The audited record / Sources

GHK-Cu references and the cited copper-peptide literature

Every study referenced across this site, with DOI and PubMed links — the verifiable record behind each finding.

How to read these GHK-Cu references

These GHK-Cu references are the peer-reviewed sources cited throughout this site, listed in the order they are numbered in the text. Each entry carries a DOI or PubMed identifier so any claim can be traced to its origin. The list spans the foundational work — Maquart's 1988 collagen dose-response and Pickart's tissue-remodeling and gene-expression reviews — through recent 2023-2025 studies on biomaterial delivery, infected-wound hydrogels, and topical anti-wrinkle formulation.

The record is deliberately bounded. Much of the foundational mechanistic and review literature originates from a single investigator and colleagues, the controlled human trials are small, and no validated human pharmacokinetic study exists. Reading the sources directly is the best way to see both what each study measured and where it stops.

  1. Maquart FX, Pickart L, Laurent M, Gillery P, Monboisse JC, Borel JP. Stimulation of collagen synthesis in fibroblast cultures by the tripeptide-copper complex glycyl-L-histidyl-L-lysine-Cu2+. FEBS Lett. 1988;238(2):343-346.
  2. Pickart L, Margolina A. Regenerative and Protective Actions of the GHK-Cu Peptide in the Light of the New Gene Data. Int J Mol Sci. 2018;19(7):1987.
  3. Pickart L, Vasquez-Soltero JM, Margolina A. GHK Peptide as a Natural Modulator of Multiple Cellular Pathways in Skin Regeneration. Biomed Res Int. 2015;2015:648108.
  4. Lee WJ, Sim HB, Jang YH, Lee SJ, Kim DW, Yim SH. Efficacy of a Complex of 5-Aminolevulinic Acid and Glycyl-Histidyl-Lysine Peptide on Hair Growth. Ann Dermatol. 2016;28(4):438-443.
  5. Hostynek JJ, Dreher F, Maibach HI. Human skin penetration of a copper tripeptide in vitro as a function of skin layer. Inflamm Res. 2011;60(1):79-86.
  6. Pickart L. The human tri-peptide GHK and tissue remodeling. J Biomater Sci Polym Ed. 2008;19(8):969-988.
  7. Lee S, Lee SM, Lee SH, Choi WK, Park SJ, Kim DY, et al. In situ photo-crosslinkable hyaluronic acid-based hydrogel embedded with GHK peptide nanofibers for bioactive wound healing. Acta Biomater. 2023.
  8. Jose S, Hughbanks ML, Binder BYK, Ingavle GC, Leach JK. Enhanced trophic factor secretion by mesenchymal stem/stromal cells with Glycine-Histidine-Lysine (GHK)-modified alginate hydrogels. Acta Biomater. 2014;10(5):1955-1964.
  9. Lane TF, Iruela-Arispe ML, Johnson RS, Sage EH. SPARC is a source of copper-binding peptides that stimulate angiogenesis. J Cell Biol. 1994;125(4):929-943.
  10. Molavi AM, Sadeghi-Avalshahr A, Nokhasteh S, Naderi-Meshkin H. Enhanced biological properties of collagen/chitosan-coated poly(epsilon-caprolactone) scaffold by surface modification with GHK-Cu peptide and 58S bioglass. Prog Biomater. 2020;9(1-2):71-82.
  11. Dymek M, Olechowska K, Hac-Wydro K, Sikora E. Liposomes as Carriers of GHK-Cu Tripeptide for Cosmetic Application. Pharmaceutics. 2023;15(10):2485.
  12. 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. J Chromatogr B Biomed Sci Appl. 1997;692(1):37-42.
  13. Bobyntsev II, Chernysheva OI, Dolgintsev ME, Smakhtin MY, Belykh AE. Anxiolytic effects of Gly-His-Lys peptide and its analogs. Bull Exp Biol Med. 2015;158(6):726-728.
  14. Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025.
  15. Chen H, Yang P, Xue P, Li S, Dan X, Li Y, Lei L, Fan X. Food-Derived Tripeptide-Copper Self-Healing Hydrogel for Infected Wound Healing. Biomater Res. 2025;29:0139.