FB2019_05, released Oct 15, 2019

Gene: Dmel\spen

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General Information
Symbol
Dmel\spen
Species
D. melanogaster
Name
split ends
Annotation Symbol
CG18497
Feature Type
protein_coding_gene
FlyBase ID
FBgn0016977
Gene Model Status
Current
Stock Availability
43 publicly available
Gene Snapshot
In progress.Contributions welcome.
Other Summaries
Also Known As
poc, l(2)03350, l(2)k08102, polycephalon, rno
Key Links
Genomic Location
Cytogenetic map
21B3-21B4
Sequence location
2L:159,032..203,397 [+]
Recombination map
2-0.5
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
GO Summary Ribbons
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the RRM Spen family. (Q8SX83)
Protein Signatures (InterPro)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
-
Predictions / Assertions
nucleic acid binding
Summaries
Pathway (FlyBase)
Positive Regulators of Wnt-TCF Signaling Pathway -
Positive regulators of canonical Wnt signaling up-regulate the pathway, resulting in the enhancement of transcriptional regulation mediated by β-catenin (arm).
Protein Function (UniProtKB)
Probable corepressor protein, which regulates different key pathways such as the EGF receptor and Wg pathways. Involved in neuronal cell fate, survival and axon guidance, cell cycle regulation and repression of head identity in the embryonic trunk. May act with the Hox gene Deformed and the EGF receptor signaling pathway. Positive regulator of the Wg pathway in larval tissues but not in embryonic tissues. May act as a transcriptional corepressor protein, which repress transcription via the recruitment of large complexes containing histone deacetylase proteins.
(UniProt, Q8SX83)
Summary (Interactive Fly)
codes for RRM motif RNA-binding protein - mutants have defects in Notch and Egf receptor signaling resulting defects in cell-fate and in axon guidance - large and small SPEN family proteins stimulate axon outgrowth during neurosecretory cell remodeling
Gene Model and Products
Number of Transcripts
8
Number of Unique Polypeptides
7

Please see the GBrowse view of Dmel\spen or the JBrowse view of Dmel\spen 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
Comments on Gene Model
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.44
Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.
Gene model reviewed during 5.45
Gene model reviewed during 5.56
Sequence Ontology: Class of Gene
gene_with_edited_transcript
(Rodriguez et al., 2012)
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
spen-RA
FBtr0078122
NM_164374
17880
5560
spen-RB
FBtr0078121
NM_079979
19604
5533
spen-RC
FBtr0078123
NM_164375
18859
5476
spen-RD
FBtr0306341
NM_001258887
17963
5505
spen-RE
FBtr0330652
NM_001272875
20622
5505
spen-RF
FBtr0330653
NM_001272876
18434
5510
spen-RG
FBtr0332336
NM_001272873
19571
5487
spen-RH
FBtr0332337
NM_001272874
17143
5458
Additional Transcript Data and Comments
Reported size (kB)
20 (northern blot)
(Wiellette et al., 1999)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
spen-PA
FBpp0077781
600.0
5560
8.90
NP_722615
AAF51535
spen-PB
FBpp0077780
597.1
5533
8.88
NP_524718
AAF51534
spen-PC
FBpp0077782
590.5
5476
8.84
NP_722616
AAN10511
spen-PD
FBpp0297434
593.5
5505
8.81
NP_001245816
AFH03493
spen-PE
FBpp0303502
593.5
5505
8.81
NP_001259804
AGB92341
spen-PF
FBpp0303503
594.0
5510
8.86
NP_001259805
AGB92342
spen-PG
FBpp0304613
591.5
5487
8.76
NP_001259802
AGB92339
spen-PH
FBpp0304614
588.5
5458
8.79
NP_001259803
AGB92340
Polypeptides with Identical Sequences

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

5505 aa isoforms: spen-PD, spen-PE
Additional Polypeptide Data and Comments
Reported size (kDa)
5476 (aa)
(Rebay et al., 2000)
5533, 5476 (aa); 597 (kD predicted)
(Wiellette et al., 1999)
Comments
External Data
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\spen using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (17 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
nucleic acid binding
inferred from electronic annotation with InterPro:IPR000504, InterPro:IPR035979
(assigned by InterPro )
(FlyBase Curators et al., 2004-)
Biological Process (15 terms)
Terms Based on Experimental Evidence (15 terms)
CV Term
Evidence
References
axon guidance
inferred from mutant phenotype
(assigned by UniProt )
(Kuang et al., 2000)
compound eye development
inferred from mutant phenotype
(Chang et al., 2008)
defense response to fungus
inferred from mutant phenotype
(Jin et al., 2008)
epidermal growth factor receptor signaling pathway
inferred from mutant phenotype
(assigned by UniProt )
(Chen and Rebay, 2000)
establishment of thoracic bristle planar orientation
inferred from mutant phenotype
(Mace and Tugores, 2004)
glial cell fate determination
inferred from mutant phenotype
(assigned by UniProt )
(Chen and Rebay, 2000)
imaginal disc-derived wing vein morphogenesis
inferred from mutant phenotype
(Mace and Tugores, 2004)
maintenance of imaginal disc-derived wing hair orientation
inferred from mutant phenotype
(Mace and Tugores, 2004)
neuroblast fate determination
inferred from mutant phenotype
(assigned by UniProt )
(Kuang et al., 2000)
peripheral nervous system development
inferred from mutant phenotype
(Mace and Tugores, 2004)
positive regulation of canonical Wnt signaling pathway
inferred from mutant phenotype
(assigned by UniProt )
(Lin et al., 2003)
inferred from genetic interaction with FLYBASE:arm; FB:FBgn0000117
(Chang et al., 2008)
inferred from genetic interaction with FLYBASE:wg; FB:FBgn0284084
(Chang et al., 2008)
regulation of transcription, DNA-templated
inferred from mutant phenotype
(assigned by UniProt )
(Wiellette et al., 1999)
segment specification
inferred from mutant phenotype
(assigned by UniProt )
(Wiellette et al., 1999)
transcription, DNA-templated
inferred from mutant phenotype
(assigned by UniProt )
(Wiellette et al., 1999)
wing disc pattern formation
inferred from mutant phenotype
(Chang et al., 2008)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
nucleus
inferred from direct assay
(assigned by UniProt )
(Lin et al., 2003, Kuang et al., 2000, Wiellette et al., 1999)
Terms Based on Predictions or Assertions (0 terms)
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
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 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 9 -- 10
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 11 -- 12
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic brain
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
wandering third instar larval stage
wing pouch | restricted
(Molnar et al., 2012)
wing hinge primordium | restricted
(Molnar et al., 2012)
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Wiellette et al., 1999)
embryonic stage 1 -- 17

Comment: reference states 0-24 hr AEL

(Wiellette et al., 1999)
embryonic stage 4 -- 17

Comment: reference states 2-24 hr AEL

(Wiellette et al., 1999)
Additional Descriptive Data
A probe specific for the long spen isoform does not detect spen transcripts in northern blots of 0-2 hr embryos, but both short and long isoforms are detected in all embryonic stages using RT-PCR.
(Wiellette et al., 1999)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
organism
(Lin et al., 2003)
embryonic stage 3 -- 17
organism
(Wiellette et al., 1999)
embryonic stage 5 -- 10
somatic cell
• nucleus
(Kuang et al., 2000)
embryonic stage 9 -- 14
presumptive embryonic/larval central nervous system
(Wiellette et al., 1999)
embryonic epidermis
(Wiellette et al., 1999)
embryonic stage 12
epidermal cell
• nucleus
(Kuang et al., 2000)
embryonic stage 13
midline glial cell
• nucleus
(Chen and Rebay, 2000)
embryonic stage 17
embryonic/larval central nervous system
• nucleus
(Kuang et al., 2000)
larval stage
ventral thoracic disc
(Lin et al., 2003)
eye disc
• nucleus
(Lin et al., 2003)
wing disc
(Lin et al., 2003)
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
day 5 of adulthood & male -- day 7 of adulthood & male
spermatozoon
(Wasbrough et al., 2010)
Additional Descriptive Data
spen proteins are expressed in most cell types starting in stage 3 embryonic nuclei. Expression continues through the rest of embryogenesis, and is concentrated in nuclei. Higher levels of expression are detected in the epidermis and central nervous system in stage 9-14 embryos.
(Wiellette et al., 1999)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
nucleus
inferred from direct assay
(assigned by UniProt )
(Lin et al., 2003, Kuang et al., 2000, Wiellette et al., 1999)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\spen in GBrowse 2
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
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, and Transgenic Constructs
Classical and Insertion Alleles ( 96 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 13 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of spen
Transgenic constructs containing regulatory region of spen
Deletions and Duplications ( 8 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal, with Scer\GAL4sli.PS
lethal, with Scer\GAL4VP16.otu
lethal (with spen14O1)
lethal (with spenC18)
lethal (with spenI55)
lethal (with spenN28)
lethal | female limited, with Scer\GAL4dome-PG14
lethal | recessive
lethal - all die before end of pupal stage, with Scer\GAL4pnr-MD237
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4elav-C155
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
fertile
Other Phenotypes
Allele
cell polarity defective | somatic clone
flightless, with Scer\GAL4Mef2.PR
immune response defective | adult stage
neuroanatomy defective | larval stage, with Scer\GAL4CCAP.PP
neuroanatomy defective | pharate adult stage, with Scer\GAL4CCAP.PP
some die during pupal stage, with Scer\GAL4pnr-MD237
visible, with Scer\GAL4nub-AC-62
visible, with Scer\GAL4pnr-MD237
visible, with Scer\GAL4sca-537.4
visible, with Scer\GAL4sev.PR
visible | somatic clone
wound healing defective
Phenotype manifest in
Allele
abdominal lateral pentascolopidial chordotonal organ lch5 | germline clone
abdominal lch5 neuron | germline clone
abdominal segmental nerve
central nervous system
chordotonal organ & axon | lateral
commissure
crossvein | somatic clone
crystal cell | third instar larval stage
dental sclerite
denticle belt, with Scer\GAL4da.G32
dorsal bridge
dorsocentral bristle | ectopic, with Scer\GAL4sca-537.4
embryonic/first instar larval cuticle, with Scer\GAL4da.G32
embryonic/larval central nervous system, with Scer\GAL4CCAP.PP
embryonic/larval somatic muscle | germline clone
embryonic epidermis
embryonic head
embryonic mesothoracic segment & sclerite | ectopic
embryonic metathoracic segment & sclerite | ectopic
embryonic thorax
embryonic thorax & cuticle
epipharyngeal sclerite
eye, with Scer\GAL4sev.PR
eye photoreceptor cell
eye photoreceptor cell | somatic clone
head & cuticle | ectopic
hemocyte | adult stage
hypostomal sclerite
intersegmental nerve
lateral process
longitudinal connective
longitudinal connective | germline clone
macrochaeta
macrochaeta | somatic clone
median tooth
mesothoracic tergum, with Scer\GAL4pnr-MD237
microchaeta | somatic clone
midline glial cell
motor neuron | embryonic stage | germline clone
mouth hooks
neurite | pharate adult stage, with Scer\GAL4CCAP.PP
ommatidium
ommatidium, with Scer\GAL4sev.PR
plasmatocyte | third instar larval stage
presumptive embryonic/larval central nervous system | germline clone
presumptive embryonic/larval peripheral nervous system
presumptive embryonic/larval peripheral nervous system | germline clone
sclerite & embryonic abdominal segment | ectopic
sclerite & embryonic head
scutellar bristle | ectopic, with Scer\GAL4sca-537.4
tormogen cell, with Scer\GAL4sca-537.4
transverse nerve
trichogen cell, with Scer\GAL4pnr-MD237
ventral midline | germline clone
ventral nerve cord commissure | germline clone
ventral thoracic disc, with Scer\GAL4ptc-559.1
wing, with Scer\GAL4nub-AC-62
wing hair | somatic clone
wing margin bristle | somatic clone
wing vein | ectopic | somatic clone
wing vein | somatic clone
Orthologs
Downloads
Download All DIOPT Orthologs
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (16)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\SPEN
10 of 15
Yes
Yes
Hsap\RBM15
3 of 15
No
No
Hsap\RBM15B
2 of 15
No
No
Hsap\BBS1
1 of 15
No
No
Hsap\NONO
1 of 15
No
No
Hsap\PSPC1
1 of 15
No
No
 
Hsap\RBM4B
1 of 15
No
No
Hsap\RBM4
1 of 15
No
No
 
Hsap\RBM14-RBM4
1 of 15
No
No
Hsap\SFPQ
1 of 15
No
No
Hsap\SRSF3
1 of 15
No
No
 
Hsap\SRSF4
1 of 15
No
No
Hsap\SRSF5
1 of 15
No
No
Hsap\SRSF6
1 of 15
No
No
 
Hsap\SRSF7
1 of 15
No
No
 
Hsap\SRSF9
1 of 15
No
No
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (15)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Spen
11 of 15
Yes
Yes
Mmus\Rbm15
3 of 15
No
No
Mmus\Rbm15b
2 of 15
No
No
Mmus\Gm21992
1 of 15
No
No
Mmus\Nono
1 of 15
No
No
Mmus\Pspc1
1 of 15
No
No
Mmus\Rbm4b
1 of 15
No
No
Mmus\Rbm4
1 of 15
No
No
Mmus\Sfpq
1 of 15
No
No
Mmus\Srsf3
1 of 15
No
No
Mmus\Srsf4
1 of 15
No
No
Mmus\Srsf5
1 of 15
No
No
Mmus\Srsf6
1 of 15
No
No
Mmus\Srsf7
1 of 15
No
No
Mmus\Srsf9
1 of 15
No
No
Rattus norvegicus (Norway rat) (15)
Rnor\Rbm15b
2 of 13
Yes
No
Rnor\Rbm15
2 of 13
Yes
No
Rnor\Spen
2 of 13
Yes
Yes
Rnor\Nono
1 of 13
No
No
Rnor\Pspc1
1 of 13
No
No
Rnor\Rbm4b
1 of 13
No
No
Rnor\Rbm4
1 of 13
No
No
Rnor\Rbm14
1 of 13
No
No
Rnor\Srsf1
1 of 13
No
No
Rnor\Srsf3
1 of 13
No
No
Rnor\Srsf4
1 of 13
No
No
Rnor\Srsf5
1 of 13
No
No
Rnor\Srsf6
1 of 13
No
No
Rnor\Srsf7
1 of 13
No
No
Rnor\Srsf9
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (6)
Xtro\rbm15
2 of 12
Yes
No
Xtro\rbm4b
1 of 12
No
No
Xtro\rbm15b
1 of 12
No
No
Xtro\sfpq
1 of 12
No
No
Xtro\srsf3
1 of 12
No
No
Xtro\srsf5
1 of 12
No
No
Danio rerio (Zebrafish) (20)
Drer\spen
9 of 15
Yes
Yes
Drer\wu:fb69f10
8 of 15
No
Yes
Drer\rbm15
2 of 15
No
No
Drer\nono
1 of 15
No
No
Drer\pspc1
1 of 15
No
No
Drer\rbm4.2
1 of 15
No
No
Drer\rbm4.3
1 of 15
No
No
Drer\rbm14b
1 of 15
No
No
Drer\rbm15b
1 of 15
No
No
Drer\sfpq
1 of 15
No
No
Drer\srsf1a
1 of 15
No
No
Drer\srsf3a
1 of 15
No
No
Drer\srsf3b
1 of 15
No
No
Drer\srsf4
1 of 15
No
No
Drer\srsf5a
1 of 15
No
No
Drer\srsf5b
1 of 15
No
No
Drer\srsf7a
1 of 15
No
No
Drer\srsf7b
1 of 15
No
No
Drer\srsf9
1 of 15
No
No
Drer\ugt2a6
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (11)
Cele\din-1
4 of 15
Yes
Yes
Cele\grld-1
2 of 15
No
No
Cele\cec-8
1 of 15
No
Yes
Cele\nono-1
1 of 15
No
No
Cele\rnp-1
1 of 15
No
No
Cele\rsp-1
1 of 15
No
No
Cele\rsp-2
1 of 15
No
No
Cele\rsp-3
1 of 15
No
No
Cele\rsp-5
1 of 15
No
No
Cele\rsp-6
1 of 15
No
No
Cele\Y57G11A.5
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (16)
Atha\FPA
4 of 9
Yes
Yes
Atha\AT4G12640
2 of 9
No
Yes
Atha\ML4
1 of 9
No
Yes
Atha\RS2Z32
1 of 9
No
No
Atha\RS2Z33
1 of 9
No
No
Atha\RS31a
1 of 9
No
No
Atha\RS31
1 of 9
No
No
Atha\RS40
1 of 9
No
No
Atha\RS41
1 of 9
No
No
Atha\RSZ21
1 of 9
No
No
Atha\RSZ22a
1 of 9
No
No
Atha\RSZ22
1 of 9
No
No
Atha\SR30
1 of 9
No
No
Atha\SR34a
1 of 9
No
No
Atha\SR34
1 of 9
No
No
Atha\SR34b
1 of 9
No
No
Saccharomyces cerevisiae (Brewer's yeast) (4)
Scer\MRN1
1 of 15
Yes
Yes
Scer\NPL3
1 of 15
Yes
No
Scer\NRD1
1 of 15
Yes
No
Scer\RNP1
1 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (4)
Spom\mug24
2 of 12
Yes
Yes
Spom\seb1
1 of 12
No
No
Spom\SPAC222.18
1 of 12
No
Yes
Spom\srp2
1 of 12
No
No
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG0919004W )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Dmel\spen
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00000014
Drosophila simulans
Dsim\GD23007
Drosophila sechellia
Dsec\GM16721
Drosophila sechellia
Dsec\GM16722
Drosophila erecta
Dere\GG24702
Drosophila yakuba
Dyak\GE16637
Drosophila ananassae
Dana\GF20711
Drosophila pseudoobscura pseudoobscura
Dpse\GA25185
Drosophila persimilis
Dper\GL25911
Drosophila willistoni
Dwil\GK23912
Drosophila virilis
Dvir\GJ24602
Drosophila mojavensis
Dmoj\GI17029
Drosophila grimshawi
Dgri\GH11539
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150794 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17016474
Glossina morsitans
Tsetse fly
Gmor\GMOY009912
Glossina morsitans
Tsetse fly
Gmor\GMOY003516
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_04764
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W07JB )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Apis florea
Little honeybee
Aflo\2147541283
Apis florea
Little honeybee
Aflo\2147488689
Apis mellifera
Western honey bee
Amel\GB47009
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP23885
Bombus terrestris
Buff-tailed bumblebee
Bter\15215225
Bombus terrestris
Buff-tailed bumblebee
Bter\15215731
Linepithema humile
Argentine ant
Lhum\LH22375
Megachile rotundata
Alfalfa leafcutting bee
Mrot\MROTU:002f63
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG003872
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_12112
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_12110
Tribolium castaneum
Red flour beetle
Tcas\TC006356
Pediculus humanus
Human body louse
Phum\PHUM127440
Rhodnius prolixus
Kissing bug
Rpro\RPRC014989
Cimex lectularius
Bed bug
Clec\CLEC002303
Cimex lectularius
Bed bug
Clec\CLEC002303-PB
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI009704
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_11845
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X07EM )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR008455
Ixodes scapularis
Black-legged tick
Isca\ISCW021726
Stegodyphus mimosarum
African social velvet spider
Smim\KFM61671.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM61670.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur01g07610
Tetranychus urticae
Two-spotted spider mite
Turt\tetur01g07600
Tetranychus urticae
Two-spotted spider mite
Turt\tetur01g07620
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_53407
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_305154
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G00NR )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Pink1L
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000004036
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000019435
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000003662
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (12)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
nito
4 of 10
nonA-l
2 of 10
nonA
2 of 10
B52
1 of 10
CG4266
1 of 10
lark
1 of 10
Rbp1-like
1 of 10
Rbp1
1 of 10
Rsf1
1 of 10
SF2
1 of 10
trv
1 of 10
x16
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    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 ( 2 )
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 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?
    SPEN; spen family transcriptional repressor
    10 of 15
    613484
        RBM15; RNA binding motif protein 15
        3 of 15
        606077
          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
          esyN Network Diagram
          Show neighbor-neighbor interactions:
          Select Layout:
          Legend:
          Protein
          RNA
          Selected Interactor(s)
          Interactions Browser
          View in Interactions Browser
          Please see the Physical Interaction reports below for full details
          RNA-protein
          Physical Interaction
          Assay
          References
          spen - stau
          anti bait coimmunoprecipitation, partial DNA sequence identification by hybridization
          (Laver et al., 2013)
          protein-protein
          Physical Interaction
          Assay
          References
          spen - Hipk
          enzymatic study, autoradiography
          (Dewald et al., 2014)
          spen - trr
          pull down, autoradiography
          (Oswald et al., 2016)
          Summary of Genetic Interactions
          esyN Network Diagram
          esyN Network Key:
          Suppression
          Enhancement
          View in Interactions Browser
          Please look at the allele data for full details of the genetic interactions
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          spen
          suppressible
          Antp
          (Wiellette et al., 1999)
          spen
          enhanceable
          Antp
          (Wiellette et al., 1999)
          spen
          enhanceable
          ck
          (Mace and Tugores, 2004)
          spen
          suppressible
          H
          (Abdelilah-Seyfried et al., 2000)
          spen
          enhanceable
          N
          (Abdelilah-Seyfried et al., 2000)
          hid|spen
          suppressible
          nito
          (Chang et al., 2008)
          spen
          enhanceable
          nito
          (Jemc and Rebay, 2006)
          spen
          suppressible
          nito
          (Jemc and Rebay, 2006)
          arm|spen
          suppressible
          nito
          (Chang et al., 2008)
          spen|wg
          suppressible
          nito
          (Chang et al., 2008)
          spen
          enhanceable
          pnt
          (Chen and Rebay, 2000)
          spen
          suppressible
          Ras85D
          (Chen and Rebay, 2000)
          spen
          suppressible
          spi
          (Chen and Rebay, 2000)
          spen
          suppressible
          tsh
          (Wiellette et al., 1999)
          Starting gene(s)
          Interaction type
          Interacting gene(s)
          Reference
          Antp
          enhanceable
          spen
          (Wiellette et al., 1999)
          aop
          enhanceable
          spen
          (Rebay et al., 2000)
          arm
          suppressible
          spen
          (Chang et al., 2008, Lin et al., 2003)
          arm|nito
          suppressible
          spen
          (Chang et al., 2008)
          CycE
          suppressible
          spen
          (Brumby et al., 2004)
          CycE
          enhanceable
          spen
          (Lane et al., 2000)
          CycE|dap
          suppressible
          spen
          (Staehling-Hampton et al., 1999)
          Dp|E2f1
          enhanceable
          spen
          (Staehling-Hampton et al., 1999)
          dpp
          suppressible
          spen
          (Lin et al., 2003)
          E2f1|Dp
          enhanceable
          spen
          (Staehling-Hampton et al., 1999)
          Egfr
          suppressible
          spen
          (Lin et al., 2003)
          H
          enhanceable
          spen
          (Abdelilah-Seyfried et al., 2000)
          hid|nito
          suppressible
          spen
          (Chang et al., 2008)
          Hipk
          suppressible
          spen
          (Dewald et al., 2014)
          ksr::tor
          enhanceable
          spen
          (Therrien et al., 2000)
          lqf
          enhanceable
          spen
          (Eun et al., 2007)
          N
          enhanceable
          spen
          (Abdelilah-Seyfried et al., 2000)
          N
          suppressible
          spen
          (Kankel et al., 2007)
          nito
          enhanceable
          spen
          (Jemc and Rebay, 2006)
          nito|wg
          suppressible
          spen
          (Chang et al., 2008)
          Raf::tor
          enhanceable
          spen
          (Dickson et al., 1996)
          Raf
          enhanceable
          spen
          (Therrien et al., 2000, Dickson et al., 1996)
          Ras85D
          suppressible
          spen
          (Chen and Rebay, 2000, Rebay et al., 2000)
          Ras85D
          enhanceable
          spen
          (Rebay et al., 2000)
          tsh
          enhanceable
          spen
          (Wiellette et al., 1999)
          wg
          suppressible
          spen
          (Chang et al., 2008)
          External Data
          Linkouts
          BioGRID - A database of protein and genetic interactions.
          DroID - A comprehensive database of gene and protein interactions.
          InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          Pathways
          Gene Group - Pathway Membership (FlyBase)
          Positive Regulators of Wnt-TCF Signaling Pathway -
          Positive regulators of canonical Wnt signaling up-regulate the pathway, resulting in the enhancement of transcriptional regulation mediated by β-catenin (arm).
          External Data
          Linkouts
          Genomic Location and Detailed Mapping Data
          Chromosome (arm)
          2L
          Recombination map
          2-0.5
          Cytogenetic map
          21B3-21B4
          Sequence location
          2L:159,032..203,397 [+]
          FlyBase Computed Cytological Location
          Cytogenetic map
          Evidence for location
          21B3-21B4
          Limits computationally determined from genome sequence between P{lacW}spenk06805&P{lacW}l(2)k13604k13604 and P{lacW}U2af38k14504
          Experimentally Determined Cytological Location
          Cytogenetic map
          Notes
          References
          21B4-21B6
          (determined by in situ hybridisation)
          (Abdelilah-Seyfried et al., 2000, Spradling et al., 1999)
          21B4-21B6
          (Rebay, 1999.9.8)
          21A4-21B6
          (Wiellette, 1999.9.18)
          21B4-21B5
          21B4--6
          (BDGP Project Members, 1994-1999)
          Experimentally Determined Recombination Data
          Location
          2-0.5
          (FlyBase, 2016)
          2-0.0
          (Comeron et al., 2012)
          2-
          (Lane et al., 2000, Kolodziej et al., 1995)
          2-1.0 +/- 0.5
          (Dickson et al., 1996)
          Left of (cM)
          (Kolodziej et al., 1995)
          Right of (cM)
          (Kolodziej et al., 1995)
          Notes
          Stocks and Reagents
          Stocks (43)
          Genomic Clones (34)
          cDNA Clones (88)
           

          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 GBrowse for alignment of the cDNAs and ESTs to the gene model.

          cDNA clones, fully sequences
          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
            GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
            Antibody Information
            Laboratory Generated Antibodies
             
            polyclonal
            (Querenet et al., 2015, Wiellette et al., 1999)
            Commercially Available Antibodies
             
            Other Information
            Relationship to Other Genes
            Source for database identify of
            Source for database merge of
            Source for merge of: poc E(Raf)2A
            (Florence and McGinnis, 1998)
            Source for merge of: spen poc l(2)k06805 l(2)03350 l(2)k07612 l(2)k08102
            (Wiellette et al., 1999)
            Source for merge of: spen CG18497
            (Bayraktaroglu, 2000.4.7)
            Source for merge of: spen yip1
            (Bayraktaroglu, 2000.11.2)
            Source for merge of: spen rno
            (Voas, 2003.6.5)
            Source for merge of: spen BcDNA:GM01870
            (FlyBase, 1996-)
            Additional comments
            Source for merge of spen BcDNA:GM01870 was a shared cDNA ( date:030728 ).
            (FlyBase, 1996-)
            Other Comments
            ChEST reveals this is a target of Mef2.
            (Junion et al., 2005)
            spen appears to be required to maintain multicellular organization.
            (Mace and Tugores, 2004)
            spen is required for wg signaling in the eye, wing and leg imaginal discs, but is not required for wg signaling in the embryo.
            (Lin et al., 2003)
            spen encodes a positively acting component of the Egfr/Ras signalling pathway.
            (Chen and Rebay, 2000)
            spen regulates development of midline glial cells.
            (Chen and Rebay, 2000)
            Maternally contributed spen protein is required for the specification of cell fate during neuronal and midline development.
            (Kuang et al., 2000)
            Identification: Identified on the basis of interaction with aop mutations.
            (Rebay, 1999)
            Mutants show a homeotic phenotype developing head-like skeletal features in the trunk.
            (Wiellette et al., 1999)
            spen does not repress ectopic head-like cuticle by repressing the expression or function of known head-determining genes. Overall orientation not stated: anon-21Ab+ spen+ kis-
            (Wiellette et al., 1999)
            spen functions in the repression of head patterning genes. spen cooperates with Antp and tsh to repress head-like sclerites in the thorax and promote thoracic identity.
            (Wiellette et al., 1999)
            Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology.
            (Roch et al., 1998)
            Identification: Enhancer trap screen designed to discover genes involved in the cellular aspects of defense mechanisms, as well as in melanotic tumor formation processes linked to blood cell disregulation.
            (Braun et al., 1997)
            Identified in a screen for modifiers of the Dfd13/Dfd3 mutant phenotype. Shows no interaction with the Pc mutant phenotype.
            (Gellon et al., 1997)
            Identified in a genetic screen for modifiers of the phl::tor12D.sev rough eye mutant phenotype.
            (Dickson et al., 1996)
            The ventral naked cuticle of T2, T3 and occasionally A1 of spen- embryos shows an apparent transformation towards head structures.
            (Harding et al., 1996)
            Identified in a screen for genes whose function is required for the function of Dfd.
            (Harding et al., 1996)
            Mutations in 12 complementation groups differentially affect lateral chordotonal axon growth, fasciculation or ventral orientation. Mutations in robo, spen, sli and los cause lch axon defasciculation.
            (Kolodziej et al., 1995)
            Origin and Etymology
            Discoverer
            Etymology
            Identification
            External Crossreferences and Linkouts ( 125 )
            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/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
            BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
            Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
            Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
            Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
            Flygut - An atlas of the Drosophila adult midgut
            GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
            InterPro - A database of protein families, domains and functional sites
            KEGG Genes - Molecular building blocks of life in the genomic space.
            modMine - A data warehouse for the modENCODE project
            Linkouts
            ApoDroso - Functional genomic database for photoreceptor development, survival and function
            BioGRID - A database of protein and genetic interactions.
            DroID - A comprehensive database of gene and protein interactions.
            DRSC - Results frm RNAi screens
            FLIGHT - Cell culture data for RNAi and other high-throughput technologies
            FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
            FlyMine - An integrated database for Drosophila genomics
            Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
            InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
            MIST (genetic) - An integrated Molecular Interaction Database
            MIST (protein-protein) - An integrated Molecular Interaction Database
            Synonyms and Secondary IDs (28)
            Reported As
            Symbol Synonym
            136/24
            (Roch et al., 1998)
            BcDNA:GM01870
            CG18497
            (Na et al., 2016, Hosono et al., 2015, Christis and Munro, 2012, Ni et al., 2010.12.1, Slack et al., 2010, Keleman et al., 2009.8.5, Quinones-Coello, 2007, Molnar et al., 2006, Norga et al., 2003)
            E(E2F)2A
            (Staehling-Hampton et al., 1999)
            E(Raf)2A
            E(Sev-CycE)2A
            (Lane et al., 2000)
            EK2-9
            (Therrien et al., 2000)
            EY2-7
            (Rebay et al., 2000)
            En(yanACT)2-7
            SPEN
            (Sofola et al., 2007)
            Spen
            (Maier, 2019, Hazegh et al., 2017, Oswald et al., 2016, Yan and Perrimon, 2015, Honti et al., 2014, Berg et al., 2012, Tokusumi et al., 2012)
            Splitends
            (Junion et al., 2005)
            l(2)03350
            (Spradling et al., 1999, Braun et al., 1997, BDGP Project Members, 1994-1999)
            l(2)k06805
            (Spradling et al., 1999, BDGP Project Members, 1994-1999)
            l(2)k07612
            (Spradling et al., 1999, BDGP Project Members, 1994-1999)
            l(2)k08102
            (Spradling et al., 1999, BDGP Project Members, 1994-1999)
            poc
            (Florence and McGinnis, 1998, Gellon et al., 1997, Harding and McGinnis, 1997, Dickson et al., 1996, Harding et al., 1996)
            rno
            (Voas, 2003.6.5, Rebay, 1999)
            spe
            (Wiellette et al., 1999)
            spen
            (Kockel et al., 2019, Landskron et al., 2018, Levis, 2018.8.30, Yu et al., 2018, Transgenic RNAi Project members, 2017-, Yao et al., 2017, Brewer-Jensen et al., 2016, Kockel et al., 2016, Na et al., 2016, Fernando et al., 2015, Gene Disruption Project members, 2015-, Querenet et al., 2015, Ashwal-Fluss et al., 2014, Dewald et al., 2014, Lye et al., 2014, Japanese National Institute of Genetics, 2012.5.21, Rodriguez et al., 2012, Grimberg et al., 2011, Guan et al., 2011, Wertheim et al., 2011, Wasbrough et al., 2010, Jin et al., 2009, Levis, 2009.3.18, Seugnet et al., 2009, Shalaby et al., 2009, Tsubouchi et al., 2009, Chang et al., 2008, Jin et al., 2008, Teleman et al., 2008, Zhao et al., 2008, Zhou et al., 2008, Doroquez et al., 2007, Eun et al., 2007, Kankel et al., 2007, Magalhaes et al., 2007, Minidorff et al., 2007, Minidorff et al., 2007, Quinones-Coello, 2007, Sofola et al., 2007, Stark et al., 2007, Zeitlinger et al., 2007, Jemc and Rebay, 2006, Mack et al., 2006, Molnar et al., 2006, Macdonald and Long, 2005, Mace et al., 2005, Mace et al., 2005)
            yip1
            Name Synonyms
            Split ends
            (Yan and Perrimon, 2015, Dewald et al., 2014, Guan et al., 2011, Jin et al., 2009, Chang et al., 2008)
            Split-ends
            (Yan and Perrimon, 2015)
            Splitends
            (Davis et al., 2011)
            polycephalon
            (Gellon et al., 1997, Harding et al., 1996)
            rhinoceros
            (Voas, 2003.6.5, Rebay, 1999)
            split ends
            (Laver et al., 2013, Shalaby et al., 2009, Zhao et al., 2008, Doroquez et al., 2007, Eun et al., 2007, Kankel et al., 2007, Jemc and Rebay, 2006, Burnette et al., 2005, Voas, 2003.6.5, Kolodziej et al., 1995)
            split-ends
            (Querenet et al., 2015, Slack et al., 2010)
            yippee interacting protein 1
            Secondary FlyBase IDs
            • FBgn0010537
            • FBgn0016056
            • FBgn0016624
            • FBgn0022059
            • FBgn0022080
            • FBgn0022115
            • FBgn0026182
            • FBgn0031230
            • FBgn0040065
            • FBgn0047291
            • FBtr0078121
            • FBpp0077780
            • FBtr0078122
            • FBpp0077781
            • FBtr0078123
            • FBpp0077782
            Datasets (0)
            Study focus (0)
            Experimental Role
            Project
            Project Type
            Title
            References (152)