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profileTobias Walther, Ph.D.

TitleProfessor of Cell Biology
InstitutionHarvard T.H. Chan School of Public Health
DepartmentCell Biology
AddressHarvard School of Public Health
Building 1, room 207
665 Huntington Ave
Boston MA 02115
Phone617/432-6017
vCardDownload vCard (login for email)
Other Positions
TitleProfessor of Genetics and Complex Diseases
InstitutionHarvard T.H. Chan School of Public Health
DepartmentGenetics and Complex Diseases


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Collapse research activities and funding
The research activities and funding listed below are automatically derived from NIH ExPORTER and other sources, which might result in incorrect or missing items. Faculty can login to make corrections and additions.
R13DK103464     (WALTHER, TOBIAS)Jul 1, 2014 - Jun 30, 2015
NIH/NIDDK
FASEB SRC on Lipid Droplets: Metabolic Consequences of the Storage of Neutral Lip
Role: Principal Investigator

R01DK101579     (FARESE, ROBERT V)May 1, 2014 - Apr 30, 2019
NIH/NIDDK
Physiology of Lipid Droplets and Triglyceride Storage
Role: Co-Principal Investigator

R01GM095982     (WALTHER, TOBIAS)Feb 15, 2012 - Dec 31, 2015
NIH/NIGMS
Cellular Functions of Plasma Membrane Organization by Eisosomes
Role: Principal Investigator

R01GM097194     (WALTHER, TOBIAS)Sep 30, 2011 - Nov 30, 2020
NIH/NIGMS
Mechanisms of Lipid Droplet Protein Targeting
Role: Principal Investigator

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Collapse selected publications
Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Faculty can login to make corrections and additions.
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  1. Gluchowski NL, Becuwe M, Walther TC, Farese RV. Lipid droplets and liver disease: from basic biology to clinical implications. Nat Rev Gastroenterol Hepatol. 2017 Apr 21. PMID: 28428634.
    View in: PubMed
  2. Gluchowski NL, Chitraju C, Picoraro JA, Mejhert N, Pinto S, Xin W, Kamin DS, Winter HS, Chung WK, Walther TC, Farese RV. Identification and characterization of a novel DGAT1 missense mutation associated with congenital diarrhea. J Lipid Res. 2017 Apr 03. PMID: 28373485.
    View in: PubMed
  3. Kory N, Grond S, Kamat SS, Li Z, Krahmer N, Chitraju C, Zhou P, Fröhlich F, Semova I, Ejsing C, Zechner R, Cravatt BF, Farese RV, Walther TC. Mice lacking lipid droplet-associated hydrolase, a gene linked to human prostate cancer, have normal cholesterol ester metabolism. J Lipid Res. 2017 Jan; 58(1):226-235. PMID: 27836991.
    View in: PubMed
  4. Fröhlich F, Olson DK, Christiano R, Farese RV, Walther TC. Proteomic and phosphoproteomic analyses of yeast reveal the global cellular response to sphingolipid depletion. Proteomics. 2016 Nov; 16(21):2759-2763. PMID: 27717283.
    View in: PubMed
  5. Wang H, Becuwe M, Housden BE, Chitraju C, Porras AJ, Graham MM, Liu XN, Thiam AR, Savage DB, Agarwal AK, Garg A, Olarte MJ, Lin Q, Fröhlich F, Hannibal-Bach HK, Upadhyayula S, Perrimon N, Kirchhausen T, Ejsing CS, Walther TC, Farese RV. Seipin is required for converting nascent to mature lipid droplets. Elife. 2016 Aug 26; 5. PMID: 27564575.
    View in: PubMed
  6. Kory N, Farese RV, Walther TC. Targeting Fat: Mechanisms of Protein Localization to Lipid Droplets. Trends Cell Biol. 2016 Jul; 26(7):535-46. PMID: 26995697; PMCID: PMC4976449 [Available on 07/01/17].
  7. Farese RV, Walther TC. Lipid droplets go nuclear. J Cell Biol. 2016 Jan 04; 212(1):7-8. PMID: 26728852; PMCID: PMC4700486.
  8. Olson DK, Fröhlich F, Farese RV, Walther TC. Taming the sphinx: Mechanisms of cellular sphingolipid homeostasis. Biochim Biophys Acta. 2016 Aug; 1861(8 Pt B):784-92. PMID: 26747648; PMCID: PMC4907851 [Available on 08/01/17].
  9. Baskin JM, Wu X, Christiano R, Oh MS, Schauder CM, Gazzerro E, Messa M, Baldassari S, Assereto S, Biancheri R, Zara F, Minetti C, Raimondi A, Simons M, Walther TC, Reinisch KM, De Camilli P. The leukodystrophy protein FAM126A (hyccin) regulates PtdIns(4)P synthesis at the plasma membrane. Nat Cell Biol. 2016 Jan; 18(1):132-8. PMID: 26571211; PMCID: PMC4689616.
  10. Fröhlich F, Petit C, Kory N, Christiano R, Hannibal-Bach HK, Graham M, Liu X, Ejsing CS, Farese RV, Walther TC. The GARP complex is required for cellular sphingolipid homeostasis. Elife. 2015 Sep 10; 4. PMID: 26357016; PMCID: PMC4600884.
  11. Siragusa M, Fröhlich F, Park EJ, Schleicher M, Walther TC, Sessa WC. Stromal cell-derived factor 2 is critical for Hsp90-dependent eNOS activation. Sci Signal. 2015 Aug 18; 8(390):ra81. PMID: 26286023; PMCID: PMC4790114.
  12. Kory N, Thiam AR, Farese RV, Walther TC. Protein Crowding Is a Determinant of Lipid Droplet Protein Composition. Dev Cell. 2015 Aug 10; 34(3):351-63. PMID: 26212136; PMCID: PMC4536137.
  13. Shibuya A, Margulis N, Christiano R, Walther TC, Barlowe C. The Erv41-Erv46 complex serves as a retrograde receptor to retrieve escaped ER proteins. J Cell Biol. 2015 Jan 19; 208(2):197-209. PMID: 25583996; PMCID: PMC4298680.
  14. Olson DK, Fröhlich F, Christiano R, Hannibal-Bach HK, Ejsing CS, Walther TC. Rom2-dependent phosphorylation of Elo2 controls the abundance of very long-chain fatty acids. J Biol Chem. 2015 Feb 13; 290(7):4238-47. PMID: 25519905; PMCID: PMC4326832.
  15. Christiano R, Nagaraj N, Fröhlich F, Walther TC. Global proteome turnover analyses of the Yeasts S. cerevisiae and S. pombe. Cell Rep. 2014 Dec 11; 9(5):1959-65. PMID: 25466257; PMCID: PMC4526151.
  16. Fröhlich F, Christiano R, Olson DK, Alcazar-Roman A, DeCamilli P, Walther TC. A role for eisosomes in maintenance of plasma membrane phosphoinositide levels. Mol Biol Cell. 2014 Sep 15; 25(18):2797-806. PMID: 25057013; PMCID: PMC4161514.
  17. Payne F, Lim K, Girousse A, Brown RJ, Kory N, Robbins A, Xue Y, Sleigh A, Cochran E, Adams C, Dev Borman A, Russel-Jones D, Gorden P, Semple RK, Saudek V, O'Rahilly S, Walther TC, Barroso I, Savage DB. Mutations disrupting the Kennedy phosphatidylcholine pathway in humans with congenital lipodystrophy and fatty liver disease. Proc Natl Acad Sci U S A. 2014 Jun 17; 111(24):8901-6. PMID: 24889630; PMCID: PMC4066527.
  18. Currie E, Guo X, Christiano R, Chitraju C, Kory N, Harrison K, Haas J, Walther TC, Farese RV. High confidence proteomic analysis of yeast LDs identifies additional droplet proteins and reveals connections to dolichol synthesis and sterol acetylation. J Lipid Res. 2014 Jul; 55(7):1465-77. PMID: 24868093; PMCID: PMC4076087.
  19. Wilfling F, Haas JT, Walther TC, Farese RV. Lipid droplet biogenesis. Curr Opin Cell Biol. 2014 Aug; 29:39-45. PMID: 24736091; PMCID: PMC4526149.
  20. Bazzini AA, Johnstone TG, Christiano R, Mackowiak SD, Obermayer B, Fleming ES, Vejnar CE, Lee MT, Rajewsky N, Walther TC, Giraldez AJ. Identification of small ORFs in vertebrates using ribosome footprinting and evolutionary conservation. EMBO J. 2014 May 02; 33(9):981-93. PMID: 24705786; PMCID: PMC4193932.
  21. Wilfling F, Thiam AR, Olarte MJ, Wang J, Beck R, Gould TJ, Allgeyer ES, Pincet F, Bewersdorf J, Farese RV, Walther TC. Arf1/COPI machinery acts directly on lipid droplets and enables their connection to the ER for protein targeting. Elife. 2014; 3:e01607. PMID: 24497546; PMCID: PMC3913038.
  22. Harrison MS, Hung CS, Liu TT, Christiano R, Walther TC, Burd CG. A mechanism for retromer endosomal coat complex assembly with cargo. Proc Natl Acad Sci U S A. 2014 Jan 07; 111(1):267-72. PMID: 24344282; PMCID: PMC3890810.
  23. Antonny B, Audhya J, l Bagnat M, von Blume J, Briggs JA, Giraudo C, Kaeser PS, Miller E, Reinisch K, Sbalzarini IF, Schuldiner M, Shen J, Takamori S, Verstreken P, Walther T. Directing traffic into the future. Dev Cell. 2013 Dec 09; 27(5):480-4. PMID: 24482802.
    View in: PubMed
  24. Thiam AR, Farese RV, Walther TC. The biophysics and cell biology of lipid droplets. Nat Rev Mol Cell Biol. 2013 Dec; 14(12):775-86. PMID: 24220094; PMCID: PMC4526153.
  25. Thiam AR, Antonny B, Wang J, Delacotte J, Wilfling F, Walther TC, Beck R, Rothman JE, Pincet F. COPI buds 60-nm lipid droplets from reconstituted water-phospholipid-triacylglyceride interfaces, suggesting a tension clamp function. Proc Natl Acad Sci U S A. 2013 Aug 13; 110(33):13244-9. PMID: 23901109; PMCID: PMC3746913.
  26. Currie E, Schulze A, Zechner R, Walther TC, Farese RV. Cellular fatty acid metabolism and cancer. Cell Metab. 2013 Aug 06; 18(2):153-61. PMID: 23791484; PMCID: PMC3742569.
  27. Krahmer N, Farese RV, Walther TC. Balancing the fat: lipid droplets and human disease. EMBO Mol Med. 2013 Jul; 5(7):973-83. PMID: 23740690; PMCID: PMC3721468.
  28. Fröhlich F, Christiano R, Walther TC. Native SILAC: metabolic labeling of proteins in prototroph microorganisms based on lysine synthesis regulation. Mol Cell Proteomics. 2013 Jul; 12(7):1995-2005. PMID: 23592334; PMCID: PMC3708181.
  29. Wilfling F, Wang H, Haas JT, Krahmer N, Gould TJ, Uchida A, Cheng JX, Graham M, Christiano R, Fröhlich F, Liu X, Buhman KK, Coleman RA, Bewersdorf J, Farese RV, Walther TC. Triacylglycerol synthesis enzymes mediate lipid droplet growth by relocalizing from the ER to lipid droplets. Dev Cell. 2013 Feb 25; 24(4):384-99. PMID: 23415954; PMCID: PMC3727400.
  30. Nguyen AD, Nguyen TA, Cenik B, Yu G, Herz J, Walther TC, Davidson WS, Farese RV. Secreted progranulin is a homodimer and is not a component of high density lipoproteins (HDL). J Biol Chem. 2013 Mar 22; 288(12):8627-35. PMID: 23364791; PMCID: PMC3605681.
  31. Krahmer N, Hilger M, Kory N, Wilfling F, Stoehr G, Mann M, Farese RV, Walther TC. Protein correlation profiles identify lipid droplet proteins with high confidence. Mol Cell Proteomics. 2013 May; 12(5):1115-26. PMID: 23319140; PMCID: PMC3650325.
  32. Haas JT, Winter HS, Lim E, Kirby A, Blumenstiel B, DeFelice M, Gabriel S, Jalas C, Branski D, Grueter CA, Toporovski MS, Walther TC, Daly MJ, Farese RV. DGAT1 mutation is linked to a congenital diarrheal disorder. J Clin Invest. 2012 Dec; 122(12):4680-4. PMID: 23114594; PMCID: PMC3533555.
  33. Moreira KE, Schuck S, Schrul B, Fröhlich F, Moseley JB, Walther TC, Walter P. Seg1 controls eisosome assembly and shape. J Cell Biol. 2012 Aug 06; 198(3):405-20. PMID: 22869600; PMCID: PMC3413353.
  34. Henriksen P, Wagner SA, Weinert BT, Sharma S, Bacinskaja G, Rehman M, Juffer AH, Walther TC, Lisby M, Choudhary C. Proteome-wide analysis of lysine acetylation suggests its broad regulatory scope in Saccharomyces cerevisiae. Mol Cell Proteomics. 2012 Nov; 11(11):1510-22. PMID: 22865919; PMCID: PMC3494197.
  35. Roczniak-Ferguson A, Petit CS, Froehlich F, Qian S, Ky J, Angarola B, Walther TC, Ferguson SM. The transcription factor TFEB links mTORC1 signaling to transcriptional control of lysosome homeostasis. Sci Signal. 2012 Jun 12; 5(228):ra42. PMID: 22692423; PMCID: PMC3437338.
  36. Farese RV, Zechner R, Newgard CB, Walther TC. The problem of establishing relationships between hepatic steatosis and hepatic insulin resistance. Cell Metab. 2012 May 02; 15(5):570-3. PMID: 22560209; PMCID: PMC3767424.
  37. Berchtold D, Piccolis M, Chiaruttini N, Riezman I, Riezman H, Roux A, Walther TC, Loewith R. Plasma membrane stress induces relocalization of Slm proteins and activation of TORC2 to promote sphingolipid synthesis. Nat Cell Biol. 2012 Apr 15; 14(5):542-7. PMID: 22504275.
    View in: PubMed
  38. Walther TC, Farese RV. Lipid droplets and cellular lipid metabolism. Annu Rev Biochem. 2012; 81:687-714. PMID: 22524315; PMCID: PMC3767414.
  39. Walther T. Tobias Walther: floating ideas on lipids. Interview by Caitlin Sedwick. J Cell Biol. 2012 Jan 23; 196(2):182-3. PMID: 22270913; PMCID: PMC3265955.
  40. Ziólkowska NE, Christiano R, Walther TC. Organized living: formation mechanisms and functions of plasma membrane domains in yeast. Trends Cell Biol. 2012 Mar; 22(3):151-8. PMID: 22245053.
    View in: PubMed
  41. Karotki L, Huiskonen JT, Stefan CJ, Ziólkowska NE, Roth R, Surma MA, Krogan NJ, Emr SD, Heuser J, Grünewald K, Walther TC. Eisosome proteins assemble into a membrane scaffold. J Cell Biol. 2011 Nov 28; 195(5):889-902. PMID: 22123866; PMCID: PMC3257569.
  42. Krahmer N, Guo Y, Wilfling F, Hilger M, Lingrell S, Heger K, Newman HW, Schmidt-Supprian M, Vance DE, Mann M, Farese RV, Walther TC. Phosphatidylcholine synthesis for lipid droplet expansion is mediated by localized activation of CTP:phosphocholine cytidylyltransferase. Cell Metab. 2011 Oct 05; 14(4):504-15. PMID: 21982710; PMCID: PMC3735358.
  43. Fei W, Shui G, Zhang Y, Krahmer N, Ferguson C, Kapterian TS, Lin RC, Dawes IW, Brown AJ, Li P, Huang X, Parton RG, Wenk MR, Walther TC, Yang H. A role for phosphatidic acid in the formation of "supersized" lipid droplets. PLoS Genet. 2011 Jul; 7(7):e1002201. PMID: 21829381; PMCID: PMC3145623.
  44. Ziólkowska NE, Karotki L, Rehman M, Huiskonen JT, Walther TC. Eisosome-driven plasma membrane organization is mediated by BAR domains. Nat Struct Mol Biol. 2011 Jun 19; 18(7):854-6. PMID: 21685922.
    View in: PubMed
  45. Bewersdorf J, Farese RV, Walther TC. A new way to look at fat. Nat Methods. 2011 Feb; 8(2):132-3. PMID: 21278725.
    View in: PubMed
  46. Walther TC, Mann M. Mass spectrometry-based proteomics in cell biology. J Cell Biol. 2010 Aug 23; 190(4):491-500. PMID: 20733050; PMCID: PMC2928005.
  47. Sourris KC, Morley AL, Koitka A, Samuel P, Coughlan MT, Penfold SA, Thomas MC, Bierhaus A, Nawroth PP, Yamamoto H, Allen TJ, Walther T, Hussain T, Cooper ME, Forbes JM. Receptor for AGEs (RAGE) blockade may exert its renoprotective effects in patients with diabetic nephropathy via induction of the angiotensin II type 2 (AT2) receptor. Diabetologia. 2010 Nov; 53(11):2442-51. PMID: 20631980; PMCID: PMC4926314.
  48. Aguilar PS, Fröhlich F, Rehman M, Shales M, Ulitsky I, Olivera-Couto A, Braberg H, Shamir R, Walter P, Mann M, Ejsing CS, Krogan NJ, Walther TC. A plasma-membrane E-MAP reveals links of the eisosome with sphingolipid metabolism and endosomal trafficking. Nat Struct Mol Biol. 2010 Jul; 17(7):901-8. PMID: 20526336; PMCID: PMC3073498.
  49. Walther TC. Keeping sphingolipid levels nORMal. Proc Natl Acad Sci U S A. 2010 Mar 30; 107(13):5701-2. PMID: 20304791; PMCID: PMC2851878.
  50. Walther TC, Olsen JV, Mann M. Yeast expression proteomics by high-resolution mass spectrometry. Methods Enzymol. 2010; 470:259-80. PMID: 20946814.
    View in: PubMed
  51. Aguilar PS, Heiman MG, Walther TC, Engel A, Schwudke D, Gushwa N, Kurzchalia T, Walter P. Structure of sterol aliphatic chains affects yeast cell shape and cell fusion during mating. Proc Natl Acad Sci U S A. 2010 Mar 02; 107(9):4170-5. PMID: 20150508; PMCID: PMC2840153.
  52. Koïtka A, Cao Z, Koh P, Watson AM, Sourris KC, Loufrani L, Soro-Paavonen A, Walther T, Woollard KJ, Jandeleit-Dahm KA, Cooper ME, Allen TJ. Angiotensin II subtype 2 receptor blockade and deficiency attenuate the development of atherosclerosis in an apolipoprotein E-deficient mouse model of diabetes. Diabetologia. 2010 Mar; 53(3):584-92. PMID: 19957160.
    View in: PubMed
  53. Krahmer N, Guo Y, Farese RV, Walther TC. SnapShot: Lipid Droplets. Cell. 2009 Nov 25; 139(5):1024-1024.e1. PMID: 19945384.
    View in: PubMed
  54. Farese RV, Walther TC. Lipid droplets finally get a little R-E-S-P-E-C-T. Cell. 2009 Nov 25; 139(5):855-60. PMID: 19945371; PMCID: PMC3097139.
  55. Fröhlich F, Walther TC. Comparing cellular proteomes by mass spectrometry. Genome Biol. 2009; 10(10):240. PMID: 19886975; PMCID: PMC2784314.
  56. Soufi B, Kelstrup CD, Stoehr G, Fröhlich F, Walther TC, Olsen JV. Global analysis of the yeast osmotic stress response by quantitative proteomics. Mol Biosyst. 2009 Nov; 5(11):1337-46. PMID: 19823750.
    View in: PubMed
  57. Choudhary C, Kumar C, Gnad F, Nielsen ML, Rehman M, Walther TC, Olsen JV, Mann M. Lysine acetylation targets protein complexes and co-regulates major cellular functions. Science. 2009 Aug 14; 325(5942):834-40. PMID: 19608861.
    View in: PubMed
  58. Fröhlich F, Moreira K, Aguilar PS, Hubner NC, Mann M, Walter P, Walther TC. A genome-wide screen for genes affecting eisosomes reveals Nce102 function in sphingolipid signaling. J Cell Biol. 2009 Jun 29; 185(7):1227-42. PMID: 19564405; PMCID: PMC2712959.
  59. Guo Y, Cordes KR, Farese RV, Walther TC. Lipid droplets at a glance. J Cell Sci. 2009 Mar 15; 122(Pt 6):749-52. PMID: 19261844; PMCID: PMC2714424.
  60. Wang H, Kakaradov B, Collins SR, Karotki L, Fiedler D, Shales M, Shokat KM, Walther TC, Krogan NJ, Koller D. A complex-based reconstruction of the Saccharomyces cerevisiae interactome. Mol Cell Proteomics. 2009 Jun; 8(6):1361-81. PMID: 19176519; PMCID: PMC2690481.
  61. Berchtold D, Walther TC. TORC2 plasma membrane localization is essential for cell viability and restricted to a distinct domain. Mol Biol Cell. 2009 Mar; 20(5):1565-75. PMID: 19144819; PMCID: PMC2649262.
  62. Stone SJ, Levin MC, Zhou P, Han J, Walther TC, Farese RV. The endoplasmic reticulum enzyme DGAT2 is found in mitochondria-associated membranes and has a mitochondrial targeting signal that promotes its association with mitochondria. J Biol Chem. 2009 Feb 20; 284(8):5352-61. PMID: 19049983; PMCID: PMC2643492.
  63. Moreira KE, Walther TC, Aguilar PS, Walter P. Pil1 controls eisosome biogenesis. Mol Biol Cell. 2009 Feb; 20(3):809-18. PMID: 19037108; PMCID: PMC2633383.
  64. Walther TC, Farese RV. The life of lipid droplets. Biochim Biophys Acta. 2009 Jun; 1791(6):459-66. PMID: 19041421; PMCID: PMC2782899.
  65. de Godoy LM, Olsen JV, Cox J, Nielsen ML, Hubner NC, Fröhlich F, Walther TC, Mann M. Comprehensive mass-spectrometry-based proteome quantification of haploid versus diploid yeast. Nature. 2008 Oct 30; 455(7217):1251-4. PMID: 18820680.
    View in: PubMed
  66. Matic I, Macek B, Hilger M, Walther TC, Mann M. Phosphorylation of SUMO-1 occurs in vivo and is conserved through evolution. J Proteome Res. 2008 Sep; 7(9):4050-7. PMID: 18707152.
    View in: PubMed
  67. Guo Y, Walther TC, Rao M, Stuurman N, Goshima G, Terayama K, Wong JS, Vale RD, Walter P, Farese RV. Functional genomic screen reveals genes involved in lipid-droplet formation and utilization. Nature. 2008 May 29; 453(7195):657-61. PMID: 18408709; PMCID: PMC2734507.
  68. Walther TC, Aguilar PS, Fröhlich F, Chu F, Moreira K, Burlingame AL, Walter P. Pkh-kinases control eisosome assembly and organization. EMBO J. 2007 Dec 12; 26(24):4946-55. PMID: 18034155; PMCID: PMC2094096.
  69. Walther TC, Brickner JH, Aguilar PS, Bernales S, Pantoja C, Walter P. Eisosomes mark static sites of endocytosis. Nature. 2006 Feb 23; 439(7079):998-1003. PMID: 16496001.
    View in: PubMed
  70. Hetzer MW, Walther TC, Mattaj IW. Pushing the envelope: structure, function, and dynamics of the nuclear periphery. Annu Rev Cell Dev Biol. 2005; 21:347-80. PMID: 16212499.
    View in: PubMed
  71. Walther TC, Askjaer P, Gentzel M, Habermann A, Griffiths G, Wilm M, Mattaj IW, Hetzer M. RanGTP mediates nuclear pore complex assembly. Nature. 2003 Aug 07; 424(6949):689-94. PMID: 12894213.
    View in: PubMed
  72. Walther TC, Alves A, Pickersgill H, Loïodice I, Hetzer M, Galy V, Hülsmann BB, Köcher T, Wilm M, Allen T, Mattaj IW, Doye V. The conserved Nup107-160 complex is critical for nuclear pore complex assembly. Cell. 2003 Apr 18; 113(2):195-206. PMID: 12705868.
    View in: PubMed
  73. Hinkle B, Slepchenko B, Rolls MM, Walther TC, Stein PA, Mehlmann LM, Ellenberg J, Terasaki M. Chromosomal association of Ran during meiotic and mitotic divisions. J Cell Sci. 2002 Dec 01; 115(Pt 23):4685-93. PMID: 12415012.
    View in: PubMed
  74. Kuersten S, Arts GJ, Walther TC, Englmeier L, Mattaj IW. Steady-state nuclear localization of exportin-t involves RanGTP binding and two distinct nuclear pore complex interaction domains. Mol Cell Biol. 2002 Aug; 22(16):5708-20. PMID: 12138183; PMCID: PMC133969.
  75. Walther TC, Pickersgill HS, Cordes VC, Goldberg MW, Allen TD, Mattaj IW, Fornerod M. The cytoplasmic filaments of the nuclear pore complex are dispensable for selective nuclear protein import. J Cell Biol. 2002 Jul 08; 158(1):63-77. PMID: 12105182; PMCID: PMC2173022.
  76. Hetzer M, Meyer HH, Walther TC, Bilbao-Cortes D, Warren G, Mattaj IW. Distinct AAA-ATPase p97 complexes function in discrete steps of nuclear assembly. Nat Cell Biol. 2001 Dec; 3(12):1086-91. PMID: 11781570.
    View in: PubMed
  77. Walther TC, Fornerod M, Pickersgill H, Goldberg M, Allen TD, Mattaj IW. The nucleoporin Nup153 is required for nuclear pore basket formation, nuclear pore complex anchoring and import of a subset of nuclear proteins. EMBO J. 2001 Oct 15; 20(20):5703-14. PMID: 11598013; PMCID: PMC125666.
  78. Hetzer M, Bilbao-Cortés D, Walther TC, Gruss OJ, Mattaj IW. GTP hydrolysis by Ran is required for nuclear envelope assembly. Mol Cell. 2000 Jun; 5(6):1013-24. PMID: 10911995.
    View in: PubMed
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