# GHK-Cu References: Cited Peer-Reviewed Studies and Literature > GHK-Cu published references: 20 primary peer-reviewed citations from FEBS Letters, IJMS, BioMed Research International, Frontiers in Pharmacology, and other journals, with DOIs and PubMed links. ## Citation Index The following citations are the published references supporting all quantitative claims and findings across this site. Each citation is numbered as it appears inline ([1], [2], …). Model type is indicated for each: in vitro (cell culture), rodent (animal model), human topical (clinical), or review/systematic. --- [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;238(2):343-346. DOI: 10.1016/0014-5793(88)80509-x PMID: 3169264 https://pubmed.ncbi.nlm.nih.gov/3169264/ Model: in vitro [2] 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. DOI: 10.3390/ijms19071987 PMID: 29986520 PMC: PMC6073405 https://pmc.ncbi.nlm.nih.gov/articles/PMC6073405/ Model: in vitro / gene array [3] Pickart L, Margolina A. Regenerative and Protective Actions of the GHK-Cu Peptide in the Light of the New Gene Data. IJMS. 2018. [Rodent collagen dressing and colon cancer cell line data cited from same publication.] DOI: 10.3390/ijms19071987 https://pmc.ncbi.nlm.nih.gov/articles/PMC6073405/ Model: rodent / in vitro [4] 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. DOI: 10.1155/2015/648108 PMID: 26236730 PMC: PMC4508379 https://pmc.ncbi.nlm.nih.gov/articles/PMC4508379/ Model: in vitro [5] 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. DOI: 10.1111/wrr.12520 PMID: 28370978 https://pubmed.ncbi.nlm.nih.gov/28370978/ Model: rodent / in vitro [6] Park J, 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. DOI: 10.18632/oncotarget.11168 PMID: 27517151 PMC: PMC5295439 https://pmc.ncbi.nlm.nih.gov/articles/PMC5295439/ Model: rodent [7] 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. DOI: 10.3390/brainsci7020020 PMID: 28212278 PMC: PMC5332963 https://pmc.ncbi.nlm.nih.gov/articles/PMC5332963/ Model: in vitro / gene array [8] 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. DOI: 10.1155/2012/324832 PMID: 22666519 PMC: PMC3359723 https://pmc.ncbi.nlm.nih.gov/articles/PMC3359723/ Model: in vitro / in silico [9] 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. DOI: 10.3389/fmolb.2022.925700 PMID: 35936787 PMC: PMC9354777 https://pmc.ncbi.nlm.nih.gov/articles/PMC9354777/ Model: rodent / in vitro [10] Lee S, Lee SM, Lee SH, et al. In situ photo-crosslinkable hyaluronic acid-based hydrogel embedded with GHK peptide nanofibers for bioactive wound healing. Acta Biomaterialia. 2023;172:159-174. DOI: 10.1016/j.actbio.2023.10.011 PMID: 37832839 https://pubmed.ncbi.nlm.nih.gov/37832839/ Model: in vitro / rodent [11] Pickart L, Vasquez-Soltero JM, Pickart FD, Majnarich J. GHK, the Human Skin Remodeling Peptide, Induces Anti-Cancer Expression of Numerous Caspase, Growth Regulatory, and DNA Repair Genes. Journal of Analytical Oncology. 2014;3(2):79-87. DOI: 10.6000/1927-7229.2014.03.02.2 https://neoplasiaresearch.com/index.php/jao/article/view/217 Model: in vitro / rodent [12] 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). DOI: 10.21926/obm.genet.2102128 https://www.lidsen.com/journals/genetics/genetics-05-02-128 Model: in vitro [13] 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. [71-woman 12-week clinical outcome data cited from same publication.] DOI: 10.1155/2015/648108 PMID: 26236730 https://pmc.ncbi.nlm.nih.gov/articles/PMC4508379/ Model: human topical [14] Mortazavi SM, Mohammadi Vadoud SA, Moghimi HR. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2024. DOI: 10.34172/bi.30071 PMID: 39963574 https://pubmed.ncbi.nlm.nih.gov/39963574/ Model: review [15] 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. DOI: 10.1001/archfaci.8.4.252 PMID: 16847171 https://pubmed.ncbi.nlm.nih.gov/16847171/ Model: human topical RCT [16] 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. Frontiers in Pharmacology. 2025. DOI: 10.3389/fphar.2025.1551843 PMID: 40672369 https://pubmed.ncbi.nlm.nih.gov/40672369/ Model: rodent [17] 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):186-190. DOI: 10.31491/apt.2024.12.158 PMID: 40823151 PMC: PMC12352503 https://pmc.ncbi.nlm.nih.gov/articles/PMC12352503/ Model: in vitro (aged mouse cells) [18] Dou Y, Lee A, Zhu L, Morton J, Ladiges W. The potential of GHK as an anti-aging peptide. Aging Pathobiology and Therapeutics. 2020;2(1):58-61. DOI: 10.31491/apt.2020.03.014 PMID: 35083444 PMC: PMC8789089 https://pubmed.ncbi.nlm.nih.gov/35083444/ Model: review [19] Ogorek K, Nowak K, Wadych E, Ruzik L, Timerbaev AR, Matczuk M. Are We Ready to Measure Skin Permeation of Modern Antiaging GHK-Cu Tripeptide Encapsulated in Liposomes? Molecules. 2025;30(1):136. DOI: 10.3390/molecules30010136 PMID: 39795193 https://pmc.ncbi.nlm.nih.gov/articles/PMC11721469/ Model: review [20] 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(16):4175-4200. DOI: 10.7150/ijms.118118 https://www.medsci.org/v22p4175.htm Model: review --- Compiled from the gene-modulation script of the published GHK-Cu record — copper-biology indexed, not prescribed.