# GHK-Cu References: The Cited Studies Behind the Dossier

> GHK-Cu references and citations: every study underpinning this copper tripeptide dossier, with authors, journal, year, DOI, and PubMed or PMC links 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](/references) in full.

## 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] 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/
[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] 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. https://pubmed.ncbi.nlm.nih.gov/11045606/
[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. https://pubmed.ncbi.nlm.nih.gov/7514608/
[9] Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025. https://pmc.ncbi.nlm.nih.gov/articles/PMC11830136/
[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. https://pubmed.ncbi.nlm.nih.gov/25900608/
[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. https://pubmed.ncbi.nlm.nih.gov/29181666/
[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. https://pubmed.ncbi.nlm.nih.gov/15803494/
[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. https://pubmed.ncbi.nlm.nih.gov/1809108/
[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. https://pubmed.ncbi.nlm.nih.gov/37832839/

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A feature dossier on the copper tripeptide GHK-Cu, set like a magazine and sourced like a citation index — every figure attributed, every gap printed in plain sight, and nothing here clinical, prescribed, or for sale.