Affinage

TNP2

Nuclear transition protein 2 · UniProt Q05952

Length
138 aa
Mass
15.6 kDa
Annotated
2026-04-28
54 papers in source corpus 24 papers cited in narrative 23 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

TNP2 is a testis-specific, zinc-containing basic nuclear protein that transiently replaces histones during spermatid chromatin condensation, serving as an essential intermediate in the histone-to-protamine transition of spermiogenesis. Its N-terminal domain harbors two novel zinc finger modules (coordinating 4 His and 4 Cys) that confer zinc-dependent, GC-rich and unmethylated CpG island DNA recognition and condensation, while its C-terminal basic domain mediates DNA binding and contains a nuclear/nucleolar localization signal; both domains cooperate for full condensation activity (PMID:7711033, PMID:8961924, PMID:10961985). TNP2 function is regulated by a cascade of post-translational modifications: Cs-PKA phosphorylation of Ser109/Thr101 promotes nuclear import while transiently reducing condensation activity, p300 acetylation of C-terminal lysines further attenuates condensation and disrupts interaction with the chaperone NPM3, and PRMT4/KMT7-mediated methylation marks chromatin-bound TP2 in elongating spermatids (PMID:11772016, PMID:14514679, PMID:19710011, PMID:25818198). Timely removal of TNP2 is accomplished by the ASB9-ELOB/C-CUL5-RBX1 ubiquitin ligase complex, which ubiquitinates TNP2 for proteasomal degradation to permit protamine deposition; loss of this degradation pathway causes TNP2 retention, defective chromatin remodeling, and male infertility in mice and humans (PMID:41915740, PMID:11385107).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 1987 High

    Identification of TNP2 as a haploid spermatid-specific basic nuclear protein with a distinctive C-terminal basic DNA-binding domain established the molecular identity and domain architecture of this chromatin transition factor.

    Evidence cDNA cloning, Northern/Southern blotting, and complete amino acid sequencing of the C-terminal peptide from mouse and rat testis

    PMID:3307778 PMID:3693351

    Open questions at the time
    • No functional assay for DNA condensation yet performed
    • Zinc binding not yet recognized
    • Regulatory modifications unknown
  2. 1990 High

    Demonstration that TP2 binds DNA with defined affinity constants and condenses DNA more effectively than TP1 established it as the primary condensation factor among transition proteins.

    Evidence Fluorescence quenching, thermal denaturation, and circular dichroism with purified TP2 and various nucleic acid substrates in vitro

    PMID:2250010

    Open questions at the time
    • Sequence specificity of condensation not yet defined
    • In vivo relevance of superior condensation ability unconfirmed
  3. 1995 High

    Discovery that TP2 is a zinc metalloprotein with two novel zinc finger modules that confer zinc-dependent preferential condensation of GC-rich DNA revealed a previously unknown mechanism for sequence-selective chromatin packaging in spermatids.

    Evidence Atomic absorption spectroscopy, 65Zn-blotting, site-directed mutagenesis, and circular dichroism with domain fragments and various polynucleotide substrates

    PMID:1930189 PMID:7711033 PMID:8076694

    Open questions at the time
    • Structural basis of zinc finger modules not resolved at atomic resolution
    • In vivo GC-rich preference not yet demonstrated
  4. 1996 High

    Showing that TP2 recognizes unmethylated CpG islands in a zinc-dependent, major-groove-reading manner and that CpG methylation abolishes this interaction revealed a mechanism linking DNA methylation status to chromatin packaging selectivity during spermiogenesis.

    Evidence Gel mobility shift assays with chelator inhibition, SssI methylase treatment, DMS methylation, and chromomycin A3 groove-competition

    PMID:8961924

    Open questions at the time
    • Biological significance of CpG selectivity for sperm epigenome not tested in vivo
    • Whether TP2 CpG recognition protects specific genomic regions is unknown
  5. 1996 High

    Quantitative immunohistochemistry established the precise temporal window of TP2 nuclear occupancy (steps 11–16) and its slightly earlier disappearance compared to TP1, defining the sequential chromatin remodeling program.

    Evidence Immunoperoxidase and immunogold localization on staged rat testis sections

    PMID:8722637

    Open questions at the time
    • Mechanism controlling differential TP1/TP2 removal kinetics unknown
    • Whether TP2 departure is a prerequisite for TP1 departure not tested
  6. 2000 High

    Mutagenesis-based identification of two novel zinc finger modules (4 His + 4 Cys) and the NLS/NoLS within the C-terminal domain defined the functional architecture of TP2 at single-residue resolution.

    Evidence Site-directed mutagenesis, 65Zn-blotting of mutants, and GFP-tagged TP2 localization in transfected COS-7 cells

    PMID:10961985

    Open questions at the time
    • COS-7 is a somatic cell; NLS function not confirmed in spermatids
    • No crystal structure of zinc finger modules
  7. 2001 High

    Tnp2 knockout mice revealed that TP2 is essential for normal acrosome attachment and sperm morphology, with background-dependent infertility and compensatory Tnp1 upregulation, demonstrating non-redundant roles of the two transition proteins.

    Evidence Homologous recombination knockout in mice, electron microscopy, fertility testing on 129/Sv and mixed backgrounds, Northern blot

    PMID:11385107

    Open questions at the time
    • Molecular basis of acrosome detachment not defined
    • Mechanism of background-dependent fertility rescue unclear
  8. 2003 High

    Identification of Cs-PKA (Cα2) as the physiological kinase phosphorylating TP2 at Ser109/Thr101, with phosphorylation promoting nuclear import into spermatid nuclei, established a regulatory switch coupling TP2 modification to its nuclear entry.

    Evidence In vitro phosphorylation with spermatid cytosol, RT-PCR for Cs-PKA, reconstituted nuclear transport assay in permeabilized round spermatids

    PMID:11772016 PMID:14514679

    Open questions at the time
    • In vivo confirmation by phospho-site knock-in mutations not performed
    • Phosphatase responsible for dephosphorylation after nuclear entry unknown
  9. 2004 High

    Systematic analysis of single and double TP-null mice demonstrated that absence of one TP causes post-translational retention (not increased synthesis) of the other, and that combined loss causes gene-dosage-dependent defects in chromatin condensation and protamine 2 processing.

    Evidence Immunohistochemistry across 9 Tnp1/Tnp2 genotype combinations, epididymal sperm analysis, ICSI

    PMID:15163613 PMID:15189834

    Open questions at the time
    • Mechanism of post-translational retention not identified
    • Whether TP removal requires a common degradation pathway not established
  10. 2005 High

    Discovery that miR-122a directly cleaves Tnp2 mRNA through its 3′-UTR via mRNA degradation (not translational repression) revealed a post-transcriptional layer limiting TP2 expression levels.

    Evidence Luciferase reporter with Tnp2 3′-UTR, miR-122a site mutagenesis, real-time RT-PCR, ribonuclease protection, and polysome fractionation

    PMID:15901636

    Open questions at the time
    • In vivo consequence of miR-122a loss on TP2 levels in spermatids not tested
    • Other miRNAs targeting Tnp2 not surveyed
  11. 2009 High

    Identification of p300/KAT3B-mediated acetylation at four C-terminal lysines that reduces DNA condensation and disrupts NPM3 interaction established acetylation as a second PTM switch regulating TP2 functional state on chromatin.

    Evidence Mass spectrometry, in vitro acetylation, circular dichroism, atomic force microscopy, and co-immunoprecipitation of TP2-NPM3

    PMID:19710011

    Open questions at the time
    • Temporal relationship between phosphorylation and acetylation in vivo not resolved
    • Deacetylase responsible for reversing acetylation not identified
  12. 2009 Medium

    Discovery that Parp2 complexes with TP2 and HSPA2 in spermatids, and that Parp2 loss causes TP2-expressing spermatid death, linked poly(ADP-ribosyl)ation to TP2 chromatin function and spermatid survival.

    Evidence In vitro protein-protein interaction, immunohistochemistry, and electron microscopy of Parp2-deficient mouse testes

    PMID:19607827

    Open questions at the time
    • Whether Parp2 directly PARylates TP2 in vivo not confirmed
    • Mechanism by which Parp2 loss triggers spermatid death not delineated
    • Single-lab finding awaiting independent confirmation
  13. 2015 High

    Mapping 19 novel PTMs and identifying PRMT4- and KMT7-mediated methylation on chromatin-bound TP2 in elongating spermatids revealed a combinatorial PTM code governing TP2 during chromatin remodeling.

    Evidence Mass spectrometry of endogenous TP2, in vitro methylation with recombinant enzymes, site mutagenesis, and modification-specific antibody immunohistochemistry

    PMID:25818198

    Open questions at the time
    • Functional consequence of individual methylation marks on condensation or degradation not tested
    • Reader proteins for TP2 methylation marks unknown
  14. 2017 High

    Demonstrating that IP6K1 maintains chromatoid body integrity to temporally repress Tnp2 translation, with Ip6k1 loss causing premature TP2 appearance and azoospermia, established the chromatoid body as a translational timing checkpoint for TP2.

    Evidence IP6K1 immunolocalization, chromatoid body analysis in Ip6k1-null mice, Western blot and RT-PCR for premature Tnp2/Prm2 expression

    PMID:28743739

    Open questions at the time
    • Molecular mechanism by which IP6K1 kinase activity maintains chromatoid body unknown
    • Whether IP6K1 acts directly on Tnp2 mRNA or through an intermediary not resolved
  15. 2026 High

    Reconstitution of the ASB9-ELOB/C-CUL5-RBX1 ubiquitin ligase complex that ubiquitinates TNP2 for proteasomal degradation, validated in Asb9-knockout mice and infertile human patients, solved the long-standing question of how TNP2 is removed to permit protamine deposition.

    Evidence Co-immunoprecipitation of CRL complex, in vitro ubiquitination assay, Asb9-knockout mouse phenotyping, and human patient analysis

    PMID:41915740

    Open questions at the time
    • Specific ubiquitination sites on TNP2 not mapped
    • Whether PTMs (acetylation, methylation) regulate ASB9 recognition of TNP2 not tested
    • Proteasomal versus non-proteasomal degradation pathways not distinguished in vivo

Open questions

Synthesis pass · forward-looking unresolved questions
  • A unified structural and temporal model integrating all PTM inputs (phosphorylation, acetylation, methylation, PARylation, ubiquitination) into a sequential regulatory cascade governing TP2 nuclear import, chromatin binding, and degradation has not been established.
  • No atomic-resolution structure of TP2 or its zinc fingers exists
  • Temporal ordering and interdependence of multiple PTMs in vivo is unknown
  • Whether CpG-selective condensation protects specific genomic loci in mature sperm remains untested

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003677 DNA binding 4 GO:0008289 lipid binding 3
Localization
GO:0005634 nucleus 4 GO:0005694 chromosome 3 GO:0005730 nucleolus 1
Pathway
R-HSA-392499 Metabolism of proteins 4 R-HSA-4839726 Chromatin organization 4 R-HSA-1474165 Reproduction 2
Partners
ASB9EP300HSPA2KMT7NPM3PARP2PRKACAPRMT4
Complex memberships
ASB9-ELOB/C-CUL5-RBX1 ubiquitin ligase complexTP2-PARP2-HSPA2 spermatid complex

Evidence

Reading pass · 23 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1987 TNP2 (TP2) encodes a small cysteine- and serine-rich basic nuclear protein with a highly basic C-terminal domain comprising ~one-third of the polypeptide and a less basic N-terminal two-thirds; a single copy gene in the mouse genome expressed specifically in haploid spermatids at high levels. cDNA cloning, Northern blot, Southern blot, amino acid sequence analysis The Journal of biological chemistry High 3693351
1987 TP2's highly basic carboxyl-terminal domain (27 residues) contains most of the basic residues and is the primary site of DNA binding, distinguishing it from core histones whose basic domain is at the amino terminus. HPLC purification, V8 protease cleavage, complete amino acid sequencing of C-terminal peptide Biochemical and biophysical research communications High 3307778
1990 TP2 binds double-stranded DNA, single-stranded DNA, and poly(rA) with defined association constants; it stabilizes DNA (raises Tm) and has superior DNA-condensing ability compared to TP1 in vitro, while unlike TP1 it does not destabilize nucleosome core particle compactness. Fluorescence quenching, thermal denaturation, circular dichroism spectroscopy, in vitro nucleic acid binding assays The Journal of biological chemistry High 2250010
1991 TP2 is a zinc metalloprotein containing two potential zinc finger motifs involving cysteine and histidine residues; native TP2 from rat elongating spermatids binds ~0.2 atoms Zn/molecule and binds 2 atoms Zn/molecule after in vitro zinc loading; cysteine residues are involved in zinc coordination. Atomic absorption spectroscopy, iodoacetamidofluorescein labeling in situ, amino acid sequence analysis Biochemical and biophysical research communications High 1930189
1992 The human PRM1, PRM2, and TNP2 genes are closely linked within a ~13–15 kb region of chromosomal DNA (16p13.13–16p13.2), with the order P1→P2→TNP2, while TNP1 maps to a different chromosome. Cosmid cloning, Southern blotting, hybridization analysis, chromosomal mapping Cytogenetics and cell genetics High 1395729 8428967
1994 Zinc binding in TP2 localizes to the N-terminal two-thirds (zinc-binding domain) involving both cysteine and histidine residues, as demonstrated by 65Zn-blotting and chemical modification; the N-terminal fragment adopts a type I beta-turn structure while the C-terminal fragment has alpha-helical character. 65Zn-blotting, Hg-affinity chromatography purification, V8 protease fragmentation, chemical modification (iodoacetic acid, diethylpyrocarbonate), circular dichroism spectroscopy FEBS letters High 8076694
1995 TP2 condenses GC-rich DNA (alternating poly(dG-dC)) preferentially and in a zinc-dependent manner; the intact protein is required for full GC-preference condensation, with neither the N-terminal zinc-binding domain nor the C-terminal basic domain alone being as effective as the full-length protein. Circular dichroism spectroscopy, EDTA chelation, V8 protease domain fragmentation, in vitro DNA condensation assays with various polynucleotides Biochemistry High 7711033
1996 TP2 recognizes a human CpG island sequence in a zinc-dependent manner; EDTA or 1,10-o-phenanthroline pretreatment abolishes complex formation; CpG methylation by SssI methylase completely abolishes TP2-CpG island interaction, whereas N-7 guanine methylation does not, indicating TP2 reads the major groove of unmethylated CpG sequences. Gel mobility shift assay, EDTA/chelator inhibition, competition with various polynucleotides, chromomycin A3 groove-binding drug competition, CpG methylation by SssI methylase, DMS methylation Biochemistry High 8961924
1996 In rat spermatids, TP1 and TP2 appear in the nucleus at step 11 coincident with chromatin condensation initiation, increase progressively through steps 11–13, then decline (TP2 disappearing by step 16, TP1 by step 17), demonstrating sequential chromatin remodeling with TP2 slightly preceding TP1 exit. Immunoperoxidase and immunogold localization, quantitative immunohistochemistry on rat testis sections Biology of reproduction High 8722637
2000 Site-directed mutagenesis identified two novel zinc finger modules in TP2 involving 4 histidine and 4 cysteine residues in a configuration defining a new class of zinc finger. The nuclear localization signal was identified as residues 87–95 (GKVSKRKAV) within the C-terminal domain, functioning as part of an extended nucleolar localization sequence (NoLS); TP2 preferentially localizes to the nucleolus in COS-7 cells. Site-directed mutagenesis, 65Zn-blotting of mutants, transfection of COS-7 cells with wild-type and mutant TP2-GFP constructs, immunofluorescence localization The Journal of biological chemistry High 10961985
2002 TP2 is phosphorylated by protein kinase A (PKA) at Ser109 and Thr101 in its C-terminal domain; phosphorylation of TP2 greatly reduces its DNA condensation property; TP2 complexed with DNA is a poor substrate for PKA; dephosphorylation by alkaline phosphatase restores DNA condensation activity. In vitro phosphorylation with testicular salt extracts and purified PKA, site-specific mutagenesis of phosphorylation sites, PKA inhibitor peptide (PKI) inhibition, phorbol ester/cGMP controls, circular dichroism DNA condensation assay, calf intestinal alkaline phosphatase dephosphorylation, in vivo phosphorylation demonstration Biochemistry High 11772016
2001 Disruption of Tnp2 in mice causes teratozoospermia with acrosome detachment from the nuclear envelope and reduced sperm motility; infertility is background-dependent (infertile on 129/Sv, fertile on mixed background); increased Tnp1 transcript in Tnp2-null testes suggests compensatory upregulation. Homologous recombination gene knockout, light and electron microscopy, fertility testing on different genetic backgrounds, Northern blot for Tnp1 transcript Molecular human reproduction High 11385107
2003 TP2 is phosphorylated by the sperm-specific PKA catalytic subunit (Cs-PKA, Cα2) at Thr101 and Ser109 in haploid spermatid cytosol; phosphorylation positively modulates NLS-dependent nuclear import of TP2 into haploid round spermatid nuclei, as demonstrated by an in vitro nuclear transport assay requiring cytoplasmic factors and ATP. In vitro phosphorylation with spermatid cytosol, RT-PCR identification of Cs-PKA, Western blot with anti-Cα1 antibodies, in vitro nuclear transport assay in permeabilized round spermatids, phosphorylation-transport correlation The Journal of biological chemistry High 14514679
2004 In Tnp1 or Tnp2 single-null mice, absence of one transition protein leads to abnormal retention of the other TP (prolonged nuclear residency) rather than increased synthesis; the elevated TP level in mutant mice is a posttranslational consequence. TPs appear in nuclei before histone displacement is complete (overlapping phases), and the absence of one TP does not affect the time of appearance or protamine expression but does affect displacement of the remaining TP. Immunohistochemistry on testis sections from Tnp1, Tnp2, and double mutant mice; comparison of protein distribution with previous biochemical analyses Biology of reproduction High 15163613
2004 Sperm from Tnp1/Tnp2 double-null mice show a gene dosage–dependent decrease in normal morphology, motility, chromatin condensation, and protamine 2 processing; some double-null sperm nuclei undergo DNA degradation during epididymal transport; both TPs show partial functional redundancy but each fulfills unique roles. Epididymal sperm analysis from 9 Tnp1/Tnp2 genotypic combinations, intracytoplasmic sperm injection (ICSI), hematoxylin/DNA fluorochrome staining, motility and morphology assessment Biology of reproduction High 15189834
2005 Mirn122a (miR-122a) directly cleaves Tnp2 mRNA via a conserved complementary site in the Tnp2 3'-UTR; base pairing of the 5'-region of Mirn122a to this site is essential for repression; the mechanism involves mRNA cleavage rather than translational inhibition. Luciferase reporter assay with Tnp2 3'-UTR, site-directed mutagenesis of Mirn122a, real-time RT-PCR, ribonuclease protection assay, polysome fractionation Biology of reproduction High 15901636
2009 TP2 is acetylated in vivo; the acetyltransferase KAT3B (p300) acetylates TP2 at four C-terminal lysine residues in vitro and in vivo; acetylation significantly reduces TP2's DNA condensation property and impedes TP2's interaction with histone chaperone NPM3. Anti-acetylated lysine immunoprecipitation, mass spectrometry, in vitro acetylation assays with p300 and PCAF, circular dichroism, atomic force microscopy, co-immunoprecipitation of TP2-NPM3 The Journal of biological chemistry High 19710011
2009 Parp2 interacts with TP2 and the transition chaperone HSPA2 in spermatids; Parp2-TP2 interaction is partly mediated by poly(ADP-ribosyl)ation; loss of Parp2 results in loss of TP2-expressing spermatids, defective chromatin condensation, and abnormal manchette microtubules, causing spermatid-specific cell death. In vitro protein-protein interaction assays, immunohistochemistry, electron microscopy of Parp2-deficient mouse testes Experimental cell research Medium 19607827
2009 In rat condensing spermatids, TP2 co-localizes preferentially with GC-rich DNA (marked by chromomycin A3 and 7-amino actinomycin D) and moves with GC-rich sequences toward the nuclear periphery during spermatid maturation; TP1 and TP2 show overlapping localization foci, suggesting concerted functional roles. Immunofluorescence co-localization with GC-selective (chromomycin A3, 7-AAD) and AT-selective (DAPI) dyes, combined immunofluorescence with anti-TP1 and anti-TP2 antibodies in rat spermatids The journal of histochemistry and cytochemistry Medium 19506090
2015 TP2 carries 19 novel post-translational modifications identified by mass spectrometry; PRMT4 (CARM1) methylates TP2 at Arg71, Arg75, and Arg92; KMT7 (Set9) methylates TP2 at Lys88 and Lys91; modification-specific antibodies show that TP2-K88me1 and TP2-R92me1 appear in elongating-to-condensing spermatids and are predominantly associated with chromatin-bound TP2. Mass spectrometry of endogenous TP2, in vitro methylation assays with PRMT4 and KMT7, site-directed mutagenesis of target residues, modification-specific antibody generation and immunohistochemistry The Journal of biological chemistry High 25818198
2017 IP6K1 is a component of the chromatoid body in round spermatids; deletion of Ip6k1 causes absence of the chromatoid body and premature translational derepression of Tnp2 and Prm2 in juvenile spermatids, resulting in abnormal spermatid elongation and azoospermia. IP6K1 immunolocalization in mouse testis, chromatoid body immunofluorescence in Ip6k1-null mice, Western blot and RT-PCR for Tnp2/Prm2 expression in mutant spermatids, histological analysis of spermatid differentiation Journal of cell science High 28743739
2001 A 74-bp promoter region (−74 to +73) of the rat Tnp2 gene is sufficient to confer testis- and spermatid-specific expression in transgenic mice; primer extension analysis identified a transcription start site 70 bp upstream of the translation start codon. Primer extension, transgenic mouse reporter assay with truncated Tnp2 promoter constructs driving a reporter gene Biochemical and biophysical research communications Medium 11716517
2026 ASB9 mediates ubiquitin-dependent proteasomal degradation of TNP2 by assembling a testis-specific Cullin-RING ligase (CRL) complex comprising TNP2-ASB9-ELOB/C-CUL5-RBX1; ASB9 deficiency causes TNP2 retention and failure of the histone-to-protamine transition, resulting in sperm head malformation and male infertility in both mice and humans. Co-immunoprecipitation to identify ASB9-TNP2-ELOB/C-CUL5-RBX1 complex, in vitro ubiquitination assay, Asb9-knockout mouse model with histological and fertility phenotyping, analysis of infertile human patients with ASB9 loss-of-function Proceedings of the National Academy of Sciences of the United States of America High 41915740

Source papers

Stage 0 corpus · 54 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2005 MicroRNA Mirn122a reduces expression of the posttranscriptionally regulated germ cell transition protein 2 (Tnp2) messenger RNA (mRNA) by mRNA cleavage. Biology of reproduction 185 15901636
2004 Abnormalities and reduced reproductive potential of sperm from Tnp1- and Tnp2-null double mutant mice. Biology of reproduction 129 15189834
2001 Teratozoospermia in mice lacking the transition protein 2 (Tnp2). Molecular human reproduction 115 11385107
2004 Nucleoprotein transitions during spermiogenesis in mice with transition nuclear protein Tnp1 and Tnp2 mutations. Biology of reproduction 95 15163613
1996 Chromatin reorganization in rat spermatids during the disappearance of testis-specific histone, H1t, and the appearance of transition proteins TP1 and TP2. Biology of reproduction 94 8722637
1987 Characterization of a cDNA clone encoding a basic protein, TP2, involved in chromatin condensation during spermiogenesis in the mouse. The Journal of biological chemistry 82 3693351
2005 Single-nucleotide polymorphisms and mutation analyses of the TNP1 and TNP2 genes of fertile and infertile human male populations. Journal of andrology 54 16291974
1995 Coordinate expression of the PRM1, PRM2, and TNP2 multigene locus in human testis. DNA and cell biology 54 7865133
2013 MicroRNA-122 influences the development of sperm abnormalities from human induced pluripotent stem cells by regulating TNP2 expression. Stem cells and development 47 23327642
1990 Interaction of spermatid-specific protein TP2 with nucleic acids, in vitro. A comparative study with TP1. The Journal of biological chemistry 46 2250010
1991 Mammalian spermatid specific protein, TP2, is a zinc metalloprotein with two finger motifs. Biochemical and biophysical research communications 40 1930189
1992 The genes for protamine 1 and 2 (PRM1 and PRM2) and transition protein 2 (TNP2) are closely linked in the mammalian genome. Cytogenetics and cell genetics 37 1395729
1993 Linkage of human spermatid-specific basic nuclear protein genes. Definition and evolution of the P1-->P2-->TP2 locus. The Journal of biological chemistry 36 8428967
1996 Zinc dependent recognition of a human CpG island sequence by the mammalian spermatidal protein TP2. Biochemistry 30 8961924
1991 Characterization of a gene encoding a basic protein of the spermatid nucleus, TNP2, and its close linkage to the protamine genes in the bull. Biological chemistry Hoppe-Seyler 30 1716912
2000 The rice Rim2 transcript accumulates in response to Magnaporthe grisea and its predicted protein product shares similarity with TNP2-like proteins encoded by CACTA transposons. Molecular & general genetics : MGG 28 11016827
2017 IP6K1 is essential for chromatoid body formation and temporal regulation of Tnp2 and Prm2 expression in mouse spermatids. Journal of cell science 27 28743739
2014 Association of TNP2 gene polymorphisms of the bta-miR-154 target site with the semen quality traits of Chinese Holstein bulls. PloS one 27 24416221
2002 Involvement of protein kinase A in the phosphorylation of spermatidal protein TP2 and its effect on DNA condensation. Biochemistry 26 11772016
1995 DNA condensation by the rat spermatidal protein TP2 shows GC-rich sequence preference and is zinc dependent. Biochemistry 26 7711033
1994 Establishment of a partially informative porcine somatic cell hybrid panel and assignment of the loci for transition protein 2 (TNP2) and protamine 1 (PRM1) to chromosome 3 and polyubiquitin (UBC) to chromosome 14. Genomics 23 7959732
2015 Mapping of Post-translational Modifications of Transition Proteins, TP1 and TP2, and Identification of Protein Arginine Methyltransferase 4 and Lysine Methyltransferase 7 as Methyltransferase for TP2. The Journal of biological chemistry 22 25818198
2000 Identification of two novel zinc finger modules and nuclear localization signal in rat spermatidal protein TP2 by site-directed mutagenesis. The Journal of biological chemistry 22 10961985
1991 Chromosomal assignment of four rat genes coding for the spermatid-specific proteins proacrosin (ACR), transition proteins 1 (TNP1) and 2 (TNP2), and protamine 1 (PRM1). Cytogenetics and cell genetics 22 1906796
1998 Extended analysis of the region encompassing the PRM1-->PRM2-->TNP2 domain: genomic organization, evolution and gene identification. The Journal of experimental zoology 19 9723181
1987 Nuclear transition protein 2 (TP2) of mammalian spermatids has a very basic carboxyl terminal domain. Biochemical and biophysical research communications 19 3307778
2009 Parp2 is required for the differentiation of post-meiotic germ cells: identification of a spermatid-specific complex containing Parp1, Parp2, TP2 and HSPA2. Experimental cell research 18 19607827
2003 Phosphorylation of rat spermatidal protein TP2 by sperm-specific protein kinase A and modulation of its transport into the haploid nucleus. The Journal of biological chemistry 18 14514679
2009 Acetylation of transition protein 2 (TP2) by KAT3B (p300) alters its DNA condensation property and interaction with putative histone chaperone NPM3. The Journal of biological chemistry 17 19710011
2001 Sperm nuclear matrix association of the PRM1-->PRM2-->TNP2 domain is independent of Alu methylation. Molecular human reproduction 17 11574659
2012 Association study of six SNPs in PRM1, PRM2 and TNP2 genes in iranian infertile men with idiopathic azoospermia. Iranian journal of reproductive medicine 15 25246894
2003 Conservation of the PRM1 --> PRM2 --> TNP2 domain. DNA sequence : the journal of DNA sequencing and mapping 14 14756422
2009 Spatiotemporal organization of AT- and GC-rich DNA and their association with transition proteins TP1 and TP2 in rat condensing spermatids. The journal of histochemistry and cytochemistry : official journal of the Histochemistry Society 13 19506090
1994 Localization of the IGHG, PRKACB, and TNP2 genes in pigs by in situ hybridization. Mammalian genome : official journal of the International Mammalian Genome Society 13 8012108
1992 The nucleotide sequence of boar transition protein 2 (TNP2) cDNA and haploid expression of the gene during spermatogenesis. Animal genetics 13 1380212
1995 Mapping the clonally unstable recombinogenic PRM1-->PRM2-->TNP2 region of human 16p13.2. DNA sequence : the journal of DNA sequencing and mapping 12 7612927
1994 Characterization of the zinc-metalloprotein nature of rat spermatidal protein TP2. FEBS letters 12 8076694
2022 How well does molecular simulation reproduce environment-specific conformations of the intrinsically disordered peptides PLP, TP2 and ONEG? Chemical science 11 35308859
2022 Investigations on the Wound Healing Potential of Tilapia Piscidin (TP)2-5 and TP2-6. Marine drugs 11 35323503
1996 Cloning of cDNA encoding rat spermatidal protein TP2 and expression in Escherichia coli. Protein expression and purification 10 8954887
2008 Comparative genomics reveals gene-specific and shared regulatory sequences in the spermatid-expressed mammalian Odf1, Prm1, Prm2, Tnp1, and Tnp2 genes. Genomics 9 18562159
2019 The association between TNP2 gene polymorphisms and Iranian infertile men with varicocele: A case-control study. International journal of reproductive biomedicine 7 31583373
1998 Hyperexpression of rat spermatidal protein TP2 in Escherichia coli by codon optimization and engineering the vector-encoded 5' UTR. Protein expression and purification 6 9675061
2008 Coe1 in Beta vulgaris L. Has a Tnp2-Domain DNA Transposase Gene within Putative LTRs and Other Retroelement-Like Features. International journal of plant genomics 5 18566682
2001 A 74-bp promoter of the Tnp2 gene confers testis- and spermatid-specific expression in transgenic mice. Biochemical and biophysical research communications 5 11716517
2022 Correlation of Novel Single Nucleotide Polymorphisms ofUSP26, TEX15, and TNP2 Genes with Male Infertility in North West of Iran. International journal of fertility & sterility 4 35103426
2022 Sequence Diversity of Tp1 and Tp2 Antigens and Population Genetic Analysis of Theileria parva in Unvaccinated Cattle in Zambia's Chongwe and Chisamba Districts. Pathogens (Basel, Switzerland) 4 35215058
2021 Search for Associations of FSHR, INHA, INHAB, PRL, TNP2 and SPEF2 Genes Polymorphisms with Semen Quality in Russian Holstein Bulls (Pilot Study). Animals : an open access journal from MDPI 4 34679903
2023 Genetic mapping and functional analysis of a classical tassel branch number mutant Tp2 in maize. Frontiers in plant science 3 37332723
2009 Fusion of a cell penetrating peptide from HIV-1 TAT to the Theileria parva antigen Tp2 enhances the stimulation of bovine CD8+ T cell responses. Veterinary immunology and immunopathology 3 19249107
2026 ASB9 promotes ubiquitin-mediated degradation of TNP2 to facilitate histone-to-protamine transition in humans and mice. Proceedings of the National Academy of Sciences of the United States of America 0 41915740
2022 Sequence variability of CatSper1 and TNP2 gene in indigenous and crossbred cattle in Bangladesh. Animal biotechnology 0 35446730
2013 Calcineurin A versus NS5A-TP2/HD domain containing 2: a case study of site-directed low-frequency random mutagenesis for dissecting target specificity of peptide aptamers. Molecular & cellular proteomics : MCP 0 23579184
1983 Influence of the thymic extract (TP2) on some humoral and cell mediated immunity reactions in normal or irradiated animals. Endocrinologie 0 6364318