Plate VII · The colophon

GHK-Cu References: Every Plate, Sourced

The complete citation register for this GHK-Cu digest — each study weighed on the pages above, here with its journal, year, DOI and PubMed or PMC link.

How this register is organized

These GHK-Cu references are the studies cited by number throughout this digest. Each was selected from the audited copper-tripeptide corpus and verified to a working DOI or PubMed/PMC record. Foundational mechanism and review work leads, followed by the skin-penetration and delivery studies, the hair-growth studies, the neuroprotective and behavioral work, and the recent 2024-2026 syntheses.

Where a finding rests on a close analog rather than GHK-Cu itself — the AHK-Cu dermal-papilla work [8] — the entry is flagged as analog context in the page where it appears, never as direct GHK-Cu efficacy. Where a result is from a combination formulation — the 5-ALA + GHK hair trial [4] — that is stated at every citation. The aim is a register you can check, not a list you have to trust.

  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.
  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.
  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.
  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. Note: tested a 5-ALA + GHK combination formulation, not pure GHK-Cu.
  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.
  6. Pickart L. The human tri-peptide GHK and tissue remodeling. J Biomater Sci Polym Ed. 2008;19(8):969-988.
  7. 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.
  8. Pyo HK, Yoo HG, Won CH, Lee SH, Kang YJ, Eun HC, Cho KH, Kim KH. The effect of tripeptide-copper complex on human hair growth in vitro. Arch Pharm Res. 2007;30(7):834-839. Note: tested AHK-Cu, an analog of GHK-Cu; cited as analog context only.
  9. 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.
  10. Dymek M, Olechowska K, Hac-Wydro K, Sikora E. Liposomes as Carriers of GHK-Cu Tripeptide for Cosmetic Application. Pharmaceutics. 2023;15(10):2485.
  11. Pickart L, Vasquez-Soltero JM, Margolina A. GHK Peptide as a Natural Modulator of Multiple Cellular Pathways in Skin Regeneration (copper coordination, stability constant log K ~16.4, and formulation chemistry). Biomed Res Int. 2015;2015:648108.
  12. 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.
  13. Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025;15:30071.
  14. Liu T, Liu Y, Zhao X, Zhang L, Wang W, Bai D, Liao Y, Wang Z, Wang M, Zhang J. Thermodynamically stable ionic liquid microemulsions pioneer pathways for topical delivery and peptide application. Bioact Mater. 2024;33:1-15.
  15. 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 HPLC with post-column derivatization. J Chromatogr B Biomed Sci Appl. 1997;692(1):37-42.