PLATE 07 · PROVENANCE
GHK-Cu references: every specimen's source
The complete cited record behind this herbarium — each study with its authors, journal, year, DOI, and PubMed or PMC link, so every pressed specimen can be traced to its plate.
How to read this reference list
These are the GHK-Cu references cited across this site, numbered to match the inline markers on every page. The record is dominated by a five-decade body of skin, wound, and gene-expression work, anchored by the foundational Maquart 1988 fibroblast collagen study [1] and the Pickart review series [2][3][6], with the human hair-count RCT [4], the human skin-penetration study [5], and the 2023-2025 neuroprotection and delivery literature [8][9][10][13] added as the most recent specimens.
Two provenance notes are catalogued here rather than hidden. A large share of the foundational 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]. And two of the cognitive-neuroprotection studies are bioRxiv preprints [8][13] — early evidence, marked as such. Each citation below links to its primary source for verification.
- 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 peptide GHK as a possible anti-aging and anti-cancer drug and its potential to retard the aging process and prevent disease (mechanism and stability context; complex stability constant log K ~16.4). Reviewed in BioMed Res Int. 2015;2015:648108. ↗
- 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. doi:10.1101/2023.11.16.567423. ↗
- Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025. ↗
- 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. ↗
- 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. ↗
- 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. ↗
- 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. doi:10.1101/2023.11.20.567908. ↗
- 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. ↗
- 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. ↗