Kristalle G. Cruz*Maureen Hill Eron*Sparsh MakhaikSergey Savinov, and Jeanne A. Hardy (2024) “A Non-Active-Site Inhibitor with Selectivity for Zika Virus NS2B-NS3 ProteaseACS Infectious Diseases. https://doi.org/10.1021/acsinfecdis.3c00330 *joint first authors pdf


Ranit Dutta*, Sparsh Makhaik*, Peiyao Zhao*, Kristalle G. Cruz, Kwang-Won Park, Hongxu Liu, Trisha L. Andrew, Jeanne A. Hardy, and S. Thayumanavan (2022) “A Colorimetric Cotton Swab for Viral Protease Detection” ACS Analytical Chemistry. https://doi.org/10.1021/acs.analchem.2c02033 *joint first authors pdf


Vishnu Raman, Nele Van Dessel, Christopher Hall, Victoria Whetherby, Samantha Whitney, Emily Kolewe, Shoshana Bloom, Abhinav Sharma, Jeanne Hardy, Mathieu Bollen, Aleyde Van Eynde, Neil S. Forbes (2021) “Intracellular delivery of protein drugs with an autonomously lysing bacterial system reduces tumor growth and metastases” Nature Communications. https://doi.org/10.1038/s41467-021-26367-9 pdf

Luam Araya*, Ishankumar Soni*, Jeanne A. Hardy, and Olivier Julien (2021) “Caspase-3 and caspase-9 substrate profiling using N-terminomics reveals unique sets of targets” ACS Chemical Biology. 16, 2280−2296 doi: 10.1021/acschembio.1c00456 *joint first authors pdf

Ishankumar V. Soni and Jeanne A. Hardy, (2021) “Caspase-9 Activation of Procaspase-3 but not Procaspase-6 is Based both on Local Context of Cleavage Site Motifs and on Sequence” ACS Biochemistry  60(37):2824-2835. pdf

Francesca Anson, Thayumanavan, and Jeanne A. Hardy (2021) “Exogenous Introduction of Initiator and Executioner Caspases Result in Different Apoptotic Outcomes” JACS Au https://doi.org/10.1021/jacsau. pdf hi res pdf

Francesca Anson, Bin Liu, Pintu Kanjilal, Peidong Wu, Jeanne A. Hardy and S. Thayumanavan (2021) “Evaluating Endosomal Escape of Caspase-3-Containing Nanomaterials Using Split GFP” Biomacromolecules DOI:Ex10.1021/acs.biomac.0c01767 pdf


Francesca Anson, Pintu Kanjilal, S. Thayumanavan and Jeanne A. Hardy  (2020) “Tracking Exogenous Intracellular Casp-3 Using Split GFP.” Protein Science. DOI:10.1002/pro.3992 pdf

Jorge L. Arias-Arias, Derek J. MacPherson, Maureen E. Hill, Jeanne A. Hardy, and Rodrigo Mora-Rodríguez (2020). “A fluorescence-activatable reporter of flavivirus NS2B-NS3 protease activity enables live imaging of infection in single cells and viral plaques.” Journal of Biological Chemistry. 295(8):2212-2226. pdf


Bach Pham, Scott J. Eron, Maureen E. Hill, Xin Li, Monifa A. Fahie, Jeanne A. Hardy, and Min Chen (2019). “A nanopore approach for analysis of caspase-7 activity in cell lysates.” Biophysical journal 117, no. 5 : 844-855. pdf

Eric S. Okerberg, Kevin B. Dagbay, Jennifer L. Green, Ishankumar Soni, Arwin Aban, Tyzoon K. Nomanbhoy, Sergey N. Savinov, Jeanne A. Hardy, and John W. Kozarich (2019). “Chemoproteomics using nucleotide acyl phosphates reveals an ATP binding site at the dimer interface of procaspase-6.” Biochemistry 58, no. 52 : 5320-5328. pdf

Derek J. MacPherson, Caitlyn L. Mills, Mary Jo Ondrechen, and Jeanne A. Hardy (2019). “Tri-arginine exosite patch of caspase-6 recruits substrates for hydrolysis.” Journal of Biological Chemistry. 294(1): 71-88. pdf

Maureen E. Hill, Muslum Yildiz, and Jeanne  A. Hardy (2019). “Cysteine disulfide traps reveal distinct conformational ensembles in Dengue Virus NS2B-NS3 protease.” Biochemistry. pdf


Jeanne A. Hardy (2018). “Fighting Kinase Resistance with Caspase Activators.” Cell Chemical Biology. 25(8),927-928. pdf

Maureen E. Hill, Anil Kumar, James A. Wells, Tom C. Hobman, Olivier Julien, and Jeanne A. Hardy (2018). “The unique cofactor region of Zika virus NS2B-NS3 protease facilitates cleavage of key host proteins.” ACS Chemical Biology.  pdf

Kristen L. Huber*, Banyuhay P. Serrano* and Jeanne A. Hardy (2018). “Caspase-9 CARD: Core Domain interactions require a properly-formed active site.” Biochemical Journal. 475(6): 1177-1196. pdf

Agne Tubeleviciute-Aydin, Libin Zhou, Gyanesh Sharma, Ishankumar V. Soni, Sergey N. 
Savinov, Jeanne A. Hardy and Andrea C. LeBlanc (2018) “Rare human Caspase-6-R65W and Caspase-6-G66R variants identify a novel regulatory region 
of Caspase-6 activity.”Scientific Reports. Article number: 4428. pdf

Scott J. Eron, Derek J. MacPherson, Kevin Dagbay and Jeanne A. Hardy (2018). “Multiple Mechanisms of Zinc-Mediated Inhibition for the Apoptotic Caspases-3, -6, -7, and -8.” ACS Chemical Biology pdf

Banyuhay P. Serrano and Jeanne A. Hardy (2018) “Phosphorylation by Protein Kinase A Disassembles the Caspase-9 Core Promoting Formation of Ordered Aggregates.” Cell Death and Differentiation. pdf


Celia Homyak, Ann Fernandez, Mollie Touve, Bo Zhao, Francesca Anson, Jeanne A. Hardy, Richard Vachet, Nathan Gianneschi, Jennifer Ross, Sankran Thayumanavan (2017) “Lipogels for Encapsulation of Hydrophilic Proteins and Hydrophobic Small Molecules.”Biomacromolecules.19(1):132-140. pdf

Banyuhay P. Serrano, Hannah S. Szydlo,Dominique Alfandari and Jeanne A. Hardy (2017) “Active-site Adjacent Phosphorylation at Tyr-397 by c-Abl Kinase Inactivates Caspase-9.”Journal of Biological Chemistry.  292(52):21352-21365pdf

Kishore Raghupathi,Scott J. Eron, Francesca Anson, Jeanne A. Hardy, Sankaran Thayumanavan (2017) “Utilizing Inverse Emulsion Polymerization to Generate Responsive Nanogels for Cytosolic Protein Delivery.” Molecular Pharmaceutics. 14(12):4515-4524. pdf

Kevin B. Dagbay and Jeanne A. Hardy (2017) “Multiple proteolytic events in caspase-6 self activation impact conformations of discrete structural regions” PNAS 114(30): E7977-86. pdf

Kevin B. Dagbay, Maureen Hill, Elizabeth Barrett* and Jeanne A. Hardy (2017) “Tumor-Associated Mutations in Caspase-6 Negatively Impact Catalytic Efficiency” Biochemistry 56(34):4568-77. pdf

Jeanne A. Hardy and Lynmarie Thompson (2017). “Mutual Mentoring to Promote Success and Satisfaction of Women Faculty in STEM.” Educational and Outreach Projects from the Cottrell Scholars Collaborative: Professional Development and Outreach ACS Books. Volume 1259. pdf

Kevin B. Dagbay, Nicolas Bolik-Coulon, Sergey N. Savinov and Jeanne A. Hardy (2017). “Caspase-6 Undergoes a Distinct Helix-Strand Interconversion Upon Substrate Binding” Journal of Biological Chemistry. 292: 4885-4897. pdf

Scott J. Eron, Kishore Raghupathi and Jeanne A. Hardy, (2017). “Dual Site Phosphorylation of Caspase-7 by PAK2 Blocks Apoptotic Activity by Two Distinct Mechanisms.” Structure (Cell Press). 25 (1): 27–39. pdf


Maureen E. Hill, Derek J. MacPherson, Peng Wu, Olivier Julien, James A. Wells, Jeanne A. Hardy, (2016). “Reprogramming Caspase-7 specificity to Caspase-6 by regio-specific mutations and selection” ACS Chemical Biology. 11(6):1603–1612. pdf

Yaning Wu, Jeannine Garnett, Dongbin Xu, Elsa R. Flores, Yunlong Zhao, Jeanne A. Hardy, Andreas Bergmann, (2015).Genetic characterization of two gain-of-function alleles of the effector caspase DrICE in Drosophila” Cell Death and Differentiation. 23(4):723-32. pdf


Judy Ventura, Scott J Eron, Daniella C González-Toro, Kishore Raghupathi, Fei Wang, Jeanne A. Hardy, Sankaran Thayumanavan, (2015). “Reactive Self-Assembly of Polymers and Proteins to Reversibly Silence a Killer Protein.” Biomacromolecules. 16(10):3161-71. pdf

Chang Soo Kim, Rubul Mout, Yunlong Zhao, Yi-Cheun Yeh, Rui Tang, Youngdo Jeong, Bradley Duncan, Jeanne A. Hardy and Vincent M. Rotello, (2015). “Co-Delivery of Protein and Small Molecule Therapeutics Using Nanoparticle-Stabilized Nanocapsules.” Bioconjug Chem. 26 (5): 950–954. pdf


Kevin Dagbay, Scott J. Eron, Banyuhay P. Serrano, Yunlong Zhao, Elih M. Velázquez-Delgado, Di Lin* and Jeanne A. Hardy, (2014). A multi-pronged approach for compiling a global map of allosteric regulation in the apoptotic caspases. In Avi Ashkenazi, Junying Yuan, Jim Wells, editors: Regulated Cell Death Part A, Methods in Enzymology, UK: Academic Press, 544: 215-249. pdf


Muslum Yildiz, Sumana Ghosh, Jeffrey A. Bell, Woody Sherman and Jeanne A. Hardy, (2013). “Allosteric inhibition of the NS2B-NS3 protease from dengue virus” ACS Chemical Biology. 8 (12): 2744–2752. pdf

Rui Tang, David J. Solfiell, Subinoy Rana, Chang Soo Kim, Rubul Mout, Elih M. Velázquez-Delgado, Apiwat Chompoosor, Zheng-Jiang Zhu, Chaekyu Kim, Bo Yan, Youngdo Jeong, Jeanne A. Hardy and Vincent M. Rotello, (2013). “Direct Delivery of Functional Proteins and Enzymes to the Cytosol Using Nanoparticle-Stabilized Nanocapsules.”ACS Nano. 7 (8): 6667–6673. pdf

Peng Wu, Samantha Nicholls and Jeanne A. Hardy, (2013). A tunable modular approach to fluorescent protease-activated reporters. Biophysical Journal. 104(7):1605-14. pdf

Samantha Nicholls and Jeanne A. Hardy, (2013). “Structural Basis of Fluorescence Quenching in Caspase Activatable-GFP.” Protein Science. 22(3), 247. Article featured as cover illustration. pdf


Elih M. Velazquez-Delgado and Jeanne A. Hardy, 2012. “Zinc-Mediated Allosteric Inhibition of Caspase-6.” Journal of Biological Chemistry. 287(43), 36000. pdf

Kristen L. Huber, Sumana Ghosh, and Jeanne A. Hardy, (2012). “Inhibition of caspase-9 by stabilized peptides targeting the dimerization interface.” Peptide Science. 98(5), 451–465. pdf

Kristen L. Huber and Jeanne A. Hardy, (2012). “Mechanism of zinc-mediated inhibition of caspase-9.” Protein Science. 21,1056-1065. Article featured as cover illustration. pdf

Elih M. Velazquez-Delgado and Jeanne A. Hardy, (2012). “Phosphorylation regulates assembly of the caspase-6 substrate-binding groove.” Structure. 20, 742–751. Article featured with a Preview. pdf

Abeer M. Jabaiah, Jennifer A. Getz, Witold A. Witkowski, Jeanne A. Hardy, Patrick S. Daugherty, (2012). Identification of protease exosite-interacting peptides that enhance substrate cleavage kinetics. Biological Chemistry. 393(9): 933-41. pdf

Gustavo E. López, Inara Colón-Díaz, Anthony Cruz, Sumana Ghosh, Samantha B. Nicholls, Usha Viswanathan, Jeanne A. Hardy, and Scott M. Auerbach, (2012). Modeling non-aqueous proton wires tethered to helical peptides: Biased proton transfer driven by helical dipoles. The Journal of Physical Chemistry A. 116(4):1283-8. pdf


Filip Jagodzinski, Jeanne A. Hardy and Ileana Streinu, (2011). “Using Rigidity Analysis To Probe Mutation-Induced Structural Changes in Proteins.” Proceedings, Bioinformatics and Biomedicine Workshops (BIBMW), 2011 IEEE International Conference (BIBM), Atlanta, USA, 2011; 432-437. pdf

Witold Witkowski and Jeanne A. Hardy, (2011). “A designed redox-controlled caspase.” Protein Science, 20, 1421-1431. Article featured as cover illustration. pdf

Samantha Nicholls, Jun Chu, Genevieve Abbruzzese, Kimberly D. Tremblay and Jeanne A. Hardy, (2011). “Mechanism of a dark-to-bright reporter of caspase activity.” Journal of Biological Chemistry, 286 (28), 24977-24986.  pdf

Sravanti Vaidya, and Jeanne A. Hardy, (2011). “Caspase-6 latent state stability relies on helical propensity.” Biochemistry 50(16), 3282-7. pdf

Sravanti Vaidya, Elih M. Velazquez-Delgado, Genevieve Abbruzzese and Jeanne A. Hardy, (2011). “Substrate-Induced Conformational Changes Occur in All Cleaved Forms of Caspase-6.” Journal of Molecular Biology, 406, 75-91. Article featured as cover illustration. pdf

Prior to 2010

Witold Witkowski and Jeanne A. Hardy, (2009). “L2’ loop is critical for caspase-7 active-site formation.” Protein Science, 18, 1459-1468. Article featured as cover illustration. pdf

Kristen L. Huber, Kevin D. Olsen* and Jeanne A. Hardy, (2009). “Robust Production of a Peptide Library using Methodological Synchronization.” Protein Expression and Purification, 67,139-147. pdf

Jeanne A. Hardy, (2007). “A Link Means a Lot: Disulfide Tethering in Structure-Based Drug Design” in Computational Approaches to Structure Based Drug Design, publishers Royal Society of Chemistry, p. 318-347. pdf

Jeanne A. Hardy, (2006). “Discovery and exploitation of allosteric sites for control of protein function.” Nanomedicine, 2(4), 291. pdf

From work prior to University of Massachusetts

Jeanne A. Hardy and James A. Wells, (2009). “Dissecting an Allosteric Switch in Caspase-7 using Chemical and Mutational Probes.” Journal of Biological Chemistry, 284(38), 26063-9. pdf

Jeanne A. Hardy and James A. Wells, (2004). “Searching for Allosteric Sites in Enzymes.” Current Opinion in Structural Biology, 14(6), 706-715. pdf

Jeanne A. Hardy, Joni Lam, Jack T. Nguyen, Thomas O’Brien, and James A. Wells, (2004). “Discovery of an allosteric site in caspases.” Proceedings of the National Academy of Science USA, 101(34), 12461-6. pdf

Daniel A. Erlanson, Stig K. Hansen, Jeanne A. Hardy, Joni Lam, Thomas O’Brien, April 11, (2003). “Methods for Identifying Allosteric Sites.” World Patent, WO 03/087051 A2.

Marco P. Cicero, Susan T. Hubl, Celia J. Harrison, Otis Littlefield, Jeanne A. Hardy, Hillary C.M. Nelson, (2001). “The wing in yeast heat shock transcription factor (HSF) DNA-binding domain is required for full activity.” Nucleic Acids Research, 29(8), 1715-23. pdf

Jeanne A. Hardy and Hillary C.M. Nelson, (2000). “Proline in an a-helical kink is required for folding kinetics but not for kinked structure, function or stability of heat shock transcription factor.” Protein Science, 9(11), 2128-2141. pdf

Jeanne A. Hardy, Scott T.R. Walsh, and Hillary C.M. Nelson, (2000). “Role of an a-Helical Bulge in the Yeast Heat Shock Transcription Factor.” Journal of Molecular Biology, 295(3), 393-409. pdf

Andrew J. Werner, Michael F. Hochella, George D. Guthrie, Jeanne A. Hardy, Ann E. Aust, J. Donald Rimstidt, (1995). “Asbestiform reibeckite (crocidolite) dissolution in the presence of Fe-chelators: Implications for mineral-induced disease.” American Mineralogist, 80 (11),1093. pdf

Jeanne A. Hardy and Ann E. Aust, (1995). “Iron in Asbestos Chemistry and Carcinogenicity.” Chemical Reviews, 95, 97-118. pdf

Jeanne A. Hardy and Ann E. Aust, (1995). “The Effect of Iron Binding on the Ability of Crocidolite Asbestos to Catalyze DNA Single-Strand Breaks.” Carcinogenesis, 16(2), 319-325. pdf