# TB-500 and Thymosin Beta-4 references — peer-reviewed source list

> Full citation list for TB-500 Doctor: every peer-reviewed paper, clinical trial, and regulatory note referenced across this site, with DOIs and PubMed/PMC/ClinicalTrials.gov URLs.

The references span 1992 (Cassimeris on the Tβ4–actin complex) through 2025 (the latest Tβ4-exosome diabetic-wound hydrogel work). Sortable by year, species, and authors.

## How to read this list

Each entry links to its DOI or to the PubMed / PMC / ClinicalTrials.gov record. The table is sortable by year, by first-author surname, and by species (in vitro, mouse, rat, pig, human). Where a single reference is a review article, the species column is marked 'review' and the entry covers ground that multiple primary papers in the table also address.

The list is the authoritative source for every claim made on this site. If a claim is not traceable to one of these citations, it is either editorial framing or it is wrong — either way, the citation table is the gate. References used in the body of this site are numbered 1–23 in the table below.

## References

[1] Cassimeris L, Safer D, Nachmias VT, Zigmond SH. Interaction of thymosin beta 4 with muscle and platelet actin: implications for actin sequestration in resting platelets. Biochemistry. 1992;31(20):4534-4537. https://pubmed.ncbi.nlm.nih.gov/1627561/
[2] Irobi E, Aguda AH, Larsson M, Guerin C, Yin HL, Burtnick LD, Blanchoin L, Robinson RC. Structural basis of actin sequestration by thymosin-β4: implications for WH2 proteins. EMBO Journal. 2004;23(18):3599-3608. https://pmc.ncbi.nlm.nih.gov/articles/PMC517612/
[3] Malinda KM, Sidhu GS, Mani H, Banaudha K, Maheshwari RK, Goldstein AL, Kleinman HK. Thymosin β4 accelerates wound healing. Journal of Investigative Dermatology. 1999;113(3):364-368. https://www.jidonline.org/article/S0022-202X(15)40595-0/fulltext
[4] Sosne G, Szliter EA, Barrett R, Kernacki KA, Kleinman H, Hazlett LD. Thymosin beta 4 promotes corneal wound healing and decreases inflammation in vivo following alkali injury. Experimental Eye Research. 2002;74(2):293-299. https://pubmed.ncbi.nlm.nih.gov/11950239/
[5] Philp D, Badamchian M, Scheremeta B, Nguyen M, Goldstein AL, Kleinman HK. Thymosin β4 and a synthetic peptide containing its actin-binding domain promote dermal wound repair in db/db diabetic mice and in aged mice. Wound Repair and Regeneration. 2003;11(1):19-24. https://onlinelibrary.wiley.com/doi/abs/10.1046/j.1524-475X.2003.11105.x
[6] Bock-Marquette I, Saxena A, White MD, DiMaio JM, Srivastava D. Thymosin β4 activates integrin-linked kinase and promotes cardiac cell migration, survival and cardiac repair. Nature. 2004;432(7016):466-472. https://pubmed.ncbi.nlm.nih.gov/15565145/
[7] Smart N, Risebro CA, Melville AAD, Moses K, Schwartz RJ, Chien KR, Riley PR. Thymosin β4 induces adult epicardial progenitor mobilization and neovascularization. Nature. 2007;445(7124):177-182. https://pubmed.ncbi.nlm.nih.gov/17108969/
[8] Hinkel R, El-Aouni C, Olson T, Horstkotte J, Mayer S, Müller S, Willhauck M, Spitzweg C, Gildehaus FJ, Münzing W, Hannappel E, Bock-Marquette I, DiMaio JM, Hatzopoulos AK, Boekstegers P, Kupatt C. Thymosin β4 is an essential paracrine factor of embryonic endothelial progenitor cell-mediated cardioprotection. Circulation. 2008;117(17):2232-2240. https://pmc.ncbi.nlm.nih.gov/articles/PMC2672916/
[9] Philp D, Nguyen M, Scheremeta B, St-Surin S, Villa AM, Orgel A, Kleinman HK, Elkin M. Thymosin β4 increases hair growth by activation of hair follicle stem cells. FASEB Journal. 2004;18(2):385-387. https://faseb.onlinelibrary.wiley.com/doi/10.1096/fj.03-0244fje
[10] Tokura Y, Nakayama Y, Fukada S, Nara N, Yamamoto H, Matsuda R, Hara T. Muscle injury-induced thymosin β4 acts as a chemoattractant for myoblasts. Journal of Biochemistry. 2011;149(1):43-48. https://pubmed.ncbi.nlm.nih.gov/20880960/
[11] Qiu P, Wheater MK, Qiu Y, Sosne G. Thymosin β4 inhibits TNF-α-induced NF-κB activation, IL-8 expression, and the sensitizing effects of its partners PINCH-1 and ILK. FASEB Journal. 2011;25(6):1815-1826. https://pmc.ncbi.nlm.nih.gov/articles/PMC3101037/
[12] Morris DC, Chopp M, Zhang L, Lu M, Zhang ZG. Thymosin β4 improves functional neurological outcome in a rat model of embolic stroke. Neuroscience. 2010;169(2):674-682. https://pmc.ncbi.nlm.nih.gov/articles/PMC2907184/
[13] Ruff D, Crockford D, Girardi G, Zhang Y. A randomized, placebo-controlled, single and multiple dose study of intravenous thymosin beta4 in healthy volunteers. Annals of the New York Academy of Sciences. 2010;1194:223-229. https://pubmed.ncbi.nlm.nih.gov/20536472/
[14] Wang X, Liu L, Qi L, Lei C, Li P, Wang Y, Liu C, Bai H, Han C, Sun Y, Liu J. A first-in-human, randomized, double-blind, single- and multiple-dose, phase I study of recombinant human thymosin β4 in healthy Chinese volunteers. Journal of Cellular and Molecular Medicine. 2021;25(17):8259-8271. https://pmc.ncbi.nlm.nih.gov/articles/PMC8419156/
[15] ReGenTree LLC / RegeneRx Biopharmaceuticals (HLB Therapeutics). Assessment of the Safety and Efficacy of RGN-259 Ophthalmic Solutions for Dry Eye Syndrome (ARISE-3). ClinicalTrials.gov identifier NCT03937882. 2020. https://clinicaltrials.gov/study/NCT03937882
[16] Sosne G, Kleinman HK, Springs C, Gross RH, Sung J, Kang S. 0.1% RGN-259 (Thymosin β4) Ophthalmic Solution Promotes Healing and Improves Comfort in Neurotrophic Keratopathy Patients in a Randomized, Placebo-Controlled, Double-Masked Phase III Clinical Trial. International Journal of Molecular Sciences. 2022;24(1):554. https://pmc.ncbi.nlm.nih.gov/articles/PMC9820614/
[17] Stark C, Taimen P, Tarkia M, Pärkkä J, Saraste A, Bizzarro V, Saraste M, Savunen T, Koskenvuo JW. Thymosin β4 protects against cardiac damage and subsequent cardiac fibrosis in mice with myocardial infarction. Cardiovascular Therapeutics. 2022;2022:1308651. https://pmc.ncbi.nlm.nih.gov/articles/PMC9187458/
[18] Yu H, Wang B, Li Z, et al. Tβ4-exosome-loaded hemostatic and antibacterial hydrogel to improve vascular regeneration and modulate macrophage polarization for diabetic wound treatment. Materials Today Bio. 2025;31:101585. https://pmc.ncbi.nlm.nih.gov/articles/PMC11893380/
[19] Lee S, Lee H, Bae S, Kim E, Oh GT, Park H, Yeon JE, Byun KS, Kim K. Targeted deletion of thymosin beta 4 in hepatic stellate cells ameliorates liver fibrosis in a transgenic mouse model. Cells. 2023;12(12):1658. https://pmc.ncbi.nlm.nih.gov/articles/PMC10297343/
[20] Esposito S, Deventer K, Goeyens L, Van Eenoo P. Doping control analysis of TB-500, a synthetic version of an active region of thymosin β4, in equine urine and plasma by liquid chromatography–mass spectrometry. Analytica Chimica Acta. 2012;753:87-95. https://www.sciencedirect.com/science/article/abs/pii/S0021967312014550
[21] Wei C, Kumar S, Kim IK, Gupta S. Systemic dosing of thymosin beta 4 before and after ischemia does not attenuate global myocardial ischemia-reperfusion injury in pigs. Frontiers in Pharmacology. 2016;7:115. https://pmc.ncbi.nlm.nih.gov/articles/PMC4853610/
[22] Goldstein AL, Hannappel E, Sosne G, Kleinman HK. Thymosin β4: a multi-functional regenerative peptide. Basic properties and clinical applications. Expert Opinion on Biological Therapy. 2012;12(1):37-51. https://pubmed.ncbi.nlm.nih.gov/22074294/
[23] Xing Y, Ye Y, Zuo H, Li Y. Progress on the function and application of thymosin β4. Frontiers in Endocrinology. 2021;12:767785. https://www.frontiersin.org/journals/endocrinology/articles/10.3389/fendo.2021.767785/full
[24] Mendias CL, Awan TM. Safety and Efficacy of Approved and Unapproved Peptide Therapies for Musculoskeletal Injuries and Athletic Performance. Sports Medicine. 2026. https://pubmed.ncbi.nlm.nih.gov/41966639/
[25] Cha HJ, Jeong MJ, Kleinman HK. Role of thymosin beta4 in tumor metastasis and angiogenesis. Journal of the National Cancer Institute. 2003;95(22):1674-1680. https://pubmed.ncbi.nlm.nih.gov/14625258/
[26] Wang WS, et al. Thymosin beta 4 is overexpressed in human pancreatic cancer cells and stimulates proinflammatory cytokine secretion and JNK activation. Cancer Biology & Therapy. 2008;7(3):419-423. https://pubmed.ncbi.nlm.nih.gov/18094619/
[27] Cooper TM, et al. Doping control analysis of TB-500, a synthetic version of an active region of thymosin β4, in equine urine and plasma by liquid chromatography-mass spectrometry. Journal of Chromatography A. 2012;1264:57-64. https://pubmed.ncbi.nlm.nih.gov/23084823/
[28] Spurney CF, et al. Evaluation of skeletal and cardiac muscle function after chronic administration of thymosin beta-4 in the dystrophin deficient mouse. PLoS One. 2010;5(1):e8976. https://pubmed.ncbi.nlm.nih.gov/20126456/
[29] Thomas A, et al. TB500/TB1000 and SGF1000: A scientific approach for a better understanding of doping-relevant peptide preparations. Drug Testing and Analysis. 2023;15(12):1411-1422. https://pubmed.ncbi.nlm.nih.gov/36482504/

---

An editorial record of the peer-reviewed Thymosin Beta-4 literature — not a clinic, not a vendor, not medical guidance.