FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\ball
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General Information
Symbol
Dmel\ball
Species
D. melanogaster
Name
ballchen
Annotation Symbol
CG6386
Feature Type
protein_coding_gene
FlyBase ID
FBgn0027889
Gene Model Status
Current
Stock Availability
12 publicly available
Enzyme Name (EC)
Gene Summary
ballchen (ball) encodes a nuclear serine/threonine kinase that regulates chromatin organization through phosphorylation of the product of baf. It contributes to stem cell maintenance in various tissues, as well as chromosome condensation and segregation in meiosis. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

nhk-1, BcDNA:LD09009 , trip, nucleosomal histone kinase-1, NHK1

Key Links
Genomic Location
Cytogenetic map
97D3-97D3
Sequence location
Recombination map
3-92
RefSeq locus
NT_033777 REGION:26864605..26866807
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (28 terms)
Molecular Function (6 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
enables histone H2AT120 kinase activity
inferred from direct assay
(Aihara et al., 2004)
enables protein binding
inferred from physical interaction with UniProtKB:P08111
(Link et al., 2019)
enables protein kinase activity
inferred from direct assay
(Lancaster et al., 2007)
enables protein kinase binding
inferred from physical interaction with UniProtKB:A8DYP0
(Katzemich et al., 2015)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables ATP binding
inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR017441
(InterPro Project Members, 2004-)
enables protein kinase activity
inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR008271
(InterPro Project Members, 2004-)
enables protein serine/threonine kinase activity
inferred from biological aspect of ancestor with PANTHER:PTN000225966
(GO Reference Genome Project, 2011-)
Biological Process (15 terms)
Terms Based on Experimental Evidence (13 terms)
CV Term
Evidence
References
involved_in chromosome condensation
inferred from mutant phenotype
(Nikalayevich and Ohkura, 2015)
involved_in female germ-line stem cell population maintenance
inferred from mutant phenotype
(Herzig et al., 2014)
involved_in female meiotic nuclear division
inferred from mutant phenotype
(Cullen et al., 2005, Ivanovska et al., 2005)
involved_in flight
inferred from mutant phenotype
(Katzemich et al., 2015)
involved_in male germ-line stem cell population maintenance
inferred from mutant phenotype
(Herzig et al., 2014)
involved_in meiotic cell cycle
inferred from mutant phenotype
(Ivanovska et al., 2005)
involved_in mitotic sister chromatid segregation
inferred from mutant phenotype
(Cullen et al., 2005)
involved_in mitotic spindle organization
inferred from mutant phenotype
(Cullen et al., 2005)
involved_in neuronal stem cell population maintenance
inferred from mutant phenotype
(Yakulov et al., 2014)
involved_in oocyte karyosome formation
inferred from mutant phenotype
(Lancaster et al., 2007)
involved_in post-translational protein modification
inferred from direct assay
(Aihara et al., 2004)
involved_in sarcomere organization
inferred from mutant phenotype
(Katzemich et al., 2015)
involved_in synaptonemal complex assembly
inferred from mutant phenotype
(Ivanovska et al., 2005)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in DNA damage response
inferred from biological aspect of ancestor with PANTHER:PTN008529751
(GO Reference Genome Project, 2011-)
involved_in signal transduction
inferred from biological aspect of ancestor with PANTHER:PTN000225966
(GO Reference Genome Project, 2011-)
Cellular Component (7 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
(Aihara et al., 2004)
located_in M band
inferred from direct assay
(Katzemich et al., 2015)
located_in nucleolus
inferred from direct assay
(Herzig et al., 2014)
located_in nucleus
inferred from direct assay
(Aihara et al., 2004)
located_in sarcomere
inferred from direct assay
(Katzemich et al., 2015)
located_in Z disc
inferred from direct assay
(Katzemich et al., 2015)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
is_active_in cytoplasm
inferred from biological aspect of ancestor with PANTHER:PTN000225966
(GO Reference Genome Project, 2011-)
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000225966
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the protein kinase superfamily. CK1 Ser/Thr protein kinase family. VRK subfamily. (Q7KRY6)
Protein Signatures (InterPro)
Catalytic Activity (EC/Rhea)
protein serine/threonine kinase activity
Summaries
Gene Snapshot
ballchen (ball) encodes a nuclear serine/threonine kinase that regulates chromatin organization through phosphorylation of the product of baf. It contributes to stem cell maintenance in various tissues, as well as chromosome condensation and segregation in meiosis. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
CK1 KINASES -
The CK1 kinases (Casein Kinase 1 family) share a highly conserved kinase domain. CK1 kinases are serine/threonine kinases with a preference for acidic substrates. The highly variable C-terminal domains are involved in regulating subcellular localization and activity. (Adapted from FBrf0132098, PMID:24825444 and PMID:15722192).
Protein Function (UniProtKB)
Serine/threonine-protein kinase involved in somatic mitosis and female meiosis (PubMed:16230526). Required for spindle organization in mitosis, and for the establishment or maintenance of meiosis-specific chromosomal configurations, including the prophase I karyosome and the metaphase I spindle (PubMed:15078818, PubMed:16301329). Specifically phosphorylates nucleosomal H2A on 'Thr-119' (PubMed:15078818). Required for the development and organization of indirect flight muscle sarcomeres by regulating the formation of M line and H zone and the correct assembly of thick and thin filaments in the sarcomere (PubMed:26251439).
(UniProt, Q7KRY6)
Summary (Interactive Fly)

multifunctional serine/threonine protein kinase - asymmetric cell division pathway - participates in proliferation control and prevents the differentiation of neuronal stem cells - acts with MASK downstream of obscurin in development of a well defined M-line and Z-disc of muscle - required for chromosome condensation in oocytes - nucleosomal H2A histone kinase

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\ball for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q7KRY6)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.47

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
ball-RA
FBtr0085095
NM_143251
2027
599
ball-RB
FBtr0085096
NM_170292
1965
599
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
ball-PA
FBpp0084465
66.0
599
10.52
Q7KRY6
NP_651508
AAF56630
ball-PB
FBpp0084466
66.0
599
10.52
Q7KRY6
NP_733171
AAN14091
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

599 aa isoforms: ball-PA, ball-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Subunit Structure (UniProtKB)

May interact with Unc-89 (via protein kinase domain 1) (PubMed:26251439). Interacts with L(2)gl (PubMed:31735666).

(UniProt, Q7KRY6)
Post Translational Modification

Phosphorylated during mitosis and female meiosis.

(UniProt, Q7KRY6)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\ball using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

1.37

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 10
organism | intense
(Yakulov et al., 2014)
embryonic stage 11
presumptive embryonic/larval central nervous system | intense
(Yakulov et al., 2014)
neuroblast of ventral nervous system | subset
(Yakulov et al., 2014)
embryonic ganglion mother cell | subset
(Yakulov et al., 2014)
late embryonic stage
embryonic brain | restricted
(Yakulov et al., 2014)
gonad
(Yakulov et al., 2014)
Additional Descriptive Data

From early syncytial cleavage to embryonic stage 10 ball-RNA is expressed ubiquitously. At embryonic stage 11 ball-transcripts are enriched in the nervous system and can be detected in most neuroblasts and a subset of ganglion mother cells. At the end of embryogenesis ball-RNA is restricted to parts of the developing brain and the gonads.

(Yakulov et al., 2014)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 3
syncytium
(Yakulov et al., 2014)
embryonic stage 7 -- 12
embryonic epidermis
(Yakulov et al., 2014)
embryonic stage 11
embryonic ganglion mother cell | subset
(Yakulov et al., 2014)
embryonic stage 11 -- 13
neuroblast of ventral nervous system | subset
(Yakulov et al., 2014)
embryonic stage 13
presumptive embryonic/larval central nervous system | intense | restricted
(Yakulov et al., 2014)
late third instar larval stage
thoracic neuroblast | intense
(Yakulov et al., 2014)
abdominal neuroblast | faint
(Yakulov et al., 2014)
adult stage | female
follicle cell of germarium
(Herzig et al., 2014)
Additional Descriptive Data

ball-protein is detected ubiquitously in the syncytial cells of the early embryo. At later stages ball-protein is expressed in epidermal cells. It becomes restricted to the developing nervous system by stage 11 when the protein is detected in most neuroblasts and a subset of ganglion mother cells. At embryonic stage 11 labeling ist stronger in cells of the peripheral part of the nervous system. During mitosis ball-protein co-localizes with chromatin.

(Yakulov et al., 2014)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
(Aihara et al., 2004)
located_in M band
inferred from direct assay
(Katzemich et al., 2015)
located_in nucleolus
inferred from direct assay
(Herzig et al., 2014)
located_in nucleus
inferred from direct assay
(Aihara et al., 2004)
located_in sarcomere
inferred from direct assay
(Katzemich et al., 2015)
located_in Z disc
inferred from direct assay
(Katzemich et al., 2015)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\ball in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 12 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 14 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ball
Transgenic constructs containing regulatory region of ball
Aberrations (Deficiencies and Duplications) ( 2 )
Inferred from experimentation ( 2 )
Gene disrupted in
Df(3R)Tl-I
(Ivanovska et al., 2005)
Df(3R)Tl-P
(Ivanovska et al., 2005)
Inferred from location ( 6 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal, with Scer\GAL4insc-Mz1407
lethal | embryonic stage
lethal | larval stage
lethal - all die before end of pupal stage
lethal - all die during pupal stage
partially lethal - majority die, with Scer\GAL4Mef2.PU
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
female semi-fertile
female sterile
female sterile, with Scer\GAL4VP16.nanos.UTR, Scer\GAL4VP16.otu, Scer\GAL4nanos.PG
female sterile (with ballE24)
female sterile (with ballE60)
female sterile (with ballE107)
female sterile (with balltrip)
male sterile
male sterile | recessive
Other Phenotypes
Allele
abnormal meiotic cell cycle
abnormal meiotic cell cycle | adult stage | female, with Scer\GAL4VP16.mat.αTub67C
abnormal mitotic cell cycle
abnormal mitotic cell cycle | third instar larval stage
abnormal neuroanatomy | larval stage
abnormal neuroanatomy | larval stage, with Scer\GAL4insc-Mz1407
abnormal neuroanatomy | larval stage | somatic clone
abnormal size | larval stage
decreased cell number | larval stage
decreased occurrence of cell division | larval stage | somatic clone
decreased occurrence of cell division | third instar larval stage
flightless, with Scer\GAL4Mef2.PU
some die during pupal stage
Phenotype manifest in
Allele
central nervous system | third instar larval stage
central nervous system | third instar larval stage (with Df(3R)ro80b)
chromosome | adult stage | female, with Scer\GAL4VP16.mat.αTub67C
chromosome | third instar larval stage
chromosome | third instar larval stage (with Df(3R)ro80b)
condensed chromosome
female germline stem cell | germline clone
female germline stem cell | larval stage
follicle stem cell | somatic clone
ganglion mother cell | larval stage, with Scer\GAL4insc-Mz1407
gonad | larval stage
imaginal disc
imaginal disc | larval stage
indirect flight muscle cell | adult stage, with Scer\GAL4Mef2.PU
karyosome
karyosome (with Df(3R)ro80b)
karyosome (with Df(3R)Tl-I)
larval brain
larval brain | larval stage
larval central nervous system
larval neuroblast | larval stage, with Scer\GAL4insc-Mz1407
male germline stem cell | germline clone
male germline stem cell | larval stage
myofibril | adult stage, with Scer\GAL4Mef2.PU
oocyte, with Scer\GAL4VP16.mat.αTub67C
oocyte | oogenesis stage S14
oocyte | oogenesis stage S14 (with Df(3R)Tl-I)
polar body nucleus | maternal effect
sarcomere | adult stage, with Scer\GAL4Mef2.PU
spindle
spindle | adult stage | female, with Scer\GAL4VP16.mat.αTub67C
spindle pole
testis
thoracic neuroblast | larval stage | somatic clone
Z disc | adult stage, with Scer\GAL4Mef2.PU
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (17)
Hsap\VRK1
12 of 14
Yes
Yes
1  
Hsap\VRK2
10 of 14
No
Yes
Hsap\VRK3
5 of 14
No
Yes
1  
Hsap\CSNK1A1L
3 of 14
No
No
Hsap\CSNK1G1
3 of 14
No
No
1  
Hsap\TTBK1
3 of 14
No
No
1  
Hsap\CSNK1D
2 of 14
No
No
6  
Hsap\CSNK1E
2 of 14
No
No
3  
Hsap\CSNK1G2
2 of 14
No
No
1  
Hsap\CSNK1G3
2 of 14
No
No
Hsap\TTBK2
2 of 14
No
No
1  
Hsap\CDK11A
1 of 14
No
No
Hsap\CDK11B
1 of 14
No
No
Hsap\CDKL5
1 of 14
No
No
2  
Hsap\CSNK1A1
1 of 14
No
No
2  
Hsap\ROCK1
1 of 14
No
No
Hsap\TPTEP2-CSNK1E
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (11)
Rnor\Vrk1
12 of 14
Yes
Yes
Rnor\Vrk2
10 of 14
No
Yes
Rnor\Vrk3
5 of 14
No
Yes
Rnor\Csnk1g1
3 of 14
No
No
Rnor\Csnk1g2
2 of 14
No
No
Rnor\Csnk1g3
2 of 14
No
No
Rnor\Ttbk1
2 of 14
No
No
Rnor\Csnk1a1
1 of 14
No
No
Rnor\Csnk1d
1 of 14
No
No
Rnor\Csnk1e
1 of 14
No
No
Rnor\Ttbk2
1 of 14
No
No
Mus musculus (laboratory mouse) (11)
Mmus\Vrk1
12 of 14
Yes
Yes
Mmus\Vrk2
10 of 14
No
Yes
Mmus\Vrk3
4 of 14
No
Yes
Mmus\Csnk1g1
3 of 14
No
No
Mmus\Ttbk1
3 of 14
No
No
Mmus\Csnk1a1
2 of 14
No
No
Mmus\Csnk1d
2 of 14
No
No
Mmus\Csnk1e
2 of 14
No
No
Mmus\Csnk1g2
2 of 14
No
No
Mmus\Csnk1g3
2 of 14
No
No
Mmus\Ttbk2
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (12)
Xtro\vrk2
7 of 13
Yes
Yes
Xtro\pde4c
6 of 13
No
No
Xtro\vrk1
6 of 13
No
Yes
Xtro\vrk3
2 of 13
No
Yes
Xtro\csnk1a1
1 of 13
No
No
Xtro\csnk1d
1 of 13
No
No
Xtro\csnk1g2
1 of 13
No
No
Xtro\dixdc1
1 of 13
No
Yes
Xtro\lvrn
1 of 13
No
No
Xtro\reg4
1 of 13
No
No
Xtro\ssr2
1 of 13
No
No
Xtro\ttbk2
1 of 13
No
No
Danio rerio (Zebrafish) (15)
Drer\vrk1
12 of 14
Yes
Yes
Drer\LOC794952
10 of 14
No
Yes
Drer\vrk2
10 of 14
No
Yes
Drer\vrk3
5 of 14
No
Yes
Drer\csnk1a1
2 of 14
No
No
Drer\csnk1da
2 of 14
No
No
Drer\csnk1db
2 of 14
No
No
Drer\csnk1e
2 of 14
No
No
Drer\csnk1g1
2 of 14
No
No
Drer\csnk1g2a
2 of 14
No
No
Drer\ttbk1a
2 of 14
No
No
Drer\ttbk1b
2 of 14
No
No
Drer\ttbk2a
2 of 14
No
No
Drer\ttbk2b
2 of 14
No
No
Drer\csnk1g2b
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (97)
Cele\vrk-1
11 of 14
Yes
Yes
Cele\C03C10.2
2 of 14
No
No
Cele\C05C12.1
2 of 14
No
No
Cele\C08F8.6
2 of 14
No
No
Cele\C09D4.3
2 of 14
No
No
Cele\C14A4.13
2 of 14
No
No
Cele\C27D8.1
2 of 14
No
No
Cele\C34B2.3
2 of 14
No
No
Cele\C39H7.1
2 of 14
No
No
Cele\C44C10.7
2 of 14
No
No
Cele\C49C8.1
2 of 14
No
No
Cele\C50F4.10
2 of 14
No
No
Cele\C55B7.10
2 of 14
No
No
Cele\C56C10.6
2 of 14
No
No
Cele\csnk-1
2 of 14
No
No
Cele\D2024.1
2 of 14
No
No
Cele\E02H4.6
2 of 14
No
No
Cele\F22F1.2
2 of 14
No
No
Cele\F25F2.1
2 of 14
No
Yes
Cele\F26A1.3
2 of 14
No
No
Cele\F33D11.7
2 of 14
No
No
Cele\F38E1.3
2 of 14
No
No
Cele\F39F10.2
2 of 14
No
No
Cele\F39F10.3
2 of 14
No
No
Cele\F41G3.5
2 of 14
No
No
Cele\F53C3.1
2 of 14
No
No
Cele\F54H5.2
2 of 14
No
No
Cele\F59A6.4
2 of 14
No
No
Cele\H05L14.1
2 of 14
No
No
Cele\K04C1.5
2 of 14
No
No
Cele\K06H7.8
2 of 14
No
No
Cele\K08H2.5
2 of 14
No
No
Cele\kin-19
2 of 14
No
No
Cele\kin-35
2 of 14
No
No
Cele\M7.7
2 of 14
No
No
Cele\R10D12.10
2 of 14
No
No
Cele\T05A7.6
2 of 14
No
No
Cele\T15B12.2
2 of 14
No
No
Cele\T19D12.5
2 of 14
No
No
Cele\tag-191
2 of 14
No
No
Cele\ttbk-3
2 of 14
No
No
Cele\ttbk-4
2 of 14
No
No
Cele\ttbk-5
2 of 14
No
No
Cele\ttbk-6
2 of 14
No
No
Cele\ttbk-7
2 of 14
No
No
Cele\W03G9.5
2 of 14
No
No
Cele\W06F12.3
2 of 14
No
No
Cele\W09C3.1
2 of 14
No
No
Cele\Y38H8A.3
2 of 14
No
No
Cele\Y39G8C.2
2 of 14
No
No
Cele\Y69F12A.1
2 of 14
No
No
Cele\Y71F9AL.2
2 of 14
No
No
Cele\ZK354.6
2 of 14
No
No
Cele\ZK507.1
2 of 14
No
No
Cele\ZK596.2
2 of 14
No
No
Cele\ZK666.8
2 of 14
No
No
Cele\B0207.7
1 of 14
No
No
Cele\B0218.5
1 of 14
No
No
Cele\C09B9.4
1 of 14
No
No
Cele\C17C3.11
1 of 14
No
No
Cele\C25H3.1
1 of 14
No
No
Cele\C28A5.6
1 of 14
No
No
Cele\C55B6.5
1 of 14
No
Yes
Cele\cdk-9
1 of 14
No
No
Cele\cdk-11.1
1 of 14
No
No
Cele\cdk-11.2
1 of 14
No
No
Cele\D2045.5
1 of 14
No
No
Cele\F16B12.5
1 of 14
No
Yes
Cele\F16B12.7
1 of 14
No
No
Cele\F22H10.1
1 of 14
No
No
Cele\F26A1.4
1 of 14
No
No
Cele\F35C11.3
1 of 14
No
No
Cele\F36H12.9
1 of 14
No
No
Cele\F56H11.6
1 of 14
No
Yes
Cele\F59E12.3
1 of 14
No
No
Cele\K09C6.2
1 of 14
No
No
Cele\K09E4.1
1 of 14
No
No
Cele\K11C4.1
1 of 14
No
No
Cele\kin-20
1 of 14
No
No
Cele\kin-33
1 of 14
No
No
Cele\M05D6.1
1 of 14
No
No
Cele\prde-1
1 of 14
No
No
Cele\R13H9.5
1 of 14
No
No
Cele\R13H9.6
1 of 14
No
No
Cele\spe-6
1 of 14
No
No
Cele\T05C12.1
1 of 14
No
No
Cele\T09B4.7
1 of 14
No
No
Cele\T11F8.4
1 of 14
No
No
Cele\ttbk-2
1 of 14
No
No
Cele\Y47G6A.13
1 of 14
No
No
Cele\Y59E9AR.10
1 of 14
No
Yes
Cele\Y65B4A.9
1 of 14
No
No
Cele\Y73B6A.2
1 of 14
No
Yes
Cele\Y111B2A.27
1 of 14
No
No
Cele\ZC373.3
1 of 14
No
No
Cele\ZC581.2
1 of 14
No
No
Cele\ZK507.3
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (10)
Agam\AgaP_AGAP001933
9 of 12
Yes
Yes
Agam\AgaP_AGAP004901
2 of 12
No
No
Agam\AgaP_AGAP011085
2 of 12
No
No
Agam\AgaP_AGAP013380
2 of 12
No
No
Agam\AgaP_AGAP000627
1 of 12
No
No
Agam\AgaP_AGAP001636
1 of 12
No
No
Agam\AgaP_AGAP003180
1 of 12
No
No
Agam\AgaP_AGAP003997
1 of 12
No
No
Agam\AgaP_AGAP008476
1 of 12
No
No
Agam\AgaP_AGAP011055
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (29)
Atha\ADK1
2 of 13
Yes
No
Atha\AT2G25760
2 of 13
Yes
No
Atha\AT3G13670
2 of 13
Yes
No
Atha\CK1
2 of 13
Yes
No
Atha\CKI1
2 of 13
Yes
No
Atha\CKL2
2 of 13
Yes
No
Atha\ckl3
2 of 13
Yes
No
Atha\ckl4
2 of 13
Yes
No
Atha\ckl5
2 of 13
Yes
No
Atha\CKL6
2 of 13
Yes
No
Atha\ckl7
2 of 13
Yes
No
Atha\ckl8
2 of 13
Yes
No
Atha\ckl10
2 of 13
Yes
No
Atha\ckl12
2 of 13
Yes
No
Atha\CKL13
2 of 13
Yes
No
Atha\AGC1.7
1 of 13
No
No
Atha\AT1G18390
1 of 13
No
No
Atha\AT1G30640
1 of 13
No
No
Atha\AT3G03930
1 of 13
No
No
Atha\AT3G03940
1 of 13
No
No
Atha\AT3G23310
1 of 13
No
No
Atha\AT4G08800
1 of 13
No
No
Atha\AT4G14350
1 of 13
No
No
Atha\AT4G33080
1 of 13
No
No
Atha\AT5G18190
1 of 13
No
No
Atha\CIPK12
1 of 13
No
No
Atha\CIPK21
1 of 13
No
No
Atha\CIPK26
1 of 13
No
No
Atha\PPCK1
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (5)
Scer\YCK1
3 of 13
Yes
No
Scer\YCK2
3 of 13
Yes
No
Scer\YCK3
3 of 13
Yes
No
Scer\HRR25
2 of 13
No
No
Scer\CBK1
1 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (5)
Spom\cki1
2 of 12
Yes
No
Spom\cki2
2 of 12
Yes
No
Spom\cki3
2 of 12
Yes
No
Spom\hhp1
2 of 12
Yes
No
Spom\hhp2
2 of 12
Yes
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:ball. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (14)
bsd
5 of 13
Asator
4 of 13
CG2577
4 of 13
CG7094
4 of 13
CG9962
4 of 13
CkIα
4 of 13
dco
4 of 13
gish
4 of 13
CG12147
3 of 13
Cdc7
2 of 13
CG5790
2 of 13
gwl
1 of 13
Myt1
1 of 13
Sik3
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 2 )
    Modifiers Based on Experimental Evidence ( 2 )
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    VRK1; VRK serine/threonine kinase 1
    12 of 14
    602168
     
    VRK2; VRK serine/threonine kinase 2
    10 of 14
    602169
        VRK3; VRK serine/threonine kinase 3
        5 of 14
        619771
             
            CSNK1A1L; casein kinase 1 alpha 1 like
            3 of 14
                CSNK1G1; casein kinase 1 gamma 1
                3 of 14
                606274
                     
                    TTBK1; tau tubulin kinase 1
                    3 of 14
                    619415
                         
                        Functional Complementation Data
                        Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
                        Interactions
                        Summary of Physical Interactions
                        Summary of Genetic Interactions
                        Interaction Browsers
                        View in FlyBase Interactions Browser   View in MIST tool (external link)
                        Please look at the allele data for full details of the genetic interactions
                        Starting gene(s)
                        Interaction type
                        Interacting gene(s)
                        Reference
                        ball
                        suppressible
                        baf
                        (Nikalayevich and Ohkura, 2015)
                        Starting gene(s)
                        Interaction type
                        Interacting gene(s)
                        Reference
                        Ankle2
                        suppressible
                        ball
                        (Link et al., 2019)
                        bam
                        suppressible
                        ball
                        (Herzig et al., 2014)
                        btl|stumps
                        suppressible
                        ball
                        (Zhu et al., 2005)
                        lok
                        enhanceable
                        ball
                        (Park and Song, 2008)
                        msps
                        suppressible
                        ball
                        (Cullen et al., 2005)
                        pbl
                        enhanceable
                        ball
                        (Gregory et al., 2007)
                        Pvr
                        suppressible
                        ball
                        (Zhu et al., 2005)
                        External Data
                        Subunit Structure (UniProtKB)
                        May interact with Unc-89 (via protein kinase domain 1) (PubMed:26251439). Interacts with L(2)gl (PubMed:31735666).
                        (UniProt, Q7KRY6 )
                        Linkouts
                        BioGRID - A database of protein and genetic interactions.
                        DroID - A comprehensive database of gene and protein interactions.
                        MIST (protein-protein) - An integrated Molecular Interaction Database
                        Pathways
                        Signaling Pathways (FlyBase)
                        Metabolic Pathways
                        FlyBase
                        External Links
                        External Data
                        Linkouts
                        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                        Class of Gene
                        Genomic Location and Detailed Mapping Data
                        Chromosome (arm)
                        3R
                        Recombination map
                        3-92
                        Cytogenetic map
                        97D3-97D3
                        Sequence location
                        FlyBase Computed Cytological Location
                        Cytogenetic map
                        Evidence for location
                        97D3-97D3
                        Limits computationally determined from genome sequence between P{lacW}scribj7B3 and P{lacW}His2AvL1602
                        Experimentally Determined Cytological Location
                        Cytogenetic map
                        Notes
                        References
                        Experimentally Determined Recombination Data
                        Location

                        3-92

                        (FlyBase, 2016)

                        3-88.9

                        (Comeron et al., 2012)
                        Left of (cM)
                        Right of (cM)
                        Notes
                        Stocks and Reagents
                        Stocks (12)
                        Genomic Clones (16)
                         

                        Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

                        cDNA Clones (112)
                         

                        Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

                        cDNA clones, fully sequenced
                        BDGP DGC clones
                        Other clones
                        Drosophila Genomics Resource Center cDNA clones

                        For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

                        cDNA Clones, End Sequenced (ESTs)
                        BDGP DGC clones
                        Other clones
                        RNAi and Array Information
                        Linkouts
                        DRSC - Results frm RNAi screens
                        Antibody Information
                        Laboratory Generated Antibodies
                         

                        polyclonal

                        (Unnikannan et al., 2020, Herzig et al., 2014)
                        Commercially Available Antibodies
                         
                        Cell Line Information
                        Publicly Available Cell Lines
                         
                          Other Stable Cell Lines
                           
                          Other Comments

                          ball is epistatic to msps and tacc mutations.

                          (Cullen et al., 2005)

                          Transfection into S2 cells of dsRNA made from templates generated with primers directed against this gene results in a significant increase in mitotic abnormalities, such as unaligned chromosomes, irregular chromosome condensation, lagging chromosomes, or chromosome bridges. Spindle morphology is sometimes disrupted, typically leading to a multipolar spindle.

                          (Cullen et al., 2005)

                          ball mutants are defective in holding chromosomes together in prophase I and forming a single unified spindle in metaphase I.

                          (Cullen et al., 2005)

                          ball phosphorylates Thr119 at the C-terminus of the His2A protein. This site is phosphorylated specifically in the nucleosome context.

                          (Aihara et al., 2004)

                          Identification: Yeast two hybrid screen for interactors with par-1.

                          (Klinge et al., 2004)

                          A hypomorphic allele shows a male sterile phenotype and causes reduced testes.

                          (Klinge et al., 2004)

                          dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

                          (Kiger et al., 2003)

                          Identification: yeast two hybrid screen for proteins that interact with the par-1 gene product.

                          (Klinge et al., 2003)

                          Male hypomorphic mutants are sterile with reduced testes.

                          (Klinge et al., 2003)
                          Relationship to Other Genes
                          Source for database merge of
                          Additional comments
                          Nomenclature History
                          Source for database identify of

                          Source for identity of: ballchen CG6386

                          (Klinge, 2004.2.25)

                          Source for identity of: ball ballchen

                          (Herzig et al., 2005)
                          Nomenclature comments
                          Etymology
                          Synonyms and Secondary IDs (15)
                          Reported As
                          Symbol Synonym
                          BALL
                          (Shaukat et al., 2021, Herzig et al., 2007)
                          Ball
                          (Harnish et al., 2021, Manring et al., 2017, Yakulov et al., 2007)
                          BcDNA:LD09009
                          (Giot et al., 2003, Cook and Carpenter, 2002)
                          CG6386
                          (Meadows, 2017.1.11, Di Cara et al., 2015, Lipinszki et al., 2015, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Ellis and Carney, 2009, Keleman et al., 2009.8.5, An et al., 2008, Ni et al., 2008, Yang et al., 2008, Cullen et al., 2005, Laviolette et al., 2005, Zhu et al., 2005, Zhu et al., 2005, Aihara et al., 2004, Brody et al., 2002)
                          NHK-1
                          (Hinnant et al., 2020, Park and Song, 2008, Cullen et al., 2005, Aihara et al., 2004)
                          NHK1
                          (Torras-Llort et al., 2020, Yakulov et al., 2007)
                          VRK
                          (Morrison et al., 2000)
                          ball
                          (Peng et al., 2024, Ecovoiu et al., 2022, Khan et al., 2021, Ostalé et al., 2021, Yamaguchi et al., 2021, Wang et al., 2020, Link et al., 2019, Zhang et al., 2018, Transgenic RNAi Project members, 2017-, Sarov et al., 2016, Katzemich et al., 2015, Kwong et al., 2015, Amcheslavsky et al., 2014, Gonzalez et al., 2014, Herzig et al., 2014, Sopko et al., 2014, Yakulov et al., 2014, Handler et al., 2013, Handler et al., 2013, Vaque et al., 2013, Habermann et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Ellis and Carney, 2009, Gunesdogan et al., 2007, Yakulov et al., 2007, Cullen et al., 2005)
                          ballchen
                          (Link and Bellen, 2020)
                          nhk-1
                          (Hughes et al., 2018, Nikalayevich and Ohkura, 2015, Lancaster et al., 2010, Brittle et al., 2007, Lancaster et al., 2007, Cullen et al., 2005, Ivanovska et al., 2005)
                          trip
                          (Pamudurti et al., 2020, Cullen et al., 2005)
                          Name Synonyms
                          Ballchen
                          (Yang et al., 2008, Yakulov et al., 2007)
                          ballchen
                          (He et al., 2014, McCracken and Locke, 2014, McGraw et al., 2008, Park and Song, 2008, Gregory et al., 2007, Gunesdogan et al., 2007, Herzig et al., 2007, Nurminsky, 2007, Klinge et al., 2003)
                          nuclear histone kinase-1
                          (Park and Song, 2008)
                          nucleosomal histone kinase-1
                          (Cullen et al., 2005, Aihara et al., 2004)
                          triplet
                          (Cullen et al., 2005)
                          Secondary FlyBase IDs
                            Datasets (0)
                            Study focus (0)
                            Experimental Role
                            Project
                            Project Type
                            Title
                            Study result (0)
                            Result
                            Result Type
                            Title
                            External Crossreferences and Linkouts ( 44 )
                            Sequence Crossreferences
                            NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                            GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                            GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                            RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                            UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                            UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                            UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                            Other crossreferences
                            AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                            DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                            EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                            FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                            FlyMine - An integrated database for Drosophila genomics
                            InterPro - A database of protein families, domains and functional sites
                            KEGG Genes - Molecular building blocks of life in the genomic space.
                            MARRVEL_MODEL - MARRVEL (model organism gene)
                            Linkouts
                            BioGRID - A database of protein and genetic interactions.
                            Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                            DroID - A comprehensive database of gene and protein interactions.
                            DRSC - Results frm RNAi screens
                            Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                            FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                            FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                            Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                            Flygut - An atlas of the Drosophila adult midgut
                            FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                            iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                            Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                            MIST (protein-protein) - An integrated Molecular Interaction Database
                            Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                            SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                            References (106)