Robert J. Tomko Jr. Ph.D.
Professor
Biosketch
Tomko Lab Webpage
Dr. Tomko completed his graduate studies in pharmacology at the University Of Pittsburgh School Of Medicine’s Drug Discovery Institute, where he became familiar with modern drug discovery and development. His thesis work probing the regulation of a short-lived protein sparked his interest in the cell’s machinery to recycle damaged or unneeded proteins. To pursue this interest, he moved to Yale University, where he was an American Cancer Society Postdoctoral Fellow in the Department of Molecular Biophysics and Biochemistry. As a postdoctoral fellow, Robert made several seminal discoveries regarding the assembly of the proteasome, a large, multisubunit protease complex that executes most of the cell’s regulatory and quality control protein degradation. In 2015, Dr. Tomko joined the faculty of Biomedical Sciences at FSU, where his group studies the structure and function of ubiquitin-proteasome system components in normal and disease states. His laboratory works on the scale of individual proteins up to whole cells, and integrates approaches spanning biochemistry, cell biology, genetics, structural biology, proteomics, biophysics, and pharmacology. Ultimately, his group aims to utilize the information gained from these basic studies of the ubiquitin-proteasome system to identify new targets and develop pharmacological modulators with potential thereapeutic benefit.
Education
American Cancer Society Postdoctoral Fellow, Yale University, Department of Molecular Biophysics and Biochemistry
Ph.D., Pharmacology and Chemical Biology, University of Pittsburgh School of Medicine and Drug Discovery Institute
B.S., Biochemistry and Molecular Biology, Lebanon Valley College
Service
2024 - Present Grants reviewer, National Science Foundation
2023 - 2024 Senator, Florida State University Senate
2022 - Present Grants reviewer, US-Israel Binational Science Foundation
2022 - Present Reviewing member, Florida Agricultural & Mechanical University Institutional Biosafety Committee
2021 - Present Editorial Board Member, the Journal of Biological Chemistry
2021 - 2022 Member, Faculty Recruitment Committee, Department of Biomedical Sciences
2019 - Present Scientific advisor, Wetware development, FSU iGEM Program
2019 - Present Member, FSU-COM Physician Assistant Program Student Progress Committee
2019 Session chair, Protein Interactions, Modifications, and Regulation Section, Experimental Biology 2019 Meeting
2019 MBPP study section member, ad hoc
2019 - Present Invited grants reviewer, Israel Science Foundation
2017 - Present Member, Graduate Policy Committee, Department of Biomedical Sciences
2016 - 2019 Florida State University College of Medicine Research Advisory Committee
2016 - 2018 Dept. of Biomedical Sciences Promotion and Tenure Committee
2015 - Present Dept. of Biomedical Sciences Faculty Search Committee
2015 - Present Dept. of Biomedical Sciences Postdoctoral Career and Mentoring Committee
Honors/Awards
2026 ZRG1 MGG-K Study Section, National Institutes of Health
2025 CSF1 Study Section, National Institutes of Health
2025 Commencement Speaker, Florida State University Physician Assistant Program Class of 2025
2025 Florida State University Developing Scholar Award (awarded to exceptionally promising FSU researchers
2025 Florida State University Distinguished Teacher Award (University's highest honor for educators)
2024 Florida State University College of Medicine Outstanding Senior Faculty Researcher Award
2021 Florida State University Undergraduate Faculty Research Mentor Award
2019 Florida State University Graduate Teaching Award
2018 Florida State University College of Medicine Outstanding Junior Faculty Researcher Award
2018 Florida State University College of Medicine Outstanding Junior Faculty Educator Award
2015 First-Year Assistant Professor Award
2014 Finalist, Earl Stadtman Symposium on Molecular Biology and Biochemistry NIH, Bethesda MD
2013 Nominee, Blavatnik Award for Young Scientists
2010 - 2013 American Cancer Society Postdoctoral Fellowship
2009 - 2010 James Hudson Brown – Alexander B. Coxe Postdoctoral Fellowship in the Medical Sciences
2007 University of Pittsburgh School of Medicine Most Well-Rounded Graduate Student Award
2002 American Chemical Society Outstanding Chemistry Major Award
2002 Southeastern PA Section American Chemical Society Award
2001 American Chemical Society Award for Outstanding Performance in Organic Chemistry
Memberships
2015 - Present, American Society for Biochemistry and Molecular Biology
2006 - Present, American Association for Cancer Research
2005 - Present, American Society for Pharmacology and Experimental Therapeutics
Courses
Medical program
BMS-6037 Foundations of Medicine II: Molecules to Mechanisms (Small group activity leader)
BMS-6041 Host Defense (Instructor)
Physician assistant program
PAS-5071 Clinical Pharmacology I (Founding course director; instructor)
PAS-5072 Clinical Pharmacology II (Founding course director; instructor)
Biomedical Sciences Ph.D. program
BMS-5525 Bioregulation (Instructor)
GMS-6003 Professional Development (Instructor)
Research Focus
Regulation of proteasome assembly and function in health and infectious disease; multidisciplinary analyses of biomedically important molecular machines; proteases and proteolysis
Publications
Dr. Tomko's full list of publications can be found HERE.
Selected publications from the Tomko lab (student authors are underlined):
Rout, S. R., Kulke, M., Droemer, M. A., Wendel, M., Cheng, T. C., Mauck, T. A., Asgari, S., Nemec, A. A., Shein, M., Sato, Y., Fukai, S., Witte, G., Tomko, R. J. Jr., Stengel, F., Zacharias, M., Schuetz, A. K., and Sakata, E. The vacuolar tauopathy-associated mutation D395G confers redox sensitivity to p97/VCP. BioRxiv (preprint). https://www.biorxiv.org/content/10.64898/2026.04.16.718620v2
Gutierrez-Morton, E., Rizkallah, R., Lawal, T., Kabbaj, M. H., Owutey, S. L., Tomko, R. J. Jr., and Wang Y. Cdc5-mediated Ulp2 phosphorylation controls the timing of polySUMOylation during the cell cycle. Journal of Cell Biology. 225(4): e202501146, 2026. https://rupress.org/jcb/article-pdf/225/4/e202501146/2027444/jcb_202501146.pdf
Maestro-Lopez, M., Cheng, T. C., Muntaner, J., Menendez, M., Alonso, M., Schweitzer, A., Ishizaka M., Tomko, R. J. Jr., Cuellar, J., Valpuesta, J. M., and Sakata, E. Structures of the 26S proteasome in complex with the Hsp70 co-chaperone Bag1 reveal a mechanism for direct substrate transfer. Science Advances. 12(8): eadz3026, 2026. https://www.science.org/doi/epdf/10.1126/sciadv.adz3026
Lawal, T., Chang, A. H., Carnley, L. G., Celoge, N. Y., Blount, T. A., Vied, C., Nemec, A. A., and Tomko, R. J. Jr. A microsporidial deubiquitinase blocks ubiquitin transfer from adenylated E1 to human UBE2K ubiquitin conjugating enzyme. BioRxiv (preprint). https://www.biorxiv.org/content/10.1101/2025.07.31.666823v1
Duke, L. C., Cone, A. S., Sun, L., Dittmer, D. P., Meckes, D. G. Jr., and Tomko, R. J. Jr. Tetraspanin CD9 alters cellular trafficking and endocytosis of tetraspanin CD63, affecting CD63 packaging into small extracellular vesicles. Journal of Biological Chemistry. 301(3): 108255, 2025 https://www.sciencedirect.com/science/article/pii/S0021925825001024?via%3Dihub
Selected for inclusion in "The Year In JBC - 2025", a collection of the most impactful papers of the year
Gutierrez-Morton, E., Haluska, C., Collins, L., Rizkallah, R., Tomko, R. J. Jr., and Wang, Y. The polySUMOylation axis promotes nucleolar release of Tof2 for mitotic exit. Cell Reports. 43(7): 114492, 2024 https://www.sciencedirect.com/science/article/pii/S2211124724008210?via%3Dihub
Laskin, G., Cabrera, A. R., Greene, N. P., Tomko, R. J. Jr., Vied, C., and Gordon, B. S. The mechanosensitive gene arrestin domain containing 2 regulates myotube diameter with direct implications for disuse atrophy with aging. Am. J. Physiol. Cell Physiol. 326(3): C768-C783, 2024 https://journals.physiology.org/doi/epdf/10.1152/ajpcell.00444.2023
Nemec, A.A. and Tomko, R. J. Jr. An unstructured proteasome inhibitor comes into focus. J. Biol. Chem. 299: 105145, 2023. PMID: 37562568.
Betancourt, D., Lawal, T., Tomko, R. J. Jr., Wiggle and Shake: Managing and Exploiting Conformational Dynamics during Proteasome Biogenesis. Biomolecules. 13(8), 1223, 2023 https://doi.org/10.3390/biom13081223
Warnock, J. L., Jobin, G. W., Kumar, S., and Tomko, R. J. Jr. Assembly chaperone Nas6 selectively destabilizes 26S proteasomes with defective regulatory particle – core particle interfaces. J Biol. Chem. 299: 102894, 2023. PMID: 36634850.
Editor’s Pick (Featured Article)
Huntsman, E. M., Cho, R., Kogan, H. V., McNamara-Bordewick, N. K., Tomko, R. J. Jr., and Snow, J. W. Proteasome inhibition is an effective treatment strategy for microsporidia infection in honey bees. Biomolecules. 11:1600, 202 PMID: 34827599.
Rossetti, M., Tomko, R. J. Jr., and Gordon, B. Androgen depletion alters the diurnal patterns to signals that regulate autophagy in the limb skeletal muscle. Mol. Cell. Biochem. 476: 959-69, 2021. PMID: 33128669.
Beesley, S., Kim, D., D'Alessandro, M., Jin, Y., Lee, K., Joo, H., Young, Y., Tomko, R. J. Jr., Kim, J. K., and Lee, C. Robustness of the circadian rhythm requires precise trafficking of the pacemaker PER protein. Proc. Nat. Acad. Sci. USA. 117: 28402-11, 2020. PMID: 33106420.
Higgins, R., Kabbaj, M-H., Sherwin, D., Howell, L.A., Hatcher, A., Tomko, R. J. Jr., and Wang, Y. The Cdc48 complex alleviates the cytotoxicity of misfolded proteins by regulating ubiquitin homeostasis. Cell Rep. 32: 107898, 2020. PMID: 32668237.
Howell, L. A., Peterson, A. K., and Tomko, R. J. Jr. Proteasome subunit alpha1 overexpression preferentially drives canonical proteasome biogenesis and enhances stress tolerance in yeast. Scientific Reports. 9: 12418, 2019. PMID: 31455793.
Nemec, A. A., Peterson A. K., Warnock, J. L., Reed, R. G., and Tomko, R. J. Jr. An Allosteric Interaction Network Promotes Conformation State-Dependent Eviction of the Nas6 Assembly Chaperone from Nascent 26S Proteasomes. Cell Reports. 26: 483-495, 2019. PMID: 30625330.
Eisele, M.†, Reed, R. G.†, Rudack, T.†, Schweitzer, A., Beck, F., Nagy, I., Pfeifer, G., Plitzko, J., Baumeister, W.,* Tomko, R. J. Jr.,*# and Sakata, E.* Expanded coverage of the 26S proteasome conformational landscape reveals mechanisms of peptidase gating. Cell Reports. 24: 1301-1315, 2018. PMID: 30067984.
†Equal contributions
*Corresponding author
#Lead contact
Featured in Trends in Biochemical Sciences 2018; 18: 30150-6.
Nemec, A. A.,* Howell, L. A.,* Peterson, A. K., Murray, M. A., and Tomko, R. J. Jr. Autophagic clearance of proteasomes in yeast requires the conserved sorting nexin Snx4. Journal of Biological Chemistry. 292: 21466-21480, 2017. PMID: 29109144.
*Equal contributions
Tomko, Jr., R.J.,* Taylor, D.W., Chen, Z.A., Wang, H.W., Rappsilber, J., and Hochstrasser, M. A Single Alpha Helix Drives Largescale Remodeling of the Proteasome Lid and Completion of Regulatory Particle Assembly. Cell. 163: 432-444, 2015.
*Corresponding Author
Tomko Lab Webpage
Dr. Tomko completed his graduate studies in pharmacology at the University Of Pittsburgh School Of Medicine’s Drug Discovery Institute, where he became familiar with modern drug discovery and development. His thesis work probing the regulation of a short-lived protein sparked his interest in the cell’s machinery to recycle damaged or unneeded proteins. To pursue this interest, he moved to Yale University, where he was an American Cancer Society Postdoctoral Fellow in the Department of Molecular Biophysics and Biochemistry. As a postdoctoral fellow, Robert made several seminal discoveries regarding the assembly of the proteasome, a large, multisubunit protease complex that executes most of the cell’s regulatory and quality control protein degradation. In 2015, Dr. Tomko joined the faculty of Biomedical Sciences at FSU, where his group studies the structure and function of ubiquitin-proteasome system components in normal and disease states. His laboratory works on the scale of individual proteins up to whole cells, and integrates approaches spanning biochemistry, cell biology, genetics, structural biology, proteomics, biophysics, and pharmacology. Ultimately, his group aims to utilize the information gained from these basic studies of the ubiquitin-proteasome system to identify new targets and develop pharmacological modulators with potential thereapeutic benefit.
American Cancer Society Postdoctoral Fellow, Yale University, Department of Molecular Biophysics and Biochemistry
Ph.D., Pharmacology and Chemical Biology, University of Pittsburgh School of Medicine and Drug Discovery Institute
B.S., Biochemistry and Molecular Biology, Lebanon Valley College
2024 - Present Grants reviewer, National Science Foundation
2023 - 2024 Senator, Florida State University Senate
2022 - Present Grants reviewer, US-Israel Binational Science Foundation
2022 - Present Reviewing member, Florida Agricultural & Mechanical University Institutional Biosafety Committee
2021 - Present Editorial Board Member, the Journal of Biological Chemistry
2021 - 2022 Member, Faculty Recruitment Committee, Department of Biomedical Sciences
2019 - Present Scientific advisor, Wetware development, FSU iGEM Program
2019 - Present Member, FSU-COM Physician Assistant Program Student Progress Committee
2019 Session chair, Protein Interactions, Modifications, and Regulation Section, Experimental Biology 2019 Meeting
2019 MBPP study section member, ad hoc
2019 - Present Invited grants reviewer, Israel Science Foundation
2017 - Present Member, Graduate Policy Committee, Department of Biomedical Sciences
2016 - 2019 Florida State University College of Medicine Research Advisory Committee
2016 - 2018 Dept. of Biomedical Sciences Promotion and Tenure Committee
2015 - Present Dept. of Biomedical Sciences Faculty Search Committee
2015 - Present Dept. of Biomedical Sciences Postdoctoral Career and Mentoring Committee
2026 ZRG1 MGG-K Study Section, National Institutes of Health
2025 CSF1 Study Section, National Institutes of Health
2025 Commencement Speaker, Florida State University Physician Assistant Program Class of 2025
2025 Florida State University Developing Scholar Award (awarded to exceptionally promising FSU researchers
2025 Florida State University Distinguished Teacher Award (University's highest honor for educators)
2024 Florida State University College of Medicine Outstanding Senior Faculty Researcher Award
2021 Florida State University Undergraduate Faculty Research Mentor Award
2019 Florida State University Graduate Teaching Award
2018 Florida State University College of Medicine Outstanding Junior Faculty Researcher Award
2018 Florida State University College of Medicine Outstanding Junior Faculty Educator Award
2015 First-Year Assistant Professor Award
2014 Finalist, Earl Stadtman Symposium on Molecular Biology and Biochemistry NIH, Bethesda MD
2013 Nominee, Blavatnik Award for Young Scientists
2010 - 2013 American Cancer Society Postdoctoral Fellowship
2009 - 2010 James Hudson Brown – Alexander B. Coxe Postdoctoral Fellowship in the Medical Sciences
2007 University of Pittsburgh School of Medicine Most Well-Rounded Graduate Student Award
2002 American Chemical Society Outstanding Chemistry Major Award
2002 Southeastern PA Section American Chemical Society Award
2001 American Chemical Society Award for Outstanding Performance in Organic Chemistry
2015 - Present, American Society for Biochemistry and Molecular Biology
2006 - Present, American Association for Cancer Research
2005 - Present, American Society for Pharmacology and Experimental Therapeutics
Medical program
BMS-6037 Foundations of Medicine II: Molecules to Mechanisms (Small group activity leader)
BMS-6041 Host Defense (Instructor)
Physician assistant program
PAS-5071 Clinical Pharmacology I (Founding course director; instructor)
PAS-5072 Clinical Pharmacology II (Founding course director; instructor)
Biomedical Sciences Ph.D. program
BMS-5525 Bioregulation (Instructor)
GMS-6003 Professional Development (Instructor)
Regulation of proteasome assembly and function in health and infectious disease; multidisciplinary analyses of biomedically important molecular machines; proteases and proteolysis
Dr. Tomko's full list of publications can be found HERE.
Selected publications from the Tomko lab (student authors are underlined):
Rout, S. R., Kulke, M., Droemer, M. A., Wendel, M., Cheng, T. C., Mauck, T. A., Asgari, S., Nemec, A. A., Shein, M., Sato, Y., Fukai, S., Witte, G., Tomko, R. J. Jr., Stengel, F., Zacharias, M., Schuetz, A. K., and Sakata, E. The vacuolar tauopathy-associated mutation D395G confers redox sensitivity to p97/VCP. BioRxiv (preprint). https://www.biorxiv.org/content/10.64898/2026.04.16.718620v2
Gutierrez-Morton, E., Rizkallah, R., Lawal, T., Kabbaj, M. H., Owutey, S. L., Tomko, R. J. Jr., and Wang Y. Cdc5-mediated Ulp2 phosphorylation controls the timing of polySUMOylation during the cell cycle. Journal of Cell Biology. 225(4): e202501146, 2026. https://rupress.org/jcb/article-pdf/225/4/e202501146/2027444/jcb_202501146.pdf
Maestro-Lopez, M., Cheng, T. C., Muntaner, J., Menendez, M., Alonso, M., Schweitzer, A., Ishizaka M., Tomko, R. J. Jr., Cuellar, J., Valpuesta, J. M., and Sakata, E. Structures of the 26S proteasome in complex with the Hsp70 co-chaperone Bag1 reveal a mechanism for direct substrate transfer. Science Advances. 12(8): eadz3026, 2026. https://www.science.org/doi/epdf/10.1126/sciadv.adz3026
Lawal, T., Chang, A. H., Carnley, L. G., Celoge, N. Y., Blount, T. A., Vied, C., Nemec, A. A., and Tomko, R. J. Jr. A microsporidial deubiquitinase blocks ubiquitin transfer from adenylated E1 to human UBE2K ubiquitin conjugating enzyme. BioRxiv (preprint). https://www.biorxiv.org/content/10.1101/2025.07.31.666823v1
Duke, L. C., Cone, A. S., Sun, L., Dittmer, D. P., Meckes, D. G. Jr., and Tomko, R. J. Jr. Tetraspanin CD9 alters cellular trafficking and endocytosis of tetraspanin CD63, affecting CD63 packaging into small extracellular vesicles. Journal of Biological Chemistry. 301(3): 108255, 2025 https://www.sciencedirect.com/science/article/pii/S0021925825001024?via%3Dihub
Selected for inclusion in "The Year In JBC - 2025", a collection of the most impactful papers of the year
Gutierrez-Morton, E., Haluska, C., Collins, L., Rizkallah, R., Tomko, R. J. Jr., and Wang, Y. The polySUMOylation axis promotes nucleolar release of Tof2 for mitotic exit. Cell Reports. 43(7): 114492, 2024 https://www.sciencedirect.com/science/article/pii/S2211124724008210?via%3Dihub
Laskin, G., Cabrera, A. R., Greene, N. P., Tomko, R. J. Jr., Vied, C., and Gordon, B. S. The mechanosensitive gene arrestin domain containing 2 regulates myotube diameter with direct implications for disuse atrophy with aging. Am. J. Physiol. Cell Physiol. 326(3): C768-C783, 2024 https://journals.physiology.org/doi/epdf/10.1152/ajpcell.00444.2023
Nemec, A.A. and Tomko, R. J. Jr. An unstructured proteasome inhibitor comes into focus. J. Biol. Chem. 299: 105145, 2023. PMID: 37562568.
Betancourt, D., Lawal, T., Tomko, R. J. Jr., Wiggle and Shake: Managing and Exploiting Conformational Dynamics during Proteasome Biogenesis. Biomolecules. 13(8), 1223, 2023 https://doi.org/10.3390/biom13081223
Warnock, J. L., Jobin, G. W., Kumar, S., and Tomko, R. J. Jr. Assembly chaperone Nas6 selectively destabilizes 26S proteasomes with defective regulatory particle – core particle interfaces. J Biol. Chem. 299: 102894, 2023. PMID: 36634850.
Editor’s Pick (Featured Article)
Huntsman, E. M., Cho, R., Kogan, H. V., McNamara-Bordewick, N. K., Tomko, R. J. Jr., and Snow, J. W. Proteasome inhibition is an effective treatment strategy for microsporidia infection in honey bees. Biomolecules. 11:1600, 202 PMID: 34827599.
Rossetti, M., Tomko, R. J. Jr., and Gordon, B. Androgen depletion alters the diurnal patterns to signals that regulate autophagy in the limb skeletal muscle. Mol. Cell. Biochem. 476: 959-69, 2021. PMID: 33128669.
Beesley, S., Kim, D., D'Alessandro, M., Jin, Y., Lee, K., Joo, H., Young, Y., Tomko, R. J. Jr., Kim, J. K., and Lee, C. Robustness of the circadian rhythm requires precise trafficking of the pacemaker PER protein. Proc. Nat. Acad. Sci. USA. 117: 28402-11, 2020. PMID: 33106420.
Higgins, R., Kabbaj, M-H., Sherwin, D., Howell, L.A., Hatcher, A., Tomko, R. J. Jr., and Wang, Y. The Cdc48 complex alleviates the cytotoxicity of misfolded proteins by regulating ubiquitin homeostasis. Cell Rep. 32: 107898, 2020. PMID: 32668237.
Howell, L. A., Peterson, A. K., and Tomko, R. J. Jr. Proteasome subunit alpha1 overexpression preferentially drives canonical proteasome biogenesis and enhances stress tolerance in yeast. Scientific Reports. 9: 12418, 2019. PMID: 31455793.
Nemec, A. A., Peterson A. K., Warnock, J. L., Reed, R. G., and Tomko, R. J. Jr. An Allosteric Interaction Network Promotes Conformation State-Dependent Eviction of the Nas6 Assembly Chaperone from Nascent 26S Proteasomes. Cell Reports. 26: 483-495, 2019. PMID: 30625330.
Eisele, M.†, Reed, R. G.†, Rudack, T.†, Schweitzer, A., Beck, F., Nagy, I., Pfeifer, G., Plitzko, J., Baumeister, W.,* Tomko, R. J. Jr.,*# and Sakata, E.* Expanded coverage of the 26S proteasome conformational landscape reveals mechanisms of peptidase gating. Cell Reports. 24: 1301-1315, 2018. PMID: 30067984.
†Equal contributions
*Corresponding author
#Lead contact
Featured in Trends in Biochemical Sciences 2018; 18: 30150-6.
Nemec, A. A.,* Howell, L. A.,* Peterson, A. K., Murray, M. A., and Tomko, R. J. Jr. Autophagic clearance of proteasomes in yeast requires the conserved sorting nexin Snx4. Journal of Biological Chemistry. 292: 21466-21480, 2017. PMID: 29109144.
*Equal contributions
Tomko, Jr., R.J.,* Taylor, D.W., Chen, Z.A., Wang, H.W., Rappsilber, J., and Hochstrasser, M. A Single Alpha Helix Drives Largescale Remodeling of the Proteasome Lid and Completion of Regulatory Particle Assembly. Cell. 163: 432-444, 2015.
*Corresponding Author