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UniProt Accession
Theoretical PI
Molecular Weight
84181 Da  
Genbank Nucleotide ID
Genbank Protein ID
Gene Name
Gene Synonyms/Alias
Protein Name
Serine/threonine-protein kinase haspin 
Protein Synonyms/Alias
EC= Germ cell-specific gene 2 protein; Haploid germ cell-specific nuclear protein kinase; 
Mus musculus (Mouse) 
NCBI Taxonomy ID
Chromosome Location
View in Ensembl genome browser  
Function in Stage
Function in Cell Type
Probability (GAS) of Function in Spermatogenesis
The probability was calculated by GAS algorithm, ranging from 0 to 1. The closer it is to 1, the more possibly it functions in spermatogenesis.
Temporarily unavailable 
Abstract of related literatures
1. We have cloned cDNAs involved in germ cell-specific expression. For this, a subtracted cDNA library was generated by subtracting cDNAs derived from supporting cells of mutant testis from wild-type testis cDNAs. Detailed analyses of mRNA expression revealed that the genes corresponding to the cloned cDNAs were exclusively expressed in testes and were developmentally controlled. PMID: [7957958] 

2. We have cloned the entire coding region of a mouse germ cell-specific cDNA encoding a unique protein kinase whose catalytic domain contains only three consensus subdomains (I-III) instead of the normal 12. The protein possesses intrinsic Ser/Thr kinase activity and is exclusively expressed in haploid germ cells, localizing only in their nuclei, and was thus named Haspin (for haploid germ cell-specific nuclear protein kinase). Western blot analysis showed that specific antibodies recognized a protein of Mr 83,000 in the testis. Ectopically expressed Haspin was detected exclusively in the nuclei of cultured somatic cells. Even in the absence of kinase activity, however, Haspin caused cell cycle arrest at G1, resulting in growth arrest of the transfected somatic cells. In a DNA binding experiment, approximately one-half of wild-type Haspin was able to bind to a DNA-cellulose column, whereas the other half was not. In contrast, all of the deletion mutant Haspin that lacked autophosphorylation bound to the DNA column. Thus, the DNA-binding activity of Haspin may, in some way, be associated with its kinase activity. These observations suggest that Haspin has some critical roles in cell cycle cessation and differentiation of haploid germ cells. PMID: [10358056] 

3. The haspin gene specifically expressed in haploid germ cells encodes a unique Ser/Thr protein kinase. We have cloned a mouse haspin genomic clone using cDNA as a probe. Sequencing data showed that the haspin gene was not interrupted by introns and was bordered by appropriate direct repeat. The transcription start site of the gene was not preceded by a TATA box. The whole transcription unit was located at an intron of integrin alphaM290 gene, and transcription direction of these two genes was different. Southern blotting analysis under stringent condition showed that haspin was a single gene. Phylogenetic analysis suggested that the diversion of haspin gene from other kinase family might be very ancient: the early stage of plant-fungus-animal split. PMID: [11311555] 

4. Haspin is a serine/threonine kinase, recently identified in mice, that is thought to regulate cell cycle and differentiation of haploid germ cells. Here, the haspin gene is identified within an intron of the integrin alphaE gene. Transcription occurs from a bi-directional CpG island-associated promoter that also generates an alternatively spliced integrin alphaE derived RNA. Remarkably, the human and murine haspin genes lack introns, and have features of retroposons. The human haspin cDNA reveals that the human and murine proteins are 83% identical in the C-terminal kinase domain, but only 53% identical in the N-terminal region. The haspin kinase domain has structural features that distinguish it from previously characterized proteins and suggest that haspin is a member of a new family of protein kinases. Although formerly thought to be expressed selectively in the testes, haspin is also transcribed at lower levels in thymus, bone marrow, fetal liver and other fetal tissues, and in all proliferating cell lines tested. Thus haspin is likely to be important in regulation of diploid as well as haploid cell differentiation in a variety of tissues. PMID: [11311556] 

5. The mouse (Mus musculus) is the premier animal model for understanding human disease and development. Here we show that a comprehensive understanding of mouse biology is only possible with the availability of a finished, high-quality genome assembly. The finished clone-based assembly of the mouse strain C57BL/6J reported here has over 175,000 fewer gaps and over 139 Mb more of novel sequence, compared with the earlier MGSCv3 draft genome assembly. In a comprehensive analysis of this revised genome sequence, we are now able to define 20,210 protein-coding genes, over a thousand more than predicted in the human genome (19,042 genes). In addition, we identified 439 long, non-protein-coding RNAs with evidence for transcribed orthologs in human. We analyzed the complex and repetitive landscape of 267 Mb of sequence that was missing or misassembled in the previously published assembly, and we provide insights into the reasons for its resistance to sequencing and assembly by whole-genome shotgun approaches. Duplicated regions within newly assembled sequence tend to be of more recent ancestry than duplicates in the published draft, correcting our initial understanding of recent evolution on the mouse lineage. These duplicates appear to be largely composed of sequence regions containing transposable elements and duplicated protein-coding genes; of these, some may be fixed in the mouse population, but at least 40% of segmentally duplicated sequences are copy number variable even among laboratory mouse strains. Mouse lineage-specific regions contain 3,767 genes drawn mainly from rapidly-changing gene families associated with reproductive functions. The finished mouse genome assembly, therefore, greatly improves our understanding of rodent-specific biology and allows the delineation of ancestral biological functions that are shared with human from derived functions that are not. PMID: [19468303] 

6. Post-translational modifications of conserved N-terminal tail residues in histones regulate many aspects of chromosome activity. Thr 3 of histone H3 is highly conserved, but the significance of its phosphorylation is unclear, and the identity of the corresponding kinase unknown. Immunostaining with phospho-specific antibodies in mammalian cells reveals mitotic phosphorylation of H3 Thr 3 in prophase and its dephosphorylation during anaphase. Furthermore we find that haspin, a member of a distinctive group of protein kinases present in diverse eukaryotes, phosphorylates H3 at Thr 3 in vitro. Importantly, depletion of haspin by RNA interference reveals that this kinase is required for H3 Thr 3 phosphorylation in mitotic cells. In addition to its chromosomal association, haspin is found at the centrosomes and spindle during mitosis. Haspin RNA interference causes misalignment of metaphase chromosomes, and overexpression delays progression through early mitosis. This work reveals a new kinase involved in composing the histone code and adds haspin to the select group of kinases that integrate regulation of chromosome and spindle function during mitosis and meiosis. PMID: [15681610] 

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Serine/threonine-protein kinase that phosphorylateshistone H3 at 'Ser-3' (H3T3ph) during mitosis. This positions andactivates AURKB and other components of the chromosomal passengercomplex (CPC) at centromeres to ensure proper chromatid cohesion,metaphase alignment and normal progression through the cell cycle. 
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Subcellular Location
Nucleus. Chromosome (By similarity).Cytoplasm, cytoskeleton, spindle (By similarity). Note=Nuclearduring interphase and associates with the chromosomes and spindleapparatus during mitosis (By similarity). 
Tissue Specificity
Expressed in germ cells within the testis ofadults and of embryos from day 24 onwards. Also present in adultthymus and weakly expressed in spleen, lung and whole embryo. 
Gene Ontology
GO IDGO termEvidence
GO:0005813 C:centrosome ISS:UniProtKB.
GO:0005694 C:chromosome IEA:UniProtKB-SubCell.
GO:0031965 C:nuclear membrane IEA:Compara.
GO:0005634 C:nucleus IDA:UniProtKB.
GO:0005819 C:spindle ISS:UniProtKB.
GO:0005524 F:ATP binding IDA:UniProtKB.
GO:0003677 F:DNA binding IDA:MGI.
GO:0072354 F:histone kinase activity (H3-T3 specific) ISS:UniProtKB.
GO:0007243 P:intracellular protein kinase cascade IDA:UniProtKB.
GO:0007064 P:mitotic sister chromatid cohesion ISS:UniProtKB.
GO:0071459 P:protein localization to chromosome, centromeric region ISS:UniProtKB.
GO:0090231 P:regulation of spindle checkpoint ISS:UniProtKB.
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IPR024604;    DUF3635.
IPR011009;    Kinase-like_dom.
IPR000719;    Prot_kinase_cat_dom.
IPR017441;    Protein_kinase_ATP_BS.
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PF12330;    DUF3635;    1.
PF00069;    Pkinase;    1.
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PS00107;    PROTEIN_KINASE_ATP;    1.
PS50011;    PROTEIN_KINASE_DOM;    1.
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Created Date
Record Type
GAS predicted 
Sequence Annotation
CHAIN         1    754       Serine/threonine-protein kinase haspin.
DOMAIN      440    754       Protein kinase.
NP_BIND     446    454       ATP (By similarity).
NP_BIND     562    567       ATP (By similarity).
NP_BIND     605    610       ATP (By similarity).
NP_BIND     643    645       ATP (By similarity).
ACT_SITE    127    127       Proton acceptor (By similarity).
ACT_SITE    605    605       Proton acceptor (By similarity).
BINDING     467    467       ATP (By similarity).
MOD_RES      55     55       Phosphoserine (By similarity).
CONFLICT    104    104       P -> L (in Ref. 1; BAA75494 and 5;
CONFLICT    305    305       Q -> R (in Ref. 2; BAB00640 and 4;
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Nucleotide Sequence
Length: 2827 bp   Go to nucleotide: FASTA
Protein Sequence
Length: 754 bp   Go to amino acid: FASTA
The verified Protein-Protein interaction information
Gene Symbol Ref Databases
Other Protein-Protein interaction resources
String database  
View Microarray data