# GHK-Cu References: The Cited Copper-Peptide Literature

> GHK-Cu references — the full cited source list behind this field guide, with DOIs and PubMed/PMC links for every collagen, gene-expression, skin, hair, wound-healing and recent study referenced across the site.

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.

## References

[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. https://pubmed.ncbi.nlm.nih.gov/3169264/
[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. https://pmc.ncbi.nlm.nih.gov/articles/PMC6073405/
[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. https://pmc.ncbi.nlm.nih.gov/articles/PMC4508379/
[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. https://pmc.ncbi.nlm.nih.gov/articles/PMC4969472/
[5] Fan C, Chen Y, Huang Q, et al. Overview of Short Peptides for Hair Loss. Biomedicines. 2026;14(4):864. https://pubmed.ncbi.nlm.nih.gov/42072405/
[6] Pickart L. The human tri-peptide GHK and tissue remodeling. J Biomater Sci Polym Ed. 2008;19(8):969-988. https://pubmed.ncbi.nlm.nih.gov/18644225/
[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. https://pubmed.ncbi.nlm.nih.gov/29181666/
[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. https://pubmed.ncbi.nlm.nih.gov/15803494/
[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. https://pubmed.ncbi.nlm.nih.gov/1809108/
[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. https://pmc.ncbi.nlm.nih.gov/articles/PMC3016279/
[11] Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025;15:30071. https://pmc.ncbi.nlm.nih.gov/articles/PMC11830136/
[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. https://pmc.ncbi.nlm.nih.gov/articles/PMC12352503/
[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. https://pmc.ncbi.nlm.nih.gov/articles/PMC12263609/
[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. https://pubmed.ncbi.nlm.nih.gov/38599632/

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A 1970s field guide to the GHK-Cu copper-peptide literature — five decades of skin, hair, wound-repair and gene-expression findings surveyed and pinned to their sources, with no clinic at the trailhead and nothing here to sell.