FlyBase Gene Report: Dmel\CASK

General Information

Symbol

Dmel\CASK

Species

D. melanogaster

Name

CASK

Annotation Symbol

CG6703

Feature Type

protein_coding_gene

FlyBase ID

FBgn0013759

Gene Model Status

Current

Stock Availability

64 publicly available

Enzyme Name (EC)

Gene Summary

CASK (CASK) encodes a member of the MAGUK family of scaffolding proteins. In epithelial tissues, the product of CASK functions as an adhesion molecule involved in cell proliferation, cytoskeletal remodeling and cell migration. In neurons, the product of CASK serves as a scaffold for a number of signaling and transmembrane molecules and is involved in synaptic development and plasticity. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Gene Group
UniProtKB
Interactive Fly

Also Known As

Camguk, Caki, cmg, CASK ortholog, Calcium/calmodulin-dependent protein kinase

Key Links

Genomic Location

Cytogenetic map

93F10-93F12

Sequence location

3R:21,783,400..21,822,674 [-]

Recombination map

3-73

RefSeq locus

NT_033777 REGION:21783400..21822674

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

Gene Ontology (GO) Annotations (30 terms)

Molecular Function (5 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

enables neurexin family protein binding

inferred from physical interaction with UniProtKB:Q9VCZ9

(Sun et al., 2009)

Terms Based on Predictions or Assertions (4 terms)

CV Term

Evidence

References

enables ATP binding

inferred from electronic annotation with InterPro:IPR000719

(InterPro Project Members, 2004-)

enables protein kinase activity

inferred from electronic annotation with InterPro:IPR000719

(InterPro Project Members, 2004-)

enables protein serine/threonine kinase activity

inferred from sequence or structural similarity with UniProtKB:O14936

(Gene Ontology Curators, 2002-)

enables signaling receptor binding

inferred from biological aspect of ancestor with PANTHER:PTN001137126

(GO Reference Genome Project, 2011-)

Biological Process (21 terms)

Terms Based on Experimental Evidence (17 terms)

CV Term

Evidence

References

involved_in adult locomotory behavior

inferred from mutant phenotype

(Slawson et al., 2011)

involved_in adult walking behavior

inferred from mutant phenotype

(Martin and Ollo, 1996)

involved_in anesthesia-resistant memory

inferred from mutant phenotype

(Malik et al., 2013)

involved_in behavioral response to ethanol

inferred from mutant phenotype

(Maiya et al., 2012)

involved_in border follicle cell migration

inferred from mutant phenotype

(Aranjuez et al., 2012)

involved_in locomotion

inferred from mutant phenotype

(Sun et al., 2009)

involved_in long-term memory

inferred from mutant phenotype

(Malik et al., 2013)

involved_in male courtship behavior

inferred from mutant phenotype

(Slawson et al., 2011)

involved_in medium-term memory

inferred from mutant phenotype

(Malik et al., 2013)

involved_in neuromuscular synaptic transmission

inferred from mutant phenotype

(Sun et al., 2009)

involved_in positive regulation of synaptic vesicle endocytosis

inferred from mutant phenotype

(Sun et al., 2009)

inferred from genetic interaction with FLYBASE:Nrx-1; FB:FBgn0038975

(Sun et al., 2009)

involved_in regulation of locomotion

inferred from mutant phenotype

(Sun et al., 2009)

inferred from genetic interaction with FLYBASE:Nrx-1; FB:FBgn0038975

(Sun et al., 2009)

involved_in regulation of neuromuscular synaptic transmission

inferred from mutant phenotype

(Sun et al., 2009)

inferred from genetic interaction with FLYBASE:Nrx-1; FB:FBgn0038975

(Sun et al., 2009)

involved_in regulation of neurotransmitter secretion

inferred from direct assay

(Zordan et al., 2005)

involved_in regulation of synaptic assembly at neuromuscular junction

inferred from mutant phenotype

(Gillespie and Hodge, 2013)

involved_in regulation of terminal button organization

inferred from mutant phenotype

(Sun et al., 2009)

involved_in synaptic vesicle endocytosis

inferred from mutant phenotype

(Sun et al., 2009)

Terms Based on Predictions or Assertions (5 terms)

CV Term

Evidence

References

involved_in intracellular protein localization

inferred from biological aspect of ancestor with PANTHER:PTN001137126

(GO Reference Genome Project, 2011-)

involved_in neurotransmitter secretion

non-traceable author statement

(Lloyd et al., 2000)

involved_in regulation of neurotransmitter secretion

inferred from biological aspect of ancestor with PANTHER:PTN001137126

(GO Reference Genome Project, 2011-)

involved_in synaptic vesicle docking

non-traceable author statement

(Lloyd et al., 2000)

involved_in synaptic vesicle targeting

non-traceable author statement

(Lloyd et al., 2000)

Cellular Component (4 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

located_in neuromuscular junction

inferred from direct assay

(Sun et al., 2009)

located_in type I terminal bouton

inferred from direct assay

(Sun et al., 2009)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

is_active_in cell-cell junction

inferred from biological aspect of ancestor with PANTHER:PTN001137125

(GO Reference Genome Project, 2011-)

is_active_in plasma membrane

inferred from biological aspect of ancestor with PANTHER:PTN001137125

(GO Reference Genome Project, 2011-)

Gene Group (FlyBase)

CALCIUM/CALMODULIN-DEPENDENT PROTEIN KINASES

Protein Family (UniProt)

Belongs to the MAGUK family. (Q24210)

Protein Signatures (InterPro)

Catalytic Activity (EC/Rhea)

protein serine/threonine kinase activity

Summaries

Gene Snapshot

Gene Group (FlyBase)

CALCIUM/CALMODULIN-DEPENDENT PROTEIN KINASES -: The Calcium/Calmodulin-dependent kinase family are serine/threonine kinases which, in general, prefer substrates containing basic residues. Many, but not all, members are regulated by calcium or calmodulin. (Adapted from FBrf0132098).

Protein Function (UniProtKB)

May regulate transmembrane proteins that bind calcium, calmodulin, or nucleotides. Functionally modulates eag potassium channels; increases eag current and whole-cell conductance. Also regulates autophosphorylation of CaMKII. In neuromuscular junctions (NMJ), plays a role in synaptic vesicle recycling and thereby in neurotransmission (PubMed:19379781). Together with Nrx-1, required for locomotion (PubMed:19379781).

(UniProt, Q24210)

Summary (Interactive Fly)

MAGUK family of scaffolding protein - guanylate domain - acts synaptically as a gain controller on the transition to calcium-independence - affects activity-dependent autophosphorylation of CaMKII - functions in epithelia as an adhesion molecule involved in cell proliferation, cytoskeletal remodeling and cell migration- serves as a scaffold for a number of signaling and transmembrane molecules and is involved in synaptic development and plasticity

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\CASK 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 Q24210)

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

Gene model reviewed during 6.19

Gene model reviewed during 5.52

Gene model reviewed during 5.56

Gene model reviewed during 6.32

A non-AUG start codon may be used for translation of one or more transcripts of this gene; based on the presence of conserved protein signatures within the 5' UTR without an in-frame AUG (FBrf0243886).

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.

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

Gene model reviewed during 5.44

Stop-codon suppression (UGA) postulated; FBrf0216884.

Gene model reviewed during 5.47

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

CASK-RA

FBtr0084162

NM_169970

3949

591

CASK-RB

FBtr0084161

NM_079717

3965

898

CASK-RD

FBtr0112872

NM_001104392

4017

591

CASK-RE

FBtr0112873

NM_001104393

2586

469

CASK-RF

FBtr0301327

NM_001170210

6544

916

CASK-RG

FBtr0301328

NM_169971

4071

609

CASK-RH

FBtr0330368

NM_001275882

4128

929

CASK-RI

FBtr0479755

NM_001382187

3164

522

Additional Transcript Data and Comments

Reported size (kB)

7.0, 4.0 (northern blot)

(Martin and Ollo, 1996)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

CASK-PA

FBpp0083560

65.2

591

6.87

FBpp0083559

100.9

898

6.20

FBpp0111785

65.2

591

6.87

FBpp0111786

53.3

469

6.47

FBpp0290542

102.5

916

6.08

FBpp0290543

66.7

609

6.67

NP_732662

AAF55920

CASK-PH

FBpp0303394

104.3

929

5.75

FBpp0428098

59.4

522

5.64

QJC18477

Polypeptides with Identical Sequences

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

591 aa isoforms: CASK-PA, CASK-PD

Additional Polypeptide Data and Comments

Reported size (kDa)

571 (aa); 63 (kD predicted)

(Martin and Ollo, 1996)

Comments

Constitutively expressed CASK protein was shown to be enzymatically active in a cotransfection assay in rat pituitary cells. Sequence analysis indicates that CASK may be a member of a new branch of CaM kinases.

(Martin and Ollo, 1996)

External Data

Subunit Structure (UniProtKB)

Interacts with eag. Interacts with CaMKII. Interacts (via PDZ domain) with Nrx-1 (via C-terminal PDZ binding motif) (PubMed:19379781).

(UniProt, Q24210)

Crossreferences

InterPro - A database of protein families, domains and functional sites

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\CASK 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).

-1.00

Transcript Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 5 -- 7

somatic cell

(Lopes et al., 2001)

embryonic stage 7

cephalic furrow

(Lopes et al., 2001)

embryonic stage 8 -- 9

ventral neurectoderm

(Lopes et al., 2001)

embryonic stage 8 -- 12

germ band

(Lopes et al., 2001)

epithelial furrow

(Lopes et al., 2001)

amnioproctodeal invagination

(Lopes et al., 2001)

posterior midgut primordium

(Lopes et al., 2001)

anterior midgut primordium

(Lopes et al., 2001)

embryonic stage 10 -- 12

ventral nerve cord primordium

Comment: >

(Lopes et al., 2001)

embryonic stage 13 -- 14

midgut primordium

(Lopes et al., 2001)

embryonic stage 13 -- third instar larval stage

larval central nervous system

(Lopes et al., 2001)

embryonic stage 17

embryonic/larval Malpighian tubule

(Lopes et al., 2001)

pupal stage -- adult stage

supraesophageal ganglion

(Lopes et al., 2001)

adult optic lobe

(Lopes et al., 2001)

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism | ubiquitous

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 4 -- 6

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 7 -- 8

organism | faint | ubiquitous

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 8 -- 12

posterior midgut primordium

(Martin and Ollo, 1996)

anterior midgut primordium

(Martin and Ollo, 1996)

embryonic stage 9 -- 10

organism | faint | ubiquitous

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 9 -- 12

midline primordium

(Kearney et al., 2004)

northern blot

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage

(Martin and Ollo, 1996)

embryonic stage 9 -- 17

presumptive embryonic/larval central nervous system | presumptive

(Martin and Ollo, 1996)

embryonic stage 11

neuroblast

(Martin and Ollo, 1996)

embryonic stage 12 -- 17

Comment: reference states 8-24 hr AEL

(Martin and Ollo, 1996)

embryonic stage 14

embryonic/larval hindgut

(Martin and Ollo, 1996)

Malpighian tubule

(Martin and Ollo, 1996)

larval stage

larval central nervous system

(Martin and Ollo, 1996)

pupal stage

central nervous system

(Martin and Ollo, 1996)

adult stage

(Martin and Ollo, 1996)

adult head

(Martin and Ollo, 1996)

oogenesis & adult stage | female

ovary

(Martin and Ollo, 1996)

adult stage & oogenesis

nurse cell

(Martin and Ollo, 1996)

adult stage & oogenesis stage S10B

polar follicle cell

(Martin and Ollo, 1996)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

wandering third instar larval stage

neuropil of larval central nervous system

(Sun et al., 2009)

larval somatic muscle cell

(Sun et al., 2009)

embryonic/larval neuromuscular junction

• terminal bouton

(Sun et al., 2009)

adult stage

lamina

(Martin and Ollo, 1996)

adult optic lobe

(Martin and Ollo, 1996)

adult brain

(Martin and Ollo, 1996)

lobula

(Martin and Ollo, 1996)

lobula plate

(Martin and Ollo, 1996)

medulla

(Martin and Ollo, 1996)

mass spectroscopy

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage

adult head

• membrane

(Aradska et al., 2015)

Additional Descriptive Data

CASK protein is expressed in the optic lobe, the neuropil of the medulla, the lobula, and the lobula plate, while weaker expression is seen in the lamina. Expression is also detected in the central brain.

(Martin and Ollo, 1996)

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in neuromuscular junction

inferred from direct assay

(Sun et al., 2009)

located_in type I terminal bouton

inferred from direct assay

(Sun et al., 2009)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\CASK in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\CASK

View exonic expression by tissue for Dmel\CASK

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

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0013759

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

42567/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0013759

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0013759

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0013759

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG6703

Flygut - An atlas of the Drosophila adult midgut

FBgn0013759

Images

FlyExpress - Embryonic expression images (BDGP data)

Show all FlyExpress processed images for Dmel\CASK

Stages(s) 1-3
Stages(s) 4-6
Stages(s) 9-10
Stages(s) 11-12
Stages(s) 13-16

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 22 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 15 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of CASK

Transgenic constructs containing regulatory region of CASK

Aberrations (Deficiencies and Duplications) ( 3 )

Inferred from experimentation ( 3 )

Gene disrupted in

Df(3R)caki^X-313

(Martin and Ollo, 1996)

Df(3R)Exel6187

(Ryder, 2004.4.26)

Df(3R)X-307

(Grillo et al., 2012)

Inferred from location ( 4 )

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

viable

viable, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

CASK^GD10585

viable, with Scer\GAL4^pnr-MD237

CASK^GD10585

Sterility

Allele

female semi-sterile, with Scer\GAL4^{VP16.mat.αTub67C}

CASK^HMC03260

female semi-sterile, with Scer\GAL4^{VP16.nanos.UTR}, Scer\GAL4^VP16.otu, Scer\GAL4^nanos.PG

fertile

Other Phenotypes

Allele

abnormal courtship behavior

CASK^X-307

abnormal courtship behavior | male

CASK^β-null

abnormal courtship behavior | male (with CASK^X-307)

CASK^X-313

abnormal courtship behavior | male (with CASK^X-313)

CASK^X-307

abnormal flight (with CASK^X-307)

CASK^X-313

abnormal flight (with CASK^X-313)

CASK^X-307

abnormal learning

CASK^β-null

abnormal learning, with Scer\GAL4^Tab2-201Y

abnormal locomotor behavior

abnormal locomotor behavior, with CASK^UAS.cBa, Scer\GAL4^C164

CASK^β-null

abnormal locomotor behavior, with CASK^β-null, Scer\GAL4^C164

CASK^UAS.cBa

abnormal locomotor behavior (with CASK^X-307)

abnormal locomotor behavior (with CASK^X-313)

abnormal locomotor behavior (with CASK^β-null)

abnormal locomotor behavior (with Df(3R)Exel6187)

CASK^β-null

abnormal locomotor behavior | dominant

CASK^β-null

abnormal locomotor behavior | late third instar larval stage (with CASK^X-307)

CASK^X-313

abnormal locomotor behavior | late third instar larval stage (with CASK^X-313)

CASK^X-307

abnormal locomotor behavior | recessive

CASK^β-null

abnormal memory

CASK^β-null

abnormal memory, with Scer\GAL4^c305a

abnormal memory, with Scer\GAL4^CaMKII.7

CASK^KK109311

abnormal memory, with Scer\GAL4^ey-OK107

abnormal neuroanatomy | larval stage | semidominant

CASK^β-null

abnormal neuroanatomy | pharate adult stage

CASK^β-null

abnormal neuroanatomy | third instar larval stage

CASK^β-null

abnormal neuroanatomy | third instar larval stage, with Scer\GAL4^Mef2.PR

abnormal neuroanatomy | third instar larval stage, with Scer\GAL4^VGlut1-OK371

CASK^KK109311

abnormal neurophysiology (with CASK^X-307)

CASK^X-313

abnormal neurophysiology (with CASK^X-313)

CASK^X-307

abnormal neurophysiology | dominant | late third instar larval stage

CASK^X-307

abnormal neurophysiology | late third instar larval stage (with CASK^X-307)

CASK^X-313

abnormal neurophysiology | late third instar larval stage (with CASK^X-313)

CASK^X-307

abnormal pain response | adult stage, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

CASK^GD10585

abnormal visual behavior (with CASK^X-307)

CASK^X-313

abnormal visual behavior (with CASK^X-313)

CASK^X-307

chemical sensitive | adult stage

decreased body size | pharate adult stage

CASK^β-null

decreased size | pharate adult stage

CASK^β-null

visible, with Scer\GAL4^pnr-MD237

CASK^GD10585

Phenotype manifest in

Allele

adult brain | pharate adult stage

CASK^β-null

adult head | pharate adult stage

CASK^β-null

border follicle cell, with Scer\GAL4^c306

CASK^GD10585

bouton | late third instar larval stage (with CASK^X-307)

CASK^X-313

bouton | late third instar larval stage (with CASK^X-313)

CASK^X-307

chaeta, with Scer\GAL4^pnr-MD237

CASK^GD10585

embryonic/larval neuromuscular junction

CASK^X-307

embryonic/larval neuromuscular junction | late third instar larval stage (with CASK^X-307)

CASK^X-313

embryonic/larval neuromuscular junction | late third instar larval stage (with CASK^X-313)

CASK^X-307

larval central nervous system | larval stage

CASK^β-null

larval CNS neuron

CASK^β-null

larval CNS neuron (with Df(3R)caki^X-313)

CASK^β-null

larval CNS neuron (with Df(3R)X-307)

CASK^β-null

neurite | larval stage

CASK^β-null

neurite | larval stage (with Df(3R)caki^X-313)

CASK^β-null

neurite | larval stage (with Df(3R)X-307)

CASK^β-null

synaptic vesicle

CASK^X-307

synaptic vesicle (with CASK^X-307)

CASK^X-313

synaptic vesicle (with CASK^X-313)

CASK^X-307

trichogen cell, with Scer\GAL4^pnr-MD237

CASK^GD10585

type Ib terminal bouton | third instar larval stage

CASK^β-null

type Ib terminal bouton | third instar larval stage, with Scer\GAL4^Mef2.PR

type Ib terminal bouton | third instar larval stage, with Scer\GAL4^VGlut1-OK371

CASK^KK109311

type Is terminal bouton | third instar larval stage

CASK^β-null

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (60)

Hsap\CASK

11 of 14

Yes

Hsap\MPP1

6 of 14

Yes

4 of 14

3 of 14

2 of 14

2 of 14

2 of 14

2 of 14

Yes

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (33)

Rnor\Cask

11 of 14

Yes

4 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Rnor\Patj

1 of 14

Rnor\Pnck

1 of 14

Rnor\Pskh1

1 of 14

Mus musculus (laboratory mouse) (38)

Mmus\Cask

11 of 14

Yes

Mmus\Mpp1

6 of 14

Yes

3 of 14

3 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (40)

Xtro\cask

7 of 13

Yes

Xtro\mpp2

2 of 13

Xtro\cacnb2

1 of 13

Xtro\cacng1

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Danio rerio (Zebrafish) (52)

Drer\caska

12 of 14

Yes

Drer\caskb

5 of 14

Yes

Drer\mpp1

5 of 14

Yes

4 of 14

3 of 14

3 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Caenorhabditis elegans (Nematode, roundworm) (16)

Cele\lin-2

13 of 14

Yes

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Anopheles gambiae (African malaria mosquito) (16)

Agam\AgaP_AGAP001683

11 of 12

Yes

3 of 12

2 of 12

2 of 12

2 of 12

2 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Arabidopsis thaliana (thale-cress) (29)

Atha\AGK2

2 of 13

Yes

Atha\AT3G06200

2 of 13

Yes

Atha\GK-1

2 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Atha\PDK1

1 of 13

Saccharomyces cerevisiae (Brewer's yeast) (3)

Scer\GUK1

2 of 13

Yes

Scer\CMK2

1 of 13

Scer\SCH9

1 of 13

Schizosaccharomyces pombe (Fission yeast) (1)

Spom\SPBC1198.05

2 of 12

Yes

Escherichia coli (enterobacterium) (1)

Ecol\gmk

2 of 11

Yes

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:CASK. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (17)

vari

5 of 13

metro

4 of 13

sdt

4 of 13

CaMKII

2 of 13

dlg1

2 of 13

Dlg5

2 of 13

oya

2 of 13

baz

1 of 13

Ca-β

1 of 13

CaMKI

1 of 13

hpo

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

pyd

1 of 13

scrib

1 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 1 )

Allele

Disease

Evidence

References

CASK^β-null

model of syndromic X-linked intellectual disability

CEA

(Tello et al., 2023, Andrew et al., 2021)

model of autism spectrum disorder

CEA

(Andrew et al., 2021)

Potential Models Based on Orthology ( 2 )

Human Ortholog

Disease

Evidence

References

CASK; calcium/calmodulin dependent serine protein kinase

model of FG syndrome

IEA

(FlyBase, 2019-)

model of syndromic X-linked intellectual disability Najm type

IEA

(FlyBase, 2019-)

Modifiers Based on Experimental Evidence ( 1 )

Allele

Disease

Interaction

References

CASK^HMC03260

exacerbates Alzheimer's disease 1

modeled by Appl^KK102543

(Silva et al., 2020)

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?

CASK; calcium/calmodulin dependent serine protein kinase

11 of 14

CALCIUM/CALMODULIN-DEPENDENT SERINE PROTEIN KINASE; CASK

MPP1; MAGUK p55 scaffold protein 1

6 of 14

305360

PALS2; protein associated with LIN7 2, MAGUK p55 family member

4 of 14

606959

MPP2; MAGUK p55 scaffold protein 2

3 of 14

600723

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.

Dmel gene

Ortholog showing functional complementation

Supporting References

CASK

Hsap\CASK

(Malik et al., 2013)

Interactions

Summary of Physical Interactions

Interaction Browsers

View in FlyBase Interactions Browser View in MIST tool (external link)

Please see the Physical Interaction reports below for full details

protein-protein

Physical Interaction

Assay

References

CASK - CaMKII

anti tag coimmunoprecipitation, western blot, pull down, anti bait coimmunoprecipitation

(Lu et al., 2003)

CASK - Nrx-1

two hybrid, pull down, western blot, anti bait coimmunoprecipitation

(Sun et al., 2009)

CASK - veli

experimental knowledge based, two hybrid

(Guruharsha et al., 2011, Bachmann et al., 2004)

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

CASK

suppressible

CaMKII

(Malik et al., 2013)

CASK

enhanceable

Nrx-1

(Sun et al., 2009)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

CaMKII

suppressible

CASK

(Gillespie and Hodge, 2013, Malik et al., 2013)

enhanceable

CASK

(Müller et al., 2005)

suppressible

(Li et al., 2012)

enhanceable

(Sun et al., 2009)

External Data

Subunit Structure (UniProtKB)

Interacts with eag. Interacts with CaMKII. Interacts (via PDZ domain) with Nrx-1 (via C-terminal PDZ binding motif) (PubMed:19379781).

(UniProt, Q24210 )

Linkouts

BioGRID - A database of protein and genetic interactions.

67530

DroID - A comprehensive database of gene and protein interactions.

FBgn0013759

MIST (genetic) - An integrated Molecular Interaction Database

42567

MIST (protein-protein) - An integrated Molecular Interaction Database

42567

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.

Q24210

Class of Gene

gene_with_edited_transcript

(Graveley et al., 2011)

gene_with_stop_codon_read_through

(Jungreis et al., 2011)

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

3-73

Cytogenetic map

93F10-93F12

Sequence location

3R:21,783,400..21,822,674 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

93F10-93F12

Limits computationally determined from genome sequence between P{PZ}tsl⁰⁰⁶¹⁷ and P{lacW}how^j5B5&P{EP}EP738

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

93F-93F

(Lloyd et al., 2000, Dimitratos et al., 1997)

93F-93F

(determined by in situ hybridisation)

(Dimitratos et al., 1997)

93E-93E

(determined by in situ hybridisation)

(Martin and Ollo, 1996)

93F6-93F14

(Martin et al., 1994)

Determined by comparing Celera genomic sequence with sequence from BDGP BAC and P1 clones. Determined by comparing Celera genomic sequence with sequence from BDGP BAC and P1 clones.

(Lloyd et al., 2000)

Experimentally Determined Recombination Data

Location

3-73

(FlyBase, 2016)

3-75.4

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (64)

Aberrations including deletions of this gene Aberrations including duplications of this gene

Bloomington

39291

w¹¹¹⁸; P{GMR64B03-GAL4}attP2

39306

w¹¹¹⁸; P{GMR64E12-GAL4}attP2

39368

w¹¹¹⁸; P{GMR65G02-GAL4}attP2

39376

w¹¹¹⁸; P{GMR65G11-GAL4}attP2

48232

w¹¹¹⁸; P{GMR64H02-GAL4}attP2

49604

w¹¹¹⁸; P{GMR64D12-GAL4}attP2

Korea

15533

P{GMR65G09-GAL4}attP2

16373

P{GMR65G11-lexA}JK22C

16385

P{GMR65G11-lexA}JK73A

16372

PBac{GMR65G11-lexA}VK00027

VDRC

v104793

P{KK109311}VIE-260B

v34184

w¹¹¹⁸; P{GD10585}v34184/TM3

More stocks available...

Genomic Clones (29)

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

cDNA Clones (89)

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.

FBgn0013759

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0013759

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Chen and Featherstone, 2011, Sun et al., 2009, Lu et al., 2003, Martin and Ollo, 1996)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

CASK function is required in the α'/β' subset of the mushroom body neurons in the adult for middle-term, anesthesia-resistant and long-term memory formation.

(Malik et al., 2013)

Caki enhances the eag potassium current by a phosphorylation-dependent mechanism.

(Marble et al., 2005)

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)

RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a phenotype when assayed in S2R⁺ cells: cells become round and detached. Kc167 cells are unaffected.

(Kiger et al., 2003)

Cloning and characterisation of Caki.

(Dimitratos et al., 1997)

The Caki protein sequence is conserved in the catalytic and calmodulin-binding domains with other type II CaM kinases. In cultured cells Caki has the ability to stimulate the transcriptional activity of the Rous sarcoma virus long terminal repeat (RSV LTR).

(Martin and Ollo, 1996)

Mutant flies, in which an encoding part of the gene is removed, display a substantial reduction in walking speed.

(Martin and Ollo, 1996)

Expressed in nurse cells.

(Martin et al., 1994)

Relationship to Other Genes

Source for database merge of

Source for merge of: Caki Cask

(Verstreken, 2005.7.11)

Source for merge of: Caki CG13413

(FlyBase Genome Annotators, 2002-2003)

Source for merge of: Caki cmg

(Dimitratos et al., 1997)

Additional comments

mir-999 is located within CASK.

(Stark et al., 2007)

Annotations CG6703, CG13412 and CG13413 merged as CG6703 in release 3 of the genome annotation.

(FlyBase Genome Annotators, 2002-2003)

Nomenclature History

Source for database identify of

Source for identity of: CASK Caki

(Griffith et al., 2010.9.7)

Nomenclature comments

Etymology

Synonyms and Secondary IDs (28)

Reported As

Symbol Synonym

CAKI

(Sun et al., 2009, Zordan et al., 2005, Bellen, 2000.12.25, Lloyd et al., 2000, Morrison et al., 2000)

CAMGUK

(Zordan et al., 2005)

CASK

(Tsakiri et al., 2025, Bhandari et al., 2024, Chvilicek et al., 2024, Loreto et al., 2024, Peng et al., 2024, Pribbenow and Owald, 2024, Garcia-Vaquero et al., 2023, Bourouliti and Skoulakis, 2022, Ecovoiu et al., 2022, Marcogliese et al., 2022, Andrew et al., 2021, Du et al., 2021, Hou and Pei, 2021, Kamiyama et al., 2021, Ostalé et al., 2021, Pang et al., 2021, Park et al., 2021, Salim et al., 2021, Silva et al., 2020, FlyBase Genome Annotators, 2019-, Hall et al., 2019, Burow et al., 2018, Wang et al., 2018, Lakhotia, 2017, Park et al., 2017, Rohde et al., 2017, Transgenic RNAi Project members, 2017-, Gene Disruption Project members, 2016-, Wang et al., 2016, Zhang et al., 2016, Gene Disruption Project members, 2015-, Grotewiel and Bettinger, 2015, Nagarkar-Jaiswal et al., 2015, Mukherjee et al., 2014, Mukherjee et al., 2014, Slawson et al., 2014, Sopko et al., 2014, Gillespie and Hodge, 2013, Malik et al., 2013, Soukup et al., 2013, Tian et al., 2013, Donelson et al., 2012, Li et al., 2012, Guan et al., 2011, Slawson et al., 2011, Griffith et al., 2010.9.7, Lloyd et al., 2000)

CASK-β

(Mukherjee et al., 2014)

CG13412

(Morrison et al., 2000, Verstreken and Bellen, 2000.9.4)

CG13413

(Whitfield, 2000.4.4)

CG6703

(Neely et al., 2011, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009, Morrison et al., 2000, Verstreken and Bellen, 2000.9.4)

CMG

(Sun et al., 2009, Marble et al., 2005)

CMG/CASK

(Morrison et al., 2000)

CaSK

(Vajente et al., 2020)

Caki

(Muhammad et al., 2015, Ashwal-Fluss et al., 2014, Maiya et al., 2012, Smibert et al., 2012, Guan et al., 2011, Jungreis et al., 2011, Neely et al., 2011, Bakal et al., 2008, Adryan et al., 2007, Stark et al., 2007, Wu et al., 2007, Wang et al., 2004)

CamGUK

(Ponting et al., 1997)

CamguK

(Guan et al., 2011)

Camguk

(Mukherjee et al., 2014, Maiya et al., 2012, Marble et al., 2005)

Cask

(Mukherjee et al., 2014)

Cmg

(Lu et al., 2003)

DLin-2

(Bachmann et al., 2004)

RE09582p

(Chen et al., 2007)

caki

(Guangming et al., 2020, Mukherjee et al., 2014, Maiya et al., 2012, Southall and Brand, 2009, Peng et al., 2007, Zordan et al., 2005, Mehren et al., 2004, Megighian et al., 2002, Martin and Ollo, 1996, Martin, 1995.12.14)

camguk

(Zordan et al., 2005)

cask

(Qiu et al., 2025, Cavieres-Lepe and Ewer, 2021, Chen et al., 2014)

cmg

(Bachmann et al., 2010, Lopes et al., 2001, Whitfield, 2000.4.4, Whitfield, 2000.7.14, Dimitratos et al., 1999, Dimitratos et al., 1997, Dimitratos et al., 1997, Dimitratos et al., 1997, Dimitratos, 1996.3.31, Dimitratos et al., 1996)

dCASK

(Lo Piccolo, 2018, Hodge et al., 2006)

Name Synonyms

CASK

CASK ortholog

(Aranjuez et al., 2012)

Calcium/calmodulin-dependent protein kinase

(Guan et al., 2011, Martin et al., 1994)

Camguk

(Neely et al., 2011, Bachmann et al., 2010, Colbran and Brown, 2004, de Lorenzo et al., 2003, Humbert et al., 2003, Lu et al., 2003, Caruana, 2002, Lee et al., 2002, de Lorenzo et al., 2001, Lee et al., 2001, Doerks et al., 2000, Dimitratos, 1996.3.31)

Camguk/CASK adaptor protein

(Marble et al., 2005)

Cask ortholog

(Wang et al., 2016)

calmodulin-dependent kinase

(Harris and Lim, 2001)

camguk

(Mehren et al., 2004, MacMullin et al., 2001, Dimitratos et al., 1997, Dimitratos et al., 1996)

Secondary FlyBase IDs

FBgn0015923
FBgn0038904
FBgn0043003

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 91 )

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.

42567

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.

Q24210

UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information

Q24210

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.

Q24210

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0013759

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

42567/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0013759

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0013759

FlyMine - An integrated database for Drosophila genomics

FBgn0013759

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

dme:Dmel_CG6703

MARRVEL_MODEL - MARRVEL (model organism gene)

42567

Linkouts

BioGRID - A database of protein and genetic interactions.

67530

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0013759

DroID - A comprehensive database of gene and protein interactions.

FBgn0013759

DRSC - Results frm RNAi screens

FBgn0013759

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0013759

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0013759

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG6703

Flygut - An atlas of the Drosophila adult midgut

FBgn0013759

FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.

FBgn0013759

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC002673

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

42567

MIST (protein-protein) - An integrated Molecular Interaction Database

42567

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

Q24210

References (171)

The following external sites may use different assemblies or annotations than FlyBase.

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