§ 05 · CITATION INDEX

references

Twenty-six citations from 1973 to 2026 — the working file's bibliography, in plain type.

how this index is organized

Citations are numbered in the order they are first introduced across the site, then sortable in the table below by year, first author, and journal. Each entry links to the original source — PubMed, PMC, the journal's DOI page, or the publisher's full-text — where available. The list is not exhaustive of fifty years of GHK-Cu literature; it covers every claim cited on this site plus the most-referenced recent reviews.

citation table

The references table renders below this paragraph as a sortable, searchable list. Use the search field to filter by author, journal, or year. Click a column header to sort. The DOI / PMID / URL column links out to the original source — PubMed Central where available, the journal's article page otherwise.

primary-source classes

The index draws from four primary-source classes. Peer-reviewed journal articles — original research and review papers indexed in PubMed or comparable databases. Conference proceedings — the 2002 American Academy of Dermatology presentation is included because it has been widely cited as a topical-cosmetic trial endpoint. Editorial reviews in clinical journals — the 2024 PRIME Journal aesthetic dermatology review summarizes IRB-approved trial data. Systematic reviews — the 2026 Dermatological Reviews publication is a current overview of copper peptides in regenerative aesthetic dermatology. Vendor white papers, marketing brochures, and Reddit threads are not represented in this index — only sources that pass the peer-review or comparable editorial-control bar.

  1. Pickart L, Margolina A. Regenerative and protective actions of the GHK-Cu peptide in the light of the new gene data. International Journal of Molecular Sciences. 2018;19(7):1987. Foundational mechanism and history review.
  2. 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 Medicine. 2012;4(8):67.
  3. Maquart FX, Pickart L, Laurent M, et al. Stimulation of collagen synthesis in fibroblast cultures by the tripeptide-copper complex glycyl-L-histidyl-L-lysine-Cu2+. FEBS Letters. 1988;238(2):343-346.
  4. Leyden J, Stephens T, Finkey MB, et al. Skin care benefits of copper peptide containing facial cream. American Academy of Dermatology 60th Annual Meeting, New Orleans, LA. 2002.
  5. Abdulghani AA, Sherr S, Shirin S, et al. Effects of topical creams containing vitamin C, a copper-binding peptide cream and melatonin compared with tretinoin on the ultrastructure of normal skin. Disease Management & Clinical Outcomes. 1998;1(4):136-141.
  6. Mulder GD, Patt LM, Sanders L, et al. Enhanced healing of ulcers in patients with diabetes by topical treatment with glycyl-L-histidyl-L-lysine copper. Wound Repair and Regeneration. 1994;2(4):259-269.
  7. Zhang Y, Zhang Y, Liu B, et al. Protective effects of GHK-Cu in bleomycin-induced pulmonary fibrosis via anti-oxidative stress and anti-inflammation pathways. Life Sciences. 2020;241:117049.
  8. Badenhorst T, Svirskis D, Wu Z. Effects of GHK-Cu on MMP and TIMP Expression, Collagen and Elastin Production, and Facial Wrinkle Parameters. Journal of Aging Science. 2016;4:166.
  9. PRIME Journal editorial review. Copper tripeptide GHK-Cu and regenerative aesthetics. PRIME Journal of Aesthetic and Anti-Aging Medicine. 2024.
  10. Jiang Y, Tsoi MF, Wong VW, et al. Synergy of GHK-Cu and hyaluronic acid on collagen IV upregulation via fibroblast and ex-vivo skin tests. Journal of Cosmetic Dermatology. 2023;22(10):2719-2727.
  11. Park JR, Lee H, Kim SI, Yang SR. The tri-peptide GHK-Cu complex ameliorates lipopolysaccharide-induced acute lung injury in mice. Oncotarget. 2016;7(36):58405-58417.
  12. Park JR, Kim SY, Yang SR, et al. Glycyl-L-histidyl-L-lysine-Cu2+ attenuates cigarette smoke-induced pulmonary emphysema and inflammation by reducing oxidative stress pathway. Frontiers in Molecular Biosciences. 2022;9:851849.
  13. 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. Aging Pathobiology and Therapeutics. 2023;5(4):137-143.
  14. 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.
  15. Bobyntsev II, Chernysheva OI, Dolgintsev ME, Smakhtin MY, Belykh AE. Anxiolytic Effects of Tripeptide Gly-His-Lys in Rats. Bulletin of Experimental Biology and Medicine. 2015;158(6):726-728.
  16. Rosenfeld M, Nickel K, Ladiges W. GHK peptide prevents sleep-deprived learning impairment in aging mice. Aging Pathobiology and Therapeutics. 2023;5(1):41-44.
  17. Arul V, Gopinath D, Gomathi K, Jayasankar R. Glycyl-histidyl-lysine binds copper(II) ions and accelerates wound healing in normal and diabetic rats. Journal of Biomedical Materials Research Part B. 2005;73B(2):383-391.
  18. Ahmed MR, Basha SH, Gopinath D, et al. Combining nerve guides, human peripheral nerve grafts, and chondroitinase to enhance nerve regeneration. Journal of the Peripheral Nervous System. 2005;10(1):17-23.
  19. Kanazawa T, Gotoh M, Ohta K, et al. Tripeptide-copper complex GHK-Cu (II) transiently improved healing outcome in a rat model of ACL reconstruction. Journal of Orthopaedic Research. 2015;33(8):1166-1175.
  20. Miller DM, DeSilva D, Pickart L, Aust SD. Protection by glycyl-L-histidyl-L-lysine against ferritin-stimulated lipid peroxidation. Advances in Experimental Medicine and Biology. 1990;264:79-84.
  21. Hsieh HJ, Hsu RJ, Lo YH, et al. Expression and Purification of Recombinant GHK Tripeptides Are Able to Protect against Acute Cardiotoxicity from Exposure to Waterborne-Copper in Zebrafish. Biomolecules. 2020;10(9):1202.
  22. He Q, Mazzola J, Ladiges W. The naturally occurring peptide GHK reverses age-related fibrosis by modulating myofibroblast function. Aging Pathobiology and Therapeutics. 2024;6(4):158.
  23. Matalka LE, Ford A, Unlap MT. Skin Anti-Aging and Biological Actions of the Plasma Tripeptide GHK-Cu. Journal of Biotechnology & Biomaterials. 2012;2(5):144.
  24. Hong Y, Downey T, Eu KW, Koh PK, Cheah PY. Effect of copper-binding peptide GHK on inhibiting metastasis-related gene expression. Clinical & Experimental Metastasis. 2010;27(2):83-90.
  25. Kang YA, Choi HR, Na JI, et al. Stem cell recovering effect of copper-free GHK in skin. Journal of Peptide Science. 2012;18(11):685-690.
  26. Copper Peptides in Regenerative Aesthetic Dermatology. Dermatological Reviews (Wiley). 2026.
  27. 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. Journal of Chromatography B: Biomedical Sciences and Applications. 1997;692(1):37-42.
  28. Hong M, Gui Y, Xu J, Zhao X, Jiang C, Zhao J, Xin X, Liu D, Tang X, Tang R, An F. Palmitoyl copper peptide and acetyl tyrosine complex enhances melanin production in both A375 and B16 cell lines. Biochemical and Biophysical Research Communications. 2025;doi:10.1016/j.bbrc.2024.151060.
  29. Miller TR, Wagner JD, Baack BR, Eisbach KJ. Effects of topical copper tripeptide complex on CO2 laser-resurfaced skin. Archives of Facial Plastic Surgery. 2006;8(4):252-259.
  30. Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2025;15:30071.
  31. Simeon A, Emonard H, Hornebeck W, Maquart FX. The tripeptide-copper complex glycyl-L-histidyl-L-lysine-Cu2+ stimulates matrix metalloproteinase-2 expression by fibroblast cultures. Life Sciences. 2000;67(18):2257-2265.
  32. 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. Oxidative Medicine and Cellular Longevity. 2012;2012:324832.
  33. Pickart L, Vasquez-Soltero JM, Margolina A. GHK Peptide as a Natural Modulator of Multiple Cellular Pathways in Skin Regeneration. BioMed Research International. 2015;2015:648108.
  34. Pickart L. The human tri-peptide GHK and tissue remodeling. Journal of Biomaterials Science, Polymer Edition. 2008;19(8):969-988.