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

> GHK-Cu references — the full cited literature behind this notebook, with PubMed and PMC links and DOIs for every collagen, gene-expression, hair, wound-healing and safety study summarized on the site.

The full citation list behind the notebook — each study with its journal, year, PMID and DOI, so any claim on this site can be checked against the source.

## How to read these GHK-Cu references

These GHK-Cu references are the complete source list for every quantitative claim in the notebook, numbered to match the inline markers on each page. Where a study is available open-access on PubMed Central, the link goes there; otherwise it points to the PubMed record. Each entry carries its DOI. The collagen dose-response [1], the gene-modulation data [2], the skin-regeneration review [3], the controlled hair-count trial [4], and the copper-handling chemistry [9] are the load-bearing clippings; the rest fill in mechanism, delivery, wound-healing models and the recent neuroprotective and colitis work. Citations are listed in full beneath this note.

## 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] Zhou XM, et al. GHK Peptide Inhibits Bleomycin-Induced Pulmonary Fibrosis in Mice by Suppressing TGFβ1/Smad-Mediated Epithelial-to-Mesenchymal Transition. Front Pharmacol. 2017;8:904. https://pmc.ncbi.nlm.nih.gov/articles/PMC5733019/
[8] Campbell JD, McDonough JE, Zeskind JE, et al. A gene expression signature of emphysema-related lung destruction and its reversal by the tripeptide GHK. Genome Med. 2012;4(8):67. https://pmc.ncbi.nlm.nih.gov/articles/PMC4064320/
[9] Pickart L, Vasquez-Soltero JM, Margolina A. The human tripeptide GHK-Cu in prevention of oxidative stress and degenerative conditions of aging: implications for cognitive health. Oxid Med Cell Longev. 2012;2012:324832. https://pmc.ncbi.nlm.nih.gov/articles/PMC3359723/
[10] 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;172:159-174. https://pubmed.ncbi.nlm.nih.gov/37832839/
[11] Jose S, Hughbanks ML, Binder BYK, Ingavle GC, Leach JK. Enhanced trophic factor secretion by mesenchymal stem/stromal cells with Glycine-Histidine-Lysine (GHK)-modified alginate hydrogels. Acta Biomater. 2014;10(5):1955-1964. https://pmc.ncbi.nlm.nih.gov/articles/PMC3976757/
[12] Molavi AM, Sadeghi-Avalshahr A, Nokhasteh S, Naderi-Meshkin H. Enhanced biological properties of collagen/chitosan-coated poly(epsilon-caprolactone) scaffold by surface modification with GHK-Cu peptide and 58S bioglass. Prog Biomater. 2020;9(1-2):71-82. https://pmc.ncbi.nlm.nih.gov/articles/PMC7289912/
[13] 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/
[14] 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/
[15] 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(5):mfae019. https://pubmed.ncbi.nlm.nih.gov/38599632/
[16] Pickart L, Thaler MM. Tripeptide in human serum which prolongs survival of normal liver cells and stimulates growth in neoplastic liver — rapid metabolism of GHK to the dipeptide HK after intravenous dosing (rat HPLC pharmacokinetic study). PMID 9187381. https://pubmed.ncbi.nlm.nih.gov/9187381/

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A hand-bound studio notebook of the GHK-Cu copper-peptide literature — every collagen, gene, and hair-count clipping pasted in and sourced to its paper, the honest gaps left in the margin, with nothing here prescribed, dispensed, or for sale.