Section 07 · sources, pinned to the page

GHK-Cu references: every clipping traced to its paper

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.

  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.
  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. 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.
  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.
  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.
  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.
  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.
  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.
  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. 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.
  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.
  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.