FRAME 05 / REFERENCES

The file, indexed.

Twenty-three primary references spanning 1988 to 2025. Each entry carries a DOI and a link to PubMed, PMC, or the publisher. Use the sort and filter controls to reorder by year, author, or journal.

How to read this index

Each numbered chip in the body of the site corresponds to one entry below. The list is rendered as a sortable, filterable table at desktop widths; on mobile it collapses to a chronologically ordered list. The DOI link, where present, points to the publisher's canonical landing page; the URL link points to PubMed, PMC, the journal article, or the publisher's open-access PDF, whichever is freely accessible.

The list spans 37 years of GHK-Cu research, from the 1988 Maquart fibroblast-collagen paper through the 2025 liposomal-delivery and tripeptide-review papers. Reading it in chronological order is itself instructive — the early frames are biochemical, the middle frames move into animal models and the first human trial, the later frames are dominated by gene-expression work and by formulation engineering.

Primary sources

The full numbered reference list lives in the table below. Where a DOI was not assigned (a few of the older or industry-published cosmetic-dermatology entries), the URL link is the authoritative source. PubMed and PMC IDs are included where available so that readers can pull the full record into their own reference managers.

  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 Letters. 1988.
  2. Wegrowski Y, Maquart FX, Borel JP. Expression of glycosaminoglycans and small proteoglycans in wounds: modulation by the tripeptide-copper complex glycyl-L-histidyl-L-lysine-Cu2+. Life Sciences. 1992.
  3. Maquart FX, Bellon G, Chaqour B, Wegrowski J, Patt LM, Trachy RE, Monboisse JC, Chastang F, Birembaut P, Gillery P, Borel JP. In vivo stimulation of connective tissue accumulation by the tripeptide-copper complex glycyl-L-histidyl-L-lysine-Cu2+ in rat experimental wounds. Journal of Clinical Investigation. 1993;92(5):2368-2376.
  4. Canapp SO Jr, Farese JP, Schultz GS, Gowda S, Ishak AM, Swaim SF, Vangilder J, Lee-Ambrose L, Martin FG. The effect of topical tripeptide-copper complex on healing of ischemic open wounds. Veterinary Surgery. 2003;32(6):515-523.
  5. Mulder GD, Patt LM, Sanders L, Rosenstock J, Altman MI, Hanley ME, Duncan GW. 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.
  6. Uno H, Kurata S. Chemical agents and peptides affect hair growth. Journal of Investigative Dermatology. 1993;101(1 Suppl):143S-147S.
  7. 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. Archives of Pharmacal Research. 2007;30(7):834-839.
  8. Pickart L, Vasquez-Soltero JM, Margolina A. GHK, the human skin remodeling peptide, induces anti-cancer expression of numerous caspase, growth regulatory, and DNA repair genes. Journal of Aging Research and Clinical Practice. 2014;3(1):45-52.
  9. Pickart L, Vasquez-Soltero JM, Margolina A. GHK and DNA: resetting the human genome to health. BioMed Research International. 2014;2014:151479.
  10. Campbell JD, McDonough JE, Zeskind JE, Hackett TL, Pechkovsky DV, Brandsma CA, Suzuki M, Gosselink JV, Liu G, Alekseyev YO, et al. A gene expression signature of emphysema-related lung destruction and its reversal by the tripeptide GHK. Genome Medicine. 2012;4(8):67.
  11. Zhang Q, Yan L, Lu J, Zhou X. Glycyl-L-histidyl-L-lysine-Cu2+ attenuates cigarette smoke-induced pulmonary emphysema and inflammation by reducing oxidative stress pathway. Frontiers in Molecular Biosciences. 2022;9:925700.
  12. Pickart L, Vasquez-Soltero JM, Margolina A. GHK-Cu may prevent oxidative stress in skin by regulating copper and modifying expression of numerous antioxidant genes. Cosmetics (MDPI). 2015;2(3):236-247.
  13. Pickart L, Margolina A. Regenerative and protective actions of the GHK-Cu peptide in the light of the new gene data. BioMed Research International. 2015;2015:648108.
  14. Pickart L, Vasquez-Soltero JM, Margolina A. The effect of the human peptide GHK on gene expression relevant to nervous system function and cognitive decline. Brain Sciences. 2017;7(2):20.
  15. Leyden J, Stephens T, Finkey MB, Barkovic S. Skin care benefits of copper peptide containing facial cream. American Academy of Dermatology Annual Meeting Abstract. 2002.
  16. Badenhorst T, Svirskis D, Merrilees M, Bolke L, 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(2):1000166.
  17. Badenhorst T, Svirskis D, Wu Z. Topically applied GHK as an anti-wrinkle peptide. BioImpacts. 2024.
  18. Mohammed YH, Yamada M, Lin LL, Grice JE, Roberts MS, Kendall MA, Prow TW. Microneedle-mediated delivery of copper peptide through skin. Pharmaceutical Research. 2015;32(6):1881-1893.
  19. Mlosek RK, et al. Are we ready to measure skin permeation of modern antiaging GHK-Cu tripeptide encapsulated in liposomes? Molecules (MDPI). 2025;30(1):136.
  20. Wang X, Liu B, Xu Q, Sun H, Shi M, Wang D, Guo M, Yu J, Zhao C, Feng B. GHK-Cu-liposomes accelerate scald wound healing in mice by promoting cell proliferation and angiogenesis. Wound Repair and Regeneration. 2017;25(2):270-278.
  21. Adnan SB, Maarof M, Fauzi MB, Md Fadilah NI. Exploring the role of tripeptides in wound healing and skin regeneration: a comprehensive review. International Journal of Medical Sciences. 2025;22:4175.
  22. Pickart L, Margolina A. Modulation of gene expression in human breast cancer MCF7 and prostate cancer PC3 cells by the human copper-binding peptide GHK-Cu. OBM Genetics. 2021;5(2):128.
  23. Pickart L, Vasquez-Soltero JM, Pickart FD, Majnarich J. Growth-modulating peptides and skin: a 30-year history. Journal of Biomaterials Science, Polymer Edition. 2008;19(8):969-988.
  24. 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.
  25. 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.
  26. 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.
  27. 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.
  28. 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.