FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\ss
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General Information
Symbol
Dmel\ss
Species
D. melanogaster
Name
spineless
Annotation Symbol
CG6993
Feature Type
protein_coding_gene
FlyBase ID
FBgn0003513
Gene Model Status
Current
Stock Availability
114 publicly available
Gene Summary
spineless (ss) encodes a protein that plays a key role in defining the distal regions of the antenna and the leg. Its stochastic expression in R7 photoreceptors also controls the expression of color Rhodopsins (the product of Rh4 vs. the product of Rh3) in the two subsets of ommatidia. [Date last reviewed: 2019-03-21] (FlyBase Gene Snapshot)
All Summaries
Also Known As

aristapedia, AHR

Key Links
Genomic Location
Cytogenetic map
89B14-89B15
Sequence location
Recombination map
3-58
RefSeq locus
NT_033777 REGION:16374434..16403690
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 (22 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
contributes_to DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from direct assay
(Kozu et al., 2006)
enables protein heterodimerization activity
inferred from physical interaction with FLYBASE:tgo; FB:FBgn0264075
(Kozu et al., 2006)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
enables nuclear receptor activity
inferred from biological aspect of ancestor with PANTHER:PTN004180845
(GO Reference Genome Project, 2011-)
enables protein dimerization activity
inferred from electronic annotation with InterPro:IPR011598, InterPro:IPR036638
(InterPro Project Members, 2004-)
enables transcription cis-regulatory region binding
inferred from biological aspect of ancestor with PANTHER:PTN004180845
(GO Reference Genome Project, 2011-)
Biological Process (14 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
involved_in antennal development
inferred from expression pattern
(Dong et al., 2002)
involved_in antennal morphogenesis
inferred from mutant phenotype
(Dong et al., 2002)
involved_in imaginal disc-derived leg segmentation
inferred from mutant phenotype
(Kozu et al., 2006)
involved_in male courtship behavior
inferred from mutant phenotype
(Kuzin et al., 2014)
involved_in memory
inferred from mutant phenotype
(Kuzin et al., 2014)
involved_in negative regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Kozu et al., 2006)
involved_in R7 cell differentiation
inferred from direct assay
(Anderson et al., 2017)
involved_in regulation of dendrite morphogenesis
inferred from mutant phenotype
(Kim et al., 2006)
involved_in regulation of R7 cell differentiation
inferred from mutant phenotype
(Thanawala et al., 2013)
involved_in specification of animal organ identity
inferred from mutant phenotype
(Adachi-Yamada et al., 2005)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
involved_in regulation of DNA-templated transcription
inferred from electronic annotation with InterPro:IPR013767
(InterPro Project Members, 2004-)
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN004180845
(GO Reference Genome Project, 2011-)
involved_in response to xenobiotic stimulus
inferred from electronic annotation with InterPro:IPR039091
(InterPro Project Members, 2004-)
involved_in xenobiotic metabolic process
inferred from electronic annotation with InterPro:IPR039091
(InterPro Project Members, 2004-)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytosol
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nucleus
inferred from direct assay
(Céspedes et al., 2010)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
part_of aryl hydrocarbon receptor complex
inferred from biological aspect of ancestor with PANTHER:PTN004180845
(GO Reference Genome Project, 2011-)
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN004180845
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
spineless (ss) encodes a protein that plays a key role in defining the distal regions of the antenna and the leg. Its stochastic expression in R7 photoreceptors also controls the expression of color Rhodopsins (the product of Rh4 vs. the product of Rh3) in the two subsets of ommatidia. [Date last reviewed: 2019-03-21]
Gene Group (FlyBase)
BASIC HELIX-LOOP-HELIX TRANSCRIPTION FACTORS -
Basic helix-loop-helix (bHLH) transcription factors are sequence-specific DNA-binding proteins that regulate transcription. They are characterized by a 60 amino acid region comprising a basic DNA binding domain followed by a HLH motif formed from two amphipathic α-helices connected by a loop. bHLH transcription factors form homo- and hetero-dimeric complexes, which bind to a E box consensus sequence. (Adapted from PMID:15186484).
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
ss: spineless
Mutations at the spineless locus display three different phenotypes: (1) The spineless phenotype is characterized by the reduction in size of all bristles; (2) the aristapedia phenotype corresponds to the transformation of distal antennal segments, specifically the arista and the distal portion of the third antennal segment, into distal mesothoracic leg segments, i.e., tarsal segments; and (3) leg-segment fusion manifested as fusion of tarsal segments on all eight legs. Homeotic tissue does not conform to developmental compartment boundaries; therefore, ss does not qualify as a selector gene (Struhl). Expression of these phenotypes varies among alleles. Some alleles show only the spineless phenotype. Aristapedia alleles, symbolized ssa, vary in expression; weak alleles show a swelling of the third antennal joint and rudimentary tarsal transformation of the base of the arista; as expression becomes more extreme, more tarsal joints are formed until in the most extreme alleles four tarsal segments and terminal claws are formed. Tarsal fusions are characteristic of extreme aristapedia alleles.
ssv: spineless-variegated
Variegates for spineless character but completely mutant for aristapedia. Male sterile. RK2A.
ss1
Bristles only a little larger than hairs; dorsocentrals least reduced; postscutellars erect. No effect on legs or aristae. Growth of bristles slows during development [Lees and Waddington, 1943, Proc. Roy. Soc. (London), Ser. B 131: 87-110]. Dominant to aristapedia phenotype of all ssa alleles. RK1.
ssa: spineless-aristapedia
thumb
ssa: spineless-aristapedia
From Bridges and Brehme, 1944, Carnegie Inst. Washington Publ. No. 552: 179.
Antennae and aristae tarsuslike; mean number of tarsal segments = 4.0; incidence of claws = 97.1% (Garcia-Bellido, 1968, Genetics 59: 487-99); classified as an intermediate allele by Struhl. Third joint of antenna like parts of a tarsal row but with broad, flat, plate-like lobes below. Tarsal segments of legs display intermediate level of fusion (Struhl). Bristles like those of a medium to slight Minute. Frequent extra dorsocentral bristles. Transformed tissue is leg tissue in every attribute tested. Transformed tarsi elicit behavioral response similar to that of normal legs when exposed to sugar solutions (Deak, 1976, Nature 260: 252-54). Regions of aristae converted into tarsi not affected by mutants affecting aristae (e.g., th and al) but are affected by those operating on tarsi (e.g., fj, d, app, and ey) [Waddington, 1939, Growth, Suppl. 1, pp. 37-44; Braun, 1940, Genetics 25: 143-49; Mglinetz and Ivanov, 1975, Genetika 11(#11): 27-33; 1976, Genetika 12(#12): 87-94]. Dissociated cells from ssa antennal disks aggregate with dissociated leg-disk cells but not with those from wild-type antennal disks (Garcia-Bellido). Antennal disks from ssa larvae give rise to leg-like structures when transplanted into wild-type hosts as do both duplicated and regenerated antennal disks formed from eye-antenna-disk fragments (Gehring and Schubiger, 1975, J. Embryol. Exp. Morph. 33: 459-69); when disks are pretreated with colchicine, the developing structures are more aristalike (Vogt, 1947, Experientia 3: 156-59). Homozygous clones of ssa tissue produce antennal leg tissue when induced before (Roberts, 1964, Genetics 49: 593-98) but not after (Postlethwait and Girton, 1974, Genetics 76: 767-74) mid-third instar. Large clones conform to anterior and posterior compartments comparable to those induced in normal mesothoracic legs [Morata and Lawrence, 1979, Dev. Biol. 70: 355-71 (fig.)]. No maternal effect; temperature independent.
ssa40a
Cold sensitive; phenotype normal in flies raised at 29; fully penetrant at 17 [mean number of tarsal segments = 2.8; incidence of tarsal claws = 1.3% as recorded by Garcia-Bellido (1968, Genetics 59: 487-99) probably in flies raised at 25]. Phenocritical stage in first half of third instar [Grigliatti and Suzuki, 1971, Proc. Nat. Acad. Sci. USA 68: 1307-11 (fig.)]. Shifts from 17 to 29 early in third instar restrict transformation to base of arista; distal extent of transformation increases as shift is effected later in development (Grigliatti and Suzuki). Opposite response to temperature for antennal transformation but not tarsal fusion reported by Mglinetz (1977, Genetika 13: 70-75). Developmental compartments demonstrated in antennal legs [Morata and Lawrence, 1979, Dev. Biol. 70: 355-71 (fig.)].
ssaB: spineless-aristapedia of Bridges
Bristles of female like a slight Minute, especially postscutellars. At 25, aristae inconspicuously thickened at base, plumed or threadlike for rest of extent. At 18 penetrance is 7% and expression weak; at 14, ssaB enhanced and resembles ssa; al causes reduction of ssaB arista at 25 but has no effect on transformed arista at 14 (Villee, 1943, J. Exp. Zool. 93: 75-98). Legs frequently have lumps at second joint of tarsi; more pronounced in male and result in doubling of sex combs, which are strung along first and second fused joints. Eyes a little flattened. Except at low temperatures, all characters slight and may overlap wild type. ssaB/ss1 has slight Minute phenotype but wild-type legs and aristae. ssaB/ssaSp like ssaSp with large tarsal aristae. RK2. Struhl (1982, Genetics 102: 737-49) and Posakony.
*ssaH (A. Hannah Alava)
Homozygous viable and fertile. Displays enlargement of base of arista. Increased number of sex-comb teeth on basitarsus of first leg in males (12-38 with mean of 24 teeth per leg).
*ssasnB (A. Hannah Alava)
Description incomplete. Extreme fusion of tarsal segments; basitarsus of male first leg has 20-50 sex-comb teeth (mean = 40); sex-comb teeth not uncommon on second leg.
ssC1
Reported to be homozygous lethal and lethal in combination with Df(3R)bxd100 (Bownes, Bournias-Vardiabasis, and Spare, 1979, Mol. Gen. Genet. 174: 67-74); however both of these combinations viable in the hands of
*ssiso53: spineless-isoallele
Homozygote is wild type. ssiso53/ssa, ssiso53/ssa63c, and ssiso53/ssa53e have thickened proximal segments of aristae, like ssaB. RK3.
Summary (Interactive Fly)

bHLH PAS domain transcription factor - plays a central role in defining the distal regions of both the antenna and leg

Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\ss 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 E1JIM6)

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

Gene model reviewed during 5.48

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

Gene model reviewed during 5.56

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
ss-RA
FBtr0083318
NM_057400
5184
884
ss-RC
FBtr0306026
NM_001260211
4019
1015
ss-RD
FBtr0306027
NM_001170158
3997
1015
Additional Transcript Data and Comments
Reported size (kB)

5.4 (northern blot)

(Duncan et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
ss-PA
FBpp0082768
97.7
884
8.06
Q9VEV9
NP_476748
AAF55308
ss-PC
FBpp0297168
111.2
1015
7.62
E1JIM6
NP_001247140
AFH06458
ss-PD
FBpp0297169
111.2
1015
7.62
E1JIM6
NP_001163629
ACZ94925
Polypeptides with Identical Sequences

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

1015 aa isoforms: ss-PC, ss-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

884 (aa)

(Duncan et al., 1998)
Comments
External Data
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\ss 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).

-0.55

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
embryonic antennal segment
(Duncan et al., 1998)
embryonic stage 8
cephalic furrow | anterior to
(Duncan et al., 1998)
embryonic stage 9 -- 10
antennal primordium
(Fisher et al., 2012)
embryonic stage 9 -- 17
presumptive embryonic/larval peripheral nervous system
(Duncan et al., 1998)
embryonic maxillary segment
(Duncan et al., 1998)
embryonic mandibular segment
(Duncan et al., 1998)
embryonic labial segment
(Duncan et al., 1998)
ventral thoracic disc primordium
(Duncan et al., 1998)
embryonic stage 11 -- 12
antennal primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic antennal sense organ
(Fisher et al., 2012)
ventral head epidermis
(Fisher et al., 2012)
second instar larval stage
antennal disc
(Duncan et al., 1998)
third instar larval stage
antennal disc
(Duncan et al., 1998)
labial disc
(Duncan et al., 1998)
genital disc
(Duncan et al., 1998)
wing disc | restricted
(Duncan et al., 1998)
eye disc morphogenetic furrow
(Duncan et al., 1998)
leg disc
(Duncan et al., 1998)
haltere disc
(Duncan et al., 1998)
eye disc
(Duncan et al., 1998)
maxillary palp | precursor
(Duncan et al., 1998)
early third instar larval stage
presumptive tarsal segment 3 -- presumptive tarsal segment 4
(Natori et al., 2012)
pupal stage
trichogen cell
(Duncan et al., 1998)
sensory mother cell
(Duncan et al., 1998)
pharate adult stage P8
photoreceptor cell R7

Comment: in 60-80% of ommatidia

(Wernet et al., 2006)
interommatidial bristle
(Wernet et al., 2006)
Additional Descriptive Data

ss expression is observed in all bristle cells as well as in one photoreceptor cell (R7) in 60-80% of all ommatidia. Levels of ss vary considerably among R7 cells, ranging from very faint to very strong in 60-80% of R7 cells.

(Wernet et al., 2006)

ss transcripts are expressed in embryos starting in stage 8 in a crescent just anterior to the cephalic furrow. As germ band extension proceeds, expression is observed in the maxillary, labial, mandibular, and antennal segments. This is followed by a patch of expression in the thoracic segments that corresponds to the leg anlage. Expression is also observed in the peripheral nervous system. ss transcripts are first detected in leg discs in 2nd instar larvae in a central ring that corresponds to the presumptive tarsal region. The ring fades by late third instar. ss is then expressed in a patch in the anterior-proximal part of the disc that gives rise to thoracic structures. Staining in the antennal disc is first detected in 2nd instar larvae and is located in an oval patch in the central portion of the disc. After disc eversion, the limits of the intense ss staining corresponds to the boundary between the second and third antennal segments. In late third instar, ss is expressed in the part of the disc corresponding to the maxillary palp anlage. In the wing disc, ss expression is in the region corresponding to the presumptive notum and the wing hinge. Expression is also observed in the haltere, genital, and labial discs and in the morphogenetic furrow in the eye disc. At pupariation, expression is observed in sensory organ precursor cells in most discs. At later stages, expression is seen in developing bristle cells but not in the associated socket cells.

(Duncan et al., 1998)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
mesothoracic leg disc
(Tsubota et al., 2008)
metathoracic leg disc
(Tsubota et al., 2008)
metathoracic leg disc | dorsal | proximal
(Tsubota et al., 2008)
third instar larval stage -- prepupal stage
mesothoracic leg disc | dorsal | proximal
(Tsubota et al., 2008)
pupal stage
mesothoracic leg disc
(Tsubota et al., 2008)
metathoracic leg disc
(Tsubota et al., 2008)
pupal stage -- adult stage
photoreceptor cell R7 of yellow ommatidium
(Johnston and Desplan, 2014)
photoreceptor cell R7 | subset
(Johnston and Desplan, 2014)
adult stage
eye photoreceptor cell | subset
(Thanawala et al., 2013)
Additional Descriptive Data

ss is expressed in photoreceptor cells R7 of yellow ommatidium (yR), where Rh4 is active and Rh3 is repressed. This group comprises 65% of all R7 cells.

(Johnston and Desplan, 2014)

ss protein is expressed in a subset of photoreceptor cells in the adult retina, in on/off manner. Expression in cells in the dorsal third region of the retina, relative to the equator is lower than in the other regions.

(Thanawala et al., 2013)

ss is expressed in the dorsal-proximal region of the 2nd thoracic and third thoracic leg discs. Expression is first detected in the mid third instar larval stage and persists at least until the early pupal stage. The distal region of the ss domain overlaps with the most proximal region of the al domain.

(Bennett et al., 2008)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in cytosol
inferred from high throughput direct assay
(Lye et al., 2014)
located_in nucleus
inferred from direct assay
(Céspedes et al., 2010)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\ss 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
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 102 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 25 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ss
Transgenic constructs containing regulatory region of ss
Aberrations (Deficiencies and Duplications) ( 37 )
Inferred from experimentation ( 37 )
Gene disrupted in
Ab(3;?)ssD114.7
(Duncan et al., 1998)
Df(3R)Abd-BS10
(Tiong et al., 1987)
Df(3R)bxd100
(Tiong et al., 1985, Struhl, 1982)
Df(3R)Exel6269
(Ryder, 2004.4.26)
Df(3R)Exel7330
(Ryder, 2004.4.26)
Df(3R)P10
(Tiong et al., 1985, Struhl, 1982)
Df(3R)P115
(Tiong et al., 1985, Struhl, 1982)
Df(3R)sbd26
(Appel, 1995)
Df(3R)sbd104
(Tiong et al., 1985, Struhl, 1984, Struhl, 1982, Lewis, 1980)
Df(3R)ss-a
(Tiong et al., 1985, Struhl, 1982)
Df(3R)ss-D114.1
(Duncan et al., 1998)
Df(3R)ss-D114.4
(Duncan et al., 1998)
Df(3R)ss-D115.8
(Duncan et al., 1998)
In(3LR)P88
(Duncan et al., 1998)
In(3LR)TM6
In(3R)ssD114.9
(Duncan et al., 1998)
In(3R)ssD115.1
(Duncan et al., 1998)
In(3R)ssD116.1
(Duncan et al., 1998)
T(1;3)ssD114.3
(Duncan et al., 1998)
T(2;3)ssD115.5
(Duncan et al., 1998)
T(Y;3)ss114.12
(Duncan et al., 1998)
T(Y;3)ssD116.2
(Duncan et al., 1998)
Tp(3;3)ss-D115.12
(Duncan et al., 1998)
Tp(3;?)ssD114.6
(Duncan et al., 1998)
Gene duplicated in
Dp(3;1)O5
Dp(3;2)P10
(Melnick et al., 1993, Struhl, 1984, Lewis, 1980, Lewis, 1978)
Dp(3;3)bxd100
Dp(3;3)sbd104
(Struhl, 1982)
Dp(3;Y)Abd-BS10
(Tiong et al., 1987)
Gene not disrupted in
Df(3R)P2
(Tiong et al., 1985)
Df(3R)P9
(Tiong et al., 1985)
Df(3R)sbd45
(Appel, 1995)
Df(3R)sbd105
(Appel, 1995)
Df(3R)Ubx109
(Tiong et al., 1985)
Tp(3;2)iab2P10
(Duncan et al., 1998)
Gene partially disrupted in
Df(3R)RS3-32
(Duncan et al., 1998)
Gene not duplicated in
Dp(3;3)S462
(Struhl, 1982)
Inferred from location ( 7 )
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 | heat sensitive, with Scer\GAL4hs.PU
lethal | recessive
partially lethal
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
viable | recessive
Sterility
Allele
female semi-sterile
fertile
fertile | recessive
male sterile
sterile
Other Phenotypes
Allele
abnormal behavior
abnormal courtship behavior
abnormal gravitaxis
abnormal memory
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4477
abnormal neuroanatomy, with Scer\GAL4ppk.PG
abnormal neuroanatomy (with ss134)
abnormal neuroanatomy (with ssD115.7)
abnormal neuroanatomy | adult stage
abnormal neuroanatomy | somatic clone
abnormal size
abnormal visual perception
decreased occurrence of cell division
flightless | recessive
increased cell death | cell autonomous | somatic clone, with Scer\GAL4Act5C.PI
increased cell death | cell non-autonomous | somatic clone, with Scer\GAL4Act5C.PI
increased occurrence of cell division
visible
visible, with Scer\GAL4GMR.PU
visible, with Scer\GAL4pnr-MD237
visible, with Scer\GAL4rn-GAL4
visible (with Df(3R)ss-D114.4)
visible (with ss1)
visible (with ss134)
visible (with ssaBR30-71)
visible (with ssabr)
visible (with ssaP88)
visible (with ssD114.9)
visible (with ssD115.7)
visible (with sssta)
visible | dominant
visible | homeotic
visible | homeotic | recessive
visible | recessive
visible | somatic clone
Phenotype manifest in
Allele
abdominal tergite, with Scer\GAL4ptc-559.1
antenna
antenna (with ss134)
antenna (with ssaBR30-71)
antenna | distal | somatic clone
antennal disc (with Df(3R)ss-D114.4)
antennal segment
antennal sense organ
arista
arista (with Df(3R)ss-D114.4)
arista | cold sensitive
arista | proximal
bract
chaeta (with ss1)
chaeta (with ss134)
chaeta (with ssaBR30-71)
chaeta (with ssaP88)
dendrite & dorsal multidendritic neuron ddaB | somatic clone
dendrite & dorsal multidendritic neuron ddaC, with Scer\GAL4ppk.PG
dendrite & dorsal multidendritic neuron ddaC | somatic clone
dendrite & dorsal multidendritic neuron ddaD | somatic clone
dendrite & dorsal multidendritic neuron ddaE
dendrite & dorsal multidendritic neuron ddaE (with ss134)
dendrite & dorsal multidendritic neuron ddaE (with ssD115.7)
dendrite & dorsal multidendritic neuron ddaE | somatic clone
dendrite & dorsal multidendritic neuron ddaF | somatic clone
dendrite & lateral multidendritic neuron ldaB | somatic clone
dendrite & tracheal innervating neuron, with Scer\GAL4477
dorsomedial papilla
eye, with Scer\GAL4GMR.long
eye, with Scer\GAL4GMR.PU
eye-antennal disc
eye photoreceptor cell, with Scer\GAL4GMR.long
eye photoreceptor cell, with Scer\GAL4GMR.PF
eye photoreceptor cell | heat sensitive, with Scer\GAL4hs.PU
femur | distal
fifth segment of antenna (with Df(3R)ss-D114.4)
fourth segment of antenna (with Df(3R)ss-D114.4)
labial sensillum
labial sensillum | somatic clone
leg
leg, with Scer\GAL4rn-GAL4
leg | distal, with Scer\GAL4ptc-559.1
leg | ectopic
leg | ectopic (with ss134)
leg | ectopic (with ssaBR30-71)
leg | ectopic | somatic clone
leg anterior compartment
leg disc (with ssD114.9)
leg disc (with ssD115.7)
leg joint
leg joint (with ssa)
leg joint (with ssD114.9)
macrochaeta
maxillary palp
maxillary palp, with Scer\GAL4ptc-559.1
mesonotum, with Scer\GAL4pnr-MD237
mesothoracic leg
mesothoracic leg disc | larval stage
mesothoracic tarsal segment
mesothoracic tarsal segment | distal | ectopic
metatarsus
metatarsus (with ssa)
metatarsus (with ssD114.9)
metathoracic leg disc | larval stage
metathoracic tarsal segment
microchaeta
ommatidium, with Scer\GAL4GMR.long
pale ommatidium | ectopic
pale ommatidium | ectopic | somatic clone
photoreceptor cell R7 of pale ommatidium, with Scer\GAL4GMR.PF
photoreceptor cell R7 of pale ommatidium, with Scer\GAL4Pan-R7
photoreceptor cell R7 of pale ommatidium | ectopic | somatic clone
photoreceptor cell R7 of pale ommatidium | heat sensitive, with Scer\GAL4hs.PU
photoreceptor cell R7 of yellow ommatidium | ectopic, with Scer\GAL4GMR.PF
photoreceptor cell R7 of yellow ommatidium | ectopic, with Scer\GAL4Pan-R7
photoreceptor cell R7 of yellow ommatidium | ectopic | heat sensitive, with Scer\GAL4hs.PU
photoreceptor cell R7 of yellow ommatidium | somatic clone
photoreceptor cell R8 of pale ommatidium
photoreceptor cell R8 of yellow ommatidium
pretarsus (with ssD114.9)
pretarsus (with ssD115.7)
pretarsus | ectopic (with Df(3R)ss-D114.4)
prothoracic leg disc | larval stage
prothoracic tarsal segment
rostral membrane, with Scer\GAL4ptc-559.1
scutellar bristle
second segment of antenna (with Df(3R)ss-D114.4)
sex comb
sex comb | ectopic (with ssa)
sex comb | ectopic (with ssD114.9)
tarsal segment
tarsal segment 2
tarsal segment 2 (with ssD114.9)
tarsal segment 2 (with ssD115.7)
tarsal segment 3
tarsal segment 3 (with ssD114.9)
tarsal segment 3 (with ssD115.7)
tarsal segment 4
tarsal segment 4 (with ssD114.9)
tarsal segment 4 (with ssD115.7)
tarsal segment, with Scer\GAL4rn-GAL4
tarsal segment (with Df(3R)ss-D114.4)
tarsal segment (with ssabr)
tarsal segment (with sssta)
tarsal segment | ectopic
tarsal segment | ectopic (with Df(3R)ss-D114.4)
third segment of antenna
third segment of antenna (with Df(3R)ss-D114.4)
third segment of antenna | distal
thorax
tibia, with Scer\GAL4ptc-559.1
tracheal branch primordium, with Scer\GAL469B
trichogen cell, with Scer\GAL4pnr-MD237
unguis | ectopic
unguis | ectopic (with Df(3R)ss-D114.4)
ventral denticle belt | medial, with Scer\GAL469B
vertical bristle
vertical bristle 2
wing
wing, with Scer\GAL4ptc-559.1
wing, with Scer\GAL4sd-SG29.1
wing disc | cell autonomous | somatic clone, with Scer\GAL4Act5C.PI
wing margin, with Scer\GAL4NP7148
yellow ommatidium
yellow ommatidium | somatic clone
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) (28)
Hsap\AHR
9 of 14
Yes
Yes
1  
Hsap\AHRR
4 of 14
No
Yes
Hsap\ARNT2
1 of 14
No
No
Hsap\ARNT
1 of 14
No
No
Hsap\ARNTL2
1 of 14
No
No
Hsap\BMAL1
1 of 14
No
No
1  
Hsap\CLOCK
1 of 14
No
No
Hsap\EPAS1
1 of 14
No
No
Hsap\HELT
1 of 14
No
No
Hsap\HEY1
1 of 14
No
No
1  
Hsap\HEY2
1 of 14
No
No
1  
Hsap\HEYL
1 of 14
No
No
Hsap\HIF1A
1 of 14
No
No
1  
Hsap\HIF3A
1 of 14
No
No
Hsap\NCOA1
1 of 14
No
No
1  
Hsap\NCOA2
1 of 14
No
No
2  
Hsap\NCOA3
1 of 14
No
No
1  
Hsap\NPAS1
1 of 14
No
No
Hsap\NPAS2
1 of 14
No
No
Hsap\NPAS3
1 of 14
No
No
Hsap\NPAS4
1 of 14
No
No
Hsap\PER1
1 of 14
No
No
1  
Hsap\PER2
1 of 14
No
No
3  
Hsap\PER3
1 of 14
No
No
Hsap\SIM1
1 of 14
No
No
Hsap\SIM2
1 of 14
No
No
1  
Hsap\USF1
1 of 14
No
No
Hsap\USF2
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (16)
Rnor\Ahr
9 of 14
Yes
Yes
Rnor\Ahrr
5 of 14
No
Yes
Rnor\Arnt2
1 of 14
No
No
Rnor\Arntl2
1 of 14
No
No
Rnor\Arntl
1 of 14
No
No
Rnor\Arnt
1 of 14
No
No
Rnor\Clock
1 of 14
No
No
Rnor\Epas1
1 of 14
No
No
Rnor\Hif1a
1 of 14
No
No
Rnor\Hif3a
1 of 14
No
No
Rnor\Ncoa2
1 of 14
No
No
Rnor\Npas1
1 of 14
No
No
Rnor\Npas2
1 of 14
No
No
Rnor\Npas4
1 of 14
No
No
Rnor\Sim1
1 of 14
No
No
Rnor\Sim2
1 of 14
No
No
Mus musculus (laboratory mouse) (18)
Mmus\Ahr
9 of 14
Yes
Yes
2  
Mmus\Ahrr
5 of 14
No
Yes
Mmus\Arnt2
1 of 14
No
No
Mmus\Arntl2
1 of 14
No
No
Mmus\Arntl
1 of 14
No
No
Mmus\Arnt
1 of 14
No
No
Mmus\Clock
1 of 14
No
No
Mmus\Epas1
1 of 14
No
No
Mmus\Hif1a
1 of 14
No
No
Mmus\Hif3a
1 of 14
No
No
Mmus\Ncoa1
1 of 14
No
No
Mmus\Ncoa2
1 of 14
No
No
Mmus\Npas1
1 of 14
No
No
Mmus\Npas2
1 of 14
No
No
Mmus\Npas3
1 of 14
No
No
Mmus\Npas4
1 of 14
No
No
Mmus\Sim1
1 of 14
No
No
Mmus\Sim2
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (15)
Xtro\ahr
4 of 13
Yes
Yes
Xtro\cep170
4 of 13
Yes
Yes
Xtro\ahrr
3 of 13
No
Yes
Xtro\arnt
1 of 13
No
No
Xtro\brd7
1 of 13
No
No
Xtro\helt
1 of 13
No
No
Xtro\hey1
1 of 13
No
No
Xtro\htr2b
1 of 13
No
No
Xtro\LOC101734390
1 of 13
No
Yes
Xtro\ncoa2
1 of 13
No
No
Xtro\npas3
1 of 13
No
No
Xtro\npas4
1 of 13
No
No
Xtro\per2
1 of 13
No
No
Xtro\reep3
1 of 13
No
No
Xtro\sim1
1 of 13
No
No
Danio rerio (Zebrafish) (28)
Drer\ahr1b
9 of 14
Yes
Yes
Drer\ahr1a
8 of 14
No
Yes
Drer\ahr2
8 of 14
No
Yes
Drer\ahrrb
4 of 14
No
Yes
Drer\ahrra
1 of 14
No
Yes
Drer\arnt2
1 of 14
No
No
Drer\arnt
1 of 14
No
No
Drer\arntl1a
1 of 14
No
No
Drer\arntl1b
1 of 14
No
No
Drer\arntl2
1 of 14
No
No
Drer\clocka
1 of 14
No
No
Drer\clockb
1 of 14
No
No
Drer\epas1a
1 of 14
No
No
Drer\epas1b
1 of 14
No
No
Drer\hif1aa
1 of 14
No
No
Drer\hif1ab
1 of 14
No
No
Drer\hif1al2
1 of 14
No
No
Drer\hif1al
1 of 14
No
No
Drer\ncoa2
1 of 14
No
No
Drer\npas1
1 of 14
No
No
Drer\npas2
1 of 14
No
No
Drer\npas3
1 of 14
No
No
Drer\npas4a
1 of 14
No
No
Drer\npas4b
1 of 14
No
No
Drer\npas4l
1 of 14
No
No
Drer\per3
1 of 14
No
No
Drer\sim1a
1 of 14
No
No
Drer\sim1b
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (5)
Cele\ahr-1
10 of 14
Yes
Yes
Cele\aha-1
1 of 14
No
No
Cele\cky-1
1 of 14
No
No
Cele\hif-1
1 of 14
No
No
Cele\hlh-34
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (15)
Agam\AgaP_AGAP010259
7 of 12
Yes
Yes
Agam\AgaP_AGAP000056
1 of 12
No
No
Agam\AgaP_AGAP000773
1 of 12
No
No
Agam\AgaP_AGAP001856
1 of 12
No
No
Agam\AgaP_AGAP002942
1 of 12
No
No
Agam\AgaP_AGAP003844
1 of 12
No
No
Agam\AgaP_AGAP003965
1 of 12
No
No
Agam\AgaP_AGAP005655
1 of 12
No
No
Agam\AgaP_AGAP005711
1 of 12
No
No
Agam\AgaP_AGAP006022
1 of 12
No
No
Agam\AgaP_AGAP007450
1 of 12
No
No
Agam\AgaP_AGAP009664
1 of 12
No
No
Agam\AgaP_AGAP009748
1 of 12
No
No
Agam\AgaP_AGAP010058
1 of 12
No
No
Agam\AgaP_AGAP012873
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
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:ss. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (12)
Clk
2 of 13
dysf
2 of 13
cwo
1 of 13
cyc
1 of 13
gce
1 of 13
Hey
1 of 13
Met
1 of 13
per
1 of 13
sima
1 of 13
sim
1 of 13
tgo
1 of 13
trh
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 1 )
    Human Ortholog
    Disease
    Evidence
    References
    AHR; aryl hydrocarbon receptor
    model of  retinitis pigmentosa 85
    IEA
    (FlyBase, 2019-)
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    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?
    AHR; aryl hydrocarbon receptor
    9 of 14
    600253
     
    AHRR; aryl hydrocarbon receptor repressor
    4 of 14
    606517
        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
        Interaction Browsers
        View in FlyBase Interactions Browser   View in MIST tool (external link)
        Please see the Physical Interaction reports below for full details
        protein-protein
        Physical Interaction
        Assay
        References
        ss - tgo
        anti tag coimmunoprecipitation, western blot, two hybrid
        (Céspedes et al., 2010, Kudo et al., 2009, Emmons et al., 1999)
        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
        ss
        enhanceable
        al
        (Mikuta and Mglinets, 1978)
        ss
        suppressible
        BEAF-32
        (Jukam et al., 2016)
        ss
        suppressible
        dan
        (Emerald et al., 2003)
        ss
        suppressible
        Dll
        (Adachi-Yamada et al., 2005)
        ss
        enhanceable
        sta
        (Melnick et al., 1993)
        ss
        suppressible
        tgo
        (Adachi-Yamada et al., 2005)
        ss|Pc
        suppressible
        trx
        (Kuzin et al., 1997)
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        al
        enhanceable
        ss
        (Mglinets and Ivanov, 1975)
        Antp
        suppressible
        ss
        (Emerald and Cohen, 2004)
        BEAF-32
        suppressible
        ss
        (Jukam et al., 2016)
        cg
        enhanceable
        ss
        (Waddington, 1953)
        ct
        enhanceable
        ss
        (Ebacher et al., 2007)
        External Data
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        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
        Reactome Metabolic Pathways - An open-source, open access, manually curated and peer-reviewed metabolic pathway database.
        External Data
        Linkouts
        Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
        Class of Gene
        Genomic Location and Detailed Mapping Data
        Chromosome (arm)
        3R
        Recombination map
        3-58
        Cytogenetic map
        89B14-89B15
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        89B14-89B15
        Limits computationally determined from genome sequence between P{PZ}gish04895&P{EP}EP3171 and P{lacW}CSN5L4032
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        89C1-89C2
        (Duncan, 1998.2.24)
        89C1-89C2
        (determined by in situ hybridisation)
        (Duncan et al., 1998)
        Experimentally Determined Recombination Data
        Location

        3-58

        (FlyBase, 2016)

        3-58.4

        (Comeron et al., 2012)

        3-58.5

        (Neel, 1942)
        Left of (cM)
        Right of (cM)
        (Struhl, 1984)
        (Holden and Suzuki, 1973)
        Notes
        Stocks and Reagents
        Stocks (114)
        Genomic Clones (27)
        cDNA Clones (14)
         

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

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

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

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

                  polyclonal

                  (Tsubota et al., 2008)
                  Commercially Available Antibodies
                   
                  Cell Line Information
                  Publicly Available Cell Lines
                   
                    Other Stable Cell Lines
                     
                      Other Comments

                      ss regulates dendrite diversity in the dendritic arborization sensory neurons.

                      (Kim et al., 2006)

                      The ss gene product is incapable of high-affinity, specific binding of 3H-TCDD or 3H-BNF.

                      (Butler et al., 2001)

                      ss and tgo proteins interact to control antennal and tarsal development.

                      (Emmons et al., 1999)

                      Loss of function alleles cause three major phenotypes: transformation of distal antenna to leg, deletion of distal leg, and reduction in the size of most bristles. Ectopic expression causes transformation of the maxillary palp and distal leg to distal antenna, and induces formation of ectopic antenna in the rostral membrane.

                      (Duncan et al., 1998)

                      ss plays a central role in defining the distal regions of both the antenna and the leg. Loss of function phenotypes suggest that ss played a central role in the evolution of distal structures in arthropod limbs.

                      (Duncan et al., 1998)

                      Mutants do not exhibit defects in the denticle belt of hairs of the larvae.

                      (Dickinson and Thatcher, 1997)

                      Results suggests that ss interacts with genes of homeotic complexes: ss gene is capable of repressing the effect of ANTC and BXC gene products on the formation of distal structure of the antennae and leg.

                      (Kuzin et al., 1997)

                      Varied sensitivities of the antenna-to-leg transformation are found both spatially and temporally.

                      (Larsen et al., 1996)

                      Mutations at the spineless locus display three different phenotypes: (1) The spineless phenotype is characterized by the reduction in size of all bristles; (2) the aristapedia phenotype corresponds to the transformation of distal antennal segments, specifically the arista and the distal portion of the third antennal segment, into distal mesothoracic leg segments, i.e., tarsal segments; and (3) leg-segment fusion manifested as fusion of tarsal segments on all eight legs. Expression of these phenotypes varies among alleles. Some alleles show only the spineless phenotype. Aristapedia alleles, symbolized ssa, vary in expression; weak alleles show a swelling of the third antennal joint and rudimentary tarsal transformation of the base of the arista; as expression becomes more extreme, more tarsal joints are formed until in the most extreme alleles four tarsal segments and terminal claws are formed. Tarsal fusions are characteristic of extreme aristapedia alleles.

                      (Lindsley and Zimm, 1992)

                      Effects of ss mutations on geotaxis measured in a Hirsch-type geotaxis maze: alleles of the "aristapedia" type show extremely positive geotaxis.

                      (McMillan and McGuire, 1992)

                      Homeotic tissue does not conform to developmental compartment boundaries; therefore, ss does not qualify as a selector gene.

                      (Struhl, 1982)
                      Relationship to Other Genes
                      Source for database merge of
                      Additional comments
                      Nomenclature History
                      Source for database identify of
                      Nomenclature comments
                      Etymology
                      Synonyms and Secondary IDs (12)
                      Reported As
                      Symbol Synonym
                      AHR
                      (Kong et al., 2021, Butler et al., 2001)
                      AhR
                      (Kudo et al., 2009)
                      CG6993
                      (Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009.3.31)
                      SS
                      (McRobert, 1991)
                      Ss
                      (Miller et al., 2021, Sigrist-Flores et al., 2021, Chen and Desplan, 2020, Ruiz-Losada et al., 2018, Friedrich et al., 2016, Viets et al., 2016, Johnston and Desplan, 2014, Johnston, 2013, Rister et al., 2013, Thanawala et al., 2013, Jukam and Desplan, 2010, Pistillo et al., 2007, Reig et al., 2007, Adachi-Yamada et al., 2006)
                      dAHR
                      (Dai et al., 2022)
                      ss
                      (Ainsworth et al., 2025, Cao et al., 2025, Balakireva et al., 2024, Gurgo et al., 2024, Prud'homme, 2024, Bravo González-Blas et al., 2023, Fisher et al., 2023, Urban et al., 2023, Deshpande et al., 2022, Liu et al., 2021, Raun et al., 2021, Chang et al., 2020, Gozalo et al., 2020, Miller et al., 2020, Ahmad and Spens, 2019, Chai et al., 2019, Courgeon and Desplan, 2019, Menon et al., 2019, Papadopoulos et al., 2019, Viets et al., 2019, Akishina et al., 2018, Heinloth et al., 2018, Palliyil et al., 2018, Anderson et al., 2017, Erceg et al., 2017, Kojima, 2017, Yan et al., 2017, Jukam et al., 2016, Miller et al., 2016, Johnston and Desplan, 2014, Kuzin et al., 2014, Williams et al., 2014, Hammonds et al., 2013, Ibrahim et al., 2013, Thanawala et al., 2013, Cole et al., 2012.7.6, Manning et al., 2012, Papadopoulos et al., 2012, Spokony, 2012.12.12, Spokony and White, 2012.11.14, Bantignies et al., 2011, Céspedes et al., 2010, Duncan et al., 2010, Long et al., 2009, Parrish et al., 2009, Johnston and Desplan, 2008, Lebreton et al., 2008, Pueyo and Couso, 2008, Tsubota et al., 2008, Ebacher et al., 2007, Emmons et al., 2007, Firth and Baker, 2007, Lidder and Desplan, 2007, Joulia et al., 2006, Kim et al., 2006, Kozu et al., 2006, Wernet et al., 2006, Reeves and Posakony, 2005, Yoshii et al., 2004)
                      ssa
                      (Lebreton et al., 2006)
                      Name Synonyms
                      Spineless
                      (Kilo et al., 2021, Heinloth et al., 2018, Copf, 2015, Wernet et al., 2015, Johnston and Desplan, 2014, Puram and Bonni, 2013, Thanawala et al., 2013, Long et al., 2009, Reig et al., 2007)
                      aristapedia
                      (Dietrich, 2003, Garcia-Bellido, 1998)
                      spineless
                      (Earl et al., 2020, Papadopoulos et al., 2019, Johnston and Desplan, 2014, Baanannou et al., 2013, Hartl et al., 2011, Takemura and Adachi-Yamada, 2011, Céspedes et al., 2010, Duncan et al., 2010, Kudo et al., 2009, Johnston and Desplan, 2008, Lebreton et al., 2008, Nicholson et al., 2008, Tsubota et al., 2008, Emmons et al., 2007, Firth and Baker, 2007, Johnston and Desplan, 2007, Lidder and Desplan, 2007, Stark et al., 2007, Crews and Brenman, 2006, Hummel and Klämbt, 2006, Joulia et al., 2006, Wernet et al., 2006, Chan et al., 2005)
                      spineless-aristapedia
                      (Greenspan, 2003)
                      Secondary FlyBase IDs
                        Datasets (0)
                        Study focus (0)
                        Experimental Role
                        Project
                        Project Type
                        Title
                        Study result (0)
                        Result
                        Result Type
                        Title
                        External Crossreferences and Linkouts ( 60 )
                        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/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                        Other crossreferences
                        AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                        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
                        FlyMine - An integrated database for Drosophila genomics
                        InterPro - A database of protein families, domains and functional sites
                        KEGG Genes - Molecular building blocks of life in the genomic space.
                        MARRVEL_MODEL - MARRVEL (model organism gene)
                        PDB - An information portal to biological macromolecular structures
                        Linkouts
                        BioGRID - A database of protein and genetic interactions.
                        DroID - A comprehensive database of gene and protein interactions.
                        DRSC - Results frm RNAi screens
                        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                        Flygut - An atlas of the Drosophila adult midgut
                        FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                        MIST (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.
                        Reactome Metabolic Pathways - An open-source, open access, manually curated and peer-reviewed metabolic pathway database.
                        References (275)