1. Do, H., Lee, W.S., Ghosh, P., Hollowell, T., Canfield, W. and Kornfeld, S: Human mannose 6-phosphate-uncovering enzyme is synthesized as a proenzyme that is activated by the endoprotease furin. J Biol Chem. 2002 Aug 16; 277(33):29737-44.
  2.  Doray, B., Bruns, K., Ghosh, P. and Kornfeld, S: Interaction of the cation-dependent mannose 6-phosphate receptor with GGA proteins. J Biol Chem. 2002 May 24; 277(21):18477-82.
  3. Doray, B., Bruns, K., Ghosh, P. and Kornfeld, S: Autoinhibition of the ligand-binding site of GGA1/3 VHS domains by an internal acidic cluster-dileucine motif. Proc Natl Acad Sci U S A. 2002 Jun 11;99(12):8072-7.
  4. Ghosh, P.*, Doray, B.*, Griffith, J., Geuze, H.J. and Kornfeld, S. :Cooperation of GGAs and AP-1 in packaging MPRs at the trans-Golgi network. Science. 2002 Sep 6;297(5587):1700-3. (*Equal contribution 1st and 2nd authors)
  5. Ghosh, P., Dahms, N.M. and Kornfeld, S: Mannose 6-phosphate receptors: new twists in the tale. Nature Rev Mol Cell Biol. 2003 Mar;4(3):202-12.
  6. Ghosh, P. and Kornfeld, S.: Phosphorylation-induced conformational changes regulate GGAs 1 and 3 function at the trans-Golgi network. J Biol Chem. 2003 Apr 18;278(16):14543-9.
  7. Ghosh, P. and Kornfeld, S.: AP-1 binding to sorting signals and release from clathrin-coated vesicles is regulated by phosphorylation. J Cell Biol. 2003 Mar 3;160(5):699-708. Epub 2003 Feb 25.
  8. Ghosh, P., Griffith, J., Geuze, H.J. and Kornfeld, S. : Mammalian GGAs act together to sort mannose 6-phosphate receptors. J Cell Biol. 2003 Nov 24;163(4):755-66.
  9. Ghosh, P. and Kornfeld, S.: GGA proteins: Key Players in Protein Sorting at the Trans-Golgi network. Eur. J Cell Biol, 2004, 83: (6)- 257-262.
  10. Ghosh, P. and Kornfeld , S. : The cytoplasmic tail of the cation-independent mannose 6-phosphate receptor contains four binding sites for AP-1. Arch Biochem Biophys. 2004 15;426(2):225-30.
  11. Ghosh P, Strong C, Naugler W, Haghighi P, Carethers JM. Peritoneal mice implicated in intestinal obstruction: report of a case and review of the literature. J Clin Gastroenterol. 2006 May-Jun;40(5):427-30.
  12. Ghosh, P, Santosa, A.C., Lin GY, Downs TM: De novo calcification of liver and nodal metastases in prostate carcinoma. Prostate Cancer Prostatic Dis. 2006 May 9. (Nature Publishing Group).
  13. Ghosh, P, Miyai, K and Chojkier, M. Gastric adenocarcinoma inducing portal hypertension: a rare presentation. World J Gastroenterol. 2007 Feb 14;13(6):960-3
  14. Ghosh, P and Jung, B: Giant Paraesophageal Hernia Causing Compression Pulmonary Atelectasis: Starved for Air when Satiated with Food. J Gastroenterol and Hepat. 2007 Apr;22(4):595.
  15. Ghosh, P., Olson, RK. and Harmon, G: Sugar Rush in the Colon: Why Incidental Findings on PET scan Justify Colonoscopy?, Amer J. Gastro, 2007 Nov;102(11):2609-10.
  16. Ghosh, P and Kum, J. Mesenteric panniculitis and sclerosing mesenteritis: a continuum of inflammation fibrosis. Clin Gastroenterol Hepatol. 2007 May;5(5):A32.
  17. Lee JK., Kum J., Ghosh, P. Gastrointestinal: Glycogenic acanthosis. J Gastroenterol Hepatol. 2007 Sep;22(9):1550.
  18. Lee JK and Ghosh, P. Non-metastatic cancer of the colon associated with pyogenic liver abscess. Am. J. Gastro, 2008 Mar; 103(3):798-9.
  19. Lee, JK., Ghosh, P., McWhorter, V., Payne, M., Olson, RK., Ramamoorthy, S., Krinsky, ML., Carethers, JM. Evidence that rectal sarcomatoid carcinoma arises from tubulo-villous adenoma: Tumorigenesis caught in the act. World J Gastroenterology, 2008 Jul 21;14(27):4389-94.
  20. Ghosh, P., Garcia-Marcos, M, Bornheimer, J. and Farquhar MG. Activation of Gai3 triggers cell migration via interaction with GIV. J. Cell Biol. 2008, Jul 28;182(2):381-93.
  21. Garcia-Marcos, M*, Ghosh, P* and Farquhar, MG. GIV/Girdin is a non-receptor GEF for Gαi that features a unique molecular target for regulating Akt activity. Proc Natl Acad Sci U S A. 2009 Mar 3;106(9):3178-83. * Equal contribution.
  22. Park, F., Lee, J.K., Madduri, G. and Ghosh, P. Generalized Megaviscera of Lupus: Refractory Intestinal Pseudoobstruction, Ureterohydronephrosis and Megacholedochus. July 28, World J Gastroenterology, 2009.
  23. Lee J.K., Carethers J.M., Ghosh, P, ArterioVenous Malformation within Jejunal Diverticulum: an unusual cause of massive gastrointestinal bleeding. Gastroenterol Res Pract. 2009;2009:384506.
  24. Ghosh, P., Carethers, J.M., Multiple "apple-core" lesions in chronic ischemic colitis: the great masquerader. Am J Gastroenterol. 2009 Nov;104(11):2870-2.
  25. Garcia-Marcos, M., Ghosh, P., Ear, J. and Farquhar MG. A structural determinant that renders G{alpha}i sensitive to activation by GIV/Girdin is required to promote cell migration. J Biol Chem. 2010 Feb 15. [Epub ahead of print]. Highlighted in "Faculty of 1000" by Dr Danny Dhanasekharan ( 
  26. Ghosh P., Beas, A., Bornheimer SJ., Garcia-Marcos M., Forry E.P., Johansson, C., Jung, B.H., Cabrera, BL., Carethers, J.M. and Farquhar MG. A Gαi-GIV molecular switch decides whether cells migrate or proliferate in response to growth factors. Mol Biol Cell., 2010 July 1;21(13):2338-2354. (Featured as Editor's Choice in Science Signaling AAAS- by Editor in Chief, Nancy Gough, entitled, "Move or Divide?" (;3/129/ec203) and also was selected for "Highlights in MBoC" and nominated for "Paper of the Year" by American Society of Cell Biology" (
  27. Lee J.K., Bhargava, V., Mittal, R.K., Ghosh, P. Achalasia, Alcohol-stasis and Acute Necrotizing Esophagitis: Connecting the Dots. Digestive Dis. and Sci. 2011 Feb; 56 (2): 612-614.
  28. Garcia-Marcos, M., Jung, B.H., Ear, J., Cabrera, B.L., Carethers, J.M., and Ghosh, P., Expression of GIV/Girdin, a Metastasis-Related Protein, Predicts Survival in Colorectal Cancer Patients. The FASEB J., 2011 Feb; 25 (2): 590-599. Featured in news article:
  29. Garcia-Marcos M., Ear, J., Farquhar, M. and Ghosh P. A GEF (GIV) and a GDI (AGS3) Regulate Autophagy by Balancing G protein Activity and Growth Factor Signals. Mol. Biol. Cell, 2011 Mar; 22 (5): 673-686.
  30. Garcia-Marcos, M., Ghosh, P, and Farquhar, MG. Molecular Mechanism of a Novel Oncogenic Mutation in GNAO. Oncogene, 2011, Feb 14.
  31. Lord, C., Bhandari, D., Menon, S., Ghassemian, M., Nycz, D., Hay, J., Ghosh, P., and Ferro-Novick, S. Sequential interactions with Sec23 control the direction of Vesicle traffic. In Press, Nature (Article), 2011. DOI: 10.1038/nature09969.
  32. Ghosh, P., Garcia-Marcos, M., and Farquhar, M.G. GIV/Girdin is a Rheostat which Fine-tunes Growth Factor Signals During Tumor progression. Cell Adhesion and Migration, May 1; Vol 5, Issue 3. 2011
  33. Garcia-Marcos, M., Kietrsunthorn, P.S., Wang, H., Ghosh, P, and Farquhar, MG. G protein binding sites on Calnuc (nucleobindin 1) and NUCB2 (nucleobindin 2) define a new class of Gαi-regulatory motifs. Journal of Biological Chemistry, 2011, June 8, DOI: 10.1074/jbc.M110.204099. 
  34. Lin, C., Ear J., Pavlova, Y., Mittal, Y., Kufareva, I., Ghassemian, M., Abagyan, R., Garcia-Marcos, M., and Ghosh, P. Tyrosine Phosphorylated GIV/Girdin Binds and Activates PI3-Kinase During Cell Migration. Science Signaling (AAAS, Washington), 2011, Sep 27;4(192):ra64. 
  35. Fukasawa, H., Obayashi, H., Schmieder, S., Lee, J., Ghosh, P., and Farquhar, M. G. Phosphorylation of Podocalyxin (Ser 415) prevents Rho A and Ezrin Activation and Disrupts its Interaction with the Actin Cytoskeleton. Am. J. Pathology, 2011, Nov;179(5):2254-65. Epub 2011 Sep 25.
  36. Mittal, Y., Pavlova, Y., Garcia-Marcos, M., and Ghosh, P. Src Homology 2-Containing Protein Tyrosine Phosphatase-1 (SHP-1) Binds and Dephosphorylates GIV/Girdin and Attenuates the GIV-PI3K-Akt Signaling Pathway. Journal of Biological Chemistry, 2011, Sep 16;286(37):32404-15. Epub 2011 Jul 28. 
  37. Garcia-Marcos, M., Kietrsunthorn PS, Pavlova Y., Adia, MA, Ghosh, P, Farquhar, MG. Functional characterization of the guanine nucleotide exchange factor (GEF) motif of GIV protein reveals a threshold effect in signaling.  Proc Natl Acad Sci USA., 2012 Feb 7; 109 (6): 1961-66. Epub 2012 Jan 25. 
  38. STAT3 Upregulates GIV/Girdin Expression, and GIV Enhances STAT3 Activation in a Positive Feedback Loop during Wound Healing and Tumor Invasion/Metastasis. Dunkel, Y., Ong, A., Notani, D., Mittal, Y., Lam, M., Mi, X., Ghosh, P., Journal of Biological Chemistry, 2012, Oct 12. 
  39. Guidelines for the use and interpretation of assays for monitoring autophagy. Klionsky DJ., Abdalla FC., ...Ghosh, P. , at al., Autophagy. 2012 Apr;8(4):445-544. PMID:22966490.
  40. Protein Kinase C-theta (PKCθ) phosphorylates and inhibits the guanidine exchange factor, GIV/Girdin. Lopez-Sanchez, I., Garcia-Marcos, M., Mittal, Y., Aznar, N., Farquhar, MG., and Ghosh, PProc Natl Acad Sci USA., 2013 2013 Apr 2;110(14):5510-5. PMID: 23509302.
  41. GIV/Girdin links VEGF Signaling to Akt survival signaling in Podocytes Independent of Nephrin. Wang H., Misaki T., Taupin, V., Eguchi A., Ghosh, P. and Farquhar M.G. J Am Soc Nephrol. 2014 Jul 10. pii: ASN.2013090985. PMID: 25012178
  42. Genome-wide mutational landscape of mucinous carcinomatosis peritonei of appendiceal origin. Hakan, Babicky ML, Ghosh, P. , Yost  S, Jepsen  K, Dai  Y, Arias  A, Samuels LM, Mose SE, Schwab RB, Peterson MR, Lowy AM, Frazer KA and Harismendy  O. Genome Med. 2014 May 29;6(5):43. doi: 10.1186/gm559. eCollection 2014. PMID: 24944587
  43. GIV/Girdin is a Central Hub for Pro-fibrogenic Signaling Networks during Liver Fibrosis. Lopez-Sanchez I., Dunkel., Y., Mittal., Y., Deminicis, S., Murayni, A., Singh, S., Kandavel, S., Seki, E., Ho, S.B., Brenner, D.A. and Ghosh, P. . Nat Commun. 2014 Jul 21;5:4451. doi: 10.1038/ncomms5451. PMID: 25043713
  44. Structural basis for activation of trimeric G proteins by multiple growth factor receptors via GIV/Girdin. Lin, C., Ear, J., Midde, K., Lopez-Sanchez, I., Aznar, N., Garcia-Marcos, M., Kufareva I., Abagyan, R., and Ghosh, P. . Mol Biol Cell. 2014 Nov 5;25(22):3654-71. doi: 10.1091/mbc.E14-05-0978. PMID: 25187647. This article has been featured online on Global Medical Discovery [ISSN 1929-8536] (
  45. GIV/ Girdin rewires multi-receptor signaling by triggering trimeric G protein activation. Garcia-Marcos, M., Ghosh, P. . and Farquhar, M.G. Journal of Biological Chemistry, Invited Mini-Review. J Biol Chem. 2015 Jan 20. pii: jbc.R114.613414. PMID: 25605737.
  46. Multi-modular Biosensors Reveal a Novel Platform for Activation of G Proteins by Growth Factor Receptors. Midde, K., Aznar N., Laederich, B., Ma, G.S., Kunkel, M., Newton, A.C. and Ghosh, P. . Proc Natl Acad Sci U S A. 2015 Feb 17. pii: 201420140. PMID: 25713130.
  47. Activation of Gai at the Golgi by GIV/Girdin Imposes Finiteness in Arf1 Signaling. Lo, I., Midde, K., Taupin, V., Lopez-Sanchez, I., Kufareva, I., Abagyan, R., Randazzo, P., Farquhar, M.G. and Ghosh, P. . Dev Cell. 2015 Apr 20;33(2):189-203. PMID: 25865347. Cited by Science STKE:
  48. Ma, G., Aznar, N., Kangrioloulos, N., Lopez-Sanchez, I., Midde, K., Dunkel Y, Muto J., Gallo R. and Ghosh, P. . Therapeutic Effects of Cell-Permeant GIV peptides that Activate G Proteins downstream of Growth Factors. Proc Natl Acad Sci U S A. 2015 Apr 29. pii: 201505543. PMID: 25926659.
  49. Aznar, N., Midde, K., Dunkel, Y., Lopez-Sanchez, I., Pavlova, Y., Marvin, A., Barbazán, J., Murray, F., Nitsche, U., Janssen, K.P., Willert, K., Goel, A., Abal, M., Garcia-Marcos, M., and Ghosh, P. . Daple is a Novel Non-Receptor GEF Required for Trimeric G protein Activation in Wnt Signaling. Elife. 2015 Jun 30;4:e07091. PMID: 26126266
  50. Ghosh, P. . G protein coupled Growth Factor Receptor Tyrosine Kinase: No longer an Oxymoron. Cell Cycle. 2015 Aug 18;14(16):2561-5. doi: 10.1080/15384101.2015.1066538. PMID: 26126131.
  51. Ghosh, P. .Heterotrimeric G proteins as Emerging Targets for Network Based Therapy in Cancer: End of a long futile campaign striking heads of a Hydra. Aging (Albany NY). 2015 Jul;7(7):469-74. PMID: 26224586
  52. Bhandari D, Lopez-Sanchez I, To A, Lo IC, Aznar N, Leyme A, Gupta V, Niesman I, Maddox AL, Garcia-Marcos M, Farquhar MG, Ghosh, P. Cyclin-dependent kinase 5 activates guanine nucleotide exchange factor GIV/Girdin to orchestrate migration-proliferation dichotomy. Proc Natl Acad Sci U S A. 2015 Sep 1;112(35):E4874-83. doi: 10.1073/pnas.1514157112. PMID: 26286990
  53. Ma, G., Lopez-Sanchez, I., Aznar, N., Midde, K., Pedram, S., Ciaraldi, T., Henry R.R. and Ghosh, P . GIV/Girdin serves as a molecular switch between insulin sensitivity and resistance. Mol Biol Cell. 2015 Nov 15;26(23):4209-23. PMID: 26378251.
  54. Lopez-Sanchez I., Kalogriopoulos, N., Kabir, F., Midde, K., Wang, HH and Ghosh, P . Focal Adhesions are Concentrated Patches for Tyrosine-Based Signal Transduction via GIV/Girdin and G proteins. Mol Biol Cell. 2015 Dec 1;26(24):4313-24.. PMID: 26446841.
  55. Lopez-Sanchez I., Ma, G.S., Pedram, S., Kalogriopoulos, N., and Ghosh, P . GIV/girdin binds exocyst subunit-Exo70 and regulates exocytosis of GLUT4 storage vesicles. Biochem Biophys Res Commun. 2015 Dec 4;468(1-2):287-93. PMID: 26514725.
  56. Aznar N., Kalogriopoulos, N., Midde, K., Lo, I and Ghosh, P . Heterotrimeric G protein signaling via GIV/Girdin: Breaking the rules of engagement, space and time. Accepted, Peer-reviewed Invited Review, In Press, BioEssays, 2016.
  57. Ghosh, P . The untapped potential of tyrosine-based G protein signaling. Peer-reviewed Invited Perspective; Pharmacol. Res., 2016 Jan 22;105:99-107. PMID: 26808081
  58. Barbazan, J., Dunkel, Y., Li, H., Nitsche, U., Janssen, K.-P., Messer, K. & Ghosh, P . Prognostic Impact of Modulators of G proteins in Circulating Tumor Cells from Patients with Metastatic Colorectal Cancer. Nat Sci Rep. 2016 Feb 26;6:22112. doi: 10.1038/srep22112. PMID: 26916336.
  59. Ghosh, P , Tie, J., Muranyi, A., Singh, S., Brunhoeber, P., Leith, K., Bowermaster, R., Liao, Z., Zhu, Y., LaFleur, B., Tran, B., Desai, J., Jones, I., Croxford, M., Jover, R., Goel, A., Waring, P., Hu, S., Teichgraeber, V., Rohr, U.-P., Ridder, R., Shanmugam, K. & Gibbs, P. Girdin (GIV) Expression as a Prognostic Marker of Recurrence in Mismatch Repair Proficient Stage II Colon Cancer. Clin Cancer Res. 2016 Mar 30. PMID: 27029492.
  60. El Ghazal R, Yin X, Johns SC, Swanson L, Macal M, Ghosh P, Zuniga EI, Fuster MM. Neoplasia. 2016 May;18(5):294-306. doi: 10.1016/j.neo.2016.04.004. [PMID: 27237321]
  61. Dunkel., Y., Dao, K., Aznar, N., Swanson, L., Lu, L., Zhu, W., Mi, X., and Ghosh, P . Prognostic impact of total and tyrosine-phosphorylated GIV/Girdin in breast cancers. FASEB J. 2016 Jul 20. [PMID: 27440794].
  62. Gupta, V., Bhandari, D., Leyme, A., Aznar, N., Midde, K., Lo, I., Ear, J., Niesman, I., López Sánchez, I., Blanco-Canosa, J.B., von Zastrow, M., Garcia-Marcos, M., Farquhar, M.G. and Ghosh, P . GIV/Girdin activates Gαi and inhibits Gαs via the same motif. Accepted, In Press, Proc Natl Acad Sci U S A, 2016.
  63. Banerjee, S., Aznar, N., Dwivedi, SKD., Chakraborty, PK., Basak, R., Mukherjee, P., Ghosh, P , and Bhattacharya, R. Mitochondrial BMI1 maintains bioenergetic homeostasis in cells. Accepted, In Press, FASEB J., 2016.
  64. Prognostic impact of total and tyrosine phosphorylated GIV/Girdin in breast cancers. Dunkel Y, Diao K, Aznar N, Swanson L, Liu L, Zhu W, Mi X, Ghosh P. FASEB J. 2016 Jul 20. pii: fj.201600500. PMID: 27440794
  65. GIV/Girdin activates Gαi and inhibits Gαs via the same motif. Gupta V, Bhandari D, Leyme A, Aznar N, Midde KK, Lo IC, Ear J, Niesman I, López-Sánchez I, Blanco-Canosa JB, von Zastrow M, Garcia-Marcos M, Farquhar MG, Ghosh P. Proc Natl Acad Sci U S A. 2016 Sep 12. pii: 201609502.
  66. AMP-activated protein kinase fortifies epithelial tight junctions during energetic stress via its effector GIV/Girdin.
    Aznar N, Patel A, Rohena CC, Dunkel Y, Joosen LP, Taupin V, Kufareva I, Farquhar MG, Ghosh P.
    Elife. 2016 Nov 4;5. pii: e20795. doi: 10.7554/eLife.20795. PMID: 27813479
  67. Biochemical, Biophysical and Cellular Techniques to Study the Guanine
    Nucleotide Exchange Factor, GIV/Girdin. Ghosh, P., Aznar, N., Swanson, L., Lo, I., Lopez-Sanchez, I., Ear, J., Rohena, C., Kalogriopoulos, N.A., Joosen, L., Dunkel, Y., Sun, N., Nguyen, P., Bhandari, D.  Current Protocols in Chemical Biology. Dec 2016.
  68. The GAPs, GEFs, and GDIs, GEMs: New kids on the heterotrimeric G protein signaling block. Ghosh, P., Rangamani, P., and Kufareva, I. Cell Cycle. Jan. 2017. PMID: Pending. 
  69. Selected Src Family Kinases Are Vital For Type I Interferon Production By Plasmacytoid Dendritic Cells. Dallari, S., Macal, M., Loureiro, ME., Jo, Y., Swanson, L., Hesser, C., Ghosh, P., and Zuniga, EI. Nature Communications. Jan 2017. PMID: Pending. 
  70. The Stress Polarity Pathway: AMPK 'GIV-es' protection against metabolic insults. Ghosh, P. Aging, 2017. PMID: Pending. 
  71. The GAPs, GEFs, GDIs and…now, GEMs: New kids on the heterotrimeric G protein signaling block. Ghosh P, Rangamani P, Kufareva I. Cell Cycle. 2017 Apr 3;16(7):607-612. doi: 10.1080/15384101.2017.1282584. Epub 2017 Mar 13. Review.
  72. A Daple-Akt feed-forward loop enhances noncanonical Wnt signals by compartmentalizing β-catenin. Aznar N, Sun N, Dunkel Y, Ear J, Buschman MD, Ghosh P. Mol Biol Cell. 2017 Dec 1;28(25):3709-3723. doi: 10.1091/mbc.E17-06-0405. Epub 2017 Oct 11.
  73. The Alzheimer's disease-protective CD33 splice variant mediates adaptive loss of function via diversion to an intracellular pool.Siddiqui SS, Springer SA, Verhagen A, Sundaramurthy V, Alisson-Silva F, Jiang W, Ghosh P, Varki A.J Biol Chem. 2017 Sep 15;292(37):15312-15320. doi: 10.1074/jbc.M117.799346. Epub 2017 Jul 26.
  74. Erythrocyte sialoglycoproteins engage Siglec-9 on neutrophils to suppress activation.Lizcano A, Secundino I, Döhrmann S, Corriden R, Rohena C, Diaz S, Ghosh P, Deng L, Nizet V, Varki A.Blood. 2017 Jun 8;129(23):3100-3110. doi: 10.1182/blood-2016-11-751636. Epub 2017 Apr 17. PMID: 28416510
  75. Convergence of Wnt, growth factor, and heterotrimeric G protein signals on the guanine nucleotide exchange factor Daple. Aznar N, Ear J, Dunkel Y, Sun N, Satterfield K, He F, Kalogriopoulos NA, Lopez-Sanchez I, Ghassemian M, Sahoo D, Kufareva I, Ghosh P. Sci Signal. 2018 Feb 27;11(519). pii: eaao4220. doi: 10.1126/scisignal.aao4220. PMID: 29487190.