Poimenidou, Maria https://orcid.org/0000-0001-7429-9218
Bobek, Jordan M. https://orcid.org/0000-0003-3387-0937
Drouillard, Donovan https://orcid.org/0000-0002-2940-3511
Harris, Tyler https://orcid.org/0009-0006-8845-7452
Solis, Elisabeth https://orcid.org/0009-0007-8900-0662
Darnell, Chad https://orcid.org/0009-0008-6947-4999
McAllister, Donna https://orcid.org/0000-0001-7332-1292
Keyes, Robert F. https://orcid.org/0009-0009-8674-6391
Ishihara-Aoki, Mayumi https://orcid.org/0000-0002-8368-8546
Aoki, Kazuhiro https://orcid.org/0000-0003-0029-4291
Sahoo, Daisy https://orcid.org/0000-0001-5210-6497
Kalyanaraman, Balaraman https://orcid.org/0000-0002-9180-8296
Smith, Brian C. https://orcid.org/0000-0001-6330-2768
Dwinell, Michael B. https://orcid.org/0000-0001-5372-633X
Funding for this research was provided by:
U.S. Department of Health & Human Services | National Institutes of Health (CA226279)
U.S. Department of Health & Human Services | National Institutes of Health (F31HL178112)
U.S. Department of Health & Human Services | National Institutes of Health (GM080202)
U.S. Department of Health & Human Services | National Institutes of Health (CA291095)
U.S. Department of Health & Human Services | National Institutes of Health (HL58012)
U.S. Department of Health & Human Services | National Institutes of Health (GM12884)
Article History
Received: 16 October 2025
Revised: 13 April 2026
Accepted: 7 May 2026
First Online: 22 May 2026
Competing interests
: MBD is a co-founder and has ownership and financial interests in Protein Foundry, LLC, and XLock Biosciences, Inc. BK has a composition-of-matter and application-of-use patent on the manufacture and use of mito-metformin in cancer (USPTO 62/779,795). Chemical modifications of mitochondria-targeted compounds, including those described herein, are covered by a provisional US Patent, U.S. Application No. 63/285,374, to MBD, RFK, and BCS. MP was supported in part by a fellowship award from the Medical College of Wisconsin Cancer Center. DD, JMB, DS, and BCS have received financial support from the NIH. All other authors declare no potential financial conflict of interest.
: All procedures and animal experiments were reviewed and approved by the Institutional Animal Care and Use Committee (IACUC) at Medical College of Wisconsin and performed in accordance with the approved Animal Use Agreement AUA000076.