Gordon Rule

Gordon Rule

Professor, Biological Sciences

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Born in Ontario, Canada, Gordon Rule earned his undergraduate degree from University of Waterloo, his master's degree from Penn State, his Ph.D. from CMU, and a post-doc in chemistry from Stanford. Rule previously served on the faculty of University of Virginia School of Medicine. He has been on the faculty of the Department of Biological Sciences at CMU since 1996. 


Post. D., Physical Chemistry, Stanford University

Ph.D., Biological Sciences, Carnegie Mellon University

M.S., Biophysics, Penn. State University, State College, PA

B.Sc., Physics, Biology, Chemistry, University of Waterloo, Waterloo, Ontario, Canada

Area Of Expertise



biophysical chemistry

Research Description

Rule began an entirely new direction of research this year on a class of enzymes called Thymidylate kinases.   Due to substantial differences between the human enzyme and the isoform found in pathogens, these enzymes are an attractive drug target. The activity of the enzyme is also required for rapid cell division, i.e. in tumors; inhibitors of the human enzyme potentiate the activity of other chemotherapeutic agents. Lastly, the human enzyme is required for the activation of AZT, an important therapeutic for the treatment of HIV.  Rule is currently searching for inhibitors of the enzyme in Plasmodium, the pathogen responsible for malaria.


Fucci, I.J., Sinha, K., Rule, G.S. (2020) Stabilization of active site dynamics leads to increased activity with 3′-azido-3′-deoxythymidine monophosphate for F105Y mutant human thymidylate kinase. ACS Omega, 5, 5, 2355-2367. http://doi.org/10.1021/acsomega.9b03766

Chen, M., Fucci, I.J., Sinha, K., Rule, G.S. (2020) dGMP Binding to Thymidylate Kinase from Plasmodium falciparum Shows Half-Site Binding and Induces Protein Dynamics at the Dimer Interface. Biochemistry, 59, 5, 694-703. https://doi.org/10.1021/acs.biochem.9b00898

Chen, M., Sinha, K., Rule, G.S., Ly, D.H. (2018) Interaction of a-Thymidine Inhibitors with Thymidylate Kinase from Plasmodium falciparum. Biochemistry, 57, 2868-2875.

Sinha, K., Rule, G.S. (2017) The Structure of Thymidylate Kinase from Candida albicans Reveals a Unique Structural Element Biochemistry, 56, 4360-4370.

Sinha, K, Jen-Jacobson, L., Rule, G.S. (2013) Divide and conquer is always best: sensitivity of methyl correlation experiments. J. Biomol NMR. 56, 331-335.

Senutovitch, N., Stanfield, R.L., Bhattacharyya, S., Rule, G.S., Wilson, I.A., Armitage, B.A., Waggoner, A.S., Berget, P.B. (2012) A Variable Light Domain Fluorogen Activating Protein Dimerizes to Activate Dimethylindole Red. Biochemistry, 51, 2471-2485.

Sinha, K., Jen-Jacobson, L., Rule, G (2011) Labeling of Threonine Methyl Groups for NMR Studies of Protein-Nucleic Acid Complexes. Biochemistry, 50, 10189-10191.

Rule, G.S, Kessler, A., Bajzek, D (2010) Molecular Visualization in STEM Education: Leveraging Jmol in an Integrated Assessment Platform. In Proceedings of World Conference on E-Learning in Corporate, Government, Healthcare, and Higher Education 2010. Chesapeake, VA: AACE.

Working Papers

Prediction of Methyl NMR chemical shifts in protein-nucleotide complexes.

University Service

Undergraduate Curriculum Committee (Biological Sciences)

            Search committee for chemistry faculty at CMU-Q.

            Search committee for assistant department head, biological sciences.

Faculty Review Committee

Professional Activities

Review of biochemistry and immunology texts (Garland, Wiley) , manuscripts in journals.

Courses Taught

03-232 Biochemistry

03-390 Molecular and Cellular Immunology

03-131 Genes, Drugs, and Diseases

03/-121/03-151 Modern Biology/Honors Modern Biology (co-taught)