From the literature
Published GHK-Cu references — cited, linked, and sourced
Every citation used across this site, with DOIs and PubMed links. GHK-Cu research from 1988 to 2026.
About this reference list
These are the published GHK-Cu references cited across this site. Every quantitative claim maps to one of these papers. The references span 1988 (first fibroblast collagen study) to 2026 (Biomedicines hair-loss review). GHK-Cu preclinical and clinical findings are cited by number in the body text; cross-reference via the ID column.
Use the table controls to sort by year, author, or journal — useful if you are tracking the chronological arc of Pickart's research program or isolating recent 2024–2025 publications. Two entries (Tucker et al. 2023) are preprints indexed in PubMed; their peer-reviewed journal publication status is noted in the table.
- 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. ↗
- Maquart FX, Bellon G, Chaqour B, et al. 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. ↗
- 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. ↗
- 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. ↗
- Park JR, Lee H, Kim SI, Yang SR. The tri-peptide GHK-Cu complex ameliorates lipopolysaccharide-induced acute lung injury in mice. Oncotarget. 2016. ↗
- Zhou XM, et al. GHK Peptide Inhibits Bleomycin-Induced Pulmonary Fibrosis in Mice by Suppressing TGFbeta1/Smad-Mediated Epithelial-to-Mesenchymal Transition. Frontiers in Pharmacology. 2017. ↗
- Pickart L, Margolina A. Topically applied GHK as an anti-wrinkle peptide: Advantages, problems and prospective. BioImpacts. 2024. ↗
- 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. ↗
- 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. ↗
- Hong Y, Downey T, Eu KW, Koh PK, Cheah PY. A 'metastasis-prone' signature for early-stage mismatch-repair proficient sporadic colorectal cancer patients and its implications for possible therapeutics. Clinical and Experimental Metastasis. 2010. ↗
- 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. Annals of Dermatology. 2016;28(4):438. ↗
- Kang YA, Choi HR, Na JI, Huh CH, Kim MJ, Youn SW, Kim KH, Park KC. Copper-GHK increases integrin expression and p63 positivity by keratinocytes. Archives of Dermatological Research. 2009. ↗
- Gruchlik A, Chodurek E, Dzierzewicz Z. Effect of GLY-HIS-LYS and its copper complex on TGF-beta secretion in normal human dermal fibroblasts. Acta Poloniae Pharmaceutica. 2014;71(6):954-958. ↗
- Hostynek JJ, Dreher F, Maibach HI. Human skin penetration of a copper tripeptide in vitro as a function of skin layer. Inflammation Research. 2010. ↗
- 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. ↗
- 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. ↗
- 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. bioRxiv (preprint). 2023. ↗
- Chen H, Yang P, et al. Food-Derived Tripeptide-Copper Self-Healing Hydrogel for Infected Wound Healing. Biomaterials Research. 2025. ↗
- Tucker M, Keely A, Park JY, et al. Intranasal GHK peptide enhances resilience to cognitive decline in aging mice. bioRxiv (preprint). 2023. ↗
- 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. ↗
- Trachy RE, Fors TD, Pickart L, Uno H. The hair follicle-stimulating properties of peptide copper complexes. Results in C3H mice. Annals of the New York Academy of Sciences. 1991. ↗
- Pickart L, Margolina A. Regenerative and Protective Actions of the GHK-Cu Peptide in the Light of the New Gene Data (wound healing synthesis). International Journal of Molecular Sciences. 2018;19(7):1987. ↗
- 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. ↗