# GHK-Cu References: The Cited Studies Behind This Research Brief

> GHK-Cu references and citations: the full list of peer-reviewed studies, reviews, and preprints behind this copper-tripeptide research brief, with DOIs and PubMed or PMC links.

Every quantitative claim in this brief maps to a numbered entry below — peer-reviewed studies, canonical reviews, and clearly marked preprints, each with a DOI and a PubMed or PMC link.

## How to read this list

These are the GHK-Cu references and citations that ground this brief. Inline markers such as [1] or [8] throughout the site resolve to the numbered entries on this page. The list mixes foundational primary studies (collagen synthesis, skin penetration, hair-count RCT), canonical reviews (tissue remodeling, skin regeneration, the 2018 gene-data paper), and clearly labeled bioRxiv preprints for the intranasal cognition work — the preprint status is stated wherever those results are cited [8][9].

Where a study used the free GHK tripeptide rather than the GHK-Cu copper chelate, the inline text says so, because copper coordination is required for most documented tissue-repair activity [6]. The full bibliographic detail, with DOIs and PubMed or PMC URLs, is rendered in the reference list.

## 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] Pickart L, Vasquez-Soltero JM, Margolina A. The Effect of the Human Peptide GHK on Gene Expression Relevant to Nervous System Function and Cognitive Decline. Brain Sci. 2017;7(2):20. https://pmc.ncbi.nlm.nih.gov/articles/PMC5332963/
[8] Tucker M, Keely A, Park JY, Rosenfeld M, Wezeman J, Mangalindan R, Ratner D, Ladiges W. Intranasal GHK peptide enhances resilience to cognitive decline in aging mice. bioRxiv (preprint). 2023. https://pmc.ncbi.nlm.nih.gov/articles/PMC10680828/
[9] Tucker M, Liao GY, Park JY, Rosenfeld M, Wezeman J, Mangalindan R, Ratner D, Darvas M, Ladiges W. Behavioral and neuropathological features of Alzheimer's disease are attenuated in 5xFAD mice treated with intranasal GHK peptide. bioRxiv (preprint). 2023. https://pmc.ncbi.nlm.nih.gov/articles/PMC10690187/
[10] Tosto R, Vecchio G, Bellia F. New Biotinylated GHK and Related Copper(II) Complex: Antioxidant and Antiglycant Properties In Vitro against Neurodegenerative Disorders. Molecules. 2023;28(18):6724. https://pubmed.ncbi.nlm.nih.gov/37764500/
[11] 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/
[12] 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;mfae019. https://pubmed.ncbi.nlm.nih.gov/38599632/
[14] 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. https://pubmed.ncbi.nlm.nih.gov/9187381/
[15] 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/
[16] 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/

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An advisory-grade brief on the GHK-Cu copper-tripeptide literature — the preclinical record weighed in full and the human-data gaps marked as gaps, with no clinic behind the memorandum and nothing here for sale.