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Volumn 94, Issue 3, 2012, Pages 597-607

Ceramide 1-phosphate stimulates proliferation of C2C12 myoblasts

Author keywords

C2C12 myoblasts; Cell cycle; Cell growth; Ceramide 1 phosphate; Myoblast proliferation

Indexed keywords

CERAMIDE 1 PHOSPHATE; CYCLIN D1; GLYCOGEN SYNTHASE KINASE 3BETA; INHIBITORY GUANINE NUCLEOTIDE BINDING PROTEIN; MAMMALIAN TARGET OF RAPAMYCIN; MITOGEN ACTIVATED PROTEIN KINASE 1; MITOGEN ACTIVATED PROTEIN KINASE 3; PHOSPHATIDYLINOSITOL 3 KINASE; PROTEIN KINASE B; RETINOBLASTOMA PROTEIN; STAUROSPORINE; THYMIDINE;

EID: 84857063449     PISSN: 03009084     EISSN: 61831638     Source Type: Journal    
DOI: 10.1016/j.biochi.2011.09.009     Document Type: Article
Times cited : (57)

References (48)
  • 1
    • 28244498646 scopus 로고    scopus 로고
    • Stem cells in postnatal myogenesis: Molecular mechanisms of satellite cell quiescence, activation and replenishment
    • DOI 10.1016/j.tcb.2005.10.007, PII S0962892405002515
    • J. Dhawan, and T.A. Rando Stem cells in postnatal myogenesis: molecular mechanisms of satellite cell quiescence, activation and replenishment Trends Cell Biol. 15 2005 666 673 (Pubitemid 41713968)
    • (2005) Trends in Cell Biology , vol.15 , Issue.12 , pp. 666-673
    • Dhawan, J.1    Rando, T.A.2
  • 2
    • 36749061251 scopus 로고    scopus 로고
    • Skeletal muscle satellite cells and adult myogenesis
    • DOI 10.1016/j.ceb.2007.09.012, PII S0955067407001342, Cell Differentiation / Cell Division, Growth and Death
    • F. Le Grand, and M.A. Rudnicki Skeletal muscle satellite cells and adult myogenesis Curr. Opin. Cell Biol. 19 2007 628 633 (Pubitemid 350216420)
    • (2007) Current Opinion in Cell Biology , vol.19 , Issue.6 , pp. 628-633
    • Le Grand, F.1    Rudnicki, M.A.2
  • 3
    • 57349160599 scopus 로고    scopus 로고
    • Pleiotropic effects of sphingolipids in skeletal muscle
    • P. Bruni, and C. Donati Pleiotropic effects of sphingolipids in skeletal muscle Cell Mol. Life Sci. 65 2008 3725 3736
    • (2008) Cell Mol. Life Sci. , vol.65 , pp. 3725-3736
    • Bruni, P.1    Donati, C.2
  • 4
    • 38549152194 scopus 로고    scopus 로고
    • Principles of bioactive lipid signalling: Lessons from sphingolipids
    • DOI 10.1038/nrm2329, PII NRM2329
    • Y.A. Hannun, and L.M. Obeid Principles of bioactive lipid signalling: lessons from sphingolipids Nat. Rev. Mol. Cell Biol. 9 2008 139 150 (Pubitemid 351158911)
    • (2008) Nature Reviews Molecular Cell Biology , vol.9 , Issue.2 , pp. 139-150
    • Hannun, Y.A.1    Obeid, L.M.2
  • 6
    • 33847408400 scopus 로고    scopus 로고
    • Inhibition of de novo ceramide synthesis upregulates phospholipase D and enhances myogenic differentation
    • DOI 10.1242/jcs.03331
    • S. Mebarek, H. Komati, F. Naro, C. Zeiller, M. Alvisi, M. Lagarde, A.F. Prigent, and G. Nemoz Inhibition of de novo ceramide synthesis upregulates phospholipase D and enhances myogenic differentiation J. Cell Sci. 120 2007 407 416 (Pubitemid 46332504)
    • (2007) Journal of Cell Science , vol.120 , Issue.3 , pp. 407-416
    • Mebarek, S.1    Komati, H.2    Naro, F.3    Zeiller, C.4    Alvisi, M.5    Lagarde, M.6    Prigent, A.-F.7    Nemoz, G.8
  • 7
    • 33746075255 scopus 로고    scopus 로고
    • Entry of muscle satellite cells into the cell cycle requires sphingolipid signaling
    • DOI 10.1083/jcb.200605028
    • Y. Nagata, T.A. Partridge, R. Matsuda, and P.S. Zammit Entry of muscle satellite cells into the cell cycle requires sphingolipid signaling J. Cell Biol. 174 2006 245 253 (Pubitemid 44079859)
    • (2006) Journal of Cell Biology , vol.174 , Issue.2 , pp. 245-253
    • Nagata, Y.1    Partridge, T.A.2    Matsuda, R.3    Zammit, P.S.4
  • 9
    • 36849029875 scopus 로고    scopus 로고
    • Sphingosine kinase activity is required for myogenic differentiation of C2C12 myoblasts
    • DOI 10.1002/jcp.21187
    • E. Meacci, F. Nuti, C. Donati, F. Cencetti, M. Farnararo, and P. Bruni Sphingosine kinase activity is required for myogenic differentiation of C2C12 myoblasts J. Cell Physiol. 214 2008 210 220 (Pubitemid 350220075)
    • (2008) Journal of Cellular Physiology , vol.214 , Issue.1 , pp. 210-220
    • Meacci, E.1    Nuti, F.2    Donati, C.3    Cencetti, F.4    Farnararo, M.5    Bruni, P.6
  • 10
    • 34548412099 scopus 로고    scopus 로고
    • 2 signaling
    • DOI 10.1016/j.febslet.2007.08.007, PII S0014579307008770
    • C. Donati, P. Nincheri, F. Cencetti, E. Rapizzi, M. Farnararo, and P. Bruni Tumor necrosis factor-alpha exerts pro-myogenic action in C2C12 myoblasts via sphingosine kinase/S1P2 signaling FEBS Lett. 581 2007 4384 4388 (Pubitemid 47368198)
    • (2007) FEBS Letters , vol.581 , Issue.23 , pp. 4384-4388
    • Donati, C.1    Nincheri, P.2    Cencetti, F.3    Rapizzi, E.4    Farnararo, M.5    Bruni, P.6
  • 11
    • 0029014026 scopus 로고
    • Short-chain ceramide-1-phosphates are novel stimulators of DNA synthesis and cell division: Antagonism by cell-permeable ceramides
    • A. Gomez-Munoz, P.A. Duffy, A. Martin, L. O'Brien, H.S. Byun, R. Bittman, and D.N. Brindley Short-chain ceramide-1-phosphates are novel stimulators of DNA synthesis and cell division: antagonism by cell-permeable ceramides Mol. Pharmacol. 47 1995 833 839
    • (1995) Mol. Pharmacol. , vol.47 , pp. 833-839
    • Gomez-Munoz, A.1    Duffy, P.A.2    Martin, A.3    O'Brien, L.4    Byun, H.S.5    Bittman, R.6    Brindley, D.N.7
  • 12
  • 13
    • 39149141339 scopus 로고    scopus 로고
    • Ceramide 1-phosphate stimulates macrophage proliferation through activation of the PI3-kinase/PKB, JNK and ERK1/2 pathways
    • P. Gangoiti, M.H. Granado, S.W. Wang, J.Y. Kong, U.P. Steinbrecher, and A. Gomez-Munoz Ceramide 1-phosphate stimulates macrophage proliferation through activation of the PI3-kinase/PKB, JNK and ERK1/2 pathways Cell. Signal. 20 2008 726 736
    • (2008) Cell. Signal. , vol.20 , pp. 726-736
    • Gangoiti, P.1    Granado, M.H.2    Wang, S.W.3    Kong, J.Y.4    Steinbrecher, U.P.5    Gomez-Munoz, A.6
  • 14
    • 21244479753 scopus 로고    scopus 로고
    • Ceramide-1-phosphate promotes cell survival through activation of the phosphatidylinositol 3-kinase/protein kinase B pathway
    • DOI 10.1016/j.febslet.2005.05.067, PII S0014579305006939
    • A. Gomez-Munoz, J.Y. Kong, K. Parhar, S.W. Wang, P. Gangoiti, M. Gonzalez, S. Eivemark, B. Salh, V. Duronio, and U.P. Steinbrecher Ceramide-1-phosphate promotes cell survival through activation of the phosphatidylinositol 3-kinase/protein kinase B pathway FEBS Lett. 579 2005 3744 3750 (Pubitemid 40897719)
    • (2005) FEBS Letters , vol.579 , Issue.17 , pp. 3744-3750
    • Gomez-Munoz, A.1    Kong, J.Y.2    Parhar, K.3    Shih, W.W.4    Gangoiti, P.5    Gonzalez, M.6    Eivemark, S.7    Salh, B.8    Duronio, V.9    Steinbrecher, U.P.10
  • 16
    • 52449119800 scopus 로고    scopus 로고
    • Ceramide kinase and the ceramide-1-phosphate/cPLA2alpha interaction as a therapeutic target
    • N.F. Lamour, and C.E. Chalfant Ceramide kinase and the ceramide-1-phosphate/cPLA2alpha interaction as a therapeutic target Curr. Drug Targets 9 2008 674 682
    • (2008) Curr. Drug Targets , vol.9 , pp. 674-682
    • Lamour, N.F.1    Chalfant, C.E.2
  • 18
    • 45449102260 scopus 로고    scopus 로고
    • Sphingosine 1-phosphate differentially regulates proliferation of C2C12 reserve cells and myoblasts
    • E. Rapizzi, C. Donati, F. Cencetti, P. Nincheri, and P. Bruni Sphingosine 1-phosphate differentially regulates proliferation of C2C12 reserve cells and myoblasts Mol. Cell. Biochem. 314 2008 193 199
    • (2008) Mol. Cell. Biochem. , vol.314 , pp. 193-199
    • Rapizzi, E.1    Donati, C.2    Cencetti, F.3    Nincheri, P.4    Bruni, P.5
  • 21
  • 22
    • 34250683438 scopus 로고    scopus 로고
    • Pertussis toxin promotes macrophage survival through inhibition of acid sphingomyelinase and activation of the phosphoinositide 3-kinase/protein kinase B pathway
    • DOI 10.1016/j.cellsig.2007.04.001, PII S0898656807001131
    • S.W. Wang, K. Parhar, K.J. Chiu, A. Tran, P. Gangoiti, J. Kong, M. Gonzalez, B. Salh, V. Duronio, U.P. Steinbrecher, and A. Gomez-Munoz Pertussis toxin promotes macrophage survival through inhibition of acid sphingomyelinase and activation of the phosphoinositide 3-kinase/protein kinase B pathway Cell. Signal. 19 2007 1772 1783 (Pubitemid 46935433)
    • (2007) Cellular Signalling , vol.19 , Issue.8 , pp. 1772-1783
    • Wang, S.W.1    Parhar, K.2    Chiu, K.J.3    Tran, A.4    Gangoiti, P.5    Kong, J.6    Gonzalez, M.7    Salh, B.8    Duronio, V.9    Steinbrecher, U.P.10    Gomez-Munoz, A.11
  • 23
    • 0025168334 scopus 로고
    • Ceramide 1-phosphate, a novel phospholipid in human leukemia (HL-60) cells. Synthesis via ceramide from sphingomyelin
    • K.A. Dressler, and R.N. Kolesnick Ceramide 1-phosphate, a novel phospholipid in human leukemia (HL-60) cells. Synthesis via ceramide from sphingomyelin J. Biol. Chem. 265 25 1990 14917 14921
    • (1990) J. Biol. Chem. , vol.265 , Issue.25 , pp. 14917-14921
    • Dressler, K.A.1    Kolesnick, R.N.2
  • 24
    • 77950606567 scopus 로고    scopus 로고
    • Use of high performance liquid chromatography-electrospray ionization-tandem mass spectrometry for the analysis of ceramide-1-phosphate levels
    • D.S. Wijesinghe, J.C. Allegood, L.B. Gentile, T.E. Fox, M. Kester, and C.E. Chalfant Use of high performance liquid chromatography-electrospray ionization-tandem mass spectrometry for the analysis of ceramide-1-phosphate levels J. Lipid Res. 51 2010 641 651
    • (2010) J. Lipid Res. , vol.51 , pp. 641-651
    • Wijesinghe, D.S.1    Allegood, J.C.2    Gentile, L.B.3    Fox, T.E.4    Kester, M.5    Chalfant, C.E.6
  • 25
    • 70350352932 scopus 로고    scopus 로고
    • The chain lenght specificity for the activation of group IV citosolic phospholipase A2 by ceramide-1-phosphate. Use of the dodecane delivery system for determining lipid specific effects
    • D.S. Wijesinghe, P. Subramanian, N.F. Lamour, L.B. Gentile, M.H. Granado, Z. Szulc, A. Bielawska, A. Gomez-Munoz, and C.E. Chalfant The chain lenght specificity for the activation of group IV citosolic phospholipase A2 by ceramide-1-phosphate. Use of the dodecane delivery system for determining lipid specific effects J. Lipid Res. 50 2009 1986 1995
    • (2009) J. Lipid Res. , vol.50 , pp. 1986-1995
    • Wijesinghe, D.S.1    Subramanian, P.2    Lamour, N.F.3    Gentile, L.B.4    Granado, M.H.5    Szulc, Z.6    Bielawska, A.7    Gomez-Munoz, A.8    Chalfant, C.E.9
  • 26
    • 2342618936 scopus 로고    scopus 로고
    • Multiple roles of the PI3K/PKB (Akt) pathway in cell cycle progression
    • J. Liang, and J.M. Slingerland Multiple roles of the PI3K/PKB (Akt) pathway in cell cycle progression Cell Cycle 2 2003 339 345
    • (2003) Cell Cycle , vol.2 , pp. 339-345
    • Liang, J.1    Slingerland, J.M.2
  • 27
    • 44749094062 scopus 로고    scopus 로고
    • Involvement of nitric oxide in the promotion of cell survival by ceramide 1-phosphate
    • P. Gangoiti, M.H. Granado, L. Arana, A. Ouro, and A. Gomez-Munoz Involvement of nitric oxide in the promotion of cell survival by ceramide 1-phosphate FEBS Lett. 582 2008 2263 2269
    • (2008) FEBS Lett. , vol.582 , pp. 2263-2269
    • Gangoiti, P.1    Granado, M.H.2    Arana, L.3    Ouro, A.4    Gomez-Munoz, A.5
  • 28
    • 1642403610 scopus 로고    scopus 로고
    • Ceramide-1-phosphate: A novel regulator of cell activation
    • DOI 10.1016/S0014-5793(04)00211-X, PII S001457930400211X
    • A. Gomez-Munoz Ceramide-1-phosphate: a novel regulator of cell activation FEBS Lett. 562 2004 5 10 (Pubitemid 38388439)
    • (2004) FEBS Letters , vol.562 , Issue.1-3 , pp. 5-10
    • Gomez-Munoz, A.1
  • 30
    • 17444431201 scopus 로고    scopus 로고
    • Phosphorylation and functional inactivation of TSC2 by Erk: Implications for tuberous sclerosis and cancer pathogenesis
    • DOI 10.1016/j.cell.2005.02.031
    • L. Ma, Z. Chen, H. Erdjument-Bromage, P. Tempst, and P.P. Pandolfi Phosphorylation and functional inactivation of TSC2 by Erk implications for tuberous sclerosis and cancer pathogenesis Cell 121 2005 179 193 (Pubitemid 40546387)
    • (2005) Cell , vol.121 , Issue.2 , pp. 179-193
    • Ma, L.1    Chen, Z.2    Erdjument-Bromage, H.3    Tempst, P.4    Pandolfi, P.P.5
  • 32
    • 0347989458 scopus 로고    scopus 로고
    • Cellular and Molecular Regulation of Muscle Regeneration
    • DOI 10.1152/physrev.00019.2003
    • S.B. Charge, and M.A. Rudnicki Cellular and molecular regulation of muscle regeneration Physiol. Rev. 84 2004 209 238 (Pubitemid 38049880)
    • (2004) Physiological Reviews , vol.84 , Issue.1 , pp. 209-238
    • Charge, S.B.P.1    Rudnicki, M.A.2
  • 33
    • 33845292007 scopus 로고    scopus 로고
    • Ceramide 1-phosphate/ceramide, a switch between life and death
    • DOI 10.1016/j.bbamem.2006.05.011, PII S0005273606001866, Sphingolipids, Apoptosis and Disease
    • A. Gomez-Munoz Ceramide 1-phosphate/ceramide, a switch between life and death Biochim. Biophys. Acta 1758 2006 2049 2056 (Pubitemid 44911057)
    • (2006) Biochimica et Biophysica Acta - Biomembranes , vol.1758 , Issue.12 , pp. 2049-2056
    • Gomez-Munoz, A.1
  • 35
    • 0027319559 scopus 로고
    • Sphingosine blocks both membrane fusion and calmodulin-dependent phosphorylation of the 100-kDa protein of chick embryonic myoblasts
    • DOI 10.1006/excr.1993.1105
    • H.S. Kim, I.H. Lee, Y.J. Jeon, C.H. Chung, and D.B. Ha Sphingosine blocks both membrane fusion and calmodulin-dependent phosphorylation of the 100-kDa protein of chick embryonic myoblasts Exp. Cell Res. 205 1993 408 411 (Pubitemid 23192150)
    • (1993) Experimental Cell Research , vol.205 , Issue.2 , pp. 408-411
    • Kim, H.S.1    Lee, I.H.2    Jeon, Y.J.3    Chung, C.H.4    Ha, D.B.5
  • 36
    • 0030722189 scopus 로고    scopus 로고
    • Regulation of distinct stages of skeletal muscle differentiation by mitogen-activated protein kinases
    • DOI 10.1126/science.278.5341.1288
    • A.M. Bennett, and N.K. Tonks Regulation of distinct stages of skeletal muscle differentiation by mitogen-activated protein kinases Science 278 1997 1288 1291 (Pubitemid 27495740)
    • (1997) Science , vol.278 , Issue.5341 , pp. 1288-1291
    • Bennett, A.M.1    Tonks, N.K.2
  • 37
    • 0035123649 scopus 로고    scopus 로고
    • Insulin signaling leading to proliferation, survival, and membrane ruffling in C2C12 myoblasts
    • DOI 10.1002/1097-4652(2001)9999:9999<::AID-JCP1058>3.0.CO;2-V
    • R. Conejo, and M. Lorenzo Insulin signaling leading to proliferation, survival, and membrane ruffling in C2C12 myoblasts J. Cell. Physiol. 187 2001 96 108 (Pubitemid 32195250)
    • (2001) Journal of Cellular Physiology , vol.187 , Issue.1 , pp. 96-108
    • Conejo, R.1    Lorenzo, M.2
  • 38
    • 0030934472 scopus 로고    scopus 로고
    • The mitogenic and myogenic actions of insulin-like growth factors utilize distinct signaling pathways
    • DOI 10.1074/jbc.272.10.6653
    • S.A. Coolican, D.S. Samuel, D.Z. Ewton, F.J. McWade, and J.R. Florini The mitogenic and myogenic actions of insulin-like growth factors utilize distinct signaling pathways J. Biol. Chem. 272 1997 6653 6662 (Pubitemid 27118151)
    • (1997) Journal of Biological Chemistry , vol.272 , Issue.10 , pp. 6653-6662
    • Coolican, S.A.1    Samuel, D.S.2    Ewton, D.Z.3    McWade, F.J.4    Florini, J.R.5
  • 39
    • 2942518272 scopus 로고    scopus 로고
    • Differential regulation of the phosphoinositide 3-kinase and MAP kinase pathways by hepatocyte growth factor vs. insulin-like growth factor-I in myogenic cells
    • DOI 10.1016/j.yexcr.2004.03.024, PII S0014482704001429
    • O. Halevy, and L.C. Cantley Differential regulation of the phosphoinositide 3-kinase and MAP kinase pathways by hepatocyte growth factor vs. insulin-like growth factor-I in myogenic cells Exp. Cell Res. 297 2004 224 234 (Pubitemid 38759575)
    • (2004) Experimental Cell Research , vol.297 , Issue.1 , pp. 224-234
    • Halevy, O.1    Cantley, L.C.2
  • 40
    • 0035203194 scopus 로고    scopus 로고
    • ERK1/2 is required for myoblast proliferation but is dispensable for muscle gene expression and cell fusion
    • N.C. Jones, Y.V. Fedorov, R.S. Rosenthal, and B.B. Olwin ERK1/2 is required for myoblast proliferation but is dispensable for muscle gene expression and cell fusion J. Cell. Physiol. 186 2001 104 115
    • (2001) J. Cell. Physiol. , vol.186 , pp. 104-115
    • Jones, N.C.1    Fedorov, Y.V.2    Rosenthal, R.S.3    Olwin, B.B.4
  • 43
    • 10044236607 scopus 로고    scopus 로고
    • The role of sphingosine and ceramide kinases in inflammatory responses
    • DOI 10.1016/j.imlet.2004.09.001, PII S0165247804002524
    • T. Baumruker, F. Bornancin, and A. Billich The role of sphingosine and ceramide kinases in inflammatory responses Immunol. Lett. 96 2005 175 185 (Pubitemid 39600773)
    • (2005) Immunology Letters , vol.96 , Issue.2 , pp. 175-185
    • Baumruker, T.1    Bornancin, F.2    Billich, A.3
  • 44
    • 0025185675 scopus 로고
    • Characterization of a ceramide kinase activity from human leukemia (HL-60) cells. Separation from diacylglycerol kinase activity
    • R.N. Kolesnick, and M.R. Hemer Characterization of a ceramide kinase activity from human leukemia (HL-60) cells. Separation from diacylglycerol kinase activity J. Biol. Chem. 265 1990 18803 18808
    • (1990) J. Biol. Chem. , vol.265 , pp. 18803-18808
    • Kolesnick, R.N.1    Hemer, M.R.2
  • 45
    • 43749105380 scopus 로고    scopus 로고
    • Regulation and traffic of ceramide 1-phosphate produced by ceramide kinase: Comparative analysis to glucosylceramide and sphingomyelin
    • A. Boath, C. Graf, E. Lidome, T. Ullrich, P. Nussbaumer, and F. Bornancin Regulation and traffic of ceramide 1-phosphate produced by ceramide kinase: comparative analysis to glucosylceramide and sphingomyelin J. Biol. Chem. 283 2008 8517 8526
    • (2008) J. Biol. Chem. , vol.283 , pp. 8517-8526
    • Boath, A.1    Graf, C.2    Lidome, E.3    Ullrich, T.4    Nussbaumer, P.5    Bornancin, F.6
  • 46
    • 0345129972 scopus 로고    scopus 로고
    • Blood borne macrophages are essential for the triggering of muscle regeneration following muscle transplant
    • DOI 10.1016/S0960-8966(98)00111-4, PII S0960896698001114
    • L. Lescaudron, E. Peltekian, J. Fontaine-Perus, D. Paulin, M. Zampieri, L. Garcia, and E. Parrish Blood borne macrophages are essential for the triggering of muscle regeneration following muscle transplant Neuromuscul. Disord. 9 1999 72 80 (Pubitemid 29170435)
    • (1999) Neuromuscular Disorders , vol.9 , Issue.2 , pp. 72-80
    • Lescaudron, L.1    Peltekian, E.2    Fontaine-Perus, J.3    Paulin, D.4    Zampieri, M.5    Garcia, L.6    Parrish, E.7
  • 47
    • 0033060101 scopus 로고    scopus 로고
    • Macrophages enhance muscle satellite cell proliferation and delay their differentiation
    • DOI 10.1002/(SICI)1097-4598(199906)22:6<724::AID-MUS9>3.0.CO;2-O
    • F. Merly, L. Lescaudron, T. Rouaud, F. Crossin, and M.F. Gardahaut Macrophages enhance muscle satellite cell proliferation and delay their differentiation Muscle Nerve 22 1999 724 732 (Pubitemid 29256841)
    • (1999) Muscle and Nerve , vol.22 , Issue.6 , pp. 724-732
    • Merly, F.1    Lescaudron, L.2    Rouaud, T.3    Crossin, F.4    Gardahaut, M.F.5
  • 48
    • 33845785972 scopus 로고    scopus 로고
    • Macrophages promote muscle membrane repair and muscle fibre growth and regeneration during modified muscle loading in mice in vivo
    • DOI 10.1113/jphysiol.2006.118265
    • J.G. Tidball, and M. Wehling-Henricks Macrophages promote muscle membrane repair and muscle fibre growth and regeneration during modified muscle loading in mice in vivo J. Physiol. 578 2007 327 336 (Pubitemid 44971129)
    • (2007) Journal of Physiology , vol.578 , Issue.1 , pp. 327-336
    • Tidball, J.G.1    Wehling-Henricks, M.2


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