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GHK-Cu References: Every Source in the Field Guide

Each number cited across this site resolves here, with the journal, year, DOI or PMID, and a direct link to the source.

How to read this list

The GHK-Cu references below are the complete source shelf for this field guide. Every inline [N] marker across the site points to one of these entries, and every quantitative claim — each dose, percentage, gene count, hair count and stability constant — traces to a numbered source here. The list mixes foundational work (the 1988 collagen dose-response, the 1991 mouse follicle study) with recent 2024-2026 studies on aged fibroblasts, colitis, neuroprotection and short-peptide hair-loss reviews. Where a paper is open-access, the link resolves to PubMed Central; otherwise it resolves to the PubMed record. Full citation details, including DOIs, are listed for each.

  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. Fan C, Chen Y, Huang Q, et al. Overview of Short Peptides for Hair Loss. Biomedicines. 2026;14(4):864.
  6. Pickart L. The human tri-peptide GHK and tissue remodeling. J Biomater Sci Polym Ed. 2008;19(8):969-988.
  7. 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.
  8. 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.
  9. 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.
  10. 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.
  11. Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025;15:30071.
  12. He Q, Mazzola J, Ladiges W. The naturally occurring peptide GHK reverses age-related fibrosis by modulating myofibroblast function. Aging Pathobiol Ther. 2024;6(4):158.
  13. Mao S, Huang J, Li J, Sun F, Zhang Q, Cheng Q, Zeng W, Lei D, Wang S, Yao J. Exploring the beneficial effects of GHK-Cu on an experimental model of colitis and the underlying mechanisms. Front Pharmacol. 2025;16:1551843.
  14. 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;16(4):mfae019.