Reference · Sources

GHK-Cu References and Citations, Listed for Verification

Every quantitative claim in this dossier maps to one of the studies below. Authors, journal, year, DOI, and a PubMed or PMC link are given so each number can be checked at the source.

How to read this list

This page is the spine of the GHK-Cu dossier. Each inline marker on /mechanism, /research, /side-effects, and /faq resolves to a numbered entry here. The list mixes foundational work (Maquart 1988, Lane 1994, Simeon 2000), the major reviews (Pickart 2008, 2015, 2018), the human penetration and hair-trial data (Hostynek 2011, Lee 2016), and the freshest synthesis (Mortazavi 2025). Where a study used the free GHK tripeptide rather than the copper complex, the dossier text says so; copper coordination is required for most reported bioactivity [7]. These are the GHK-Cu references and citations in full.

  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. Simeon A, Emonard H, Hornebeck W, Maquart FX. The tripeptide-copper complex glycyl-L-histidyl-L-lysine-Cu2+ stimulates matrix metalloproteinase-2 expression by fibroblast cultures. Life Sci. 2000;67(18):2257-2265.
  8. 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.
  9. Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025.
  10. 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.
  11. Sever'yanova LA, Dolgintsev ME. Effects of Tripeptide Gly-His-Lys in Pain-Induced Aggressive-Defensive Behavior in Rats. Bull Exp Biol Med. 2017;164(2):140-143.
  12. Arul V, Kartha R, Jayakumar R. Biotinylated GHK peptide incorporated collagenous matrix: A novel biomaterial for dermal wound healing in rats. J Biomed Mater Res B Appl Biomater. 2005;73(2):383-391.
  13. Trachy RE, Fors TD, Pickart L, Uno H. The hair follicle-stimulating properties of peptide copper complexes. Results in C3H mice. Ann N Y Acad Sci. 1991;642:468-469.
  14. 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.