

Subject Item: _:vb45456566
rdf:type: n2:Context
rdf:value: Indeed, ROS have been proven to cross cell membranes, to spread through gap junctions [69–71] and to enter into the cell through specific membrane aquaporin channels [71,>>72<<]. Therefore, ROS behave as an efficient paracrine signal that ultimately will result in Upd activation.
n2:mentions: n3:17105724

Subject Item: _:vb45456567
rdf:type: n2:Context
rdf:value: In addition to JNK, ROS are stressors involved in p38 activation [>>73<<]. ROS may activate the p38 pathway through the oxidative modification of intracellular kinases such as redox-sensitive activating protein-1 ASK1 [74].
n2:mentions: n3:23849857

Subject Item: _:vb45456568
rdf:type: n2:Context
rdf:value: ROS may activate the p38 pathway through the oxidative modification of intracellular kinases such as redox-sensitive activating protein-1 ASK1 [>>74<<]. We showed here that not only JNK but also p38 is required for regeneration. Moreover, the p38a 1 allele seems to particularly affect upd expression and regeneration. This concurs with the finding that Drosophila p38a is more susceptible
n2:mentions: n3:11266364

Subject Item: _:vb45456569
rdf:type: n2:Context
rdf:value: This concurs with the finding that Drosophila p38a is more susceptible to environmental stressors, such as oxidative stress [>>18<<]. However, other p38 kinases could contribute to tissue regeneration. Indeed, heterozygous alleles of the p38 activating kinase lic, which normally do not show patterning defects after rpr-mediated ablation, can result in incomplete
n2:mentions: n3:15514678

Subject Item: _:vb45456570
rdf:type: n2:Context
rdf:value: In addition, we cannot discard that p38c, which has been recently found involved in intestinal immune homeostasis [>>75<<], may also function in imaginal disc regeneration.
n2:mentions: n3:25254641

Subject Item: _:vb45456571
rdf:type: n2:Context
rdf:value: Beneficial ROS production is an ubiquitous reaction associated with inflammatory responses to wounding [>>4<<,6,76]. Recent findings show that ROS produced in dynamic epithelia operate as a tuning mechanism for reorganization of epithelia [77]. Therefore, it could be that changes in mechanical stress generated during wounding and epithelial
n2:mentions: n3:19494811

Subject Item: _:vb45456572
rdf:type: n2:Context
rdf:value: Beneficial ROS production is an ubiquitous reaction associated with inflammatory responses to wounding [4,>>6<<,76]. Recent findings show that ROS produced in dynamic epithelia operate as a tuning mechanism for reorganization of epithelia [77]. Therefore, it could be that changes in mechanical stress generated during wounding and epithelial
n2:mentions: n3:23314862

Subject Item: _:vb45456573
rdf:type: n2:Context
rdf:value: Beneficial ROS production is an ubiquitous reaction associated with inflammatory responses to wounding [4,6,>>76<<]. Recent findings show that ROS produced in dynamic epithelia operate as a tuning mechanism for reorganization of epithelia [77]. Therefore, it could be that changes in mechanical stress generated during wounding and epithelial disruption
n2:mentions: n3:23394834

Subject Item: _:vb45456574
rdf:type: n2:Context
rdf:value: Recent findings show that ROS produced in dynamic epithelia operate as a tuning mechanism for reorganization of epithelia [>>77<<]. Therefore, it could be that changes in mechanical stress generated during wounding and epithelial disruption (mechanical stretching) results in ROS production. Some dead cells were also found after physical injury. Thus, a partial
n2:mentions: n3:24486154

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