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The full citation list, paper-clipped in order of reference across the site. Click through to the primary source.

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How this list is organized

Numbered in order of first appearance across the site. DOIs link to the primary publication where available. Where a citation is a review or summary, it is marked as such in the citation line. Where a result is from a press release describing presented data, that is also marked. The list is not exhaustive of the GHK-Cu literature; it covers every reference used elsewhere on this site.

Primary citations

See the references_index for the full numbered list with DOI and PMC links.

  1. PubChem. Copper Tripeptide-1 (Glycyl-L-histidyl-L-lysine copper(II) complex). National Library of Medicine. ↗
  2. Zhang Q, Yan L, Lu J, Zhou X. Glycyl-L-histidyl-L-lysine-Cu2+ attenuates cigarette smoke-induced pulmonary emphysema and inflammation by reducing oxidative stress pathway. Frontiers in Molecular Biosciences. 2022. ↗
  3. He Q, Mazzola J, Ladiges W. The naturally occurring peptide GHK reverses age-related fibrosis by modulating myofibroblast function. Aging Pathobiology and Therapeutics. 2024. ↗
  4. Mao S, Huang J, Li J, et al. Exploring the beneficial effects of GHK-Cu on an experimental model of colitis and the underlying mechanisms. Frontiers in Pharmacology. 2025. ↗
  5. Pickart L, Margolina A. Regenerative and Protective Actions of the GHK-Cu Peptide in the Light of the New Gene Data. International Journal of Molecular Sciences. 2018;19(7):1987. ↗
  6. Xu Y, Xu C, He L, et al. Copper peptide-incorporated 3D-printed silk-based scaffolds promote vascularized bone regeneration. Chemical Engineering Journal. 2022. ↗
  7. Chen H, Yang P, Xue P, et al. Food-Derived Tripeptide-Copper Self-Healing Hydrogel for Infected Wound Healing. Biomaterials Research. 2025. ↗
  8. Jiang Y, Wu Y, Wang K, et al. Synergy of GHK-Cu and hyaluronic acid on collagen IV upregulation via fibroblast and ex-vivo skin tests. Journal of Cosmetic Dermatology. 2023. ↗
  9. Min JH, Sarlus H, Harris RA. Glycyl-l-histidyl-l-lysine prevents copper- and zinc-induced protein aggregation and central nervous system cell death in vitro. Metallomics. 2024. ↗
  10. Yaman Agaoglu FB, et al. Interaction of GHK Tripeptide with Receptors Targeted in Some Cancer Studies: A Theoretical Approach with Molecular Docking. Hittite Journal of Science and Engineering. 2023. ↗
  11. Pickart L, Vasquez-Soltero JM, Margolina A. Modulation of Gene Expression in Human Breast Cancer MCF7 and Prostate Cancer PC3 Cells by the Human Copper-Binding Peptide GHK-Cu. OBM Genetics. 2021. ↗
  12. Pickart L, Vasquez-Soltero JM, Margolina A. The Effect of the Human Peptide GHK on Gene Expression Relevant to Nervous System Function and Cognitive Decline. Brain Sciences. 2017;7(2):20. ↗
  13. Bobyntsev II, Chernysheva OI, Dolgintsev ME, Smakhtin MY, Belykh AE. Influence of Gly-His-Lys peptide on anxiety-related behavior in rats. Bulletin of Experimental Biology and Medicine. 2015. ↗
  14. Campbell JD, McDonough JE, Zeskind JE, et al. A gene expression signature of emphysema-related lung destruction and its reversal by the tripeptide GHK. Genome Medicine. 2012;4(8):67. ↗
  15. Arul V, Kartha R, Jayakumar R. Glycyl histidyl lysine-Cu(II) chelate-loaded collagen films for accelerated wound healing in diabetic rats. Journal of Biomedical Materials Research Part B. 2007. ↗
  16. Stefaniak E, Plonka D, Drew SC, et al. Intermediate Cu(II)-Thiolate Species in the Reduction of Cu(II)GHK by Glutathione: A Handy Chelate for Biological Cu(II) Reduction. Inorganic Chemistry. 2021. ↗
  17. Carey W, et al. Epigenetic mechanisms activated by GHK-Cu increase skin collagen density in clinical trial. EurekAlert press release citing presented clinical data, 2024. ↗
  18. Pickart L, Vasquez-Soltero JM, Margolina A. GHK Peptide as a Natural Modulator of Multiple Cellular Pathways in Skin Regeneration. BioMed Research International. 2015. ↗
  19. U.S. Food and Drug Administration. List of Bulk Drug Substances for Use in Compounding Under Section 503A of the Federal Food, Drug, and Cosmetic Act — Category 2 and historical category status. Cited for the April 2026 removal of injectable GHK-Cu. ↗
  20. Adnan SB, et al. Exploring the Role of Tripeptides in Wound Healing and Skin Regeneration: A Comprehensive Review. International Journal of Medical Sciences. 2025. ↗
  21. Pickart L, Margolina A. Skin Regenerative and Anti-Cancer Actions of Copper Peptides. Cosmetics (MDPI). 2018;5(2):29. ↗
  22. Mohammed YH, Yamada M, Lin LL, et al. Microneedle-Mediated Delivery of Copper Peptide Through Skin. Pharmaceutical Research. 2015. ↗
  23. Endo T, Miyagi M, Ujiie A. Simultaneous determination of glycyl-L-histidyl-L-lysine and its metabolite, L-histidyl-L-lysine, in rat plasma by high-performance liquid chromatography with post-column derivatization. Journal of Chromatography B: Biomedical Sciences and Applications. 1997. ↗
  24. Miller TR, Wagner JD, Baack BR, Eisbach KJ. Effects of topical copper tripeptide complex on CO2 laser-resurfaced skin. Archives of Facial Plastic Surgery. 2006;8(4):252-259. ↗
  25. Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025;15:30071. ↗
  26. Hong M, Gui Y, Xu J, et al. Palmitoyl copper peptide and acetyl tyrosine complex enhances melanin production in both A375 and B16 cell lines. Biochemical and Biophysical Research Communications. 2025. ↗
  27. Pickart L. The human tri-peptide GHK and tissue remodeling. Journal of Biomaterials Science, Polymer Edition. 2008;19(8):969-988. ↗