In Vitro Immunodetection of Prothymosin Alpha in Normal and Pathological Conditions

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Abstract

Prothymosin alpha (ProTα) is a highly acidic polypeptide, ubiquitously expressed in almost all mammalian cells and tissues and consisting of 109 amino acids in humans. ProTα is known to act both, intracellularly, as an anti-apoptotic and proliferation mediator, and extracellularly, as a biologic response modifier mediating immune responses similar to molecules termed as “alarmins”. Antibodies and immunochemical techniques for ProTα have played a leading role in the investigation of the biological role of ProTα, several aspects of which still remain unknown and contributed to unraveling the diagnostic and therapeutic potential of the polypeptide. This review deals with the so far reported antibodies along with the related immunodetection methodology for ProTα (immunoassays as well as immunohistochemical, immunocytological, immunoblotting, and immunoprecipitation techniques) and its application to biological samples of interest (tissue extracts and sections, cells, cell lysates and cell culture supernatants, body fluids), in health and disease states. In this context, literature information is critically discussed, and some concluding remarks are presented.

Keywords: Prothymosin alpha, antibodies, immunodetection, immunoassay, immunohistochemistry, immunocytology, immunoblotting, disease biomarker.

[1]
Haritos, A.A.; Goodall, G.J.; Horecker, B.L. Prothymosin alpha: isolation and properties of the major immunoreactive form of thymosin alpha 1 in rat thymus. Proc. Natl. Acad. Sci. USA, 1984, 81(4), 1008-1011.
[http://dx.doi.org/10.1073/pnas.81.4.1008] [PMID: 6583693]
[2]
Goldstein, A.L.; Slater, F.D.; White, A. Preparation, assay, and partial purification of a thymic lymphocytopoietic factor (thymosin). Proc. Natl. Acad. Sci. USA, 1966, 56(3), 1010-1017.
[http://dx.doi.org/10.1073/pnas.56.3.1010] [PMID: 5230175]
[3]
Goldstein, A.L.; Low, T.L.; McAdoo, M.; McClure, J.; Thurman, G.B.; Rossio, J.; Lai, C.Y.; Chang, D.; Wang, S.S.; Harvey, C.; Ramel, A.H.; Meienhofer, J. Thymosin alpha1: isolation and sequence analysis of an immunologically active thymic polypeptide. Proc. Natl. Acad. Sci. USA, 1977, 74(2), 725-729.
[http://dx.doi.org/10.1073/pnas.74.2.725] [PMID: 265536]
[4]
Haritos, A.A.; Blacher, R.; Stein, S.; Caldarella, J.; Horecker, B.L. Primary structure of rat thymus prothymosin alpha. Proc. Natl. Acad. Sci. USA, 1985, 82(2), 343-346.
[http://dx.doi.org/10.1073/pnas.82.2.343] [PMID: 3855555]
[5]
Yialouris, P.P.; Evangelatos, G.P.; Soteriadis-Vlahos, C.; Heimer, E.P.; Felix, A.M.; Tsitsiloni, O.E.; Haritos, A.A. The identification of prothymosin alpha-like material in vertebrate lymphoid organs by a radioimmunoassay for the N-terminal decapeptide. J. Immunol. Methods, 1988, 106(2), 267-275.
[http://dx.doi.org/10.1016/0022-1759(88)90207-4] [PMID: 3339257]
[6]
Aniello, F.; Branno, M.; De Rienzo, G.; Ferrara, D.; Palmiero, C.; Minucci, S. First evidence of prothymosin alpha in a non-mammalian vertebrate and its involvement in the spermatogenesis of the frog Rana esculenta. Mech. Dev., 2002, 110(1-2), 213-217.
[http://dx.doi.org/10.1016/S0925-4773(01)00569-X] [PMID: 11744386]
[7]
De Rienzo, G.; Di Sena, R.; Ferrara, D.; Palmiero, C.; Chieffi Baccari, G.; Minucci, S. Temporal and spatial localization of prothymosin alpha transcript in the Harderian gland of the frog, Rana esculenta. J. Exp. Zool., 2002, 292(7), 633-639.
[http://dx.doi.org/10.1002/jez.10097] [PMID: 12115928]
[8]
Zhang, B.C.; Sun, L. Tongue sole (Cynoglossus semilaevis) prothymosin alpha: Cytokine-like activities associated with the intact protein and the C-terminal region that lead to antiviral immunity via Myd88-dependent and -independent pathways respectively. Dev. Comp. Immunol., 2015, 53(1), 96-104.
[http://dx.doi.org/10.1016/j.dci.2015.07.004] [PMID: 26162512]
[9]
Samara, P.; Karachaliou, C.E.; Ioannou, K.; Papaioannou, N.E.; Voutsas, I.F.; Zikos, C.; Pirmettis, I.; Papadopoulos, M.; Kalbacher, H.; Livaniou, E.; Tsitsilonis, O.E.; Voelter, W. Prothymosin Alpha: An Alarmin and More..... Curr. Med. Chem., 2017, 24(17), 1747-1760.
[http://dx.doi.org/10.2174/0929867324666170518110033] [PMID: 28521686]
[10]
Eschenfeldt, W.H.; Berger, S.L. The human prothymosin alpha gene is polymorphic and induced upon growth stimulation: evidence using a cloned cDNA. Proc. Natl. Acad. Sci. USA, 1986, 83(24), 9403-9407.
[http://dx.doi.org/10.1073/pnas.83.24.9403] [PMID: 3467312]
[11]
Gomez-Marquez, J.; Rodríguez, P. Prothymosin alpha is a chromatin-remodelling protein in mammalian cells. Biochem. J., 1998, 333(Pt 1), 1-3.
[http://dx.doi.org/10.1042/bj3330001] [PMID: 9639554]
[12]
Tsitsiloni, O.E.; Yialouris, P.P.; Sekeri-Pataryas, K.; Haritos, A.A. Prothymosin alpha is not a nuclear polypeptide. Experientia, 1989, 45(4), 332-334.
[http://dx.doi.org/10.1007/BF01957467] [PMID: 2707372]
[13]
Enkemann, S.A.; Ward, R.D.; Berger, S.L. Mobility within the nucleus and neighboring cytosol is a key feature of prothymosin-alpha. J. Histochem. Cytochem., 2000, 48(10), 1341-1355.
[http://dx.doi.org/10.1177/002215540004801005] [PMID: 10990488]
[14]
Jiang, X.; Kim, H.E.; Shu, H.; Zhao, Y.; Zhang, H.; Kofron, J.; Donnelly, J.; Burns, D.; Ng, S.C.; Rosenberg, S.; Wang, X. Distinctive roles of PHAP proteins and prothymosin-alpha in a death regulatory pathway. Science, 2003, 299(5604), 223-226.
[http://dx.doi.org/10.1126/science.1076807] [PMID: 12522243]
[15]
Karapetian, R.N.; Evstafieva, A.G.; Abaeva, I.S.; Chichkova, N.V.; Filonov, G.S.; Rubtsov, Y.P.; Sukhacheva, E.A.; Melnikov, S.V.; Schneider, U.; Wanker, E.E.; Vartapetian, A.B. Nuclear oncoprotein prothymosin alpha is a partner of Keap1: implications for expression of oxidative stress-protecting genes. Mol. Cell. Biol., 2005, 25(3), 1089-1099.
[http://dx.doi.org/10.1128/MCB.25.3.1089-1099.2005] [PMID: 15657435]
[16]
Ioannou, K.; Samara, P.; Livaniou, E.; Derhovanessian, E.; Tsitsilonis, O.E. Prothymosin alpha: a ubiquitous polypeptide with potential use in cancer diagnosis and therapy. Cancer Immunol. Immunother., 2012, 61(5), 599-614.
[http://dx.doi.org/10.1007/s00262-012-1222-8] [PMID: 22366887]
[17]
Reclos, G.J.; Baxevanis, C.N.; Sfagos, C.; Papageorgiou, C.; Tsokos, G.C.; Papamichail, M. Multiple sclerosis: II. Effects of prothymosin alpha on the autologous and allogeneic MLR in patients with multiple sclerosis. Clin. Exp. Immunol., 1987, 70(2), 336-344.
[PMID: 2962790]
[18]
Baxevanis, C.N.; Reclos, G.J.; Papamichail, M.; Tsokos, G.C. Prothymosin alpha restores the depressed autologous and allogeneic mixed lymphocyte responses in patients with systemic lupus erythematosus. Immunopharmacol. Immunotoxicol., 1987, 9(4), 429-440.
[http://dx.doi.org/10.3109/08923978709035224] [PMID: 2963855]
[19]
Gusella, G.L.; Teixeira, A.; Aberg, J.; Uversky, V.N.; Mosoian, A. Prothymosin-α Variants Elicit Anti-HIV-1 Response via TLR4 Dependent and Independent Pathways. PLoS One, 2016, 11(6), e0156486.
[http://dx.doi.org/10.1371/journal.pone.0156486] [PMID: 27310139]
[20]
Maeda, S.; Sasaki, K.; Halder, S.K.; Fujita, W.; Ueda, H. Neuroprotective DAMPs member prothymosin alpha has additional beneficial actions against cerebral ischemia-induced vascular damages. J. Pharmacol. Sci., 2016, 132(1), 100-104.
[http://dx.doi.org/10.1016/j.jphs.2016.05.006] [PMID: 27543170]
[21]
Sarandeses, C.S.; Covelo, G.; Díaz-Jullien, C.; Freire, M. Prothymosin alpha is processed to thymosin alpha 1 and thymosin alpha 11 by a lysosomal asparaginyl endopeptidase. J. Biol. Chem., 2003, 278(15), 13286-13293.
[http://dx.doi.org/10.1074/jbc.M213005200] [PMID: 12554742]
[22]
Enkemann, S.A.; Wang, R.H.; Trumbore, M.W.; Berger, S.L. Functional discontinuities in prothymosin alpha caused by caspase cleavage in apoptotic cells. J. Cell. Physiol., 2000, 182(2), 256-268.
[http://dx.doi.org/10.1002/(SICI)1097-4652(200002)182:2<256:AID-JCP15>3.0.CO;2-N] [PMID: 10623890]
[23]
Evstafieva, A.G.; Belov, G.A.; Kalkum, M.; Chichkova, N.V.; Bogdanov, A.A.; Agol, V.I.; Vartapetian, A.B. Prothymosin alpha fragmentation in apoptosis. FEBS Lett., 2000, 467(2-3), 150-154.
[http://dx.doi.org/10.1016/S0014-5793(00)01139-X] [PMID: 10675528]
[24]
Skopeliti, M.; Iconomidou, V.A.; Derhovanessian, E.; Pawelec, G.; Voelter, W.; Kalbacher, H.; Hamodrakas, S.J.; Tsitsilonis, O.E. Prothymosin alpha immunoactive carboxyl-terminal peptide TKKQKTDEDD stimulates lymphocyte reactions, induces dendritic cell maturation and adopts a beta-sheet conformation in a sequence-specific manner. Mol. Immunol., 2009, 46(5), 784-792.
[http://dx.doi.org/10.1016/j.molimm.2008.09.014] [PMID: 18976813]
[25]
Fortis, S.P.; Anastasopoulou, E.A.; Voutsas, I.F.; Baxevanis, C.N.; Perez, S.A.; Mahaira, L.G. Potential Prognostic Molecular Signatures in a Preclinical Model of Melanoma. Anticancer Res., 2017, 37(1), 143-148.
[http://dx.doi.org/10.21873/anticanres.11299] [PMID: 28011484]
[26]
Tsitsilonis, O.E.; Bekris, E.; Voutsas, I.F.; Baxevanis, C.N.; Markopoulos, C.; Papadopoulou, S.A.; Kontzoglou, K.; Stoeva, S.; Gogas, J.; Voelter, W.; Papamichail, M. The prognostic value of alpha-thymosins in breast cancer. Anticancer Res., 1998, 18(3A), 1501-1508.
[PMID: 9673361]
[27]
Magdalena, C.; Dominguez, F.; Loidi, L.; Puente, J.L. Tumour prothymosin alpha content, a potential prognostic marker for primary breast cancer. Br. J. Cancer, 2000, 82(3), 584-590.
[http://dx.doi.org/10.1054/bjoc.1999.0968] [PMID: 10682670]
[28]
Tzai, T.S.; Tsai, Y.S.; Shiau, A.L.; Wu, C.L.; Shieh, G.S.; Tsai, H.T. Urine prothymosin-alpha as novel tumor marker for detection and follow-up of bladder cancer. Urology, 2006, 67(2), 294-299.
[http://dx.doi.org/10.1016/j.urology.2005.08.034] [PMID: 16461079]
[29]
Jou, Y.C.; Tung, C.L.; Tsai, Y.S.; Shen, C.H.; Syue-Yi, C.; Shiau, A.L.; Tsai, H.T.; Wu, C.L.; Tzai, T.S. Prognostic relevance of prothymosin-alpha expression in human upper urinary tract transitional cell carcinoma. Urology, 2009, 74(4), 951-957.
[http://dx.doi.org/10.1016/j.urology.2008.11.060] [PMID: 19679339]
[30]
Freire, M.; Díaz-Jullien, C.; Covelo, G. Developments in prothymosin α research.Recent Research and Development in Proteins; Pandalai, S.G., Ed.; Transworld Research Network: Kerala, India, 2002, Vol. 1, pp. 257-276.
[31]
MacRaild, C.A.; Richards, J.S.; Anders, R.F.; Norton, R.S. Antibody Recognition of Disordered Antigens. Structure, 2016, 24(1), 148-157.
[http://dx.doi.org/10.1016/j.str.2015.10.028] [PMID: 26712277]
[32]
Sukhacheva, E.A.; Evstafieva, A.G.; Fateeva, T.V.; Shakulov, V.R.; Efimova, N.A.; Karapetian, R.N.; Rubtsov, Y.P.; Vartapetian, A.B. Sensing prothymosin alpha origin, mutations and conformation with monoclonal antibodies. J. Immunol. Methods, 2002, 266(1-2), 185-196.
[http://dx.doi.org/10.1016/S0022-1759(02)00098-4] [PMID: 12133636]
[33]
Kijogi, C.M.; Khayeka-Wandabwa, C.; Sasaki, K.; Tanaka, Y.; Kurosu, H.; Matsunaga, H.; Ueda, H. Subcellular dissemination of prothymosin alpha at normal physiology: immunohistochemical vis-a-vis western blotting perspective. BMC Physiol., 2016, 16, 2.
[http://dx.doi.org/10.1186/s12899-016-0021-4] [PMID: 26932824]
[34]
Evstafieva, A.G.; Belov, G.A.; Rubtsov, Y.P.; Kalkum, M.; Joseph, B.; Chichkova, N.V.; Sukhacheva, E.A.; Bogdanov, A.A.; Pettersson, R.F.; Agol, V.I.; Vartapetian, A.B. Apoptosis-related fragmentation, translocation, and properties of human prothymosin alpha. Exp. Cell Res., 2003, 284(2), 211-223.
[http://dx.doi.org/10.1016/S0014-4827(02)00047-2] [PMID: 12651154]
[35]
Su, B.H.; Tseng, Y.L.; Shieh, G.S.; Chen, Y.C.; Shiang, Y.C.; Wu, P.; Li, K.J.; Yen, T.H.; Shiau, A.L.; Wu, C.L. Prothymosin α overexpression contributes to the development of pulmonary emphysema. Nat. Commun., 2013, 4, 1906.
[http://dx.doi.org/10.1038/ncomms2906] [PMID: 23695700]
[36]
Klimentzou, P.; Paravatou-Petsotas, M.; Zikos, C.; Beck, A.; Skopeliti, M.; Czarnecki, J.; Tsitsilonis, O.; Voelter, W.; Livaniou, E.; Evangelatos, G.P. Development and immunochemical evaluation of antibodies Y for the poorly immunogenic polypeptide prothymosin alpha. Peptides, 2006, 27(1), 183-193.
[http://dx.doi.org/10.1016/j.peptides.2005.07.002] [PMID: 16150512]
[37]
Härtle, S.; Magor, K.E.; Göbel, T.W.; Davison, F.; Kaspers, B. Structure and Evolution of Avian Immunoglobulins.Avian Immunology; 2nd ed; Schat, K.A.; Kaspers, B.; Kaiser, P., Eds.; Academic Press: Boston., 2014, pp. 103-120.
[38]
Suzuki, S.; Takahashi, S.; Takahashi, S.; Takeshita, K.; Hikosaka, A.; Wakita, T.; Nishiyama, N.; Fujita, T.; Okamura, T.; Shirai, T. Expression of prothymosin alpha is correlated with development and progression in human prostate cancers. Prostate, 2006, 66(5), 463-469.
[http://dx.doi.org/10.1002/pros.20385] [PMID: 16353248]
[39]
Leys, C.M.; Nomura, S.; LaFleur, B.J.; Ferrone, S.; Kaminishi, M.; Montgomery, E.; Goldenring, J.R. Expression and prognostic significance of prothymosin-alpha and ERp57 in human gastric cancer. Surgery, 2007, 141(1), 41-50.
[http://dx.doi.org/10.1016/j.surg.2006.05.009] [PMID: 17188166]
[40]
Tsai, Y.S.; Jou, Y.C.; Lee, G.F.; Chen, Y.C.; Shiau, A.L.; Tsai, H.T.; Wu, C.L.; Tzai, T.S. Aberrant prothymosin-alpha expression in human bladder cancer. Urology, 2009, 73(1), 188-192.
[http://dx.doi.org/10.1016/j.urology.2008.05.031] [PMID: 18692879]
[41]
Kobayashi, D.; Kumagai, J.; Morikawa, T.; Wilson-Morifuji, M.; Wilson, A.; Irie, A.; Araki, N. An integrated approach of differential mass spectrometry and gene ontology analysis identified novel proteins regulating neuronal differentiation and survival. Mol. Cell. Proteomics, 2009, 8(10), 2350-2367.
[http://dx.doi.org/10.1074/mcp.M900179-MCP200] [PMID: 19525549]
[42]
Kashat, L.; So, A.K.; Masui, O.; Wang, X.S.; Cao, J.; Meng, X.; Macmillan, C.; Ailles, L.E.; Siu, K.W.; Ralhan, R.; Walfish, P.G. Secretome-based identification and characterization of potential biomarkers in thyroid cancer. J. Proteome Res., 2010, 9(11), 5757-5769.
[http://dx.doi.org/10.1021/pr100529t] [PMID: 20873772]
[43]
Matsunaga, H.; Ueda, H. Stress-induced non-vesicular release of prothymosin-α initiated by an interaction with S100A13, and its blockade by caspase-3 cleavage. Cell Death Differ., 2010, 17(11), 1760-1772.
[http://dx.doi.org/10.1038/cdd.2010.52] [PMID: 20467443]
[44]
Bowick, G.C.; Soman, K.V.; Wang, H.; Aronson, J.F.; Luxon, B.A.; Lomas, L.O.; Gorenstein, D.G.; Herzog, N.K. Proteomic analysis of Pichindé virus infection identifies differential expression of prothymosin-alpha. J. Biomed. Biotechnol., 2010, 2010-->, 2010.
[45]
Tripathi, S.C.; Matta, A.; Kaur, J.; Grigull, J.; Chauhan, S.S.; Thakar, A.; Shukla, N.K.; Duggal, R.; Choudhary, A.R.; Dattagupta, S.; Sharma, M.C.; Ralhan, R.; Siu, K.W. Overexpression of prothymosin alpha predicts poor disease outcome in head and neck cancer. PLoS One, 2011, 6(5), e19213.
[http://dx.doi.org/10.1371/journal.pone.0019213] [PMID: 21573209]
[46]
Halder, S.K.; Ueda, H. Regional distribution and cell type-specific subcellular localization of Prothymosin alpha in brain. Cell. Mol. Neurobiol., 2012, 32(1), 59-66.
[http://dx.doi.org/10.1007/s10571-011-9734-x] [PMID: 21750924]
[47]
Ferrara, D.; Pariante, P.; Di Matteo, L.; Serino, I.; Oko, R.; Minucci, S. First evidence of prothymosin α localization in the acrosome of mammalian male gametes. J. Cell. Physiol., 2013, 228(8), 1629-1637.
[http://dx.doi.org/10.1002/jcp.24332] [PMID: 23359453]
[48]
Tsai, Y.S.; Jou, Y.C.; Tung, C.L.; Lin, C.T.; Shen, C.H.; Chen, S.Y.; Tsai, H.T.; Lai, C.L.; Wu, C.L.; Tzai, T.S. Loss of nuclear prothymosin-α expression is associated with disease progression in human superficial bladder cancer. Virchows Arch., 2014, 464(6), 717-724.
[http://dx.doi.org/10.1007/s00428-014-1578-6] [PMID: 24733561]
[49]
Zhang, M.; Cui, F.; Lu, S.; Lu, H.; Jiang, T.; Chen, J.; Zhang, X.; Jin, Y.; Peng, Z.; Tang, H. Increased expression of prothymosin-α, independently or combined with TP53, correlates with poor prognosis in colorectal cancer. Int. J. Clin. Exp. Pathol., 2014, 7(8), 4867-4876.
[PMID: 25197357]
[50]
Ha, S.Y.; Song, D.H.; Hwang, S.H.; Cho, S.Y.; Park, C.K. Expression of prothymosin alpha predicts early recurrence and poor prognosis of hepatocellular carcinoma. HBPD INT, 2015, 14(2), 171-177.
[http://dx.doi.org/10.1016/S1499-3872(14)60326-X] [PMID: 25865690]
[51]
Chauhan, S.S.; Kaur, J.; Kumar, M.; Matta, A.; Srivastava, G.; Alyass, A.; Assi, J.; Leong, I.; MacMillan, C.; Witterick, I.; Colgan, T.J.; Shukla, N.K.; Thakar, A.; Sharma, M.C.; Siu, K.W.; Walfish, P.G.; Ralhan, R. Prediction of recurrence-free survival using a protein expression-based risk classifier for head and neck cancer. Oncogenesis, 2015, 4, e147.
[http://dx.doi.org/10.1038/oncsis.2015.7] [PMID: 25893634]
[52]
Wierzbicka-Tutka, I.; Sokołowski, G.; Bałdys-Waligórska, A.; Adamek, D.; Radwańska, E.; Gołkowski, F. Prothymosin-alpha and Ki-67 expression in pituitary adenomas. Postepy Hig. Med. Dosw., 2016, 70(0), 1117-1123.
[http://dx.doi.org/10.5604/17322693.1223796] [PMID: 27892895]
[53]
Barbeito, P.; Sarandeses, C.S.; Díaz-Jullien, C.; Muras, J.; Covelo, G.; Moreira, D.; Freire-Cobo, C.; Freire, M. Prothymosin α interacts with SET, ANP32A and ANP32B and other cytoplasmic and mitochondrial proteins in proliferating cells. Arch. Biochem. Biophys., 2017, 635, 74-86.
[http://dx.doi.org/10.1016/j.abb.2017.10.014] [PMID: 29106904]
[54]
Chen, K.; Xiong, L.; Yang, Z.; Huang, S.; Zeng, R.; Miao, X. Prothymosin-α and parathymosin expression predicts poor prognosis in squamous and adenosquamous carcinomas of the gallbladder. Oncol. Lett., 2018, 15(4), 4485-4494.
[http://dx.doi.org/10.3892/ol.2018.7824] [PMID: 29541218]
[55]
Vlachoyiannopoulos, P.G.; Frillingos, S.; Tzioufas, A.G.; Seferiadis, K.; Moutsopoulos, H.M.; Tsolas, O. Circulating antibodies to prothymosin alpha in systemic lupus erythematosus. Clin. Immunol. Immunopathol., 1989, 53(2 Pt 1), 151-160.
[http://dx.doi.org/10.1016/0090-1229(89)90045-7] [PMID: 2791337]
[56]
Cordero, O.J.; Sarandeses, C.S.; Nogueira, M. Binding of 125I-prothymosin alpha to lymphoblasts through the non-thymosin alpha 1 sequence. Life Sci., 1996, 58(20), 1757-1770.
[http://dx.doi.org/10.1016/0024-3205(96)00157-9] [PMID: 8637400]
[57]
Haritos, A.A.; Horecker, B.L. A radioimmunoassay for thymosin alpha 1 that detects the native polypeptide, prothymosin alpha. J. Immunol. Methods, 1985, 81(2), 199-205.
[http://dx.doi.org/10.1016/0022-1759(85)90204-2] [PMID: 2410515]
[58]
Dominguez, F.; Magdalena, C.; Cancio, E.; Roson, E.; Paredes, J.; Loidi, L.; Zalvide, J.; Fraga, M.; Forteza, J.; Regueiro, B.J. Tissue concentrations of prothymosin alpha: a novel proliferation index of primary breast cancer. Eur. J. Cancer, 1993, 29A(6), 893-897.
[http://dx.doi.org/10.1016/S0959-8049(05)80433-2] [PMID: 8387320]
[59]
Loidi, L.; Vidal, A.; Zalvide, J.B.; Puente, J.L.; Reyes, F.; Domínguez, F. Development of ELISA to estimate thymosin alpha1, the N terminus of prothymosin alpha, in human tumors. Clin. Chem., 1997, 43(1), 59-63.
[http://dx.doi.org/10.1093/clinchem/43.1.59] [PMID: 8990223]
[60]
Costopoulou, D.; Leondiadis, L.; Czarnecki, J.; Ferderigos, N.; Ithakissios, D.S.; Livaniou, E.; Evangelatos, G.P. Direct ELISA method for the specific determination of prothymosin alpha in human specimens. J. Immunoassay, 1998, 19(4), 295-316.
[http://dx.doi.org/10.1080/01971529808005487] [PMID: 9840299]
[61]
Samara, P.; Kalbacher, H.; Ioannou, K.; Radu, D.L.; Livaniou, E.; Promponas, V.J.; Voelter, W.; Tsitsilonis, O. Development of an ELISA for the quantification of the C-terminal decapeptide prothymosin α(100-109) in sera of mice infected with bacteria. J. Immunol. Methods, 2013, 395(1-2), 54-62.
[http://dx.doi.org/10.1016/j.jim.2013.06.011] [PMID: 23831611]
[62]
Samara, P.; Miriagou, V.; Zachariadis, M.; Mavrofrydi, O.; Promponas, V.J.; Dedos, S.G.; Papazafiri, P.; Kalbacher, H.; Voelter, W.; Tsitsilonis, O. A fragment of the alarmin prothymosin α as a novel biomarker in murine models of bacteria-induced sepsis. Oncotarget, 2017, 8(30), 48635-48649.
[http://dx.doi.org/10.18632/oncotarget.18149] [PMID: 28611290]
[63]
Wu, C.G.; Habib, N.A.; Mitry, R.R.; Reitsma, P.H.; van Deventer, S.J.; Chamuleau, R.A. Overexpression of hepatic prothymosin alpha, a novel marker for human hepatocellular carcinoma. Br. J. Cancer, 1997, 76(9), 1199-1204.
[http://dx.doi.org/10.1038/bjc.1997.533] [PMID: 9365169]
[64]
Karachaliou, C-E.; Klimetzou, P.; Paravatou-Petsotas, M.; Tsitsilonis, O.; Voelter, W.; Kalbacher, H.; Zikos, C.; Livaniou, E. Development of an IgG/IgY sandwich-ELISA for the bioactive polypeptide Prothymosin alpha. Peptides 2018 Proceedings Volume of the 35th EPS; European Peptide Society, 2018, pp. 230-232.
[65]
Clinton, M.; Frangou-Lazaridis, M.; Panneerselvam, C.; Horecker, B.L. Prothymosin alpha and parathymosin: mRNA and polypeptide levels in rodent tissues. Arch. Biochem. Biophys., 1989, 269(1), 256-263.
[http://dx.doi.org/10.1016/0003-9861(89)90107-0] [PMID: 2916841]
[66]
Tsitsiloni, O.E.; Stiakakis, J.; Koutselinis, A.; Gogas, J.; Markopoulos, C.; Yialouris, P.; Bekris, S.; Panoussopoulos, D.; Kiortsis, V.; Voelter, W. Expression of alpha-thymosins in human tissues in normal and abnormal growth. Proc. Natl. Acad. Sci. USA, 1993, 90(20), 9504-9507.
[http://dx.doi.org/10.1073/pnas.90.20.9504] [PMID: 8415730]
[67]
Tsitsiloni, O.E.; Heimer, E.; Felix, A.; Yialouris, P.P.; Vamvoukakis, J.; Voelter, W.; Haritos, A.A. Radioimmunoassays for the C-terminus of prothymosin alpha and the N-terminus of parathymosin alpha for the measurement of the levels of alpha-thymosins in human cancer. J. Immunol. Methods, 1994, 169(2), 163-171.
[http://dx.doi.org/10.1016/0022-1759(94)90260-7] [PMID: 7510758]
[68]
Panneerselvam, C.; Haritos, A.A.; Caldarella, J.; Horecker, B.L. Prothymosin alpha in human blood. Proc. Natl. Acad. Sci. USA, 1987, 84(13), 4465-4469.
[http://dx.doi.org/10.1073/pnas.84.13.4465] [PMID: 3474615]
[69]
Conteas, C.N.; Mutchnick, M.G.; Palmer, K.C.; Weller, F.E.; Luk, G.D.; Naylor, P.H.; Erdos, M.R.; Goldstein, A.L.; Panneerselvam, C.; Horecker, B.L. Cellular levels of thymosin immunoreactive peptides are linked to proliferative events: evidence for a nuclear site of action. Proc. Natl. Acad. Sci. USA, 1990, 87(9), 3269-3273.
[http://dx.doi.org/10.1073/pnas.87.9.3269] [PMID: 2375791]
[70]
Jou, Y.C.; Tsai, Y.S.; Hsieh, H.Y.; Chen, S.Y.; Tsai, H.T.; Chen, K.J.; Wang, S.T.; Shiau, A.L.; Wu, C.L.; Tzai, T.S. Plasma thymosin-α1 level as a potential biomarker in urothelial and renal cell carcinoma. Urol. Oncol., 2013, 31(8), 1806-1811.
[http://dx.doi.org/10.1016/j.urolonc.2012.03.011] [PMID: 22609059]
[71]
Su, Y.C.; Ou, H.Y.; Wu, H.T.; Wu, P.; Chen, Y.C.; Su, B.H.; Shiau, A.L.; Chang, C.J.; Wu, C.L. Prothymosin-α Overexpression Contributes to the Development of Insulin Resistance. J. Clin. Endocrinol. Metab., 2015, 100(11), 4114-4123.
[http://dx.doi.org/10.1210/jc.2015-2277] [PMID: 26348351]
[72]
Ueda, H.; Fujita, R.; Yoshida, A.; Matsunaga, H.; Ueda, M. Identification of prothymosin-alpha1, the necrosis-apoptosis switch molecule in cortical neuronal cultures. J. Cell Biol., 2007, 176(6), 853-862.
[http://dx.doi.org/10.1083/jcb.200608022] [PMID: 17353361]
[73]
Lorey, M.B.; Rossi, K.; Eklund, K.K.; Nyman, T.A.; Matikainen, S. Global Characterization of Protein Secretion from Human Macrophages Following Non-canonical Caspase-4/5 Inflammasome Activation. Mol. Cell. Proteomics, 2017, 16(4)(Suppl. 1), S187-S199.
[http://dx.doi.org/10.1074/mcp.M116.064840] [PMID: 28196878]
[74]
Garcia-Caballero, T.; Dominguez, F.; Roson, E.; Gallego, R.; Zalvide, J.; Forteza, J.; Beiras, A. Distribution of prothymosin alpha in rat and human adrenal cortex. Anat. Rec., 1994, 239(1), 88-94.
[http://dx.doi.org/10.1002/ar.1092390110] [PMID: 7913591]
[75]
Roson, E.; Garcia-Caballero, G.; Heimer, E.P.; Felix, A.M.; Dominguez, F. Cellular distribution of prothymosin alpha and parathymosin in rat thymus and spleen. J. Histochem. Cytochem., 1990, 38(12), 1889-1894.
[http://dx.doi.org/10.1177/38.12.2254650] [PMID: 2254650]
[76]
Pawlikowski, M.; Winczyk, K. Immunohistochemical detection of prothymosin alpha in pituitary adenomas--a new marker of tumor recurrence? Folia Histochem. Cytobiol., 2009, 47(4), 559-562.
[PMID: 20430720]
[77]
Pawlikowski, M.; Radek, M.; Kunert-Radek, J.; Jaranowska, M.; Świętosławski, J.; Winczyk, K. Overexpression of prothymosin alpha is related to pituitary adenoma recurrence but not to adenoma invasiveness and proliferation. Endokrynol. Pol., 2014, 65(5), 382-386.
[http://dx.doi.org/10.5603/EP.2014.0053] [PMID: 25301489]
[78]
Klimentzou, P.; Drougou, A.; Fehrenbacher, B.; Schaller, M.; Voelter, W.; Barbatis, C.; Paravatou-Petsotas, M.; Livaniou, E. Immunocytological and preliminary immunohistochemical studies of prothymosin alpha, a human cancer-associated polypeptide, with a well-characterized polyclonal antibody. J. Histochem. Cytochem., 2008, 56(11), 1023-1031.
[http://dx.doi.org/10.1369/jhc.2008.950956] [PMID: 18711212]
[79]
Letsas, K.P.; Frangou-Lazaridis, M.; Skyrlas, A.; Tsatsoulis, A.; Malamou-Mitsi, V. Transcription factor-mediated proliferation and apoptosis in benign and malignant thyroid lesions. Pathol. Int., 2005, 55(11), 694-702.
[http://dx.doi.org/10.1111/j.1440-1827.2005.01899.x] [PMID: 16271081]
[80]
Ferrara, D.; Izzo, G.; Liguori, L.; d'Istria, M.; Aniello, F.; Minucci, S. Evidence for the involvement of prothymosin alpha in the spermatogenesis of the frog Rana esculenta J Exp Zool A Ecol Genet Physiol, 2009, 311, 1-10.
[81]
Pariante, P.; Dotolo, R.; Venditti, M.; Ferrara, D.; Donizetti, A.; Aniello, F.; Minucci, S. Prothymosin alpha expression and localization during the spermatogenesis of Danio rerio. Zygote, 2016, 24(4), 583-593.
[http://dx.doi.org/10.1017/S0967199415000568] [PMID: 26450176]
[82]
Castro, J.M.; Barcia, M.G. Localization of prothymosin alpha in the nucleus. Biochem. Biophys. Res. Commun., 1996, 224(1), 140-146.
[http://dx.doi.org/10.1006/bbrc.1996.0997] [PMID: 8694801]
[83]
Barbini, L.; Gonzalez, R.; Dominguez, F.; Vega, F. Apoptotic and proliferating hepatocytes differ in prothymosin alpha expression and cell localization. Mol. Cell. Biochem., 2006, 291(1-2), 83-91.
[http://dx.doi.org/10.1007/s11010-006-9200-0] [PMID: 16845491]
[84]
Gou, L.T.; Tong, A.P.; Yan, F.; Yuan, Z.; He, F.; Wang, W.; Zhou, Y.; Chen, L.J.; Tang, M.H.; Yang, J.L. Altered protein-expressing profile in hPNAS4-induced apoptosis in A549 human lung adenocarcinoma cells. J. Cell. Biochem., 2009, 108(5), 1211-1219.
[http://dx.doi.org/10.1002/jcb.22353] [PMID: 19795389]
[85]
Franco, F.J.; Diaz, C.; Barcia, M.; Arias, P.; Gomez-Marquez, J.; Soriano, F.; Mendez, E.; Freire, M. Synthesis and apparent secretion of prothymosin alpha by different subpopulations of calf and rat thymocytes. Immunology, 1989, 67(2), 263-268.
[PMID: 2787781]
[86]
Hannappel, E.; van Kampen, M. Determination of thymosin beta 4 in human blood cells and serum. J. Chromatogr. A, 1987, 397, 279-285.
[http://dx.doi.org/10.1016/S0021-9673(01)85010-X] [PMID: 3654821]
[87]
Tan, W.K.Y.; Purnamawati, K.; Pakkiri, L.S.; Tan, S.H.; Yang, X.; Chan, M.Y.; Drum, C.L. Sources of variability in quantifying circulating thymosin beta-4: literature review and recommendations. Expert Opin. Biol. Ther., 2018, 18(sup1), 141-147.
[http://dx.doi.org/10.1080/14712598.2018.1448382] [PMID: 29502471]
[88]
Mandaliti, W.; Nepravishta, R.; Sinibaldi Vallebona, P.; Pica, F.; Garaci, E.; Paci, M. Thymosin α1 Interacts with Exposed Phosphatidylserine in Membrane Models and in Cells and Uses Serum Albumin as a Carrier. Biochemistry, 2016, 55(10), 1462-1472.
[http://dx.doi.org/10.1021/acs.biochem.5b01345] [PMID: 26909491]