Gene Symbol: Tnp2
Description: transition protein 2
Alias: TP2, Tp-2, nuclear transition protein 2, transition protein 2 (alt.)
Species: mouse
Products:     Tnp2

Top Publications

  1. Quenet D, Mark M, Govin J, Van Dorsselear A, Schreiber V, Khochbin S, et al. Parp2 is required for the differentiation of post-meiotic germ cells: identification of a spermatid-specific complex containing Parp1, Parp2, TP2 and HSPA2. Exp Cell Res. 2009;315:2824-34 pubmed publisher
    ..protein-protein interaction assays, we show that Parp2 interacts significantly with both the transition protein TP2 and the transition chaperone HSPA2, whereas Parp1 binds weakly to HSPA2...
  2. Zhao M, Shirley C, Mounsey S, Meistrich M. Nucleoprotein transitions during spermiogenesis in mice with transition nuclear protein Tnp1 and Tnp2 mutations. Biol Reprod. 2004;71:1016-25 pubmed
    ..The transition nuclear proteins, TP1 and TP2, and the protamines, PRM1 and PRM2, are the major nuclear proteins involved in this process...
  3. Keime S, Heitland K, Kumm S, Schlosser M, Hroch N, Holtz W, et al. Characterization of four genes encoding basic proteins of the porcine spermatid nucleus and close linkage of three of them. Biol Chem Hoppe Seyler. 1992;373:261-70 pubmed
    ..In contrast to TNP2 the gene for TNP1 is well conserved among species, indicating a non species-specific function in the chromatin ..
  4. Malla A, Bhandari R. IP6K1 is essential for chromatoid body formation and temporal regulation of Tnp2 and Prm2 expression in mouse spermatids. J Cell Sci. 2017;130:2854-2866 pubmed publisher
    ..juvenile spermatids from Ip6k1-/- mice display premature expression of the transition protein TNP2 and the protamine PRM2 due to translational derepression...
  5. Held T, Barakat A, Mohamed B, Paprotta I, Meinhardt A, Engel W, et al. Heat-shock protein HSPA4 is required for progression of spermatogenesis. Reproduction. 2011;142:133-44 pubmed publisher
    ..These results provide evidence that HSPA4 is required for normal spermatogenesis. ..
  6. Nayernia K, Meinhardt A, Drabent B, Adham I, Muller C, Steckel M, et al. Synergistic effects of germ cell expressed genes on male fertility in mice. Cytogenet Genome Res. 2003;103:314-20 pubmed
    ..Single null mutations in mouse genes encoding four male germ cell proteins, transition protein 2 (Tnp2), proacrosin (Acr), histone H1.1 (H1...
  7. Nayernia K, Drabent B, Adham I, Möschner M, Wolf S, Meinhardt A, et al. Male mice lacking three germ cell expressed genes are fertile. Biol Reprod. 2003;69:1973-8 pubmed
    ..1-/- (70.7% vs. 29.3%) but not with sperm from triple knockout mice Tnp-2-/-/Acr-/-/H1t-/- (53.6% vs. 46.4%). These results are consistent with a model that suggests that some sperm proteins play a role during sperm competition. ..
  8. Meetei A, Ullas K, Rao M. Identification of two novel zinc finger modules and nuclear localization signal in rat spermatidal protein TP2 by site-directed mutagenesis. J Biol Chem. 2000;275:38500-7 pubmed
    Spermatidal protein TP2, which appears transiently during stages 12-16 of mammalian spermiogenesis, is a DNA condensing zinc metalloprotein with a preference to GC-rich DNA...
  9. Schluter G, Celik A, Obata R, Schlicker M, Hofferbert S, Schlung A, et al. Sequence analysis of the conserved protamine gene cluster shows that it contains a fourth expressed gene. Mol Reprod Dev. 1996;43:1-6 pubmed
    ..sequence of the entire human PGC (25 kb in total) and the 3' portion of the rat protamine cluster (PRM2 and TNP2 genes and intergenic region)...

Scientific Experts

More Information


  1. Suganuma R, Yanagimachi R, Meistrich M. Decline in fertility of mouse sperm with abnormal chromatin during epididymal passage as revealed by ICSI. Hum Reprod. 2005;20:3101-8 pubmed
    ..We performed ICSI with testicular, caput, and cauda epididymal sperm from infertile Tnp1-/-Tnp2+/- mutant mice, which have a minimal level of transition nuclear proteins and are sterile by natural mating...
  2. Dai L, Tsai Morris C, Sato H, Villar J, Kang J, Zhang J, et al. Testis-specific miRNA-469 up-regulated in gonadotropin-regulated testicular RNA helicase (GRTH/DDX25)-null mice silences transition protein 2 and protamine 2 messages at sites within coding region: implications of its role in germ cell development. J Biol Chem. 2011;286:44306-18 pubmed publisher
    ..Failure of expression of Transition protein 2 (TP2) and Protamine 2 (Prm2) proteins (chromatin remodelers, essential for spermatid elongation and completion of ..
  3. Martins R, Ostermeier G, Krawetz S. Nuclear matrix interactions at the human protamine domain: a working model of potentiation. J Biol Chem. 2004;279:51862-8 pubmed
    ..nuclear matrix is considered the primary candidate regulating the opening of the multigenic PRM1 --> PRM2 --> TNP2 human protamine domain...
  4. Chioccarelli T, Cacciola G, Altucci L, Lewis S, Simon L, Ricci G, et al. Cannabinoid receptor 1 influences chromatin remodeling in mouse spermatids by affecting content of transition protein 2 mRNA and histone displacement. Endocrinology. 2010;151:5017-29 pubmed publisher
    ..We demonstrated that CNR1 activation regulated chromatin remodeling of spermatids by either increasing Tnp2 levels or enhancing histone displacement...
  5. Burnicka Turek O, Shirneshan K, Paprotta I, Grzmil P, Meinhardt A, Engel W, et al. Inactivation of insulin-like factor 6 disrupts the progression of spermatogenesis at late meiotic prophase. Endocrinology. 2009;150:4348-57 pubmed publisher
    ..These results demonstrate that INSL6 is required for the progression of spermatogenesis. ..
  6. Levesque D, Veilleux S, Caron N, Boissonneault G. Architectural DNA-binding properties of the spermatidal transition proteins 1 and 2. Biochem Biophys Res Commun. 1998;252:602-9 pubmed
    ..The two major transition proteins of the elongating spermatids, namely TP1 and TP2, were expressed and purified using a bacterial expression system...
  7. Zhao M, Shirley C, Yu Y, Mohapatra B, Zhang Y, Unni E, et al. Targeted disruption of the transition protein 2 gene affects sperm chromatin structure and reduces fertility in mice. Mol Cell Biol. 2001;21:7243-55 pubmed
    ..To investigate the role of TP2, we generated mice with a targeted deletion of its gene, Tnp2. Spermatogenesis in Tnp2 null mice was almost normal, with testis weights and epididymal sperm counts being ..
  8. Topaloglu O, Schluter G, Nayernia K, Engel W. A 74-bp promoter of the Tnp2 gene confers testis- and spermatid-specific expression in transgenic mice. Biochem Biophys Res Commun. 2001;289:597-601 pubmed
    ..In this study, the 5' regulatory region of rat Tnp2 gene was investigated by primer extension analysis and transgenic mice study...
  9. Nayernia K, Drabent B, Meinhardt A, Adham I, Schwandt I, Muller C, et al. Triple knockouts reveal gene interactions affecting fertility of male mice. Mol Reprod Dev. 2005;70:406-16 pubmed
    ..of five genes that were expressed in meiotic and haploid spermatogenic cells in mice, transition protein 2 (Tnp2), proacrosin (Acr), histone H1.1 (H1...
  10. Hussain S, Tuorto F, Menon S, Blanco S, Cox C, Flores J, et al. The mouse cytosine-5 RNA methyltransferase NSun2 is a component of the chromatoid body and required for testis differentiation. Mol Cell Biol. 2013;33:1561-70 pubmed publisher
    ..Specific NSun2- and Dnmt2-methylated tRNAs decreased in abundance when both methyltransferases were deleted, suggesting that RNA methylation pathways play an essential role in male germ cell differentiation. ..
  11. Shawlot W, Vazquez Chantada M, Wallingford J, Finnell R. Rfx2 is required for spermatogenesis in the mouse. Genesis. 2015;53:604-611 pubmed publisher
    ..Our results indicate that Rfx2 is not essential for embryonic development in the mouse but is required for spermatogenesis. genesis 53:604-611, 2015. © 2015 Wiley Periodicals, Inc. ..
  12. Shum E, Jones S, Shao A, Dumdie J, Krause M, Chan W, et al. The Antagonistic Gene Paralogs Upf3a and Upf3b Govern Nonsense-Mediated RNA Decay. Cell. 2016;165:382-95 pubmed publisher
    ..Mice conditionally lacking UPF3A exhibit "hyper" NMD and display defects in embryogenesis and gametogenesis. Our results support a model in which UPF3A serves as a molecular rheostat that directs developmental events. ..
  13. Choi Y, Aizawa A, Hecht N. Genomic analysis of the mouse protamine 1, protamine 2, and transition protein 2 gene cluster reveals hypermethylation in expressing cells. Mamm Genome. 1997;8:317-23 pubmed
  14. Zhao M, Shirley C, Hayashi S, Marcon L, Mohapatra B, Suganuma R, et al. Transition nuclear proteins are required for normal chromatin condensation and functional sperm development. Genesis. 2004;38:200-13 pubmed
    ..TPs and their importance for sperm development, we generated mice lacking both TPs, since mice lacking only TP1 or TP2 were fertile. Our results indicated that TP1 and TP2 had partially complemented each other...
  15. Yang R, Tsai Morris C, Kang J, Dufau M. Elucidation of RNA binding regions of gonadotropin-regulated testicular RNA helicase (GRTH/DDX25) to transcripts of a chromatin remodeling protein essential for spermatogenesis. Horm Mol Biol Clin Investig. 2015;22:119-30 pubmed publisher
    ..Transition protein 2 (Tp2), which associates with GRTH and is required for spermatid elongation, failed to express in GRTH null mice with ..
  16. Yu Z, Raabe T, Hecht N. MicroRNA Mirn122a reduces expression of the posttranscriptionally regulated germ cell transition protein 2 (Tnp2) messenger RNA (mRNA) by mRNA cleavage. Biol Reprod. 2005;73:427-33 pubmed
    ..One highly conserved sequence in the 3'-untranslated region (UTR) of transition protein 2 (Tnp2) mRNA, a testis-specific and posttranscriptionally regulated mRNA in postmeiotic germ cells, is complementary to ..
  17. Zhong J, Peters A, Lee K, Braun R. A double-stranded RNA binding protein required for activation of repressed messages in mammalian germ cells. Nat Genet. 1999;22:171-4 pubmed
    ..The timing of Prbp expression suggests that it may function as a chaperone in the assembly of specific translationally regulated ribonucleoprotein particles. ..
  18. Adham I, Nayernia K, Burkhardt Göttges E, Topaloglu O -, Dixkens C, Holstein A, et al. Teratozoospermia in mice lacking the transition protein 2 (Tnp2). Mol Hum Reprod. 2001;7:513-20 pubmed
    It is believed that the transition proteins (Tnp1 and Tnp2) participate in the removal of the nucleohistones and in the initial condensation of the spermatid nucleus...
  19. Kleene K, Flynn J. Characterization of a cDNA clone encoding a basic protein, TP2, involved in chromatin condensation during spermiogenesis in the mouse. J Biol Chem. 1987;262:17272-7 pubmed
    ..homologue of a protein involved in the initial phases of condensation of chromatin during spermiogenesis in rats, TP2, based on similarities in the sequence of the carboxyl terminus, composition, molecular weight, and electrophoretic ..
  20. Engel W, Keime S, Kremling H, Hameister H, Schluter G. The genes for protamine 1 and 2 (PRM1 and PRM2) and transition protein 2 (TNP2) are closely linked in the mammalian genome. Cytogenet Cell Genet. 1992;61:158-9 pubmed
    The genes for two protamines (PRM1 and PRM2) and for two transition proteins (TNP1 and TNP2) have been characterized in several mammalian species...
  21. Tseden K, Topaloglu O, Meinhardt A, Dev A, Adham I, Muller C, et al. Premature translation of transition protein 2 mRNA causes sperm abnormalities and male infertility. Mol Reprod Dev. 2007;74:273-9 pubmed
    ..It is believed that transition protein 2 (Tnp2) is necessary for maintaining the normal processing of protamines and, consequently, the completion of chromatin ..
  22. Shirley C, Hayashi S, Mounsey S, Yanagimachi R, Meistrich M. Abnormalities and reduced reproductive potential of sperm from Tnp1- and Tnp2-null double mutant mice. Biol Reprod. 2004;71:1220-9 pubmed
    ..Epididymal sperm from the nine possible Tnp1 and Tnp2 null genotypes demonstrated a general decrease in normal morphology, motility, chromatin condensation, and degree ..
  23. Xing W, Krishnamurthy H, Sairam M. Role of follitropin receptor signaling in nuclear protein transitions and chromatin condensation during spermatogenesis. Biochem Biophys Res Commun. 2003;312:697-701 pubmed
    ..In the adult, absence of FSHR signaling prolongs retention of mono-uH2A, leading to impair transition of basic nucleoproteins and chromatin remodeling during mouse spermatogenesis. ..
  24. Barral S, Morozumi Y, Tanaka H, Montellier E, Govin J, de Dieuleveult M, et al. Histone Variant H2A.L.2 Guides Transition Protein-Dependent Protamine Assembly in Male Germ Cells. Mol Cell. 2017;66:89-101.e8 pubmed publisher
    ..Hence, the current view of histone-to-nucleoprotamine transition should be revisited and include an additional step with H2A.L.2 assembly prior to the action of TPs and Prms. ..
  25. Urano A, Endoh M, Wada T, Morikawa Y, Itoh M, Kataoka Y, et al. Infertility with defective spermiogenesis in mice lacking AF5q31, the target of chromosomal translocation in human infant leukemia. Mol Cell Biol. 2005;25:6834-45 pubmed
    ..These results may have clinical implications in the understanding of human male infertility. ..
  26. Vourekas A, Zheng K, Fu Q, Maragkakis M, Alexiou P, Ma J, et al. The RNA helicase MOV10L1 binds piRNA precursors to initiate piRNA processing. Genes Dev. 2015;29:617-29 pubmed publisher
    ..Our results support a model in which MOV10L1 RNA helicase activity promotes unwinding and funneling of the single-stranded piRNA precursor transcripts to the endonuclease that catalyzes the first cleavage step of piRNA processing. ..
  27. Cantù C, Valenta T, Hausmann G, Vilain N, Aguet M, Basler K. The Pygo2-H3K4me2/3 interaction is dispensable for mouse development and Wnt signaling-dependent transcription. Development. 2013;140:2377-86 pubmed publisher
  28. Kleene K, Gerstel J, Shih D. Nucleotide sequence of the gene encoding mouse transition protein 2. Gene. 1990;95:301-2 pubmed
    ..Since the genes encoding protamines 1 and 2 and transition protein 1 in mammals contain a single intron in the same position, protamines and transition proteins appear to be evolutionarily related. ..
  29. Pradeepa M, Manjunatha S, Sathish V, Agrawal S, Rao M. Involvement of importin-4 in the transport of transition protein 2 into the spermatid nucleus. Mol Cell Biol. 2008;28:4331-41 pubmed a unique chromatin-remodeling process in which histones are replaced by transition protein 1 (TP1), TP2, and TP4, which are further replaced by protamines...
  30. Nayernia K, Vauti F, Meinhardt A, Cadenas C, Schweyer S, Meyer B, et al. Inactivation of a testis-specific Lis1 transcript in mice prevents spermatid differentiation and causes male infertility. J Biol Chem. 2003;278:48377-85 pubmed
    ..These results provide evidence for an essential and hitherto uncharacterized role of the Lis1 protein in spermatogenesis, particularly in the differentiation of spermatids into spermatozoa. ..
  31. Nelson J, Krawetz S. Linkage of human spermatid-specific basic nuclear protein genes. Definition and evolution of the P1-->P2-->TP2 locus. J Biol Chem. 1993;268:2932-6 pubmed
    ..P1 gene to isolate a human cosmid clone which contains the genes for protamines P1 and P2 and transition protein TP2. Sequence and hybridization analysis of this cosmid showed that the TP2 gene resided 3' of the P2 gene at a ..
  32. Moore C, Lee J, Birren B, Stetten G, Baxter L, Reeves R. Integration of cytogenetic with recombinational and physical maps of mouse chromosome 16. Genomics. 1999;59:1-5 pubmed
    ..A direct application of these probes is demonstrated by identifying mice trisomic for distal Chr 16 using FISH analysis of interphase nuclei. ..