FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\Dark
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General Information
Symbol
Dmel\Dark
Species
D. melanogaster
Name
Death-associated APAF1-related killer
Annotation Symbol
CG6829
Feature Type
protein_coding_gene
FlyBase ID
FBgn0263864
Gene Model Status
Current
Stock Availability
13 publicly available
Gene Summary
Death-associated APAF1-related killer (Dark) encodes an essential component of the apoptosome. The products of Dark and Dronc form a pivotal holoenzyme required for apoptotic cell death with the products of Dark acting as an adaptor of the multimeric complex. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

ark, hac-1, dapaf-1, Apaf-1, Apaf-1-related killer

Key Links
Genomic Location
Cytogenetic map
53E4-53E4
Sequence location
Recombination map
2-81
RefSeq locus
NT_033778 REGION:17020027..17026703
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 (24 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
enables ADP binding
inferred from direct assay
(Tian et al., 2024)
enables ATP binding
inferred from direct assay
(Yuan et al., 2011)
enables protein binding
inferred from physical interaction with UniProtKB:Q9XYF4
(Quinn et al., 2000)
enables structural molecule activity
inferred from direct assay
(Yuan et al., 2011)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables ADP binding
inferred from electronic annotation with InterPro:IPR002182
(InterPro Project Members, 2004-)
enables CARD domain binding
inferred from sequence model
(Yuan et al., 2011)
Biological Process (18 terms)
Terms Based on Experimental Evidence (18 terms)
CV Term
Evidence
References
involved_in apoptotic process
inferred from genetic interaction with FLYBASE:hid; FB:FBgn0003997
(Srivastava et al., 2007)
inferred from direct assay
(Zhou et al., 1999)
involved_in autophagic cell death
inferred from mutant phenotype
(Mills et al., 2006)
involved_in central nervous system formation
inferred from mutant phenotype
(Mills et al., 2006)
involved_in chaeta development
inferred from mutant phenotype
(Kanuka et al., 2005)
involved_in compound eye retinal cell programmed cell death
inferred from mutant phenotype
(Leulier et al., 2006)
involved_in dendrite morphogenesis
inferred from mutant phenotype
(Williams et al., 2006)
involved_in melanization defense response
inferred from mutant phenotype
(Obata et al., 2014)
involved_in negative regulation of humoral immune response
inferred from mutant phenotype
(Obata et al., 2014)
involved_in neuron cellular homeostasis
inferred from genetic interaction with FLYBASE:Ank2; FB:FBgn0261788
(Keller et al., 2011)
involved_in positive regulation of apoptotic process involved in development
inferred from mutant phenotype
(Quinn et al., 2000)
involved_in positive regulation of glial cell apoptotic process
inferred from mutant phenotype
(Srivastava et al., 2007)
involved_in programmed cell death
inferred from direct assay
(Mills et al., 2006)
involved_in response to gamma radiation
inferred from direct assay
(Mills et al., 2006)
involved_in response to starvation
inferred from mutant phenotype
(Obata et al., 2014)
involved_in S-adenosylmethionine cycle
inferred from mutant phenotype
(Obata et al., 2014)
involved_in salivary gland histolysis
inferred from mutant phenotype
(Akdemir et al., 2006, Leulier et al., 2006)
involved_in sperm individualization
inferred from mutant phenotype
(Arama et al., 2006, Huh et al., 2004)
involved_in triglyceride homeostasis
inferred from mutant phenotype
(Obata et al., 2014)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
part_of apoptosome
inferred from direct assay
(Yuan et al., 2011)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Death-associated APAF1-related killer (Dark) encodes an essential component of the apoptosome. The products of Dark and Dronc form a pivotal holoenzyme required for apoptotic cell death with the products of Dark acting as an adaptor of the multimeric complex. [Date last reviewed: 2019-03-07]
Gene Group (FlyBase)
APOPTOSOME -
The apoptosome is a ring-like multisubunit complex assembled upon receiving a pro-apoptotic signal. In Drosophila, Dark assembles into two stacked rings of eight subunits each. This scaffold binds ADP and an initiator caspase (Dronc in D.mel) which cleaves effector caspase(s). (Adapted from FBrf0212736).
Protein Function (UniProtKB)
Main component of the apoptosome, an adapter protein complex that mediates activation of the caspase cascade in programmed cell death initiated by the intrinsic apoptosis pathway (PubMed:10984473, PubMed:16310803, PubMed:21220123, PubMed:25644603, PubMed:27916517). Recruits the initiator caspase Dronc, promoting its autocatalytic cleavage and activation; triggers the caspase cascade upstream of the effector caspases Drice and dcp-1 (PubMed:10619022, PubMed:10619023, PubMed:10984473, PubMed:21220123, PubMed:25644603). Effector of reaper, grim and hid induced cell killing (PubMed:10559939, PubMed:10619023, PubMed:16645639). Produces at least two isoforms that may activate distinct caspases by separate pathways (PubMed:10619023). Nucleotide binding protein with strong specificity for dATP; dATP binding is not essential, but promotes homooligomerization to form the apoptosome complex (PubMed:10619022, PubMed:10619023, PubMed:16310803, PubMed:21220123, PubMed:25644603, PubMed:27916517). There is currently no evidence of dATPase activity and structural analysis of the nucleotide binding pocket suggests negligible to no dATPase activity (Probable). Caspase activation by the apoptosome was initially thought to be dependent on binding of cytochrome c released from mitochondria, as is the case for vertebrate orthologs (PubMed:10619022, PubMed:10619023). Apoptosis induction by the Dark apoptosome appears to be cytochrome c independent (PubMed:11901172). Required for stress-induced apoptosis, for example in response to radiation (UV or gamma) or treatment with macromolecular synthesis inhibitors like cycloheximide and actinomycin D (PubMed:11901172, PubMed:16533943). Required for most, but not all programmed cell death during embryonic development (PubMed:16645639). Involved in developmental cell number regulation by apoptosis, particularly during neuronal and sensory organ development (PubMed:10559939, PubMed:10619022, PubMed:10619023, PubMed:16533943, PubMed:16645639). Involved in reduction of midline glial cell numbers by programmed cell death during embryogenesis (PubMed:16645639). During metamorphosis involved in removal of larval salivary glands by programmed cell death, but not of the larval midgut (PubMed:16533943).
(UniProt, Q7KLI1)
Summary (Interactive Fly)

CED-4 domain protein that links mitochondrial events to activation of caspases during programd cell death

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

Please see the JBrowse view of Dmel\Dark 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 Q7KLI1)

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
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

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

Gene model reviewed during 5.50

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Dark-RA
FBtr0086967
NM_166208
4883
531
Dark-RB
FBtr0086968
NM_166207
4824
1440
Additional Transcript Data and Comments
Reported size (kB)

7.0 (northern blot)

(Kanuka et al., 1999)

5 (northern blot)

(Rodriguez et al., 1999)

5.5 (unknown)

(Zhou et al., 1999)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Dark-PA
FBpp0086121
62.3
531
8.15
Q9U8R4
NP_725638
AAM68488
Dark-PB
FBpp0086122
165.6
1440
6.08
Q7KLI1
NP_725637
AAF57916
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)

1440, 531 (aa)

(Kanuka et al., 1999)

1440 (aa)

(Rodriguez et al., 1999, Zhou et al., 1999)
Comments
External Data
Subunit Structure (UniProtKB)

Exists as an auto-inhibited monomer; this state is maintained by intramolecular interactions between the NB-ARC and WD repeat domains (PubMed:25644603, PubMed:38381783). In vitro, at high protein concentrations or in the presence of non-physiologically high dATP concentrations, assembles into homooctameric rings 300A in diameter to form the apoptosome complex; two rings can interact face to face (via CARD domains with some involvement of the WD repeat beta-propellors) to form a double ring structure (PubMed:16310803, PubMed:21220123, PubMed:25644603, PubMed:27916517). The octameric ring may represent a non-functional or off-pathway conformation (Probable). Under physiological conditions oligomerization results in formation of a single unsealed ring or partial helical spiral structure; this structure may be an assembly intermediate (PubMed:38381783). Apoptosome assembly is stimulated by dATP binding (PubMed:16310803, PubMed:25644603, PubMed:38381783). Interacts (via CARD and WD domains) with the initiator caspase Dronc (via CARD domain); the interaction stimulates Dark oligomerization to form the apoptosome and brings pairs of Dronc molecules together on the apoptosome to facilitate their dimerization and activation by autocatalytic cleavage (PubMed:10984473, PubMed:21220123, PubMed:25644603, PubMed:38381783). Assembly of the apoptosome does not require cytochrome c binding (PubMed:16310803, PubMed:25644603). After autocatalytic cleavage Dronc caspase domains dissociates from the apoptosome but the CARD domain remains associated (PubMed:25644603). Interacts (via N-terminus) with Dredd (PubMed:10559939).

(UniProt, Q7KLI1)
Post Translational Modification

May be AMPylated on Lys-251.

(UniProt, Q7KLI1)
Domain

Possesses an N-terminal caspase activation and recruitment domain (CARD) (PubMed:16310803, PubMed:21220123, PubMed:25644603, PubMed:27916517). At high protein concentrations in vitro the CARD domain mediates the interaction of two octameric apoptosome rings to form a double ring structure (PubMed:21220123).

Possesses a NB-ARC domain (nucleotide-binding adapter shared by APAF-1, certain R gene products and CED-4) that forms the inner hub of the apoptosome ring and is the main mediator of oligomerization (PubMed:21220123, PubMed:25644603, PubMed:27916517). The interface of the NB-ARC domain and a neighboring helical region forms a binding pocket that can accommodate a nucleotide tri-phosphate with specificity for dATP (PubMed:21220123, PubMed:27916517). In its monomeric non-activated state the binding pocket may be associated with ADP (PubMed:38381783).

(UniProt, Q7KLI1)
Crossreferences
InterPro - A database of protein families, domains and functional sites
PDB - An information portal to biological macromolecular structures
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\Dark 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).

-0.81

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4
organism | ubiquitous
(Zhou et al., 1999)
embryonic stage 4 -- 6
organism | faint | ubiquitous
(Fisher et al., 2012)
embryonic stage 7
ventral neurectoderm
(Zhou et al., 1999)
anterior transverse furrow & cephalic furrow & posterior transverse furrow & ventral furrow
(Zhou et al., 1999)
embryonic stage 7 -- 8
organism | faint | ubiquitous
(Fisher et al., 2012)
embryonic stage 9 -- 10
organism | faint | ubiquitous
(Fisher et al., 2012)
embryonic stage 10 -- 11
procephalic neurogenic region
(Zhou et al., 1999)
head mesoderm
(Zhou et al., 1999)
embryonic stage 11 -- 12
head mesoderm
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
embryonic stage 13 -- 16
organism | faint | ubiquitous
(Fisher et al., 2012)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Kanuka et al., 1999)
embryonic stage & larval stage & pupal stage
(Kanuka et al., 1999)
embryonic stage -- adult stage
(Rodriguez et al., 1999)
embryonic stage 1 -- 17
organism
(Rodriguez et al., 1999)
third instar larval stage
organism
(Rodriguez et al., 1999)
eye disc
(Kanuka et al., 1999)
wing disc
(Kanuka et al., 1999)
prepupal stage
organism
(Rodriguez et al., 1999)
Additional Descriptive Data

The longer form of Dark transcript appears to be the predominantly expressed form. It is detected in embyros, larvae, pupae, and S2 cells on northern blots. It is expressed ubiquitously in embryos and in larval eye and wing discs.

(Kanuka et al., 1999)

The short form of Dark transcript appears to be the minor form which is detected in embryos.

(Kanuka et al., 1999)

Dark transcripts are detected at all stages assayed on northern blots. Levels are elevated in times associated with histolysis of larval tissues (third instar larvae and early pupae). Levels are lowest in embryos and adults.

(Rodriguez et al., 1999)

Dark transcripts are broadly distributed in preblastoderm embryos. By stage 7, transcripts are predominantly located in the ventral neurogenic region and around several invagination furrows. In stages 10-11, the highest Dark transcript levels are found in the ectoderm and mesoderm of the procephalic region. Later, expression is observed in a segmentally repeated pattern. Dark expression is induced by X-ray or UV irradiation.

(Zhou et al., 1999)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
part_of apoptosome
inferred from direct assay
(Yuan et al., 2011)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Dark 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
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
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
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 47 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 17 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Dark
Transgenic constructs containing regulatory region of Dark
Aberrations (Deficiencies and Duplications) ( 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
lethal (with Dark82)
lethal (with Darke03004)
lethal (with Df(2R)BSC359)
lethal | pupal stage
lethal | recessive
lethal - all die before end of pupal stage
lethal - all die before end of pupal stage, with Scer\GAL4αTub84B.PL
lethal - all die before end of pupal stage (with Dark1)
lethal - all die before end of pupal stage (with Dark2)
lethal - all die before end of pupal stage (with Dark3)
lethal - all die before end of pupal stage (with Dark4)
lethal - all die before end of pupal stage | partially
lethal - all die before end of pupal stage | recessive
lethal - all die during embryonic stage | recessive
partially lethal
partially lethal - majority die | larval stage
partially lethal - majority die | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable (with DarkL46)
viable (with DarkP46)
Sterility
Allele
female sterile
male sterile
Other Phenotypes
Allele
abnormal cell death
abnormal cell death, with Scer\GAL4Hml.PG
abnormal cell death (with Dark82)
abnormal cell death (with DarkCD4)
abnormal cell death | partially
abnormal cell death | somatic clone
abnormal developmental rate
abnormal immune response, with Scer\GAL4GMR11F02
abnormal neuroanatomy
abnormal neuroanatomy | adult stage
abnormal neuroanatomy | adult stage, with Scer\GAL4CCAP.PP
abnormal neuroanatomy | adult stage (with Dark82)
abnormal neuroanatomy | adult stage (with DarkCD4)
abnormal neuroanatomy | larval stage
abnormal size | third instar larval stage
abnormal starvation stress response
decreased cell death
decreased cell death | adult stage
decreased cell death | adult stage, with Scer\GAL4CCAP.PP
decreased cell death | adult stage (with Dark82)
decreased cell death | adult stage (with DarkCD4)
decreased cell death | conditional
decreased cell death | embryonic stage
decreased cell death | embryonic stage | germline clone
decreased cell death | embryonic stage | recessive
decreased cell death | recessive
decreased fecundity | male | recessive
decreased fecundity | recessive
increased cell death (with Dark82)
increased cell death (with DarkN28)
increased cell death (with DarkP46)
increased cell number | adult stage, with Scer\GAL4esg.PU
melanotic mass phenotype (with DarkL46)
melanotic mass phenotype (with DarkP46)
melanotic mass phenotype | recessive
melanotic necrosis
melanotic necrosis, with Scer\GAL4Bx-MS1096
melanotic necrosis, with Scer\GAL4GMR11F02
melanotic necrosis | adult stage, with Ncra\QFQF2.R11F02
radiation resistant
short lived, with Ncra\QFQF2.R11F02
some die during embryonic stage | recessive
some die during pupal stage
some die during pupal stage, with Scer\GAL4αTub84B.PL
some die during pupal stage (with Dark1)
some die during pupal stage (with Dark2)
some die during pupal stage (with Dark3)
some die during pupal stage (with Dark4)
some die during pupal stage | partially
some die during pupal stage | recessive
visible
visible | recessive
Phenotype manifest in
Allele
adult gut, with Scer\GAL4esg.PU
anterior scutellar bristle
chaeta | increased number, with Scer\GAL4αTub84B.PL
chordotonal organ precursor cell
cystic bulge
dendrite
embryo
embryo | dorsal closure stage
embryonic/larval salivary gland
embryonic brain
embryonic epidermis
embryonic head
embryonic neuroblast
eo neuron | increased number
eye disc
eye photoreceptor cell
imaginal disc epithelial cell | somatic clone
interommatidial precursor cell | pharate adult stage P9 | increased number, with Scer\GAL4GMR.PF
larval brain
larval central nervous system
larval digestive system
larval digestive system (with Darkk11502)
larval digestive system (with DarkP803-Δ15)
larval digestive system (with Df(2R)BSC49)
larval dorsal multidendritic neuron ddaC
larval ventral nerve cord
larval ventral nerve cord | larval stage
larval ventral nerve cord commissure
macrochaeta
macrochaeta | increased number
male germline cell (with Dark82)
male germline cell (with DarkN28)
male germline cell (with DarkP46)
melanotic mass
melanotic mass (with Darkk11502)
melanotic mass (with DarkP803-Δ15)
melanotic mass (with Df(2R)BSC49)
midline glial cell | ectopic
midline glial cell | increased number
nucleus
ommatidium
ommatidium | cell autonomous | somatic clone
peptidergic neuron | adult stage | ectopic
peptidergic neuron | adult stage | ectopic, with Scer\GAL4CCAP.PP
peptidergic neuron | adult stage | ectopic (with Dark82)
peptidergic neuron | adult stage | ectopic (with DarkCD4)
pigment cell
posterior scutellar bristle
posterior scutellar bristle | increased number
presumptive embryonic/larval central nervous system
salivary gland
salivary gland | pupal stage, with Scer\GAL434B
scutellar bristle | ectopic
sensory mother cell | increased number
spermatid
thecogen cell | increased number
waste bag
wing
wing, with Ncra\QFQF2.R11F02
wing, with Scer\GAL4Bx-MS1096
wing, with Scer\GAL4GMR11F02
wing, with Scer\GAL4αTub84B.PL
wing (with DarkL46)
wing (with DarkP46)
wing | progressive
wing | somatic clone
wing disc
wing disc | third instar larval stage
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) (1)
Hsap\APAF1
4 of 14
Yes
Yes
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) (1)
Rnor\Apaf1
4 of 14
Yes
Yes
Mus musculus (laboratory mouse) (1)
Mmus\Apaf1
2 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (1)
Xtro\apaf1
2 of 13
Yes
Yes
Danio rerio (Zebrafish) (1)
Drer\apaf1
4 of 14
Yes
Yes
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP010737
6 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (49)
Atha\AT1G12290
1 of 13
Yes
Yes
Atha\AT1G15890
1 of 13
Yes
Yes
Atha\AT1G50180
1 of 13
Yes
Yes
Atha\AT1G51480
1 of 13
Yes
Yes
Atha\AT1G52660
1 of 13
Yes
Yes
Atha\AT1G53350
1 of 13
Yes
Yes
Atha\AT1G58390
1 of 13
Yes
Yes
Atha\AT1G58400
1 of 13
Yes
Yes
Atha\AT1G58410
1 of 13
Yes
Yes
Atha\AT1G58602
1 of 13
Yes
Yes
Atha\AT1G58807
1 of 13
Yes
Yes
Atha\AT1G58848
1 of 13
Yes
Yes
Atha\AT1G59124
1 of 13
Yes
Yes
Atha\AT1G59218
1 of 13
Yes
Yes
Atha\AT1G59780
1 of 13
Yes
Yes
Atha\AT1G61180
1 of 13
Yes
Yes
Atha\AT1G61190
1 of 13
Yes
Yes
Atha\AT1G61300
1 of 13
Yes
Yes
Atha\AT1G61310
1 of 13
Yes
Yes
Atha\AT1G62630
1 of 13
Yes
Yes
Atha\AT1G63350
1 of 13
Yes
Yes
Atha\AT1G63360
1 of 13
Yes
Yes
Atha\AT3G14460
1 of 13
Yes
Yes
Atha\AT3G14470
1 of 13
Yes
Yes
Atha\AT3G15700
1 of 13
Yes
Yes
Atha\AT3G46710
1 of 13
Yes
Yes
Atha\AT3G46730
1 of 13
Yes
Yes
Atha\AT4G10780
1 of 13
Yes
Yes
Atha\AT4G16095
1 of 13
Yes
Yes
Atha\AT4G27190
1 of 13
Yes
Yes
Atha\AT4G27220
1 of 13
Yes
Yes
Atha\AT5G05400
1 of 13
Yes
Yes
Atha\AT5G35450
1 of 13
Yes
Yes
Atha\AT5G43730
1 of 13
Yes
Yes
Atha\AT5G43740
1 of 13
Yes
Yes
Atha\AT5G47250
1 of 13
Yes
Yes
Atha\AT5G47260
1 of 13
Yes
Yes
Atha\AT5G48620
1 of 13
Yes
Yes
Atha\AT5G63020
1 of 13
Yes
Yes
Atha\CW9
1 of 13
Yes
Yes
Atha\LOV1
1 of 13
Yes
Yes
Atha\RFL1
1 of 13
Yes
Yes
Atha\RPM1
1 of 13
Yes
Yes
Atha\RPP8
1 of 13
Yes
Yes
Atha\RPP13
1 of 13
Yes
Yes
Atha\RPS2
1 of 13
Yes
Yes
Atha\RPS5
1 of 13
Yes
Yes
Atha\SUMM2
1 of 13
Yes
Yes
Atha\ZAR1
1 of 13
Yes
Yes
Saccharomyces cerevisiae (Brewer's yeast) (0)
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Dark. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
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 ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 4 )
    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?
    APAF1; apoptotic peptidase activating factor 1
    4 of 14
    602233
        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
        Dark
        enhanceable
        Debcl
        (Galindo et al., 2009)
        Dark
        enhanceable
        Dronc
        (Chew et al., 2004)
        Dark
        suppressible
        foxo
        (Obata et al., 2014)
        Dark
        enhanceable
        Gnmt
        (Obata et al., 2014)
        Dark
        suppressible
        pcm
        (Waldron et al., 2015)
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Ank2
        suppressible
        Dark
        (Keller et al., 2011)
        Dcp-1
        suppressible
        Dark
        (Sinenko, 2017, Zhou et al., 1999)
        Debcl
        suppressible
        Dark
        (Colussi et al., 2000)
        Diap1
        suppressible
        Dark
        (Sinenko, 2017, Leulier et al., 2006, Cullen and McCall, 2004, Rodriguez et al., 2002)
        Diap1|hid
        suppressible
        Dark
        (Rodriguez et al., 2002)
        Dp
        suppressible
        Dark
        (Moon et al., 2005)
        Dronc
        enhanceable
        Dark
        (Domingues and Ryoo, 2012, Verghese et al., 2012)
        Dronc
        suppressible
        Dark
        (Sinenko, 2017, Quinn et al., 2000)
        Dyro
        suppressible
        Dark
        (Shima et al., 2025)
        E2f1
        suppressible
        Dark
        (Morris et al., 2006)
        egr
        suppressible
        Dark
        (Moreno et al., 2002)
        Gnmt
        enhanceable
        Dark
        (Obata et al., 2014)
        grim
        suppressible
        Dark
        (Sinenko, 2017, Rodriguez et al., 1999)
        grim|Diap1
        suppressible
        Dark
        (Rodriguez et al., 2002)
        hid
        suppressible
        Dark
        (Srivastava et al., 2007, Leulier et al., 2006, Rodriguez et al., 1999)
        hpo
        enhanceable
        Dronc|Dark
        (Verghese et al., 2012)
        hpo|Dronc
        enhanceable
        Dark
        (Verghese et al., 2012)
        Idh
        suppressible
        Dark
        (Reitman et al., 2015)
        IKKε
        enhanceable
        Dark
        (Oshima et al., 2006)
        mad2|bsk
        suppressible
        Dark
        (Wong et al., 2014)
        ninaE
        suppressible
        Dark
        (Griciuc et al., 2014)
        p53
        suppressible
        Dark
        (Fan et al., 2010)
        pcm
        suppressible
        Dark
        (Waldron et al., 2015)
        Rac1
        suppressible
        Dark
        (Geisbrecht and Montell, 2004)
        rpr
        suppressible
        Dark
        (Sinenko, 2017, Kanuka et al., 1999, Rodriguez et al., 1999)
        rpr
        suppressible
        Usp15-31|Dark
        (Sinenko, 2017)
        rpr
        suppressible
        Usp8|Dark
        (Sinenko, 2017)
        synr
        suppressible
        Dark
        (Sulekh et al., 2024)
        YME1L
        suppressible
        Dark
        (Qi et al., 2016)
        External Data
        Subunit Structure (UniProtKB)
        Exists as an auto-inhibited monomer; this state is maintained by intramolecular interactions between the NB-ARC and WD repeat domains (PubMed:25644603, PubMed:38381783). In vitro, at high protein concentrations or in the presence of non-physiologically high dATP concentrations, assembles into homooctameric rings 300A in diameter to form the apoptosome complex; two rings can interact face to face (via CARD domains with some involvement of the WD repeat beta-propellors) to form a double ring structure (PubMed:16310803, PubMed:21220123, PubMed:25644603, PubMed:27916517). The octameric ring may represent a non-functional or off-pathway conformation (Probable). Under physiological conditions oligomerization results in formation of a single unsealed ring or partial helical spiral structure; this structure may be an assembly intermediate (PubMed:38381783). Apoptosome assembly is stimulated by dATP binding (PubMed:16310803, PubMed:25644603, PubMed:38381783). Interacts (via CARD and WD domains) with the initiator caspase Dronc (via CARD domain); the interaction stimulates Dark oligomerization to form the apoptosome and brings pairs of Dronc molecules together on the apoptosome to facilitate their dimerization and activation by autocatalytic cleavage (PubMed:10984473, PubMed:21220123, PubMed:25644603, PubMed:38381783). Assembly of the apoptosome does not require cytochrome c binding (PubMed:16310803, PubMed:25644603). After autocatalytic cleavage Dronc caspase domains dissociates from the apoptosome but the CARD domain remains associated (PubMed:25644603). Interacts (via N-terminus) with Dredd (PubMed:10559939).
        (UniProt, Q7KLI1 )
        Linkouts
        DroID - A comprehensive database of gene and protein interactions.
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Pathways
        Signaling Pathways (FlyBase)
        Metabolic Pathways
        FlyBase
        External Links
        External Data
        Linkouts
        KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
        Class of Gene
        Genomic Location and Detailed Mapping Data
        Chromosome (arm)
        2R
        Recombination map
        2-81
        Cytogenetic map
        53E4-53E4
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        53E4-53E4
        Limits computationally determined from genome sequence between P{lacW}l(2)k09837k09837 and P{lacW}l(2)k11405k11405&P{EP}EP641
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        54A2-54A2
        (Oh et al., 2003)
        53F-53F
        (Kanuka, 1999.5.20, Kanuka et al., 1999, Zhou, 1999.10.19)
        53F1-53F1
        (Rodriquez, 1999.6.25)
        53F1-53F2
        (BDGP Project Members, 1994-1999)
        Experimentally Determined Recombination Data
        Location

        2-81

        (FlyBase, 2016)

        2-84.9

        (Comeron et al., 2012)
        Left of (cM)
        Right of (cM)
        Notes
        Stocks and Reagents
        Stocks (13)
        Genomic Clones (22)
        cDNA Clones (20)
         

        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
          Antibody Information
          Laboratory Generated Antibodies
           
          Commercially Available Antibodies
           
          Cell Line Information
          Publicly Available Cell Lines
           
            Other Stable Cell Lines
             
              Other Comments

              Gene expression is increased in response to the presence of two copies of Scer\GAL4hs.PB.

              (Liu and Lehmann, 2008)

              Ark mutant hemocytes are completely resistant to a Smac mimetic (which antagonises inhibitor of apoptosis proteins) or cycloheximide.

              (Akdemir et al., 2006)

              Expression of Ark in the post-embryonic central nervous system is necessary to reverse organismal lethality and to produce a viable adult.

              (Akdemir et al., 2006)

              Ark is required for programmed cell death.

              (Akdemir et al., 2006)

              Removal of Ark results in profound developmental defects and lethality.

              (Mills et al., 2006)

              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)

              1 allele of l(2)SH0173 recovered in a P-insertion screen.

              (Oh et al., 2003)

              Loss of Ark function results in pronounced inhibition of stress-induced apoptosis, but not rpr or grim induced apoptosis.

              (Zimmermann et al., 2002)

              Loss of function mutations of Ark show reduced cell death and caspase activities in embryos.

              (Kanuka et al., 1999)

              Loss of function mutations in Ark attenuate programmed cell deaths during development, resulting in hyperplasia of the central nervous system and other abnormalities including melanotic tumours and defective wings.

              (Rodriguez et al., 1999)

              Ark is required for normal cell death during embryonic development.

              (Zhou et al., 1999)

              Loss of Ark function causes reduced cell death.

              (Zhou et al., 1999)

              Transcription unit identified during molecular analysis of the RhoGEF2 region.

              (Hacker and Perrimon, 1998)
              Relationship to Other Genes
              Source for database merge of

              Source for merge of: Ark anon-53Fa

              (Rodriguez, 1999.7.30)

              Source for merge of: Ark Hac1

              (FlyBase, 1996-)

              Source for merge of: l(2)SH0173 Ark

              (Link et al., 2007)
              Additional comments

              Source for merge of Ark Hac1 was sequence comparison ( date:991227 ).

              (FlyBase, 1996-)
              Nomenclature History
              Source for database identify of

              Source for identity of: Dark Ark

              (Steller et al., 2014.7.9)
              Nomenclature comments

              Changed from 'Ark' to 'Dark' to reflect common usage in the literature.

              (Steller et al., 2014.7.9)
              Etymology
              Synonyms and Secondary IDs (52)
              Reported As
              Symbol Synonym
              APAF-1
              (Griciuc et al., 2014)
              APAF1
              (Chew et al., 2009)
              ARK
              (Arya and White, 2015, Bergmann, 2010, Reddy et al., 2010, Cakouros et al., 2008, Srivastava et al., 2007, Zimmermann et al., 2002)
              Apaf
              (Reitman et al., 2015, Wong et al., 2014)
              Apaf-1
              (Waldron et al., 2015, Fabian and Brill, 2012, Keller et al., 2011, Yuan et al., 2011, Pret et al., 2008, Arama et al., 2006)
              Apaf1
              (Reitman et al., 2015, Domingues and Ryoo, 2008)
              Ark
              (Yuan et al., 2019, Mondal et al., 2018, Nakajima and Kuranaga, 2017, Dabrowska et al., 2016, Denton and Kumar, 2015, Waldron et al., 2015, Wang et al., 2015, Ihry and Bashirullah, 2014, Shklyar et al., 2013, Yacobi-Sharon et al., 2013, Chakrabarti et al., 2012, Hazelett et al., 2012, Ihry et al., 2012, Izutsu et al., 2012, Kagey et al., 2012, Suissa et al., 2011, Xu et al., 2009, Dorstyn and Kumar, 2008, Hou et al., 2008, Liu and Lehmann, 2008, Bodai et al., 2007, Link et al., 2007, Martin et al., 2007, Minakhina et al., 2007, Sevrioukov et al., 2007, Minakhina and Steward, 2006)
              CG6829
              (National Institute of Genetics Fly Stocks, 2022-, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Lee et al., 2003)
              D-Apaf-1
              (Bergmann et al., 2003)
              DARK
              (Scopelliti et al., 2025, Sentürk et al., 2020, Koerver et al., 2016, Reitman et al., 2015, Ziraldo and Ma, 2015, Adrain, 2006, Akdemir et al., 2006, Oshima, 2006, Oshima et al., 2006, Dorstyn et al., 2004, Moreno and Basler, 2003, Martin, 2002, Moreno et al., 2002, Hu and Yang, 2000, Rodriquez, 1999.6.25)
              Dapaf-1
              (Dick and Megeney, 2013, Arama et al., 2006, Mendes et al., 2006, Baehrecke, 2003, Igaki et al., 2002)
              Dapaf-1/HAC-1
              (Kanuka et al., 2005)
              Dark
              (Hounsell and Fan, 2025, Patterson et al., 2025, Sproston et al., 2025, Xiu et al., 2025, Yadav et al., 2025, Farodoye et al., 2024, Jang et al., 2024, Lacin et al., 2024, Sulekh et al., 2024, Tian et al., 2024, Umargamwala et al., 2024, Cordes et al., 2023, Heigwer et al., 2023, Huang et al., 2023, Nakano et al., 2023, Nowaskie et al., 2023, Ciesielski et al., 2022, Dziedziech and Theopold, 2022, Mast et al., 2022, Moore et al., 2022, Tarikere et al., 2022, Xu et al., 2022, Buhlman et al., 2021, Hounsell and Fan, 2021, Vrailas-Mortimer et al., 2021, Fujisawa et al., 2020, Kosakamoto et al., 2020, Nakajima et al., 2020, Xu et al., 2020, Bieser et al., 2019, Meltzer et al., 2019, Stamm et al., 2019, Tettamanti and Casartelli, 2019, Diwanji and Bergmann, 2018, Dorstyn et al., 2018, Xu et al., 2018, Kamber Kaya et al., 2017, Kawamoto et al., 2016, Melzer and Broemer, 2016, Nicolson et al., 2015, Sakamaki et al., 2015, Griciuc et al., 2014, Steller et al., 2014.7.9, D'Brot et al., 2013, Denton et al., 2013, Dick and Megeney, 2013, Lee et al., 2013, Yuan and Akey, 2013, Chen et al., 2012, Darding and Meier, 2012, Domingues and Ryoo, 2012, Fabian and Brill, 2012, Vecchio et al., 2012, Verghese et al., 2012, Kuranaga, 2011, Miura, 2011, Yuan et al., 2011, Kim et al., 2010, Yang et al., 2010, Chew et al., 2009, Li and Cadigan, 2008, Pret et al., 2008, Chew et al., 2007, Lannan et al., 2007, Link et al., 2007, Akdemir et al., 2006, Arama et al., 2006, Leulier et al., 2006, Mendes et al., 2006, Kanuka et al., 2005, Kornbluth and White, 2005, Purves et al., 2005, Danial and Korsmeyer, 2004, McCall, 2004, Muro et al., 2004, Baehrecke, 2003, Cagan, 2003, Claveria and Torres, 2003, Lawen, 2003, Meier and Silke, 2003, Van Hoof and Goris, 2003, Adams and Cory, 2002, Igaki et al., 2002, Adams and Cory, 2001, Kimbrell and Beutler, 2001, Chen and Abrams, 2000, Gaumer et al., 2000, Kumar, 2000, Abrams, 1999, White, 1999)
              Dark/Apaf-I
              (Agnès et al., 2007)
              Dark/Dapaf-1/HAC1
              (Igaki and Miura, 2004)
              Dark/Hac-1/dApaf-1
              (Brachmann and Cagan, 2003)
              Dark/Hac-1/dApaf1
              (Leulier et al., 2006)
              Hac-1
              (Mendes et al., 2006, Kornbluth and White, 2005, Baehrecke, 2003, Gorski and Marra, 2002, Bialik and Gottlieb, 2000)
              Hac-1/Dark
              (Steller, 2000)
              Hac1
              (Darding and Meier, 2012, Zhou, 1999.10.19)
              T1
              (Hacker and Perrimon, 1998)
              anon-53Fa
              apaf-1
              (Shapiro et al., 2008, Kuranaga and Miura, 2007)
              apaf1
              (Richardson and Portela, 2018, Shapiro and Ryoo, 2008)
              arc
              (Lee et al., 2001)
              ark
              (Gerritsma et al., 2019, Jalvingh et al., 2014, Kuang et al., 2014, Li et al., 2013, Woodfield et al., 2013, Yacobi-Sharon et al., 2013, Chinchore et al., 2012, Florentin and Arama, 2012, Lee et al., 2011, Fan and Bergmann, 2010, Fan et al., 2010, Denton et al., 2009, Schauer et al., 2009, Berry and Baehrecke, 2007, Peterson et al., 2007, Srivastava et al., 2007, Arama et al., 2006, Herz et al., 2006, Mendes et al., 2006, Mills et al., 2006, Muro et al., 2006, Williams et al., 2006, Hay et al., 2004, Li and White, 2003, Gorski and Marra, 2002, Lee et al., 2002, Lee and Baehrecke, 2001, Lee et al., 2001, Simon et al., 2001, Thummel, 2001, Baehrecke, 2000, Lee et al., 2000, Vernooy et al., 2000)
              dAPAF-1
              (Kornbluth and White, 2005)
              dApaf-1
              (Gorski and Marra, 2002)
              dApaf-1/DARK/HAC-1
              (Meier et al., 2000)
              dApaf1
              (Reitman et al., 2015, Darding and Meier, 2012, Li et al., 2002)
              dArk
              (Qi et al., 2016, Morris et al., 2006, Moskalev et al., 2006, Moskalev and Zainullin, 2004, Moskalev, 2003)
              dapaf
              (Arama et al., 2003)
              dapaf-1
              (Frigo et al., 2023, Miura, 2011, Kuranaga and Miura, 2007, Zhou and Steller, 2003, Vernooy et al., 2000, Kanuka et al., 1999)
              dapaf-1L
              (Kanuka, 1999.5.20)
              dapaf-1S
              (Kanuka, 1999.5.20)
              dark
              (Braun et al., 2025, Lassetter et al., 2023, Prakash et al., 2022, Blanco et al., 2020, Ojha and Tapadia, 2020, Shinoda et al., 2019, Garcia-Garcia et al., 2017, Kang et al., 2017, Napoletano et al., 2017, Wang and Baker, 2015, Obata et al., 2014, Obata et al., 2014, Denton et al., 2013, Lee et al., 2013, Lee et al., 2013, Keller et al., 2011, Khammari et al., 2011, Winbush and Weeks, 2011, Li et al., 2010, Roignant and Treisman, 2010, Wang et al., 2010, Umemori et al., 2009, Joza et al., 2008, Shapiro and Ryoo, 2008, Shapiro et al., 2008, Kuranaga and Miura, 2007, Link et al., 2007, McCall et al., 2007, Yin et al., 2007, Akdemir et al., 2006, Choi et al., 2006, Peterson et al., 2006, Williams et al., 2006, Yu et al., 2006, Moon et al., 2005, Muro et al., 2005, Sang et al., 2005, Srivastava et al., 2005, Cakouros et al., 2004, Chew et al., 2004, Geisbrecht and Montell, 2004, Kumar and Cakouros, 2004, Arama et al., 2003, Bergmann et al., 2003, Ditzel et al., 2003, Kumar et al., 2003, Zhou and Steller, 2003, Dorstyn et al., 2002, Li et al., 2002, Muro et al., 2002, Peterson et al., 2002, Richardson and Kumar, 2002, Rodriguez et al., 2002, Colussi et al., 2000, Quinn et al., 2000, Rodriguez et al., 2000, Vernooy et al., 2000, Rodriguez, 1999.7.30, Rodriguez et al., 1999)
              dark/dapaf-1/hac-1
              (Kumar and Doumanis, 2000)
              dark/hac-1/dapaf-1
              (Tittel and Steller, 2000)
              dpf
              (Ciesielski et al., 2022)
              hac-1
              (Kuranaga and Miura, 2007, Zhang et al., 2006, Gottfried et al., 2004, Arama et al., 2003, Zhou and Steller, 2003, Zhou and Steller, 2002, Vernooy et al., 2000, Zhou et al., 1999)
              hac1
              (Shapiro and Ryoo, 2008)
              l(2)SH0173
              Name Synonyms
              Apaf-1 related
              (Fan and Bergmann, 2010)
              Apaf-1 related killer
              (Begun et al., 2007, Srivastava et al., 2007, Yu et al., 2006, Sang et al., 2005, Rodriquez, 1999.6.25)
              Apaf-1-Related Killer
              (Umemori et al., 2009)
              Apaf-1-related killer
              (Chinchore et al., 2012, Xu et al., 2009, Cakouros et al., 2008, Dorstyn and Kumar, 2008, Morris et al., 2006)
              Apaf-1-related-killer
              Dark
              (Colon-Plaza and Su, 2022, Cheng et al., 2017)
              Death-associated APAF1-related killer
              (Steller et al., 2014.7.9)
              Drosophila Apaf-1-related killer
              (Adrain, 2006, Morris et al., 2006, Rodriguez et al., 1999)
              Transcription unit 1
              (Hacker and Perrimon, 1998)
              apaf1-related killer
              (Domingues and Ryoo, 2012)
              dark
              (Galindo et al., 2009)
              lethal (2) SH0173
              Secondary FlyBase IDs
              • FBgn0065914
              • FBgn0024252
              • FBgn0028733
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 62 )
              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
              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.
              BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
              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
              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)
              PDB - An information portal to biological macromolecular structures
              Linkouts
              Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
              DroID - A comprehensive database of gene and protein interactions.
              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
              iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
              Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
              KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
              MIST (genetic) - An integrated Molecular Interaction Database
              MIST (protein-protein) - An integrated Molecular Interaction Database
              Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
              References (318)