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As a result, there is an urgent need to identify risk factors that contribute to HCC initiation and provide targetable vulnerabilities to improve patient survival. While myriad risk factors are known, elevated copper (Cu) levels in HCC patients and the incidence of hepatobiliary malignancies in Wilson disease patients, which exhibit hereditary liver Cu overload, suggests the possibility that metal accumulation promotes malignant transformation. Here we found that expression of the Cu transporter genes ATP7A, ATP7B, SLC31A1, and SLC31A2 was significantly altered in liver cancer samples and were associated with elevated Cu levels in liver cancer tissue and cells. Further analysis of genomic copy number data revealed that alterations in Cu transporter gene loci correlate with poorer survival in HCC patients. Genetic loss of the Cu importer SLC31A1 (CTR1) or pharmacologic suppression of Cu decreased the viability, clonogenic survival, and anchorage-independent growth of human HCC cell lines. Mechanistically, CTR1 knockdown or Cu chelation decreased glycolytic gene expression and downstream metabolite utilization and as a result forestalled tumor cell survival after exposure to hypoxia, which mimics oxygen deprivation elicited by transarterial embolization, a standard-of-care therapy used for patients with unresectable HCC. Taken together, these findings established an association between altered Cu homeostasis and HCC and suggest that limiting Cu bioavailability may provide a new treatment strategy for HCC by restricting the metabolic reprogramming necessary for cancer cell survival.<\/jats:p>","DOI":"10.1039\/d0mt00156b","type":"journal-article","created":{"date-parts":[[2020,11,2]],"date-time":"2020-11-02T14:24:03Z","timestamp":1604327043000},"page":"1995-2008","update-policy":"https:\/\/doi.org\/10.1039\/rsc_crossmark_policy","source":"Crossref","is-referenced-by-count":123,"title":["Altered copper homeostasis underlies sensitivity of hepatocellular carcinoma to copper chelation"],"prefix":"10.1093","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5849-6649","authenticated-orcid":false,"given":"Caroline I","family":"Davis","sequence":"first","affiliation":[{"name":"Biochemistry and Molecular Biophysics Graduate Group, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xingxing","family":"Gu","sequence":"additional","affiliation":[{"name":"Department of Cancer Biology, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ryan M","family":"Kiefer","sequence":"additional","affiliation":[{"name":"Medical Degree Program, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"},{"name":"Penn Image-Guided Interventions Laboratory, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7786-0242","authenticated-orcid":false,"given":"Martina","family":"Ralle","sequence":"additional","affiliation":[{"name":"Department of Molecular and Medical Genetics, Oregon Health and Science University, Portland, OR, 97239, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Terence P","family":"Gade","sequence":"additional","affiliation":[{"name":"Department of Cancer Biology, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"},{"name":"Penn Image-Guided Interventions Laboratory, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"},{"name":"Abramson Family Cancer Research Institute, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"},{"name":"Department of Radiology, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA 19104, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2786-2802","authenticated-orcid":false,"given":"Donita C","family":"Brady","sequence":"additional","affiliation":[{"name":"Department of Cancer Biology, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"},{"name":"Abramson Family Cancer Research Institute, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2020,11,4]]},"reference":[{"key":"2021121004362601800_cit1","first-page":"7","volume-title":"CA-Cancer J. 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