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_:b389835344 , _:b389835345 , _:b389835346 , _:b389835347 , _:b389835348 , _:b389835349 , _:b389835350 , _:b389835351 , _:b389835336 , _:b389835337 , _:b389835338 , _:b389835339 , _:b389835340 , _:b389835341 , _:b389835342 , _:b389835343 , _:b389835328 , _:b389835329 , _:b389835330 , _:b389835331 , _:b389835332 , _:b389835333 , _:b389835334 , _:b389835335 ; ns1:pmcid "PMC0" ; bibo:doi "10.1371%2Fjournal.ppat.1000196" . @prefix ns4: . ns4:contains _:b4304219 , _:b4304291 , _:b4304259 , _:b4304274 . _:b4304219 rdf:type ns4:Section . @prefix dc: . _:b4304219 dc:title "introduction" ; ns4:contains _:b4304256 , _:b4304257 , _:b4304258 , _:b4304220 , _:b4304221 , _:b4304222 , _:b4304223 , _:b4304232 , _:b4304233 , _:b4304234 , _:b4304235 , _:b4304236 , _:b4304237 , _:b4304238 , _:b4304239 , _:b4304224 , _:b4304225 , _:b4304226 , _:b4304227 , _:b4304228 , _:b4304229 , _:b4304230 , _:b4304231 , _:b4304248 , _:b4304249 , _:b4304250 , _:b4304251 , _:b4304252 , _:b4304253 , _:b4304254 , _:b4304255 , _:b4304240 , _:b4304241 , _:b4304242 , _:b4304243 , _:b4304244 , _:b4304245 , _:b4304246 , _:b4304247 . _:b4304220 rdf:type ns1:Context ; rdf:value "This most likely occurs via binding to the RNA-sensor retinoic acid inducible gene (RIG-I) and inhibition of RIG-I-mediated signaling in response to viral RNA [>>1<<],[2]. On the other hand NS1 inhibits maturation [3],[4] and nuclear export of host mRNAs [5]. Other functions of the multifunctional protein include block of activation of the dsRNA-activated protein kinase PKR by direct interaction [6] or" ; ns1:mentions . _:b4304221 rdf:type ns1:Context ; rdf:value "This most likely occurs via binding to the RNA-sensor retinoic acid inducible gene (RIG-I) and inhibition of RIG-I-mediated signaling in response to viral RNA [1],[>>2<<]. On the other hand NS1 inhibits maturation [3],[4] and nuclear export of host mRNAs [5]. Other functions of the multifunctional protein include block of activation of the dsRNA-activated protein kinase PKR by direct interaction [6] or" ; ns1:mentions . _:b4304222 rdf:type ns1:Context ; rdf:value "On the other hand NS1 inhibits maturation [>>3<<],[4] and nuclear export of host mRNAs [5]." ; ns1:mentions . _:b4304223 rdf:type ns1:Context ; rdf:value "On the other hand NS1 inhibits maturation [3],[>>4<<] and nuclear export of host mRNAs [5]." ; ns1:mentions . _:b4304224 rdf:type ns1:Context ; rdf:value "On the other hand NS1 inhibits maturation [3],[4] and nuclear export of host mRNAs [>>5<<]. Other functions of the multifunctional protein include block of activation of the dsRNA-activated protein kinase PKR by direct interaction [6] or activation of the phosphatidylinositol-3 kinase PI3K/Akt pathway to prevent premature" ; ns1:mentions . _:b4304225 rdf:type ns1:Context ; rdf:value "Other functions of the multifunctional protein include block of activation of the dsRNA-activated protein kinase PKR by direct interaction [>>6<<] or activation of the phosphatidylinositol-3 kinase PI3K/Akt pathway to prevent premature apoptosis induction [7],[8]." ; ns1:mentions . _:b4304226 rdf:type ns1:Context ; rdf:value "of the multifunctional protein include block of activation of the dsRNA-activated protein kinase PKR by direct interaction [6] or activation of the phosphatidylinositol-3 kinase PI3K/Akt pathway to prevent premature apoptosis induction [>>7<<],[8]." ; ns1:mentions . _:b4304227 rdf:type ns1:Context ; rdf:value "multifunctional protein include block of activation of the dsRNA-activated protein kinase PKR by direct interaction [6] or activation of the phosphatidylinositol-3 kinase PI3K/Akt pathway to prevent premature apoptosis induction [7],[>>8<<]." ; ns1:mentions . _:b4304228 rdf:type ns1:Context ; rdf:value "IFN are among the first molecules synthesized in response to viral infections [>>9<<]. The IFN family includes three classes. Type I comprises the well known IFN\u03B1 and IFN\u03B2." ; ns1:mentions . _:b4304229 rdf:type ns1:Context ; rdf:value "All classes of IFN bind to different receptors and are structurally not related [>>10<<],[11]. Type I IFN belong to the key cytokines produced by influenza A virus-infected epithelial cells [12],[13]. The antiviral activity of type I IFN is mediated by a set of IFN-induced genes (ISGs)." ; ns1:mentions . _:b4304230 rdf:type ns1:Context ; rdf:value "All classes of IFN bind to different receptors and are structurally not related [10],[>>11<<]. Type I IFN belong to the key cytokines produced by influenza A virus-infected epithelial cells [12],[13]. The antiviral activity of type I IFN is mediated by a set of IFN-induced genes (ISGs)." ; ns1:mentions . _:b4304231 rdf:type ns1:Context ; rdf:value "Type I IFN belong to the key cytokines produced by influenza A virus-infected epithelial cells [>>12<<],[13]. The antiviral activity of type I IFN is mediated by a set of IFN-induced genes (ISGs)." ; ns1:mentions . _:b4304232 rdf:type ns1:Context ; rdf:value "Type I IFN belong to the key cytokines produced by influenza A virus-infected epithelial cells [12],[>>13<<]. The antiviral activity of type I IFN is mediated by a set of IFN-induced genes (ISGs)." ; ns1:mentions . _:b4304233 rdf:type ns1:Context ; rdf:value "Binding of IFN\u03B1/\u03B2 to its receptor is the initial step in this signaling process, followed by activation of the JAK family and subsequent activation of STAT proteins [>>14<<]." ; ns1:mentions . _:b4304234 rdf:type ns1:Context ; rdf:value "At the same time the receptor-bound JAK1 phosphorylates STAT1 at Y701 [>>15<<],[16]." ; ns1:mentions . _:b4304235 rdf:type ns1:Context ; rdf:value "At the same time the receptor-bound JAK1 phosphorylates STAT1 at Y701 [15],[>>16<<]." ; ns1:mentions . _:b4304236 rdf:type ns1:Context ; rdf:value "The phosphorylated transcription factors dimerise and bind to IRF-9 [>>17<<]. The newly formed heterotrimer, called IFN-stimulated gene factor 3 (ISGF3), translocates into the nucleus and binds to IFN-stimulated response elements (ISRE), to initiate gene transcription of ISGs." ; ns1:mentions . _:b4304237 rdf:type ns1:Context ; rdf:value "Treatment of cells with type I IFN up-regulates expression of an array of genes including SP110, IRF-1 and many others [>>18<<]. Among these ISGs the 2\u2032, 5\u2032 -oligoadenylate synthetase 1 (OAS1), the Mx proteins and the dsRNA-activated protein kinase (PKR) are described to directly interfere with viral replication [19]. Both, PKR and the OAS1/RNaseL system are" ; ns1:mentions . _:b4304238 rdf:type ns1:Context ; rdf:value "Among these ISGs the 2\u2032, 5\u2032 -oligoadenylate synthetase 1 (OAS1), the Mx proteins and the dsRNA-activated protein kinase (PKR) are described to directly interfere with viral replication [>>19<<]. Both, PKR and the OAS1/RNaseL system are capable of inhibiting cellular and viral translation." ; ns1:mentions . _:b4304239 rdf:type ns1:Context ; rdf:value "The large T-antigen of murine polyomavirus (MPyV) binds to JAK1 and inhibits downstream signaling [>>20<<], whereas the VP24 of Ebola virus (EBOV) binds to karyopherina-1 thereby blocking nuclear accumulation of STAT1 [21]." ; ns1:mentions . _:b4304240 rdf:type ns1:Context ; rdf:value "The large T-antigen of murine polyomavirus (MPyV) binds to JAK1 and inhibits downstream signaling [20], whereas the VP24 of Ebola virus (EBOV) binds to karyopherina-1 thereby blocking nuclear accumulation of STAT1 [>>21<<]." ; ns1:mentions . _:b4304241 rdf:type ns1:Context ; rdf:value "All members contain a central SH2 domain, an N-terminus of variable length and sequence and a C-terminal 40 amino-acid module called SOCS box [>>22<<]. The SOCS box is necessary for recruitment of the ubiquitin transferase system and for stabilization and/or degradation of SOCS proteins [23]\u2013[25]. The N-terminus contains a kinase inhibitory region (KIR), which functions as pseudo" ; ns1:mentions . _:b4304242 rdf:type ns1:Context ; rdf:value "The SOCS box is necessary for recruitment of the ubiquitin transferase system and for stabilization and/or degradation of SOCS proteins [>>23<<]\u2013[25]. The N-terminus contains a kinase inhibitory region (KIR), which functions as pseudo substrate for the JAK [26]. SOCS-1 and SOCS-3 differ in their mode of action. For inhibition of the kinase activity of JAKs, SOCS-1 binds directly" ; ns1:mentions . _:b4304243 rdf:type ns1:Context ; rdf:value "The SOCS box is necessary for recruitment of the ubiquitin transferase system and for stabilization and/or degradation of SOCS proteins [23]\u2013[>>25<<]. The N-terminus contains a kinase inhibitory region (KIR), which functions as pseudo substrate for the JAK [26]. SOCS-1 and SOCS-3 differ in their mode of action. For inhibition of the kinase activity of JAKs, SOCS-1 binds directly to the" ; ns1:mentions . _:b4304244 rdf:type ns1:Context ; rdf:value "The N-terminus contains a kinase inhibitory region (KIR), which functions as pseudo substrate for the JAK [>>26<<]. SOCS-1 and SOCS-3 differ in their mode of action. For inhibition of the kinase activity of JAKs, SOCS-1 binds directly to the activation loop of JAKs [26]\u2013[28]. In contrast, SOCS-3 first binds to the receptor [29],[30]." ; ns1:mentions . _:b4304245 rdf:type ns1:Context ; rdf:value "For inhibition of the kinase activity of JAKs, SOCS-1 binds directly to the activation loop of JAKs [>>26<<]\u2013[28]. In contrast, SOCS-3 first binds to the receptor [29],[30]." ; ns1:mentions . _:b4304246 rdf:type ns1:Context ; rdf:value "For inhibition of the kinase activity of JAKs, SOCS-1 binds directly to the activation loop of JAKs [26]\u2013[>>28<<]. In contrast, SOCS-3 first binds to the receptor [29],[30]." ; ns1:mentions . _:b4304247 rdf:type ns1:Context ; rdf:value "For inhibition of the kinase activity of JAKs, SOCS-1 binds directly to the activation loop of JAKs [26]\u2013[28]. In contrast, SOCS-3 first binds to the receptor [>>29<<],[30]." ; ns1:mentions . _:b4304248 rdf:type ns1:Context ; rdf:value "For inhibition of the kinase activity of JAKs, SOCS-1 binds directly to the activation loop of JAKs [26]\u2013[28]. In contrast, SOCS-3 first binds to the receptor [29],[>>30<<]." ; ns1:mentions . _:b4304249 rdf:type ns1:Context ; rdf:value "Induction of SOCS-3 gene transcription by viruses was reported for HSV-1, HCV [>>31<<]\u2013[33] and for respiratory viruses, such as SARS and RSV [34],[35]. The level of induction of SOCS-3 by HSV-1 seems to determine whether infection turns to acute or persistent progression [31]." ; ns1:mentions . _:b4304250 rdf:type ns1:Context ; rdf:value "Induction of SOCS-3 gene transcription by viruses was reported for HSV-1, HCV [31]\u2013[>>33<<] and for respiratory viruses, such as SARS and RSV [34],[35]. The level of induction of SOCS-3 by HSV-1 seems to determine whether infection turns to acute or persistent progression [31]." ; ns1:mentions . _:b4304251 rdf:type ns1:Context ; rdf:value "Induction of SOCS-3 gene transcription by viruses was reported for HSV-1, HCV [31]\u2013[33] and for respiratory viruses, such as SARS and RSV [>>34<<],[35]. The level of induction of SOCS-3 by HSV-1 seems to determine whether infection turns to acute or persistent progression [31]." ; ns1:mentions . _:b4304252 rdf:type ns1:Context ; rdf:value "Induction of SOCS-3 gene transcription by viruses was reported for HSV-1, HCV [31]\u2013[33] and for respiratory viruses, such as SARS and RSV [34],[>>35<<]. The level of induction of SOCS-3 by HSV-1 seems to determine whether infection turns to acute or persistent progression [31]." ; ns1:mentions . _:b4304253 rdf:type ns1:Context ; rdf:value "The level of induction of SOCS-3 by HSV-1 seems to determine whether infection turns to acute or persistent progression [>>31<<]. For HCV it has been suggested that upregulation of SOCS-3 may contribute to the non-responsiveness of HCV patients to IFN therapy [33], [36]\u2013[38]. Elevated SOCS-3 mRNA levels during RSV infection were linked to Th2 cell-mediated immune" ; ns1:mentions . _:b4304254 rdf:type ns1:Context ; rdf:value "For HCV it has been suggested that upregulation of SOCS-3 may contribute to the non-responsiveness of HCV patients to IFN therapy [>>33<<], [36]\u2013[38]." ; ns1:mentions . _:b4304255 rdf:type ns1:Context ; rdf:value "For HCV it has been suggested that upregulation of SOCS-3 may contribute to the non-responsiveness of HCV patients to IFN therapy [33], [>>36<<]\u2013[38]. Elevated SOCS-3 mRNA levels during RSV infection were linked to Th2 cell-mediated immune disease as atopic dermatitis and asthma [39],[40]." ; ns1:mentions . _:b4304256 rdf:type ns1:Context ; rdf:value "For HCV it has been suggested that upregulation of SOCS-3 may contribute to the non-responsiveness of HCV patients to IFN therapy [33], [36]\u2013[>>38<<]. Elevated SOCS-3 mRNA levels during RSV infection were linked to Th2 cell-mediated immune disease as atopic dermatitis and asthma [39],[40]." ; ns1:mentions . _:b4304257 rdf:type ns1:Context ; rdf:value "Elevated SOCS-3 mRNA levels during RSV infection were linked to Th2 cell-mediated immune disease as atopic dermatitis and asthma [>>39<<],[40]." ; ns1:mentions . _:b4304258 rdf:type ns1:Context ; rdf:value "Elevated SOCS-3 mRNA levels during RSV infection were linked to Th2 cell-mediated immune disease as atopic dermatitis and asthma [39],[>>40<<]." ; ns1:mentions . _:b4304259 rdf:type ns4:Section ; dc:title "materials and methods" ; ns4:contains _:b4304264 , _:b4304265 , _:b4304266 , _:b4304267 , _:b4304268 , _:b4304269 , _:b4304270 , _:b4304271 , _:b4304260 , _:b4304261 , _:b4304262 , _:b4304263 , _:b4304272 , _:b4304273 . _:b4304260 rdf:type ns1:Context ; rdf:value "The human NS1 deficient influenza virus mutant \u0394NS1 and its isogenic wild type variant were propagated and used as described earlier [>>7<<],[67]. It should be noted that this isogenic wild type strain as described by Garcia-Sastre et al. [68] is different from the PR8 (Giessen variant) used in the other experiments and in many previous studies [52],[67]. While both variants" ; ns1:mentions . _:b4304261 rdf:type ns1:Context ; rdf:value "The human NS1 deficient influenza virus mutant \u0394NS1 and its isogenic wild type variant were propagated and used as described earlier [7],[>>67<<]. It should be noted that this isogenic wild type strain as described by Garcia-Sastre et al. [68] is different from the PR8 (Giessen variant) used in the other experiments and in many previous studies [52],[67]. While both variants are of" ; ns1:mentions . _:b4304262 rdf:type ns1:Context ; rdf:value "It should be noted that this isogenic wild type strain as described by Garcia-Sastre et al. [>>68<<] is different from the PR8 (Giessen variant) used in the other experiments and in many previous studies [52],[67]." ; ns1:mentions . _:b4304263 rdf:type ns1:Context ; rdf:value "It should be noted that this isogenic wild type strain as described by Garcia-Sastre et al. [68] is different from the PR8 (Giessen variant) used in the other experiments and in many previous studies [>>52<<],[67]. While both variants are of the PR8 Mount Sinai type they exhibit different replication properties. The human alveolar epithelial cell line A549 and the corresponding A549 IFNAR II-1 shRNA and A549-nt-pLKO.1-2 cells (generated at the" ; ns1:mentions . _:b4304264 rdf:type ns1:Context ; rdf:value "It should be noted that this isogenic wild type strain as described by Garcia-Sastre et al. [68] is different from the PR8 (Giessen variant) used in the other experiments and in many previous studies [52],[>>67<<]. While both variants are of the PR8 Mount Sinai type they exhibit different replication properties. The human alveolar epithelial cell line A549 and the corresponding A549 IFNAR II-1 shRNA and A549-nt-pLKO.1-2 cells (generated at the RKI," ; ns1:mentions . _:b4304265 rdf:type ns1:Context ; rdf:value "To score for production of viral plaques the overlay was stained for 1 h using 1 ml neutral red in PBS per well [>>69<<]." ; ns1:mentions . _:b4304266 rdf:type ns1:Context ; rdf:value "A monoclonal antibody directed against the viral NS1 was generated at the IMV, Muenster, Germany [>>70<<]. A monoclonal anti-Myc-tag antibody to detect Myc-M1 was kindly provided by Viktor Wixler. IMV, Muenster, Germany. All secondary antibodies were purchased from Amersham and diluted 1\u22362500 in TBS-T. Secondary antibodies were incubated for" ; ns1:mentions . _:b4304267 rdf:type ns1:Context ; rdf:value "To ascertain changes in expression of the gene of interest the differences between expression of GAPDH and the gene of interest was calculated using the 2\u2212\u0394\u0394CT method [>>71<<]. For quantitative real time Brilliant QPCR SYBR Green Mastermix (Stratagene) was used according to manufacturer's instructions. The fragment of interest was amplified in 40 cycles. The following primers were used (see Table 1 for identity" ; ns1:mentions . _:b4304268 rdf:type ns1:Context ; rdf:value "The pCFG5-EGZ retroviral vector used for transfection [>>72<<] as well as the constructs to express dominant negative MKK6 (MKK6Ala) or IKK2 (IKK2KD) have been described earlier [52],[73]." ; ns1:mentions . _:b4304269 rdf:type ns1:Context ; rdf:value "The pCFG5-EGZ retroviral vector used for transfection [72] as well as the constructs to express dominant negative MKK6 (MKK6Ala) or IKK2 (IKK2KD) have been described earlier [>>52<<],[73]. The Phoenix amphotropic retroviral producer cells (a gift from G. Nolan, Stanford, CA) [74] were cultured in Dulbecco's modified Eagle's medium containing 10% fetal bovine serum, 100 units/ml penicillin and 100 mg/ml streptomycin." ; ns1:mentions . _:b4304270 rdf:type ns1:Context ; rdf:value "The pCFG5-EGZ retroviral vector used for transfection [72] as well as the constructs to express dominant negative MKK6 (MKK6Ala) or IKK2 (IKK2KD) have been described earlier [52],[>>73<<]. The Phoenix amphotropic retroviral producer cells (a gift from G. Nolan, Stanford, CA) [74] were cultured in Dulbecco's modified Eagle's medium containing 10% fetal bovine serum, 100 units/ml penicillin and 100 mg/ml streptomycin." ; ns1:mentions . _:b4304271 rdf:type ns1:Context ; rdf:value "Nolan, Stanford, CA) [>>74<<] were cultured in Dulbecco's modified Eagle's medium containing 10% fetal bovine serum, 100 units/ml penicillin and 100 mg/ml streptomycin." ; ns1:mentions . _:b4304272 rdf:type ns1:Context ; rdf:value "Generation of MKK6Ala or IKKKD expressing producer cells as well as transduction of A549 cells to stably express these transgenes was performed as previously described [>>52<<],[53]." ; ns1:mentions . _:b4304273 rdf:type ns1:Context ; rdf:value "Generation of MKK6Ala or IKKKD expressing producer cells as well as transduction of A549 cells to stably express these transgenes was performed as previously described [52],[>>53<<]." ; ns1:mentions . _:b4304274 rdf:type ns4:Section ; dc:title "results" ; ns4:contains _:b4304288 , _:b4304289 , _:b4304290 , _:b4304284 , _:b4304285 , _:b4304286 , _:b4304287 , _:b4304280 , _:b4304281 , _:b4304282 , _:b4304283 , _:b4304276 , _:b4304277 , _:b4304278 , _:b4304279 , _:b4304275 . _:b4304275 rdf:type ns1:Context ; rdf:value "Several protein tyrosine phosphatases (PTPs) are known to mediate dephosphorylation of both, JAKs and STATs [>>41<<]. In order to investigate whether influenza A virus activates phosphatases that subsequently target JAKs or STATs, we treated infected or uninfected A549 cells with the well-known tyrosine phosphatase inhibitor sodium vanadate [42],[43]." ; ns1:mentions . _:b4304276 rdf:type ns1:Context ; rdf:value "In order to investigate whether influenza A virus activates phosphatases that subsequently target JAKs or STATs, we treated infected or uninfected A549 cells with the well-known tyrosine phosphatase inhibitor sodium vanadate [>>42<<],[43]. Uninfected cells or cells infected with PR8 for 10 h were incubated with increasing amounts of this compound 10 min prior to stimulation with IFN\u03B2. This time point of infection was chosen since we observed considerable inhibition of" ; ns1:mentions . _:b4304277 rdf:type ns1:Context ; rdf:value "In order to investigate whether influenza A virus activates phosphatases that subsequently target JAKs or STATs, we treated infected or uninfected A549 cells with the well-known tyrosine phosphatase inhibitor sodium vanadate [42],[>>43<<]. Uninfected cells or cells infected with PR8 for 10 h were incubated with increasing amounts of this compound 10 min prior to stimulation with IFN\u03B2." ; ns1:mentions . _:b4304278 rdf:type ns1:Context ; rdf:value "SOCS proteins are described to have high affinity for JAK and STAT proteins and to inhibit the transmission of IFN\u03B1 and IFN\u03B2 induced signaling [>>44<<],[45]. To examine whether expression of SOCS genes is induced in influenza virus infected cells, A549 cells were infected with PR8 for different time points. Subsequently total RNA was analyzed for the amount of SOCS-1 and SOCS-3 mRNA by" ; ns1:mentions . _:b4304279 rdf:type ns1:Context ; rdf:value "SOCS proteins are described to have high affinity for JAK and STAT proteins and to inhibit the transmission of IFN\u03B1 and IFN\u03B2 induced signaling [44],[>>45<<]. To examine whether expression of SOCS genes is induced in influenza virus infected cells, A549 cells were infected with PR8 for different time points. Subsequently total RNA was analyzed for the amount of SOCS-1 and SOCS-3 mRNA by means" ; ns1:mentions . _:b4304280 rdf:type ns1:Context ; rdf:value "In contrast to cellular RNA, influenza viral RNA carries a triphosphate group at its 5\u2032 terminus that was previously shown to be a major pathogen pattern that triggers cellular signaling [>>46<<]. To verify that indeed the viral 5\u2032 triphosphate RNA in the pool of RNAs from infected cells is the major trigger for induction of SOCS-3 expression, RNA from infected or uninfected cells was treated with phosphatase to remove the 5\u2032" ; ns1:mentions . _:b4304281 rdf:type ns1:Context ; rdf:value "The MKK/p38 mitogen activated protein kinase (MAPK) pathway [>>47<<]\u2013[49] as well as the I\u03BAB kinase (IKK)/nuclear factor of \u03BAB (NF-\u03BAB) cascade [50]\u2013[52] are both known to be activated by RNA or influenza virus infection and to be involved in the control of SOCS-3 expression." ; ns1:mentions . _:b4304282 rdf:type ns1:Context ; rdf:value "The MKK/p38 mitogen activated protein kinase (MAPK) pathway [47]\u2013[>>49<<] as well as the I\u03BAB kinase (IKK)/nuclear factor of \u03BAB (NF-\u03BAB) cascade [50]\u2013[52] are both known to be activated by RNA or influenza virus infection and to be involved in the control of SOCS-3 expression." ; ns1:mentions . _:b4304283 rdf:type ns1:Context ; rdf:value "The MKK/p38 mitogen activated protein kinase (MAPK) pathway [47]\u2013[49] as well as the I\u03BAB kinase (IKK)/nuclear factor of \u03BAB (NF-\u03BAB) cascade [>>50<<]\u2013[52] are both known to be activated by RNA or influenza virus infection and to be involved in the control of SOCS-3 expression." ; ns1:mentions . _:b4304284 rdf:type ns1:Context ; rdf:value "The MKK/p38 mitogen activated protein kinase (MAPK) pathway [47]\u2013[49] as well as the I\u03BAB kinase (IKK)/nuclear factor of \u03BAB (NF-\u03BAB) cascade [50]\u2013[>>52<<] are both known to be activated by RNA or influenza virus infection and to be involved in the control of SOCS-3 expression." ; ns1:mentions . _:b4304285 rdf:type ns1:Context ; rdf:value "These mutants have been previously shown to efficiently block p38 or NF-\u03BAB signaling, respectively [>>52<<]\u2013[54]. To monitor SOCS-3 gene induction, wild type, vector or mutant expressing cell lines were infected with PR8 (Figure 6A) or stimulated with RNA from virally infected or uninfected A549 cells (Figure 6C). Induction of IFN\u03B2 mRNA was" ; ns1:mentions . _:b4304286 rdf:type ns1:Context ; rdf:value "These mutants have been previously shown to efficiently block p38 or NF-\u03BAB signaling, respectively [52]\u2013[>>54<<]. To monitor SOCS-3 gene induction, wild type, vector or mutant expressing cell lines were infected with PR8 (Figure 6A) or stimulated with RNA from virally infected or uninfected A549 cells (Figure 6C). Induction of IFN\u03B2 mRNA was" ; ns1:mentions . _:b4304287 rdf:type ns1:Context ; rdf:value "stably transduced with empty vector, dominant negative MKK6Ala or IKK2KD were either infected with PR8 for 3 hours (MOI\u200A=\u200A5) (A and B) or with the influenza A virus mutant \u0394NS1 and the corresponding isogenic wild type virus (G and H) [>>74<<] or transfected for 3 hours with RNA from infected or uninfected A549 cells (C\u2013F)." ; ns1:mentions . _:b4304288 rdf:type ns1:Context ; rdf:value "The NS1 protein is known to block RNA dependent signaling and NF\u03BAB activation [>>55<<]. Accordingly, infection of cells with the mutant virus resulted in a more pronounced and sustained, albeit delayed induction of SOCS-3 (Figure 6G) if compared to infection with the isogenic wild type, that is a very poor inducer of SOCS-3" ; ns1:mentions . _:b4304289 rdf:type ns1:Context ; rdf:value "To further assess a functional role of SOCS-3 in virus-induced suppression of STAT1 phosphorylation we analyzed mouse cells with a targeted deletion of the SOCS-3 gene [>>56<<]. Wild type and SOCS-3 deficient mouse embryonic fibroblasts (MEF) were infected for different time points with PR8." ; ns1:mentions . _:b4304290 rdf:type ns1:Context ; rdf:value "These genes are described as type I IFN-induced genes [>>18<<]. Indeed mRNA levels of all three representative ISGs were elevated in SOCS-3 knock out versus wild type cells at almost every time point during the course of infection." ; ns1:mentions . _:b4304291 rdf:type ns4:Section ; dc:title "discussion" ; ns4:contains _:b4304296 , _:b4304297 , _:b4304298 , _:b4304299 , _:b4304300 , _:b4304301 , _:b4304302 , _:b4304303 , _:b4304292 , _:b4304293 , _:b4304294 , _:b4304295 , _:b4304304 , _:b4304305 , _:b4304306 , _:b4304307 , _:b4304308 , _:b4304309 , _:b4304310 , _:b4304311 . _:b4304292 rdf:type ns1:Context ; rdf:value "Influenza virus propagation is sensitive to IFN activities and therefore, like other viral pathogens, these viruses do not only induce type I IFN but also antagonize the production and effects of these cytokines at the same time [>>55<<]. For influenza A and B viruses, this is accomplished through their non-structural NS1 proteins that are structurally related polypeptides of 26 kDa (A/NS1) and 32 kDa (B/NS1), which are abundantly expressed in infected cells [55]." ; ns1:mentions . _:b4304293 rdf:type ns1:Context ; rdf:value "For influenza A and B viruses, this is accomplished through their non-structural NS1 proteins that are structurally related polypeptides of 26 kDa (A/NS1) and 32 kDa (B/NS1), which are abundantly expressed in infected cells [>>55<<]. NS1 proteins predominantly act on the level of IFN gene induction in infected cells by obstructing RIG-I-dependent signaling through interaction with cellular factor(s) and/or sequestration of RNAs generated during virus replication" ; ns1:mentions . _:b4304294 rdf:type ns1:Context ; rdf:value "NS1 proteins predominantly act on the level of IFN gene induction in infected cells by obstructing RIG-I-dependent signaling through interaction with cellular factor(s) and/or sequestration of RNAs generated during virus replication [>>1<<],[2],[57]. Some NS1 proteins were also described to inhibit the maturation of cellular pre-mRNAs raising the possibility that this activity additionally reduces production of IFN\u03B1/\u03B2 in infected cells [58],[59]." ; ns1:mentions . _:b4304295 rdf:type ns1:Context ; rdf:value "proteins predominantly act on the level of IFN gene induction in infected cells by obstructing RIG-I-dependent signaling through interaction with cellular factor(s) and/or sequestration of RNAs generated during virus replication [1],[>>2<<],[57]. Some NS1 proteins were also described to inhibit the maturation of cellular pre-mRNAs raising the possibility that this activity additionally reduces production of IFN\u03B1/\u03B2 in infected cells [58],[59]." ; ns1:mentions . _:b4304296 rdf:type ns1:Context ; rdf:value "predominantly act on the level of IFN gene induction in infected cells by obstructing RIG-I-dependent signaling through interaction with cellular factor(s) and/or sequestration of RNAs generated during virus replication [1],[2],[>>57<<]. Some NS1 proteins were also described to inhibit the maturation of cellular pre-mRNAs raising the possibility that this activity additionally reduces production of IFN\u03B1/\u03B2 in infected cells [58],[59]." ; ns1:mentions . _:b4304297 rdf:type ns1:Context ; rdf:value "Some NS1 proteins were also described to inhibit the maturation of cellular pre-mRNAs raising the possibility that this activity additionally reduces production of IFN\u03B1/\u03B2 in infected cells [>>58<<],[59]. While NS1 also interferes with the activity of some ISGs, such as the dsRNA dependent kinase PKR [5],[60], so far no type I IFN antagonistic mechanism was described for influenza viruses that act on the level of IFN signaling rather" ; ns1:mentions . _:b4304298 rdf:type ns1:Context ; rdf:value "Some NS1 proteins were also described to inhibit the maturation of cellular pre-mRNAs raising the possibility that this activity additionally reduces production of IFN\u03B1/\u03B2 in infected cells [58],[>>59<<]. While NS1 also interferes with the activity of some ISGs, such as the dsRNA dependent kinase PKR [5],[60], so far no type I IFN antagonistic mechanism was described for influenza viruses that act on the level of IFN signaling rather than" ; ns1:mentions . _:b4304299 rdf:type ns1:Context ; rdf:value "While NS1 also interferes with the activity of some ISGs, such as the dsRNA dependent kinase PKR [>>5<<],[60], so far no type I IFN antagonistic mechanism was described for influenza viruses that act on the level of IFN signaling rather than gene induction." ; ns1:mentions . _:b4304300 rdf:type ns1:Context ; rdf:value "While NS1 also interferes with the activity of some ISGs, such as the dsRNA dependent kinase PKR [5],[>>60<<], so far no type I IFN antagonistic mechanism was described for influenza viruses that act on the level of IFN signaling rather than gene induction." ; ns1:mentions . _:b4304301 rdf:type ns1:Context ; rdf:value "While it was reported in the literature that expression of SOCS proteins can be induced upon stimulation with IFN [>>61<<] we could not detect any significant gene induction by IFN\u03B2 in A549 cells." ; ns1:mentions . _:b4304302 rdf:type ns1:Context ; rdf:value "We hypothesized previously that the incomplete inhibition conferred by NS1 is an indication that the virus exploits the remaining signaling activities for efficient replication [>>52<<],[62],[63]." ; ns1:mentions . _:b4304303 rdf:type ns1:Context ; rdf:value "Recently it was reported that IFN-induced gene expression responses are potentiated in cells, which lack the NF-\u03BAB factors p50 or p65 [>>64<<]. Although these authors described an inhibitory binding of NF-\u03BAB transcription factors to some IFN-induced gene, this mechanism might be cell type dependent since we could not observe similar effects in the cell types used here (data not" ; ns1:mentions . _:b4304304 rdf:type ns1:Context ; rdf:value "Thus, given the NF-\u03BAB dependent induction of SOCS-3 described in the present manuscript, we provide an additional molecular mechanism that may explain the phenomenon described by Wei et al. [>>64<<]." ; ns1:mentions . _:b4304305 rdf:type ns1:Context ; rdf:value "for beneficial effects of SOCS-3 gene expression on viral replication came from studies using the HCV core protein as a replacement for the influenza A viral NS1 in the context of infections with a NS1 deficient influenza virus [>>33<<]. One of the hallmark responses of HCV core expression is a rapid induction of SOCS-3 expression." ; ns1:mentions . _:b4304306 rdf:type ns1:Context ; rdf:value "Given the role of SOCS-3 described here, it was not surprising that HCV core could partially rescue growth of the NS1 deficient virus [>>33<<]." ; ns1:mentions . _:b4304307 rdf:type ns1:Context ; rdf:value "While this manuscript was in preparation it was demonstrated by Pothlichet et al. that influenza A virus-induced SOCS-1 and SOCS-3 upregulation requires a TLR-3-independent, RIG-I/MAVS-dependent pathway [>>65<<]. Moreover, over-expression of SOCS-1 and SOCS-3 in infected cells revealed that both molecules inhibit antiviral responses." ; ns1:mentions . _:b4304308 rdf:type ns1:Context ; rdf:value "It is well known that the capability of type I IFNs to induce SOCS proteins is strongly cell type specific [>>31<<]. While in some cell types SOCS-3 expression appears to be type I IFN dependent (e.g. fetal liver cells) [31] it is clearly independent of IFN in other cell types [66]. Recently it was shown that SOCS-3 is not significantly induced by IFN\u03B1" ; ns1:mentions . _:b4304309 rdf:type ns1:Context ; rdf:value "While in some cell types SOCS-3 expression appears to be type I IFN dependent (e.g. fetal liver cells) [>>31<<] it is clearly independent of IFN in other cell types [66]." ; ns1:mentions . _:b4304310 rdf:type ns1:Context ; rdf:value "While in some cell types SOCS-3 expression appears to be type I IFN dependent (e.g. fetal liver cells) [31] it is clearly independent of IFN in other cell types [>>66<<]. Recently it was shown that SOCS-3 is not significantly induced by IFN\u03B1 in A549 cells [18], the major cell type used in our study. Evidence that cell type specificities may be the cause of discrepancy is additionally provided by the fact" ; ns1:mentions . _:b4304311 rdf:type ns1:Context ; rdf:value "Recently it was shown that SOCS-3 is not significantly induced by IFN\u03B1 in A549 cells [>>18<<], the major cell type used in our study." ; ns1:mentions . _:b389835269 rdf:type ns1:RelevantBibliographicResource . @prefix xsd: . _:b389835269 ns1:RelevantScore "22"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b389835270 rdf:type ns1:RelevantBibliographicResource ; ns1:RelevantScore "16"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b389835271 rdf:type ns1:RelevantBibliographicResource ; ns1:RelevantScore "12"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b389835272 rdf:type ns1:RelevantBibliographicResource ; ns1:RelevantScore "12"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b389835273 rdf:type ns1:RelevantBibliographicResource ; ns1:RelevantScore "10"^^xsd:nonNegativeInteger ; ns1:hasRelevantPaperId . _:b389835274 rdf:type ns1:RelevantBibliographicResource ; 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