// 07 · REFERENCES
GHK-Cu References and Citation Index
Every study cited on this site, indexed with DOI and PubMed/PMC links — the source record behind each GHK-Cu benchmark reported here.
How to read these GHK-Cu references
These GHK-Cu references are the complete source list for the numbers reported across this site. Each inline [N] marker on the research, skin, hair, dosage and FAQ pages resolves to an entry below, with a DOI and a PubMed or PMC URL so any claim can be checked at the primary source. The list spans the foundational work — Maquart's 1988 fibroblast collagen study [1] and Pickart's tissue-remodeling and gene reviews [2][6] — through the 2024-2025 colitis, neuroprotection and anti-wrinkle literature [13][14][15]. The same citations underpin the GHK-Cu wound-healing research and the GHK-Cu safety and copper toxicity discussions elsewhere on the site.
Two cautions belong with the list. A large share of the foundational GHK-Cu mechanistic and review literature originates from a single investigator, Loren Pickart (1938-2023), and colleagues, so independent replication of the broader gene-expression and anti-aging claims is limited [2][6]. And the genome-wide signature is bioinformatic — derived largely from Connectivity Map analyses that still need protein-level in vivo validation [2]. The independent emphysema-gene-reversal [9] and the non-Pickart fibrosis [8], colitis [14] and neuroprotection [15] studies are noted here because they widen that evidentiary base.
Regulatory status of the cited compound
For context on the record below: there is no FDA- or EMA-approved therapeutic indication for GHK-Cu by any route. Topical Copper Tripeptide-1 is a legal cosmetic ingredient with a long safety record; injectable, oral and other systemic uses are unapproved and research-only [6]. One topical wound-healing trial (CuHeal, NCT07437586) has been registered, but there are no completed Phase 2/3 trials for systemic or injectable GHK-Cu. The references are the published research record — not evidence of an approved use.
- 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. ↗
- 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. ↗
- 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. ↗
- 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. ↗
- 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. ↗
- Pickart L. The human tri-peptide GHK and tissue remodeling. J Biomater Sci Polym Ed. 2008;19(8):969-988. ↗
- 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. ↗
- Zhou XM, et al. GHK Peptide Inhibits Bleomycin-Induced Pulmonary Fibrosis in Mice by Suppressing TGFbeta1/Smad-Mediated Epithelial-to-Mesenchymal Transition. Front Pharmacol. 2017;8:904. ↗
- 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. ↗
- 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. ↗
- 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. ↗
- 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. ↗
- Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025. ↗
- 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. ↗
- 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. ↗