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Upon cytokine binding to their cognate receptor, the receptor-associated Jaks are activated and in turn phosphorylate tyrosine residues in the receptor cytoplasmic domain.</csml:comment>
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Upon cytokine binding to their cognate receptor, the receptor-associated Jaks are activated and in turn phosphorylate tyrosine residues in the receptor cytoplasmic domain.</csml:comment>
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This event provides a docking site for proteins with Src homology 2 domains, one important class of which is the Stat family of transcription factors.</csml:comment>
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With seven members in all (Stat1, Stat2, Stat3, Stat4, Stat5a, Stat5b, and Stat6), these DNA-binding proteins provide a rapid membrane to nucleus mechanism for regulation of gene expression.</csml:comment>
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With seven members in all (Stat1, Stat2, Stat3, Stat4, Stat5a, Stat5b, and Stat6), these DNA-binding proteins provide a rapid membrane to nucleus mechanism for regulation of gene expression.</csml:comment>
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<csml:comment type="text">PMID: 18400190
As an example of the overall importance of cytokine-cytokine receptor-Jak-Stat pathway signaling in thymic T cell development, IL-7 signaling ensures development of appropriate lymphocyte numbers.

PMID: 18400190
Stat5 signaling from cytokines that use γc is essential for T cell development.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Stat5 signaling from cytokines that use γc is essential for T cell development.

PMID: 18400190
Socs3 is an important inhibitor of cytokines that use gp130 (IL-23R and IL-6R), whereas Socs1 is anticipated to inhibit any cytokines that use gamma-c (IL-7, IL-21 as shown).</csml:comment>
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<csml:comment type="text">PMID: 18400190, 16418296
IL-7 activates Stat5a and Stat5b, and deletion of the locus encoding Stat5a and Stat5b also results in a severe SCID phenotype.

PMID: 18400190
Socs3 is an important inhibitor of cytokines that use gp130 (IL-23R and IL-6R), whereas Socs1 is anticipated to inhibit any cytokines that use gamma-c (IL-7, IL-21 as shown).</csml:comment>
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<csml:comment type="text">PMID: 18400190, 16418296
IL-7 activates Stat5a and Stat5b, and deletion of the locus encoding Stat5a and Stat5b also results in a severe SCID phenotype.

PMID: 18400190
Socs3 is an important inhibitor of cytokines that use gp130 (IL-23R and IL-6R), whereas Socs1 is anticipated to inhibit any cytokines that use gamma-c (IL-7, IL-21 as shown).</csml:comment>
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<csml:comment type="text">PMID: 18400190
Stat5 signaling from cytokines that use γc is essential for T cell development.

PMID: 18400190, 16455959
Deficiency of gamma-c or Jak3 causes a failure to produce Foxp3-positive regulatory T cells.

PMID: 18400190
On one hand, Socs1 regulates T cell development by inhibiting gamma-c signaling.</csml:comment>
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<csml:comments>
<csml:comment type="text">PMID: 18400190
Stat5 signaling controls the development of FoxP3-positive Treg cells in part through the direct activation of Foxp3 gene expression.

PMID: 18400190
Stat5 appears to have very direct effects on Treg cells in that these transcription factors bind directly to the Foxp3 gene.</csml:comment>
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<csml:comment type="text">PMID: 18400190
IL-12 activates Stat4, whereas IL-4 activates Stat6.</csml:comment>
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<csml:comment type="text">PMID: 18400190
IL-12 activates Stat4, whereas IL-4 activates Stat6.</csml:comment>
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<csml:comment type="text">PMID: 18400190
These fates are driven by the cytokine milieu with IL-12 driving Th1 cell differentiation and IL-4 promoting Th2 cell differentiation.

PMID: 18400190, 11121747
Stat4- and Stat6-deficient mice have impaired Th1 and Th2 cell responses, respectively.

PMID: 18400190, 12242445
Depending upon the circumstance, type I IFN signaling may enhance or inhibit Th1 cell responses.</csml:comment>
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<csml:comment type="text">Indirect</csml:comment>
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<csml:comment type="text">PMID: 18400190
These fates are driven by the cytokine milieu with IL-12 driving Th1 cell differentiation and IL-4 promoting Th2 cell differentiation.

PMID: 18400190, 11121747
Stat4- and Stat6-deficient mice have impaired Th1 and Th2 cell responses, respectively.</csml:comment>
</csml:comments>
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<csml:comment type="text">PMID: 18400190
In this regard, it is important to note that type I IFNs also activate Stat4.</csml:comment>
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<csml:comments>
<csml:comment type="text"/>
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<csml:comment type="text">PMID: 18400190
Stat5 appears to have very direct effects on Treg cells in that these transcription factors bind directly to the Foxp3 gene.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Stat5 appears to have very direct effects on Treg cells in that these transcription factors bind directly to the Foxp3 gene.</csml:comment>
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<csml:comment type="text">PMID: 18400190, 17227828
Accordingly, deficiency of both Stat5a and Stat5b also leads to loss of Treg cells and inability to induce Treg cells in vitro.

PMID: 18400190
In both cases, cytokines that use gamma-c are important drivers of Treg cell development.</csml:comment>
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<csml:comment type="text"/>
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<csml:comment type="text">PMID: 18400190, 17581589, 17581537
Th17 cells can be generated from naive CD4+ T cells by IL-6 and TGF-beta but can also produce another cytokine IL-21, which promotes IL-17 production in an autocrine-paracrine manner.</csml:comment>
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<csml:comment type="text">PMID: 18400190
IL-6, IL-21, and IL-23 all activate Stat3 via their cognate receptors.</csml:comment>
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<csml:comment type="text">PMID: 18400190
IL-6, IL-21, and IL-23 all activate Stat3 via their cognate receptors.</csml:comment>
</csml:comments>
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<csml:comment type="text">PMID: 18400190
IL-6, IL-21, and IL-23 all activate Stat3 via their cognate receptors.</csml:comment>
</csml:comments>
</csml:process>
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<csml:comments>
<csml:comment type="text"/>
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</csml:connector>
<csml:connector id="c68" name="c68" refID="e32" type="cso30:c:InputProcess">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
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</csml:connectorSimulationProperty>
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</csml:connector>
<csml:connector id="c66" name="c66" refID="e34" type="cso30:c:OutputProcess">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
<csml:parameter key="stoichiometry" value="1"/>
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</csml:connectorSimulationProperty>
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</csml:toolSpecificGraphics>
</csml:shape>
</csml:viewProperty>
<csml:comments>
<csml:comment type="text"/>
</csml:comments>
</csml:connector>
<csml:processSimulationProperty>
<csml:priority value="0"/>
<csml:firing firingOnce="false" firingStyle="csml-firingStyle:and" type="csml-variable:Boolean" value="true"/>
<csml:delay delayStyle="nodelay" value="0.0"/>
<csml:processKinetic calcStyle="csml-calcStyle:speed" fast="false" kineticStyle="csml-kineticStyle:mass">
<csml:parameter key="coefficient1" value="0.1"/>
<csml:parameter key="coefficient2" value="1.0"/>
</csml:processKinetic>
</csml:processSimulationProperty>
<csml:viewProperty viewID="">
<csml:position position="auto" positionID="default" x="680.0" y="1098.0"/>
<csml:shape shapeID="default" visible="true">
<csml:image fileFormat="svg" filePath="" fileType="resourceFile" height="23.0" outlineColor="0 0 0 0" resourceID="10005" width="23.0"/>
<csml:toolSpecificGraphics/>
</csml:shape>
</csml:viewProperty>
<csml:biologicalProperty refBiologicalEventID="cso30:i:ME_Binding" refCellComponentID="cso30:i:CC_Nucleoplasm"/>
<csml:comments>
<csml:comment type="text">PMID: 18400190, 16698929, 17884812
However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
</csml:comments>
</csml:process>
<csml:process id="p26" name="p26" type="cso30:c:ProcessBiological">
<csml:connector id="c65" name="c65" refID="e31" type="cso30:c:InputProcess">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
<csml:parameter key="stoichiometry" value="1"/>
</csml:connectorKinetic>
</csml:connectorSimulationProperty>
<csml:viewProperty viewID="">
<csml:position position="auto" positionID="default" x="529.0" y="1089.0"/>
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</csml:toolSpecificGraphics>
</csml:shape>
</csml:viewProperty>
<csml:comments>
<csml:comment type="text"/>
</csml:comments>
</csml:connector>
<csml:connector id="c69" name="c69" refID="e33" type="cso30:c:InputProcess">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
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</csml:connector>
<csml:connector id="c67" name="c67" refID="e35" type="cso30:c:OutputProcess">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
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</csml:toolSpecificGraphics>
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<csml:comments>
<csml:comment type="text"/>
</csml:comments>
</csml:connector>
<csml:processSimulationProperty>
<csml:priority value="0"/>
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<csml:processKinetic calcStyle="csml-calcStyle:speed" fast="false" kineticStyle="csml-kineticStyle:mass">
<csml:parameter key="coefficient1" value="0.1"/>
<csml:parameter key="coefficient2" value="1.0"/>
</csml:processKinetic>
</csml:processSimulationProperty>
<csml:viewProperty viewID="">
<csml:position position="auto" positionID="default" x="544.0" y="1104.0"/>
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<csml:biologicalProperty refBiologicalEventID="cso30:i:ME_Binding" refCellComponentID="cso30:i:CC_Nucleoplasm"/>
<csml:comments>
<csml:comment type="text">PMID: 18400190, 16698929, 17884812
However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
</csml:comments>
</csml:process>
<csml:process id="p27" name="p27" type="cso30:c:ProcessBiological">
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<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
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</csml:connectorKinetic>
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<csml:comments>
<csml:comment type="text"/>
</csml:comments>
</csml:connector>
<csml:connector id="c63" name="c63" refID="e31" type="cso30:c:OutputProcess">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
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</csml:connectorSimulationProperty>
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</csml:viewProperty>
<csml:comments>
<csml:comment type="text"/>
</csml:comments>
</csml:connector>
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<csml:priority value="0"/>
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<csml:processKinetic calcStyle="csml-calcStyle:speed" fast="false" kineticStyle="csml-kineticStyle:mass">
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</csml:processKinetic>
</csml:processSimulationProperty>
<csml:viewProperty viewID="">
<csml:position position="auto" positionID="default" x="443.0" y="1151.0"/>
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<csml:toolSpecificGraphics/>
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<csml:biologicalProperty refBiologicalEventID="cso30:i:ME_Translocation" refCellComponentID="cso30:i:CC_Cytosol"/>
<csml:comments>
<csml:comment type="text">PMID: 18400190, 16698929, 17884812
However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
</csml:comments>
</csml:process>
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<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
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</csml:viewProperty>
<csml:comments>
<csml:comment type="text">Indirect</csml:comment>
</csml:comments>
</csml:connector>
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<csml:connectorSimulationProperty>
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<csml:comments>
<csml:comment type="text"/>
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<csml:biologicalProperty refBiologicalEventID="cso30:i:ME_GeneExpression" refCellComponentID="cso30:i:CC_Nucleoplasm"/>
<csml:comments>
<csml:comment type="text">PMID: 18400190, 16698929, 17884812
However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
</csml:comments>
</csml:process>
<csml:process id="p29" name="p29" type="cso30:c:ProcessBiological">
<csml:connector id="c73" name="c73" refID="e35" type="cso30:c:InputAssociation">
<csml:connectorSimulationProperty>
<csml:connectorFiring connectorFiringStyle="csml-connectorFiringStyle:threshold" value="0"/>
<csml:connectorKinetic>
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</csml:connectorKinetic>
</csml:connectorSimulationProperty>
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<csml:comments>
<csml:comment type="text">Indirect</csml:comment>
</csml:comments>
</csml:connector>
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<csml:connectorSimulationProperty>
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<csml:comments>
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However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
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Accordingly, selective deletion of Stat3 in T cells abrogates Th17 cell differentiation in part because the expression of RORgammat and RORalpha, two nuclear hormone receptors essential for Th17 cell development, is also abrogated.

PMID: 18400190
IL-23 and IL-6 enforce Th17 cell development via the direct or indirect induction of Rorc and Rora expression.</csml:comment>
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Accordingly, selective deletion of Stat3 in T cells abrogates Th17 cell differentiation in part because the expression of RORgammat and RORalpha, two nuclear hormone receptors essential for Th17 cell development, is also abrogated.

PMID: 18400190
IL-23 and IL-6 enforce Th17 cell development via the direct or indirect induction of Rorc and Rora expression.</csml:comment>
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Accordingly, selective deletion of Stat3 in T cells abrogates Th17 cell differentiation in part because the expression of RORgammat and RORalpha, two nuclear hormone receptors essential for Th17 cell development, is also abrogated.

PMID: 18400190
IL-23 and IL-6 enforce Th17 cell development via the direct or indirect induction of Rorc and Rora expression.

PMID: 18400190, 17363300
Parenthetically, it is interesting to note that IL-2 acting through Stat5 inhibits Th17 cell differentiation.</csml:comment>
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<csml:comment type="text">PMID: 18400190, 17363300
Parenthetically, it is interesting to note that IL-2 acting through Stat5 inhibits Th17 cell differentiation.</csml:comment>
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<csml:comment type="text">PMID: 18400190
whereas IL-6 and TGF-beta-1 are important for the differentiation of Th17 cells, IL-6 alone so far appears to be capable of inducing IL-22.</csml:comment>
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<csml:comment type="text">PMID: 18400190
whereas IL-6 and TGF-beta-1 are important for the differentiation of Th17 cells, IL-6 alone so far appears to be capable of inducing IL-22.</csml:comment>
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<csml:comment type="text">PMID: 18400190
An additional complexity of Th17 T cells in inflammation concerns the Stat3-activating cytokine IL-22.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Importantly, the genes encoding the Socs proteins are direct targets of Stat proteins; the Jak-Stat cascades thereby control their own signaling output by feedback inhibition.</csml:comment>
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<csml:comment type="text">PMID: 18400190, 12076535, 10051596
Each of the eight Socs proteins have two major domains, an SH2 domain and a Socs box that complexes with elongins B and C, a cullin and Rbx2, to form a E3 ubiquitin ligase.</csml:comment>
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The Socs SH2 domains bind phosphorylated tyrosine residues in their substrates. The best characterized Socs substrates are specific tyrosine residues in the cytoplasmic tails of cytokine receptors.</csml:comment>
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The current model of Socs function postulates that the E3 activity of a Socs protein will target the substrate to be ubiquitinated and then directed to the proteosome for degradation.</csml:comment>
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The current model of Socs function postulates that the E3 activity of a Socs protein will target the substrate to be ubiquitinated and then directed to the proteosome for degradation.</csml:comment>
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The G-CSFR is responsible for transducing the signals from G-CSF via four tyrosine residues located in the cytoplasmic tail of the receptor.</csml:comment>
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G-CSFR signaling via the cytoplasmic tyrosines activates numerous signaling molecules including Stat5, Stat3, and MAP kinases.</csml:comment>
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G-CSFR signaling via the cytoplasmic tyrosines activates numerous signaling molecules including Stat5, Stat3, and MAP kinases.

PMID: 18400190, 12133942
Indeed, Socs3 binds to one of the tyrosine residues in the G-CSFR (Y729) and restricts the amplitude of Stat3 signaling.</csml:comment>
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G-CSFR signaling via the cytoplasmic tyrosines activates numerous signaling molecules including Stat5, Stat3, and MAP kinases.</csml:comment>
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Indeed, Socs3 binds to one of the tyrosine residues in the G-CSFR (Y729) and restricts the amplitude of Stat3 signaling.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Thus, loss of Socs3 causes increased G-CSFR signaling leading to increased neutrophil numbers, whereas loss of Stat3 (and failure to induce Socs3 expression) also leads to increased neutrophil numbers.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Thus, loss of Socs3 causes increased G-CSFR signaling leading to increased neutrophil numbers, whereas loss of Stat3 (and failure to induce Socs3 expression) also leads to increased neutrophil numbers.

PMID: 18400190
Socs3 expression is strongly induced by IL-10, along with the Stat3-dependent genes whose products regulate the anti-inflammatory signaling system (“anti-inflammatory response” AIR gene whose identity has yet to be determined) illustrated as inhibiting the expression at the transcriptional level of classic pro-inflammatory genes.</csml:comment>
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Socs3 expression is strongly induced by IL-10, along with the Stat3-dependent genes whose products regulate the anti-inflammatory signaling system (“anti-inflammatory response” AIR gene whose identity has yet to be determined) illustrated as inhibiting the expression at the transcriptional level of classic pro-inflammatory genes.</csml:comment>
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Socs3 expression is strongly induced by IL-10, along with the Stat3-dependent genes whose products regulate the anti-inflammatory signaling system (“anti-inflammatory response” AIR gene whose identity has yet to be determined) illustrated as inhibiting the expression at the transcriptional level of classic pro-inflammatory genes.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Gp130 has multiple tyrosine residues in its cytoplasmic tail, all of which bar one, Y757, serve as docking sites for Stat proteins (especially Stat3)

PMID: 18400190
Socs3 is an important inhibitor of cytokines that use gp130 (IL-23R and IL-6R), whereas Socs1 is anticipated to inhibit any cytokines that use gamma-c (IL-7, IL-21 as shown).</csml:comment>
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<csml:comment type="text">PMID: 18400190
Gp130 has multiple tyrosine residues in its cytoplasmic tail, all of which bar one, Y757, serve as docking sites for Stat proteins (especially Stat3)

PMID: 18400190
Socs3 is an important inhibitor of cytokines that use gp130 (IL-23R and IL-6R), whereas Socs1 is anticipated to inhibit any cytokines that use gamma-c (IL-7, IL-21 as shown).</csml:comment>
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However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
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However, Stat3 also directly regulates the expression of Il21 and Il17.</csml:comment>
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Y757 docks the SH2 domain of Socs3 and is by far the best-characterized Socs-cytokine receptor interaction.</csml:comment>
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Y757 docks the SH2 domain of Socs3 and is by far the best-characterized Socs-cytokine receptor interaction.</csml:comment>
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<csml:comment type="text">PMID: 18400190, 18276893
IL-22 induces the expression of antimicrobial proteins, including S1008A, a zinc and manganese chelating protein that deprives bacteria of essential cations.</csml:comment>
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Socs1, unlike the broad expression of Socs3, is activated predominantly by interferon signaling, although other cytokines such as IL-4 also activate Socs1 expression but in a cell-type-dependent way.</csml:comment>
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Because TLR signaling induces autocrine-paracrine IFN-alpha-beta production, this might underlie the cause of LPS sensitivity in Socs1−/−; Ifng−/− mice.</csml:comment>
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Because TLR signaling induces autocrine-paracrine IFN-alpha-beta production, this might underlie the cause of LPS sensitivity in Socs1−/−; Ifng−/− mice.</csml:comment>
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Mansell et al. have demonstrated that Socs1 can bind to and regulate the degradation of Mal (also known as TIRAP), an adaptor molecule specifically associated with TLR2 and TLR4 signaling.</csml:comment>
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Previous studies had shown that the tyrosine kinase Btk phosphorylates Mal, providing binding sites for Socs1.</csml:comment>
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Mansell et al. have demonstrated that Socs1 can bind to and regulate the degradation of Mal (also known as TIRAP), an adaptor molecule specifically associated with TLR2 and TLR4 signaling.</csml:comment>
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The chemokine CXCL12 activates CXCR4 and induces phosphorylation of FAK, a ubiquitous tyrosine kinase, in addition to other pathways.

PMID: 18400190, 12163560
Therefore, the assignment of FAK as a substrate of Socs3 potentially explains previous data that suggested that Socs3 directly regulates CXCR4 to inhibit its activity.</csml:comment>
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<csml:comment type="text">PMID: 18400190
The chemokine CXCL12 activates CXCR4 and induces phosphorylation of FAK, a ubiquitous tyrosine kinase, in addition to other pathways.

PMID: 18400190
Le et al. have shown that Socs3 is crucial to regulate the amounts of CXCL12-activated phospho-FAK because Socs3-deficient B cells have increased amount of phospho-FAK.</csml:comment>
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<csml:comment type="text">PMID: 18400190, 12387736
Link and colleagues have also postulated that signals from the G-CSFR negatively regulate CXCL12 amounts in the bone marrow, thereby allowing escape of neutrophils into the circulation.</csml:comment>
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<csml:comment type="text">PMID: 18400190
Addressing this question is complicated by the diversity of stimuli and downstream signaling molecules that contribute to myeloid IL-10 production including TLR agonists via TLRs and the p38 MAP kinase pathways and zymosan via the dectin and ERK pathways, to list a few.</csml:comment>
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Addressing this question is complicated by the diversity of stimuli and downstream signaling molecules that contribute to myeloid IL-10 production including TLR agonists via TLRs and the p38 MAP kinase pathways and zymosan via the dectin and ERK pathways, to list a few.</csml:comment>
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Addressing this question is complicated by the diversity of stimuli and downstream signaling molecules that contribute to myeloid IL-10 production including TLR agonists via TLRs and the p38 MAP kinase pathways and zymosan via the dectin and ERK pathways, to list a few.</csml:comment>
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Addressing this question is complicated by the diversity of stimuli and downstream signaling molecules that contribute to myeloid IL-10 production including TLR agonists via TLRs and the p38 MAP kinase pathways and zymosan via the dectin and ERK pathways, to list a few.</csml:comment>
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<csml:comment type="text">PMID: 18400190, 18400189
Aspects of this complex regulatory pathway including the role of IL-27 in driving IL-10 production via Stat1 and Stat3 are covered in the review by Li and Flavell, (2008).</csml:comment>
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