Peer-reviewed Publications

  1. 2021, Peptide-based Human Bladder Cancer Trial – Nat Comm
  2. 2020, Bladder Cancer Therapy Without Toxicity – Int J Cancer
  3. 2020, Origins of Cell Division – Mol Biol Evol.
  4. 2019, Targeting of Heat Shock Proteins – J Mol Biol
  5. 2016, Targeting of Nucleotide-binding Proteins – Oncogene
  6. 2016, Protein-dependent Membrane Interaction – Sci Reports
  7. 2015, Receptor-independent Membrane Remodeling – Sci Reports
  8. 2015, Targeting The Molecular Motor F-ATP Synthase – J Mol Biol
  9. 2014, Colon Cancer Treatment – Gut
  10. 2013, Colon Cancer Highlight – Nature Reviews Gastroenterology & Hepatology
  11. 2013, Cell Death Through Ion Channel Activation – PLoS ONE
  12. 2012, Review-Functional Properties And Therapeutic Potential – Future Oncol
  13. 2012, HAMLET Structure – PLoS ONE
  14. 2011, Apoptosis-like Death in Bacteria – PLoS ONE
  15. 2010, Review, Structure – FEBS Journal
  16. 2010, Lipid Membranes And Perturbs Their Structure And Integrity – PLoS ONE
  17. 2010, Treatment Of Bladder Cancer – J Urology
  18. 2008, Review – Ann Med
  19. 2007, Study Of Human Bladder Cancer- Int J Cancer
  20. 2007, Histone Acetylation – Cancer Res
  21. 2007, Review-HAMLET, Protein Folding, And Tumor Cell Death – Bioch Bioph Res Com
  22. 2006, Species Variation Of Alpha-lactalbumins – Bioch Bioph Res Com
  23. 2006, Cell Death Is Not By Apoptosis – Apoptosis
  24. 2005, Review Advances
  25. 2005, Review Advances – J Nutri
  26. 2004, Clinical Study Of Human Skin Papillomas – N Engl J Med
  27. 2004, Brain Tumor Therapy – Cancer Res
  28. 2003, HAMLET Targets Nuclear Histones – J Biol Chem
  29. 2003, Defining Oleic Acid As The Lipid Cofactor – Protein Sci
  30. 2000, The Definition Of The HAMLET Complex – PNAS
  31. 2000, Antibacterial Effects – Mol Microbiol
  32. 1995, The Discovery Of HAMLET’s Effect – PNAS


    1. Antonín Brisuda, James C. S. Ho, Pancham S. Kandiyal, Justin T-Y. Ng, Ines Ambite, Daniel S. C. Butler, Jaromir Háček, Murphy Lam Yim Wan, Thi Hien Tran, Aftab Nadeem, Tuan Hiep Tran, Anna Hastings, Petter Storm, Daniel L. Fortunati, Parisa Esmaeili, Hana Novotna, Jakub Horňák, Y. G. Mu , K. H. Mok, Marek Babjuk & Catharina Svanborg. Nat Comm 12, 3427 (2021).
    2. Hien Tran T, Ambite I, Butler D, Wan MLY, Tran HT, Höglund U, Babjuk M and Svanborg C. 2020. Bladder cancer therapy without toxicity – A dose-escalation study of alpha1-oleate. Int J Cancer 147(9):2479–2492.
    3. Hansen JS, Tran TH, Cavalera M, Paul S, Chaudhuri A, Lindkvist-Petersson K, Ho J and Svanborg C. 2020. Peptide-oleate complexes create novel membrane-bound compartments. Mol Biol Evol 37(11):3083-3093.
    4. Nadeem A, Ho J, Tran, TH, Paul S, Granqvist V, Despretz N and Svanborg C. 2019. Beta-sheet-specific interactions with heat shock proteins define a mechanism of delayed tumor cell death in response to HAMLET. J Mol Biol 431(14): 2612-27.
    5. Ho CS J., Nadeem A, Rydström A, Puthia M and Svanborg C. 2016. Targeting of nucleotide-binding proteins by HAMLET – a conserved tumor cell death mechanism. Oncogene 35, 897-907.
    6. Chaudhuri A, Prasanna X, Agiru P, Chakraborty H, Rydström A, Ho JC, Svanborg C, Sengupta D, Chattopadhyay A. 2016. Protein-dependent Membrane Interaction of A Partially Disordered Protein Complex with Oleic Acid: Implications for Cancer Lipidomics. Sci Reports 6:35015.
    7. Nadeem A, Ho CS J, Sanborn J, Rydström A, Gettel DL, Ngassam VN, Klausen TK, Pedersen SF, Lam M, Parikh AN and Svanborg C. 2015. Protein receptor-independent plasma membrane remodeling by HAMLET; a tumoricidal protein-lipid complex. Sci Reports 5:16432.
    8. Ho CS J, Sielaff H, Nadeem A, Svanborg C and Gruber G. 2015. The molecular motor F-ATP synthase is targeted by the tumoricidal protein HAMLET. J Mol Biol 427(10):1866-74.
    9. Puthia M, Storm P, Nadeem A, Hsiung S, Svanborg C. 2014. Prevention and treatment of colon cancer by peroal administration of HAMLET (human α-lactalbumin made lethal to tumour cells). Gut 63: 131-42.
    10. Smith K. 2013. Therapy: HAMLET takes a leading role on the colorectal cancer stage. Nat Rev Gastroenterol Hepatol 10:126.
    11. Storm P, Kjaer Klausen T, Trulsson M, Ho CS J, Dosnon M, Westergren T, Chao YX, Rydstrom A, Yang H, Pedersen SF, Svanborg C. 2013. A Unifying Mechanism for Cancer Cell Death through Ion Channel Activation by HAMLET. PLoS ONE 8(3): e58578.
    12. James Ho CS1, Anna Rydström1, Maria Trulsson1, Johannes Bålfors1, Petter Storm1, Manoj Puthia1, Aftab Nadeem1 & Catharina Svanborg. Hamlet: functional properties and therapeutic potential. Review future Oncol (2012) 8(10), 1301-1313.
    13. Ho CS J, Rydstrom A, Manimekalai MSS, Svanborg C, Grüber G. 2012. Low resolution solution structure of HAMLET and the importance of its alpha-domains in tumoricidal activity. PLoS ONE 7(12): e53051.
    14. Hakansson A, Roche-Hakansson H, Mossberg AK and Svanborg C. 2011. Apoptosis-like death in bacteria induced by HAMLET, a human milk lipid-protein complex. PLoS ONE 3:e17717.
    15. Ho CS J, Storm P, Rydström A, Bowen B, Alsin F, Sullivan L, Ambite I, Hun Mok K, Northen T and Svanborg C. 2013. Lipids as Tumoricidal Components of Human Alpha-lactalbumin Made Lethal to Tumor Cells (HAMLET); Unique and Shared Effects on Signaling and Death. J Biol Chem doi:10.1074/jbc.M113.468405.
    16. Mossberg AK, Puchades M, Halskau O, Baumann A, Lanekoff I, Chao Y, Martinez A, Svanborg C and Karlsson R. 2010. HAMLET interacts with lipid membranes and perturbs their structure and integrity. PLoS ONE 2:e9384.
    17. Mossberg AK, Hou Y, Svensson M, Holmqvist B and Svanborg C. 2010. HAMLET treatment delays bladder cancer development. J. Urology 183:1590.
    18. Xie Y, Min S, Harte NP, Kirk H, O’Brien JE, Voorheis HP, Svanborg C, Hun Mok K. 2013. Electrostatic interactions play an essential role in the binding of oleic acid with α-lactalbumin in the HAMLET-like complex: A study using charge-specific chemical modifications. Proteins 81(1): 1-17.
    19. Baumann A, Gjerde AU, Ying M, Svanborg C, Holmsen H, Glomm WR, Martinez A, Halskau O. 2012. HAMLET forms annular oligomers when depostited with phospholipid monolayers. J Mol Biol 418(1-2): 90-102.
    20. Storm P, Aits S, Puthia MK, Urbano A, Northen T, Powers S, Bowen B, Chao Y, Reindl W, Lee DY, Sullivan NL, Zhang J, Trulsson M, Yang H, Watson JD and Svanborg C. 2011. Conserved features of cancer cells define their sensitivity of HAMLET-induced death; c-Myc and glycolysis. Oncogene 30, 4765–4779.
    21. Trulsson M, Yu H, Gisselsson L, Chao Y, Urbano A, Aits S, Mossberg AK and Svanborg, C. 2011. HAMLET binding to α-actinin facilitates tumor cell detachment. PLoS ONE 3:e17179.
    22. Pettersson-Kastberg J, Mossberg AK, Trulsson M, Yong YJ, Min S, Lim Y, O’Brien JE, Svanborg C, Mok KH. 2009. Alpha-Lactalbumin, engineered to be nonnative and inactive, kills tumor cells when in complex with oleic acid: a new biological function resulting from partial unfolding. J Mol Biol 394(5):994.
    23. Gustafsson, L., Aits, S., Onnerfjord, P., Trulsson, M., Storm, P., Svanborg, C. 2009. Changes in proteasome structure and function caused by HAMLET in tumor cells. PLoS ONE 4:e5229.
    24. Aits, S., Gustafsson, L., Hallgren, O., Brest, P., Gustafsson, M., Trulsson, M., Mossberg, A. K., Simon, H.-U., Mograbi, B. and Svanborg, C. 2009. HAMLET (human alpha-lactalbumin made lethal to tumor cells) triggers autophagic tumor cell death. Int J Cancer 124:1008.
    25. Brest, P., Gustafsson, M., Mossberg, A. K., Gustafsson, L., Duringer, C., Hamiche, A., Svanborg, C. 2007. Histone deacetylase inhibitors promote the tumoricidal effect of HAMLET. Cancer Res 67:11327.
    26. Mossberg, A. K., Wullt, B., Gustafsson, L., Mansson, W., Ljunggren, E., Svanborg, C. 2007. Bladder cancers respond to intravesical instillation of HAMLET (human alpha-lactalbumin made lethal to tumor cells). Int J Cancer 121:1352.
    27. Pettersson, J., Mossberg, A.-K., Svanborg. C. 2006. alpha-Lactalbumin species variation, HAMLET formation, and tumor cell death. Biochem Biophys Res Commun 345:260.
    28. Hallgren, O., Gustafsson, L., Irjala, H., Selivanova, G., Orrenius, S., Svanborg, C. 2006. HAMLET triggers apoptosis but tumor cell death is independent of caspases, Bcl-2 and p53. Apoptosis 11:221.
    29. Fast, J., Mossberg, A. K., Nilsson, H., Svanborg, C., Akke, M., Linse, S. 2005. Compact oleic acid in HAMLET. FEBS Lett 579:6095.
    30. Fast, J., Mossberg, A. K., Svanborg, C., Linse, S. 2005. Stability of HAMLET–a kinetically trapped alpha-lactalbumin oleic acid complex. Protein Sci 14:329.
    31. Gustafsson, L., I. Leijonhufvud, A. Aronsson, A. K. Mossberg, and C. Svanborg. 2004. Treatment of skin papillomas with topical alpha-lactalbumin-oleic acid. N Engl J Med350:2663.
    32. Baltzer, A., C. Svanborg, and R. Jaggi. 2004. Apoptotic cell death in the lactating mammary gland is enhanced by a folding variant of alpha-lactalbumin. Cell Mol Life Sci61:1221.
    33. Casbarra, A., L. Birolo, G. Infusini, F. Dal Piaz, M. Svensson, P. Pucci, C. Svanborg, and G. Marino. 2004. Conformational analysis of HAMLET, the folding variant of human alpha-lactalbumin associated with apoptosis. Protein Sci 13:1322.
    34. Fischer, W., L. Gustafsson, A. K. Mossberg, J. Gronli, S. Mork, R. Bjerkvig, and C. Svanborg. 2004. Human alpha-lactalbumin made lethal to tumor cells (HAMLET) kills human glioblastoma cells in brain xenografts by an apoptosis-like mechanism and prolongs survival. Cancer Res 64:2105.
    35. Duringer, C., A. Hamiche, L. Gustafsson, H. Kimura, and C. Svanborg. 2003. HAMLET Interacts with Histones and Chromatin in Tumor Cell Nuclei. J Biol Chem 278:42131.
    36. Svensson, M., J. Fast, A. K. Mossberg, C. Duringer, L. Gustafsson, O. Hallgren, C. L. Brooks, L. Berliner, S. Linse, and C. Svanborg. 2003. Alpha-lactalbumin unfolding is not sufficient to cause apoptosis, but is required for the conversion to HAMLET (human alpha-lactalbumin made lethal to tumor cells). Protein Sci 12:2794.
    37. Svensson, M., A. K. Mossberg, J. Pettersson, S. Linse, and C. Svanborg. 2003. Lipids as cofactors in protein folding: stereo-specific lipid-protein interactions are required to form HAMLET (human alpha-lactalbumin made lethal to tumor cells). Protein Sci 12:2805.
    38. Kohler, C., V. Gogvadze, A. Hakansson, C. Svanborg, S. Orrenius, and B. Zhivotovsky. 2001. A folding variant of human alpha-lactalbumin induces mitochondrial permeability transition in isolated mitochondria. Eur J Biochem 268:186.
    39. Svensson, M., A. Hakansson, A. K. Mossberg, S. Linse, and C. Svanborg. 2000. Conversion of alpha-lactalbumin to a protein inducing apoptosis. Proc Natl Acad Sci USA 97:4221.
    40. Hakansson, A., M. Svensson, A. K. Mossberg, H. Sabharwal, S. Linse, I. Lazou, B. Lonnerdal, and C. Svanborg. 2000. A folding variant of alpha-lactalbumin with bactericidal activity against Streptococcus pneumoniae. Mol Microbiol 35:589.
    41. Kohler, C., A. Hakansson, C. Svanborg, S. Orrenius, and B. Zhivotovsky. 1999. Protease activation in apoptosis induced by MAL. Exp Cell Res 249:260.
    42. Svensson, M., H. Sabharwal, A. Hakansson, A. K. Mossberg, P. Lipniunas, H. Leffler, C. Svanborg, and S. Linse. 1999. Molecular characterization of alpha-lactalbumin folding variants that induce apoptosis in tumor cells. J Biol Chem 274:6388.
    43. Hakansson, A., J. Andreasson, B. Zhivotovsky, D. Karpman, S. Orrenius, C. Svanborg. 1999. Multimeric alpha-lactalbumin from human milk induces apoptosis through a direct effect on cell nuclei. Exp Cell Res 246:451.
    44. Hakansson, A., Zhivotovsky, B., Orrenius, S., Sabharwal, H., Svanborg, C. 1995. Apoptosis induced by a human milk protein. Proc Natl Acad Sci USA 92:8064.


R1.       Svensson, M., C. Duringer, O. Hallgren, A. K. Mossberg, A. Hakansson, S. Linse, and C. Svanborg. 2002. Hamlet–a complex from human milk that induces apoptosis in tumor cells but spares healthy cells. Adv Exp Med Biol 503:125.

R2.       Svanborg, C., H. Agerstam, A. Aronson, R. Bjerkvig, C. Duringer, W. Fischer, L. Gustafsson, O. Hallgren, I. Leijonhuvud, S. Linse, A. K. Mossberg, H. Nilsson, J. Pettersson, and M. Svensson. 2003. HAMLET kills tumor cells by an apoptosis-like mechanism–cellular, molecular, and therapeutic aspects. Adv Cancer Res 88:1.

R3.       Duringer, C., Hallgren, O., Gustafsson, L., Pettersson, J., Mossberg, A. K., Manolov, T. and Svanborg, C. 2005. HAMLET; a novel tool to identify apoptotic pathways in tumor cells. in Sluyser, M. (Ed), Application of apoptosis to cancer treatment., Springer, pp. 223-246.

R4.       Gustafsson, L., Hallgren, O., Mossberg, A. K., Pettersson, J., Fischer, W., Aronsson, A. and Svanborg, C. 2005. HAMLET Kills Tumor Cells by Apoptosis: Structure, Cellular Mechanisms, and Therapy. J Nutr 135:1299.

R5.       Mok, K. H., Pettersson, J., Orrenius, S. and Svanborg, C. 2007. HAMLET, protein folding, and tumor cell death. Biochem Biophys Res Commun 354:1.

 R6.       Hallgren, O., Aits, S., Brest, P., Gustafsson, L., Mossberg, A. K., Wullt, B. and Svanborg, C. 2008. Apoptosis and tumor cell death in response to HAMLET (human alpha-lactalbumin made lethal to tumor cells). Adv Exp Med Biol 606:217.

R7.       Pettersson-Kastberg, J., Aits, S., Gustafsson L., Mossberg A., Storm, P., Trulsson, M., Persson, F., Mok, K. H. and Svanborg, C. 2009. Can misfolded proteins be beneficial? The HAMLET case. Ann med 41:162.

R8.       Svanborg, C., Aits, S., Brest, P., Gustafsson, L., Mok, K., Moreillhon, C., Mossberg, A., Pettersson, J., Storm, P. and Trulsson, M. 2009. HAMLET, a tumoricidal molecular complex from human milk. BMC Proceedings 3(Suppl 5):S11

R9.       Mossberg, A. K., Mok, K. H., Morozova-Roche, L. A. and Svanborg, C. 2010. Structure and function of human a-lactalbumin made lethal to tumor cell (HAMLET)-type complexes. FEBS Journal 277:4614.

P10.     Min S, Meehan J, Sullivan LM, Harte NP, Xie Y, Davey GP, Svanborg C, Brodkorb A, Mok KH. (2012) Alternatively folded proteins with unexpected beneficial functions. Biochem Soc Trans. 40(4):746-751.

P11.     Ho CS J, Rydström A, Trulsson M, Bålfors J, Storm P, Puthia M, Nadeem A and Svanborg C. 2012. HAMLET: functional properties and therapeutic potential. Future Oncol. 8(10), 1301-1313.

P12.    Ho JCS, Nadeem A and Svanborg C. 2017. HAMLET – A protein-lipid complex with broad tumoricidal activity. Biochem Biophys Res Commun 482 (3), 454–458.