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type=\u0022text\/css\u0022 rel=\u0022stylesheet\u0022 href=\u0022\/\/d282kpwvnogo5m.cloudfront.net\/sites\/default\/files\/advagg_css\/css__ce2QY63WIanKyr8eSq7eavr1XQRRmFD6ZSmwpyJi8lM__zXwFqpqmxrZOXXcd_TpBQpjuELbmIP9wBR5UuTDWAO4__YJWWMMdfCJuAFm5cUEp88OsodhO3ZA-2lzRfoBsSlk4.css\u0022 media=\u0022all\u0022 \/\u003E\n\u003Clink rel=\u0027stylesheet\u0027 type=\u0027text\/css\u0027 href=\u0027\/sites\/all\/modules\/contrib\/panels\/plugins\/layouts\/onecol\/onecol.css\u0027 \/\u003E\u003C\/head\u003E\u003Cbody\u003E\u003Cdiv class=\u0022panels-ajax-tab-panel panels-ajax-tab-panel-sageoa-tab-art\u0022\u003E\u003Cdiv class=\u0022panel-display panel-1col clearfix\u0022 \u003E\n  \u003Cdiv class=\u0022panel-panel panel-col\u0022\u003E\n    \u003Cdiv\u003E\u003Cdiv class=\u0022panel-pane pane-highwire-markup\u0022 \u003E\n  \n      \n  \n  \u003Cdiv class=\u0022pane-content\u0022\u003E\n    \u003Cdiv class=\u0022highwire-markup\u0022\u003E\u003Cdiv xmlns=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022 id=\u0022content-block-markup\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cdiv class=\u0022article fulltext-view \u0022\u003E\u003Cspan class=\u0022highwire-journal-article-marker-start\u0022\u003E\u003C\/span\u003E\u003Cdiv class=\u0022section abstract\u0022 id=\u0022abstract-1\u0022\u003E\u003Ch2\u003ESummary\u003C\/h2\u003E\n            \u003Cp id=\u0022p-1\u0022\u003ENew research findings can now explain the complete pathway for the regulation of thrombopoietin production, which regulates the production of platelets. Platelets lacking sialic acid due to aging are removed by the Ashwell-Morell receptor. The signaling pathways Janus kinase 2 and signal transducer and activator of transcription 3 then stimulates the production of thrombopoietin within the hepatocyte nucleus.\u003C\/p\u003E\n         \u003C\/div\u003E\u003Cul class=\u0022kwd-group\u0022\u003E\u003Cli class=\u0022kwd\u0022\u003EAshwell-Morell receptor\u003C\/li\u003E\u003Cli class=\u0022kwd\u0022\u003Edesialyted platelets\u003C\/li\u003E\u003Cli class=\u0022kwd\u0022\u003Ehuman hepatoma cells\u003C\/li\u003E\u003Cli class=\u0022kwd\u0022\u003Ein vitro\u003C\/li\u003E\u003C\/ul\u003E\u003Cdiv class=\u0022section\u0022 id=\u0022sec-1\u0022\u003E\n         \n         \u003Cp id=\u0022p-2\u0022\u003EThe Ashwell-Morell receptor (AMR) on the hepatocyte binds and removes desialylated platelets as they circulate and age in the blood. After binding to the AMR, the desialylated platelets induce hepatic expression of thrombopoietin (TPO) mRNA, thereby regulating platelet production. Renata Grozovsky, PhD, Brigham and Women\u2019s Hospital, Boston, Massachusetts, USA, presented data from a study [Grozovsky R et al. \u003Cem\u003ENat Medicine.\u003C\/em\u003E 2015] showing the TPO expression by way of the AMR is further regulated by Janus kinase 2 (JAK2) and signal transducer and activator of transcription 3 (STAT3) protein signaling in vivo and in vitro.\u003C\/p\u003E\n         \u003Cp id=\u0022p-3\u0022\u003EIn 1994, 4 studies identified ligands that significantly increased platelet levels [Bartley TD et al. \u003Cem\u003ECell.\u003C\/em\u003E 1994; de Sauvage FJ et al. \u003Cem\u003ENature.\u003C\/em\u003E 1994; Kuter DJ et al. \u003Cem\u003EProc Natl Acad Sci USA\u003C\/em\u003E. 1994; Lok S et al. \u003Cem\u003ENature.\u003C\/em\u003E 1994]. The substance was subsequently identified as TPO, a glycoprotein hormone produced by the liver and kidney that in humans is encoded by the TPO gene. TPO regulates platelet production, supporting the survival, proliferation, and differentiation of platelet precursors and bone marrow megakaryocytes. The complete understanding of this pathway remained elusive until Dr Grozovsky\u2019s research.\u003C\/p\u003E\n         \u003Cp id=\u0022p-4\u0022\u003EIn the in vitro treatment, desialylated platelets were shown to stimulate TPO production in human hepatoma cells (HepG2). The data indicate that following platelet desialylation, platelet ingestion by the AMR, TPO mRNA and the TPO protein significantly increase within 6 hours (\u003Cem\u003EP\u2009\u003C\/em\u003E\u0026lt;\u2009.01) [Grozovsky R et al. \u003Cem\u003ENat Medicine\u003C\/em\u003E 2015].\u003C\/p\u003E\n         \u003Cp id=\u0022p-5\u0022\u003EDesialylated human platelets were also shown to activate JAK2-STAT3 signaling in HepG2 cells (\u003Ca id=\u0022xref-fig-1-1\u0022 class=\u0022xref-fig\u0022 href=\u0022#F1\u0022\u003EFigure 1\u003C\/a\u003E). Phospho-STAT3 was translocated into the nucleus and TPO protein expression increased as well. Both of the latter were prevented by pharmacological JAK inhibition.\u003C\/p\u003E\n         \u003Cdiv id=\u0022F1\u0022 class=\u0022fig pos-float  odd\u0022\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F1.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022Desialylated Human Platelets Stimulate JAK2 and STAT3 Phosphorylation in HepG2 CellsJACK2, Janus kinase 2; STAT3, activator of transcription 3.*P\u0026#x2009;\u0026amp;lt;\u0026#x2009;.05, **P\u0026#x2009;\u0026amp;lt;\u0026#x2009;.01Reproduced with permission from R Grozovsky, PhD.\u0022 class=\u0022fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1042707281\u0022 data-figure-caption=\u0022\u0026amp;lt;div xmlns=\u0026amp;quot;http:\/\/www.w3.org\/1999\/xhtml\u0026amp;quot;\u0026amp;gt;Desialylated Human Platelets Stimulate JAK2 and STAT3 Phosphorylation in HepG2 CellsJACK2, Janus kinase 2; STAT3, activator of transcription 3.*\u0026amp;lt;em\u0026amp;gt;P\u0026amp;lt;\/em\u0026amp;gt;\u0026#x2009;\u0026amp;amp;lt;\u0026#x2009;.05, **\u0026amp;lt;em\u0026amp;gt;P\u0026amp;lt;\/em\u0026amp;gt;\u0026#x2009;\u0026amp;amp;lt;\u0026#x2009;.01Reproduced with permission from R Grozovsky, PhD.\u0026amp;lt;\/div\u0026amp;gt;\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cimg class=\u0022fragment-image\u0022 alt=\u0022Figure 1.\u0022 src=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F1.medium.gif\u0022\/\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u00220 first\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F1.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Figure 1.\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00221\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F1.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00222 last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/11644\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022 xmlns:xhtml=\u0022http:\/\/www.w3.org\/1999\/xhtml\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003EFigure 1.\u003C\/span\u003E \n               \u003Cp id=\u0022p-6\u0022 class=\u0022first-child\u0022\u003EDesialylated Human Platelets Stimulate JAK2 and STAT3 Phosphorylation in HepG2 Cells\u003C\/p\u003E\n               \u003Cp id=\u0022p-7\u0022\u003EJACK2, Janus kinase 2; STAT3, activator of transcription 3.\u003C\/p\u003E\n               \u003Cp id=\u0022p-8\u0022\u003E*\u003Cem\u003EP\u003C\/em\u003E\u2009\u0026lt;\u2009.05, **\u003Cem\u003EP\u003C\/em\u003E\u2009\u0026lt;\u2009.01\u003C\/p\u003E\n               \u003Cp id=\u0022p-9\u0022\u003EReproduced with permission from R Grozovsky, PhD.\u003C\/p\u003E\n            \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\n         \u003Cp id=\u0022p-10\u0022\u003EIn the in vivo treatment, AMR-deficient mice had increased platelet count, survival, and loss of sialic acid compared with wild type mice showing that removal of desialylated platelets by the AMR regulated in vivo platelet survival.\u003C\/p\u003E\n         \u003Cp id=\u0022p-11\u0022\u003EIn livers isolated from Asgr2-null mice, TPO mRNA expression decreased by 40%, while TPO mRNA expression in livers from mice lacking sialyltransferase (St3gal4-null) was increased by 30%, compared with wild type mice. In St3gal4-null mice, desialylated platelet clearance is increased and specifically mediated by the AMR. This indicates that desialylated platelet uptake by the AMR regulates liver TPO levels.\u003C\/p\u003E\n         \u003Cp id=\u0022p-12\u0022\u003EWhen desialylated platelets isolated from St3gal4\u003Cem\u003E-\u003C\/em\u003Enull or Asgr2-null mice are infused into wild type mice, hepatic TPO mRNA levels increase as early as 12 hours after infusion, while desialylated platelets infused into Asgr2-null mice have no effect on TPO mRNA synthesis. Further, treatment with JAK inhibitors block the desialylated platelet stimulated TPO mRNA increase in wild type mice (\u003Ca id=\u0022xref-fig-2-1\u0022 class=\u0022xref-fig\u0022 href=\u0022#F2\u0022\u003EFigure 2\u003C\/a\u003E).\u003C\/p\u003E\n         \u003Cdiv id=\u0022F2\u0022 class=\u0022fig pos-float  odd\u0022\u003E\u003Cdiv class=\u0022highwire-figure\u0022\u003E\u003Cdiv class=\u0022fig-inline-img-wrapper\u0022\u003E\u003Cdiv class=\u0022fig-inline-img\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F2.large.jpg?width=800\u0026amp;height=600\u0026amp;carousel=1\u0022 title=\u0022JAK Inhibitors Block TPO mRNA Increase in Wild Type MiceJAK, Janus kinase; TPO, thrombopoietin.Reproduced with permission from R Grozovsky, PhD.\u0022 class=\u0022fragment-images colorbox-load\u0022 rel=\u0022gallery-fragment-images-1042707281\u0022 data-figure-caption=\u0022JAK Inhibitors Block TPO mRNA Increase in Wild Type MiceJAK, Janus kinase; TPO, thrombopoietin.Reproduced with permission from R Grozovsky, PhD.\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003E\u003Cimg class=\u0022fragment-image\u0022 alt=\u0022Figure 2.\u0022 src=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F2.medium.gif\u0022\/\u003E\u003C\/a\u003E\u003C\/div\u003E\u003C\/div\u003E\u003Cul class=\u0022highwire-figure-links inline\u0022\u003E\u003Cli class=\u00220 first\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F2.large.jpg?download=true\u0022 class=\u0022highwire-figure-link highwire-figure-link-download\u0022 title=\u0022Download Figure 2.\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload figure\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00221\u0022\u003E\u003Ca href=\u0022http:\/\/d282kpwvnogo5m.cloudfront.net\/content\/spmdc\/14\/55\/4\/F2.large.jpg\u0022 class=\u0022highwire-figure-link highwire-figure-link-newtab\u0022 target=\u0022_blank\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EOpen in new tab\u003C\/a\u003E\u003C\/li\u003E\u003Cli class=\u00222 last\u0022\u003E\u003Ca href=\u0022\/highwire\/powerpoint\/11645\u0022 class=\u0022highwire-figure-link highwire-figure-link-ppt\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EDownload powerpoint\u003C\/a\u003E\u003C\/li\u003E\u003C\/ul\u003E\u003C\/div\u003E\u003Cdiv class=\u0022fig-caption\u0022\u003E\u003Cspan class=\u0022fig-label\u0022\u003EFigure 2.\u003C\/span\u003E \n               \u003Cp id=\u0022p-13\u0022 class=\u0022first-child\u0022\u003EJAK Inhibitors Block TPO mRNA Increase in Wild Type Mice\u003C\/p\u003E\n               \u003Cp id=\u0022p-14\u0022\u003EJAK, Janus kinase; TPO, thrombopoietin.\u003C\/p\u003E\n               \u003Cp id=\u0022p-15\u0022\u003EReproduced with permission from R Grozovsky, PhD.\u003C\/p\u003E\n            \u003Cdiv class=\u0022sb-div caption-clear\u0022\u003E\u003C\/div\u003E\u003C\/div\u003E\u003C\/div\u003E\n         \u003Cp id=\u0022p-16\u0022\u003EThe accumulated data show that circulating desialylated platelets by way of the AMR and JAK2-STAT3 signaling stimulated TPO production, providing a physiological feedback mechanism to regulate plasma TPO levels and platelet production in vivo and in vitro. The complete understanding of this feedback mechanism illuminates the pathophysiology of platelet diseases, such as thrombocythemia and immune thrombocytopenia. This feedback has clinical implications, however, as the administration of JAK1\/2 inhibitors can cause thrombocytopenia in some groups of patients.\u003C\/p\u003E\n      \u003C\/div\u003E\u003Cul class=\u0022copyright-statement\u0022\u003E\u003Cli class=\u0022fn\u0022 id=\u0022copyright-statement-1\u0022\u003E\u00a9 2014 SAGE Publications\u003C\/li\u003E\u003C\/ul\u003E\u003Cspan class=\u0022highwire-journal-article-marker-end\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Cspan id=\u0022related-urls\u0022\u003E\u003C\/span\u003E\u003C\/div\u003E\u003Ca href=\u0022http:\/\/mdc.sagepub.com\/content\/14\/55\/4.abstract\u0022 class=\u0022hw-link hw-link-article-abstract\u0022 data-icon-position=\u0022\u0022 data-hide-link-title=\u00220\u0022\u003EView Summary\u003C\/a\u003E\u003C\/div\u003E  \u003C\/div\u003E\n\n  \n  \u003C\/div\u003E\n\u003C\/div\u003E\n  \u003C\/div\u003E\n\u003C\/div\u003E\n\u003C\/div\u003E\u003Cscript type=\u0022text\/javascript\u0022 src=\u0022http:\/\/mdc.sagepub.com\/sites\/all\/modules\/highwire\/highwire\/plugins\/highwire_markup_process\/js\/highwire_figures.js?nzo942\u0022\u003E\u003C\/script\u003E\n\u003Cscript type=\u0022text\/javascript\u0022 src=\u0022http:\/\/mdc.sagepub.com\/sites\/all\/modules\/highwire\/highwire\/plugins\/highwire_markup_process\/js\/highwire_openurl.js?nzo942\u0022\u003E\u003C\/script\u003E\n\u003C\/body\u003E\u003C\/html\u003E"}