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Gene Dmel\cos

General Information
Symbol	Dmel\cos	Species	D. melanogaster
Name	costa	Annotation symbol	CG1708
Feature type	protein_coding_gene	FlyBase ID	FBgn0000352
Gene Model Status	Current	Stock availability	6 publicly available
Also Known As	cos2, Cos-2, l(2)161/28, Cos2
Genomic Location
Chromosome (arm)	2R	Recombination map	2-57
Cytogenetic map	43B1-43B2	Sequence location	2R:3,269,669..3,274,617 [-]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents modENCODE GBrowse	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene costa is referred to in FlyBase by the symbol Dmel\cos (CG1708, FBgn0000352). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: smoothened binding; microtubule binding; protein binding; protein kinase binding; transcription factor binding. There is experimental evidence that it is involved in the biological process: smoothened signaling pathway; regulation of protein stability; negative regulation of sequence-specific DNA binding transcription factor activity; imaginal disc-derived wing morphogenesis; cytoplasmic sequestering of transcription factor; negative regulation of transcription factor import into nucleus; positive regulation of hh target transcription factor activity. 89 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; adult segment; organ system; adult mesothoracic segment; thoracic segment; imaginal precursor; dorsal thoracic disc; vagina; vulva; larval thorax; epiproct; non-connected developing system. It has one annotated transcript and one annotated polypeptide. Protein features are: Kinesin, motor domain. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of low expression. Peak expression observed within 00-12 hour embryonic stages, in adult female stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult head, larval/adult central nervous system, larval trachea, adult ovary, larval carcass. Comments on Affy2 ProbeSet: ProbeSet 1633554_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of cos. Gene sequence location is 2R:3269669..3274617.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	cos: costa (P. Simpson) Homozygotes, hemizygotes, and heteroallelic heterozygotes for class I alleles die as embryos with a normal cuticular phenotype. When derived from homozygous ovarian clones, however, abnormal deletion-duplication patterns in each segment are observed; posterior rows of abdominal denticles are replaced by mirror-image duplications of the anterior rows, including the segmental boundary, which is thereby duplicated; thoracic denticle belts missing; cos1 especially severely affected in this regard, including loss of entire segments. Heterozygotes derived from homozygous ovarian clones show slightly abnormal segmental patterning. Flies heterozygous for class I alleles or cos deficiencies and at the same time heterozygous for Cos alleles display pattern duplications of wings and halteres. Class II alleles are hemizygous lethal but homozygous viable. They display wing and haltere duplications indistinguishable from those found for cos1/Cos1 heterozygotes as described by Whittle. Flies heterozygous for class III alleles and Cos show moderately reduced viability and occasional wing duplications; flies homozygous for class III alleles show reduced viability and have wild-type phenotype, but enhance the pattern duplications associated with Cos/+; hemizygotes for class III alleles or heterozygotes with class I alleles are lethal or nearly so in the presence of Cos, the survivors invariably exhibiting pattern duplications of wings and halteres.
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2011_10
FB2012_01	Sequence features BKN26270 HFA07598
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 43B1-43B2   Limits computationally determined from genome sequence between P{lacW}vimark16722 and P{lacW}cosk16101
Experimentally Determined Cytological Location
Cytogenetic map Notes References 43B1-43B2   (determined by in situ hybridisation)   (Spradling et al., 1999) 43B2-43C2     (Sisson et al., 1997) 43A1-43C2   (determined by in situ hybridisation)   (Scott, 1992) 43B1-43B2     (BDGP Project Members, 1994-1999) 43B-43B   (determined by in situ hybridisation)   (Torok et al., 1993) 42E-43E   Placed in 42E--43E by deficiency mapping (details unspecified).   (Liu and Montell, 1999)
Experimentally Determined Recombination Data
Location	2-57 (Grau and Simpson, 1987)
Left of (cM)	cn (Grau and Simpson, 1987)
Right of (cM)	pk (Grau and Simpson, 1987)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\cos for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) cos-RA FBtr0089015  NM_057744   4772   1201
Additional Transcript Data & Comments
Reported size (kB)	4.9 (northern blot) (Sisson et al., 1997)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein cos-PA   FBpp0088087   133.1   1201   5.64     NP_477092   AAF59270
Additional Polypeptide Data & Comments
Reported size (kDa)	1201 (aa); 175 (kD observed); 133 (kD predicted) (Sisson et al., 1997)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Kinesin, motor domain (IPR001752)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA441088   AA735169   AE013599 AAF59270   AC005452   AF019250 AAB66813   AI456015   AI515298   AI515829   AQ034038   AW943304   BI376165   BI376178   BI607787   BI616096   BI628040   BT001504 AAN71259   BT015259 AAT94488   BT044167   CL610224   CO265212   CO269217   CO273030   CO276296   CO293072   CO306013   CO313286   CO321998   CO323177   EC063700   EC064671   EC071187   EC075013   EC079056   EC088050   EC198861   EC223386   EC238867
UniProtKB/Swiss-Prot
UniProtKB/TrEMBL	O16844
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage & larval stage     (Sisson et al., 1997)
Additional Descriptive Data
	cos transcripts are detected at high levels during the first 4 hours of embryogenesis, at moderate levels between 8 and 12 hours and at low levels in later embryos. They are also detected in third instar larvae on northern blots. (Sisson et al., 1997)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage | early     (Sisson et al., 1997) embryonic stage   segment | anterior   (Sisson et al., 1997) embryonic stage 9 -- 12   organism | segment polarity expression pattern   (Sisson et al., 1997) larval stage   wing disc | anterior   (Sisson et al., 1997)
Additional Descriptive Data
	In syncytial embryos, cos protein is distributed uniformly in the cortical cytoplasm. In the late syncytial embryo, it accumulates between and apical to nuclei. From the surface, the pattern looks like a honeycomb. Throughout cellularization, cos protein is associated with furrow canals and during late cellularization is associated with expanded furrow canals. In cellular blastoderm embryos and during gastrulation, cos protein is in the cytoplasm and at the periphery of all cells. A striped pattern of cos protein is observed beginning in late stage 9 and decaying in stage 12. Each stripe is continuous along the dorsal/ventral axis both in the ectoderm and the underlying mesoderm. The stripes appear to form in the anterior compartment of each segment. In wing discs, cos protein levels are elevated in the anterior compartment (Sisson et al., 1997)
Marker for
Subcellular Localization
CV Term	cytoplasm (Sisson et al., 1997)
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. modENCODE Temporal Expression Data for FBgn0000352    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0000352 Very low expression bin max Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of low expression. Peak expression observed within 00-12 hour embryonic stages, in adult female stages. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0000352 expression level Developmental Stage   Expression Level embryo 00-02hr    2583 embryo 02-04hr    3758 embryo 04-06hr    2968 embryo 06-08hr    1995 embryo 08-10hr    1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day    1466 adult female 30day    1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (2583) embryo 02-04hr  (3758) embryo 04-06hr  (2968) embryo 06-08hr  (1995) embryo 08-10hr  (1771) embryo 10-12hr  (1100) embryo 12-14hr  (963) embryo 14-16hr  (1114) embryo 16-18hr  (985) embryo 18-20hr  (641) embryo 20-22hr  (1148) embryo 22-24hr  (1054) larva L1  (518) larva L2  (302) larva L3 12hr old  (222) larva L3 puffstage 1-2  (282) larva L3 puffstage 3-6  (594) larva L3 puffstage 7-9  (718) white prepupae new  (781) white prepupae 12hr  (1062) white prepupae 24hr  (1327) pupae 2d postWPP  (701) pupae 3d postWPP  (357) pupae 4d postWPP  (248) adult male 01day  (218) adult male 05day  (222) adult male 30day  (247) adult female 01day  (760) adult female 05day  (1466) adult female 30day  (1493) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (2583) embryo 02-04hr  (3758) embryo 04-06hr  (2968) embryo 06-08hr  (1995) embryo 08-10hr  (1771) embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day  1466 adult female 30day  1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    2583 embryo 02-04hr    3758 embryo 04-06hr    2968 embryo 06-08hr    1995 embryo 08-10hr    1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day    1466 adult female 30day    1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    2583 embryo 02-04hr    3758 embryo 04-06hr    2968 embryo 06-08hr    1995 embryo 08-10hr    1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day    1466 adult female 30day    1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0000352 expression level Developmental Stage   Expression Level embryo 00-02hr    2583 embryo 02-04hr    3758 embryo 04-06hr    2968 embryo 06-08hr    1995 embryo 08-10hr    1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day    1466 adult female 30day    1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (2583) embryo 02-04hr  (3758) embryo 04-06hr  (2968) embryo 06-08hr  (1995) embryo 08-10hr  (1771) embryo 10-12hr  (1100) embryo 12-14hr  (963) embryo 14-16hr  (1114) embryo 16-18hr  (985) embryo 18-20hr  (641) embryo 20-22hr  (1148) embryo 22-24hr  (1054) larva L1  (518) larva L2  (302) larva L3 12hr old  (222) larva L3 puffstage 1-2  (282) larva L3 puffstage 3-6  (594) larva L3 puffstage 7-9  (718) white prepupae new  (781) white prepupae 12hr  (1062) white prepupae 24hr  (1327) pupae 2d postWPP  (701) pupae 3d postWPP  (357) pupae 4d postWPP  (248) adult male 01day  (218) adult male 05day  (222) adult male 30day  (247) adult female 01day  (760) adult female 05day  (1466) adult female 30day  (1493) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (2583) embryo 02-04hr  (3758) embryo 04-06hr  (2968) embryo 06-08hr  1995 embryo 08-10hr  1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day  1466 adult female 30day  1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    2583 embryo 02-04hr    3758 embryo 04-06hr    2968 embryo 06-08hr    1995 embryo 08-10hr    1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day    1466 adult female 30day    1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    2583 embryo 02-04hr    3758 embryo 04-06hr    2968 embryo 06-08hr    1995 embryo 08-10hr    1771 embryo 10-12hr    1100 embryo 12-14hr    963 embryo 14-16hr    1114 embryo 16-18hr    985 embryo 18-20hr    641 embryo 20-22hr    1148 embryo 22-24hr    1054 larva L1    518 larva L2    302 larva L3 12hr old    222 larva L3 puffstage 1-2    282 larva L3 puffstage 3-6    594 larva L3 puffstage 7-9    718 white prepupae new    781 white prepupae 12hr    1062 white prepupae 24hr    1327 pupae 2d postWPP    701 pupae 3d postWPP    357 pupae 4d postWPP    248 adult male 01day    218 adult male 05day    222 adult male 30day    247 adult female 01day    760 adult female 05day    1466 adult female 30day    1493 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Developmental Stage   Expression Level embryo 00-02hr     embryo 02-04hr     embryo 04-06hr     embryo 06-08hr     embryo 08-10hr     embryo 10-12hr     embryo 12-14hr     embryo 14-16hr     embryo 16-18hr     embryo 18-20hr     embryo 20-22hr     embryo 22-24hr     larva L1     larva L2     larva L3 12hr old     larva L3 puffstage 1-2     larva L3 puffstage 3-6     larva L3 puffstage 7-9     white prepupae new     white prepupae 12hr     white prepupae 24hr     pupae 2d postWPP     pupae 3d postWPP     pupae 4d postWPP     adult male 01day     adult male 05day     adult male 30day     adult female 01day     adult female 05day     adult female 30day     FlyAtlas Anatomical Expression Data for FBgn0000352    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0000352 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult head, larval/adult central nervous system, larval trachea, adult ovary, larval carcass. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Linear, scaled to maximum FBgn0000352 expression level Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0000352 expression level Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    440.875 Larval Midgut    43.5 Larval Hindgut    52.1 Larval Malpighian Tubules    34.9 Larval Fat Body    90.2 Larval Salivary Gland    51.5 Larval Trachea    241.85 Larval Carcass    128.95 Adult Head    106.7 Adult Eye    69.725 Adult Brain    362.2 Adult Thoracic-Abdominal Ganglion    180.2 Adult Crop    68.4 Adult Midgut    41 Adult Hindgut    34.9 Adult Malpighian Tubules no informative data Adult Fat Body    42.4 Adult Salivary Gland no informative data Adult Heart    61.525 Adult VirginFemale Spermatheca    69.4 Adult InseminatedFemale Spermatheca    65.7 Adult Ovary    429 Adult Testis    21.7 Adult Male Accessory Gland    34 Adult Carcass no informative data Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules no informative data Adult Fat Body     Adult Salivary Gland no informative data Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass no informative data FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    106.7   NA  Eye    69.725   NA  Brain    362.2   440.875  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    180.2   NA  Crop    68.4   43.5  Midgut    41   52.1  Hindgut    34.9   34.9  Malpighian Tubules    no informative data   90.2  Fat Body    42.4   51.5  Salivary Gland    no informative data   NA  Heart    61.525   241.85  Trachea    NA   NA  VirginFemale Spermatheca    69.4   NA  InseminatedFemale Spermatheca    65.7   NA  Ovary    429   NA  Testis    21.7   NA  Male Accessory Gland    34   128.95  Carcass    no informative data modENCODE Temporal Expression Data (Graveley et al., 2011) FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010) mE2_34_mRNA_expression_cluster_17
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000352 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG1708-RA
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal cosk16128 lethal | embryonic stage cos3 lethal | embryonic stage | germline clone | maternal effect | recessive cos2R-183-20 lethal | embryonic stage | rescuable maternal effect cos1 cos2 cos5 cos7 lethal | partially (with cos3) cos11 lethal | partially (with cos11) cos3 lethal | pupal stage cos7 lethal | recessive cos1 cos2 cos2R-183-20 cos3 cos4 cos5 cos6 cos8 cos9 cos10 cos12 cos14 cos15 cos16 cos52B7 cosk16101 cosk16122 cosk16128 semi-viable | recessive cos7 viable cos11 Other Phenotypes Allele hyperplasia cosk16128 visible, with Scer\GAL4Bx-MS1096 cosS182N.Scer\UAS.T:Avic\GFP cosS182N.Scer\UAS cosS182T.Scer\UAS.T:Avic\GFP cosScer\UAS.T:Avic\GFP visible, with Scer\GAL4ptc-559.1 cosCSBD.Scer\UAS.T:Ivir\HA1 visible | dominant cosk16101 visible | recessive cos7 visible | somatic clone cosunspecified wild-type cos+t6.1 Phenotype manifest in Allele 1st posterior cell, with Scer\GAL4ptc-559.1 cosCSBD.Scer\UAS.T:Ivir\HA1 border follicle cell | ectopic cos52B7 border follicle cell | ectopic | somatic clone cos52B7 border follicle cell | somatic clone cos52B7 border follicle cell | somatic clone | supernumerary cos52B7 clasper cosV1 cosV5 denticle belt | embryonic stage cos1 cos7 denticle belt | embryonic stage | germline clone cos2 cos5 egg chamber | somatic clone, with Scer\GAL4Act5C.PP cosScer\UAS.cHa embryonic/first instar larval cuticle | germline clone cos1 cos2 cos5 embryonic abdominal segment | germline clone | maternal effect cos2R-183-20 embryonic epidermis cos5 embryonic epidermis | germline clone cos1 cos2 embryonic segment | germline clone cos1 cos2 cos5 embryonic segment | germline clone | maternal effect cos2R-183-20 epiproct cosV1 cosV5 female gonopod | dorsal cosV1 cosV5 female gonopod | ventral cosV1 cosV5 follicle cell | ectopic | somatic clone cos1 genital arch cosV1 cosV5 germarium | somatic clone, with Scer\GAL4Act5C.PP cosScer\UAS.cHa haltere cos7 hypandrial process cosV1 cosV5 hypoproct cosV1 cosV5 imaginal disc cosk16101 cosk16128 interfollicle cell | somatic clone, with Scer\GAL4Act5C.PP cosScer\UAS.cHa lateral plate cosV1 cosV5 oocyte | somatic clone cos1 polar follicle cell | ectopic | somatic clone cos52B7 cosunspecified pupal epidermis cos5 sex comb | ectopic | somatic clone cos1 vulva cosV1 cosV5 wing cos1 cos7 cosk16101 wing (with cos3) cos11 wing (with cos11) cos3 wing | anterior, with Scer\GAL4Bx-MS1096 cosS182N.Scer\UAS cosS182N.Scer\UAS.T:Avic\GFP wing | anterior compartment cosV1 wing | ectopic, with Scer\GAL4Bx-MS1096 cosS182N.Scer\UAS cosS182N.Scer\UAS.T:Avic\GFP wing | posterior compartment cosV1 wing | somatic clone cosunspecified wing blade, with Scer\GAL4Bx-MS1096 cosS182T.Scer\UAS.T:Avic\GFP cosScer\UAS.T:Avic\GFP wing disc cos2 wing disc | anterior | somatic clone, with Scer\GAL4Act5C.PP cosS182N.Scer\UAS cosS182N.Scer\UAS.T:Avic\GFP wing vein cos1 wing vein L3, with Scer\GAL4Bx-MS1096 cosS182T.Scer\UAS.T:Avic\GFP wing vein L3 | ectopic | somatic clone cosunspecified wing vein L3 | proximal, with Scer\GAL4ptc-559.1 cosCSBD.Scer\UAS.T:Ivir\HA1 wing vein L4, with Scer\GAL4Bx-MS1096 cosS182T.Scer\UAS.T:Avic\GFP wing vein L4 | proximal, with Scer\GAL4ptc-559.1 cosCSBD.Scer\UAS.T:Ivir\HA1
Classical Alleles ( 28 )
For All Classical Alleles Show
Allele of cos Class Mutagen Stocks Known lesion cos5 loss of function allele ethyl methanesulfonate 3 -- cosk16101 P-element activity 2 Yes cosEY08735 P-element activity 1 -- cos2 amorphic allele - genetic evidence, loss of function allele X ray 0 Yes cos10 ethyl methanesulfonate 0 -- cos11 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- cos12 ethyl methanesulfonate 0 -- cos14 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes cos15 ethyl methanesulfonate 0 -- cos16 ethyl methanesulfonate 0 -- cos1 loss of function allele ethyl methanesulfonate 0 -- cos2R-183-20 ethyl methanesulfonate 0 -- cos31 0 -- cos3 loss of function allele X ray 0 -- cos4 loss of function allele ethyl methanesulfonate 0 -- cos52B7 ethyl methanesulfonate 0 -- cos6 loss of function allele ethyl methanesulfonate 0 -- cos7 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- cos8 ethyl methanesulfonate 0 -- cos9 ethyl methanesulfonate 0 -- cosk16122 P-element activity 0 -- cosk16128 P-element activity 0 -- cosunspecified 0 -- cosV1 wild-type allele, loss of function allele spontaneous 0 Yes cosV2 loss of function allele spontaneous 0 -- cosV3 loss of function allele spontaneous 0 -- cosV4 loss of function allele spontaneous 0 -- cosV5 loss of function allele ethyl methanesulfonate 0 --
Alleles Carried on Transgenic Constructs ( 61 )
For All Alleles Carried on Transgenic Constructs Show
Allele of cos Class Mutagen Stocks Known lesion cos+t6.1 in vitro construct - genomic fragment 0 Yes cos+t6.5 in vitro construct - genomic fragment 0 Yes cos1-1201.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cos1-605.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cos1-751.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cos1-900.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cos115-1201.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cos501-800.Act5C.T:Ivir\HA1,T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cos6.5.T:Avic\GFP in vitro construct 0 Yes cos6.5ΔC.T:Avic\GFP in vitro construct 0 Yes cos605-1201.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cos900-1201.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosAct5C.L.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosAct5C.PC in vitro construct - regulatory fusion 0 Yes cosAct5C.T:Disc\RFP-mRFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosAct5C.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosAct5C.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosAct5C.T:Rren\LUC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosAct5C.Z.T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes coscargo.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosCC.Scer\UAS.T:Ivir\HA1 in vitro construct - deletion 0 Yes cosCSBD.Act5C.T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosCSBD.Act5C.T:Ivir\HA1,T:HIV-1\nes in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosCSBD.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosCSBD.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosCT1.Scer\UAS.T:Ivir\HA1 in vitro construct - deletion 0 Yes cosCT2.Scer\UAS.T:Ivir\HA1 in vitro construct - deletion 0 Yes cosdsRNA.621-841 in vitro construct - RNAi 0 Yes cosdsRNA.cGa in vitro construct - RNAi 0 Yes cosdsRNA.cLa in vitro construct - RNAi 0 Yes cosGD9280 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes cosKK106270 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes cosM.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosMB.Scer\UAS.T:Ivir\HA1 in vitro construct - deletion 0 Yes cosmotor.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosneck.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosS182N.Scer\UAS.T:Avic\GFP in vitro construct - amino acid replacement 0 Yes cosS182N.Scer\UAS in vitro construct - amino acid replacement 0 Yes cosS182T.Scer\UAS.T:Avic\GFP in vitro construct - amino acid replacement 0 Yes cosS182T.Scer\UAS in vitro construct - amino acid replacement 0 Yes cosS572A.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis 0 Yes cosS572D.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion in vitro construct - site directed mutagenesis 0 Yes cosScer\UAS.cHa in vitro construct - regulatory fusion 0 Yes cosScer\UAS.cRa in vitro construct - regulatory fusion 0 Yes cosScer\UAS.cUa in vitro construct - regulatory fusion 0 Yes cosScer\UAS.T:Avic\GFP in vitro construct - coding region fusion 0 Yes cosScer\UAS.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosScer\UAS.T:Zzzz\FLAG in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosstalk.Act5C.T:Hsap\MYC in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosΔC.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosΔC.Scer\UAS.T:Avic\GFP,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosΔC.Scer\UAS.T:Ivir\HA1 in vitro construct - site directed mutagenesis 0 Yes cosΔcargo.Act5C in vitro construct - deletion in vitro construct - regulatory fusion 0 Yes cosΔMotor.Scer\UAS.T:Avic\GFP,T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes cosΔMotor.Scer\UAS.T:Zzzz\His6 in vitro construct - regulatory fusion in vitro construct - deletion 0 Yes cosΔN.Act5C.T:Ivir\HA1,T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosΔN.Act5C.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes cosΔN1.Scer\UAS.T:Ivir\HA1 in vitro construct - deletion 0 Yes cosΔN2.Scer\UAS.T:Ivir\HA1 in vitro construct - deletion 0 Yes cosΔNC.Scer\UAS.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes cosΔNeck.Scer\UAS.T:Avic\GFP in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)CA58 (Grau and Simpson, 1987) Df(2R)cn-S5 (Heitzler et al., 1993) Df(2R)cos-2 (Heitzler et al., 1993) Df(2R)cos-3 (Heitzler et al., 1993) Df(2R)Drlrv3 (Heitzler et al., 1993) Df(2R)Drlrv5 (Heitzler et al., 1993) Df(2R)Drlrv7 (Heitzler et al., 1993) Df(2R)Drlrv15 (Heitzler et al., 1993) Df(2R)Drlrv17 (Heitzler et al., 1993) Df(2R)Drlrv18 (Heitzler et al., 1993) Df(2R)Drlrv20 (Heitzler et al., 1993) Df(2R)Drlrv21 (Heitzler et al., 1993, Sisson et al., 1997) Df(2R)Drlrv24 (Heitzler et al., 1993) Df(2R)Drlrv25 (Heitzler et al., 1993) Df(2R)Drlrv26 (Heitzler et al., 1993) Df(2R)Drlrv28 (Heitzler et al., 1993) Df(2R)Drlrv30 (Heitzler et al., 1993) Df(2R)DX1 (Heitzler et al., 1993) Df(2R)DX14 (Heitzler et al., 1993) Df(2R)nap16 (Spradling et al., 1999, BDGP Project Members, 1994-1999) Df(2R)Npk-rv1 (Roote and Ashburner, 2005.6) Df(2R)pk78k (Spradling et al., 1999, Grau and Simpson, 1987, BDGP Project Members, 1994-1999) Df(2R)pk78s (Grau and Simpson, 1987, BDGP Project Members, 1994-1999) Df(2R)sple-D2 (Heitzler et al., 1993) Df(2R)sple-D3 (Heitzler et al., 1993) Df(2R)sple-J1 (Heitzler et al., 1993) Df(2R)sple-J2 (Heitzler et al., 1993) Df(2R)sple-N3 (Heitzler et al., 1993) Df(2R)ST1 (Spradling et al., 1999, Grau and Simpson, 1987, BDGP Project Members, 1994-1999)
Not disrupted in	Df(2R)cn-S6 (Heitzler et al., 1993) Df(2R)cn-S8 (Heitzler et al., 1993) Df(2R)Drlrv22 (Heitzler et al., 1993) Df(2R)EW60 (Heitzler et al., 1993, Sisson et al., 1997) Df(2R)H23 (Heitzler et al., 1993) Df(2R)nap2 (Heitzler et al., 1993) Df(2R)NCX5 (Heitzler et al., 1993) Df(2R)NCX8 (Heitzler et al., 1993) Df(2R)NCX9 (Heitzler et al., 1993) Df(2R)NCX10 (Heitzler et al., 1993) Df(2R)NCX11 (Heitzler et al., 1993, Sisson et al., 1997) Df(2R)NCX13 (Heitzler et al., 1993) Df(2R)Npk-rv2 (Roote and Ashburner, 2005.6) Df(2R)Npk-rv4 (Roote and Ashburner, 2005.6) Df(2R)pk-N5 (Heitzler et al., 1993) Df(2R)pwn5 (Heitzler et al., 1993) Df(2R)sple-D1 (Heitzler et al., 1993, Sisson et al., 1997) Df(2R)tor-rx6 (Heitzler et al., 1993)
Duplicated in	Dp(2;3)P32 (Grau and Simpson, 1987)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD9280} NA P{KK106270} NA P{UAS-Cos2.CC.HA} NA P{UAS-Cos2.CT1.HA} NA P{UAS-Cos2.CT2.HA} NA P{UAS-Cos2.FLAG} NA P{UAS-Cos2.MB.HA} NA P{UAS-Cos2.W} NA P{UAS-Cos2.ΔC.HA} NA P{UAS-Cos2.ΔN1.HA} NA P{UAS-Cos2.ΔN2.HA} NA P{UAS-Cos2.ΔNC.HA} NA P{UAS-cos.GFP} NA P{UAS-cos.H} NA P{UAS-cos.HA-CSBD} NA P{UAS-cos.R} NA P{UAS-cos.S182N} NA P{UAS-cos.S182T} NA P{UAS-cos.S182T-GFP} NA P{UAS-cos.U} NA P{UAS-cos.ΔC-GFP} NA P{UAS-cos.ΔMotor} NA P{UAS-cos.ΔMotor-GFP} NA P{UAS-HA-Cos2.S572A} NA P{UAS-HA-Cos2.S572D} NA characterization construct P{Act5C-mRFP-cos} NA P{cos.K6.1} NA P{cos.K6.5} NA P{cos.K6.5-GFP} NA P{cos.K6.5ΔC-GFP} NA P{kinco.HA} NA
Insertions
	Type of insertions Name Expression data insertion of enhancer trap P{lacW}cosk16101 No P{lacW}cosk16122 No P{lacW}cosk16128 No P{lacW}l(2)SH0394SH0394 No insertion of mobile activating element P{EPgy2}cosEY08735 NA P{GSV1}C371C371 NA miscellaneous insertions ?{}cosV1 NA
Gene Ontology: Function, Process & Cellular Component ( 26 unique terms )
Terms Based on Experimental Evidence ( 17 terms )
Molecular Function
CV term Evidence References
microtubule binding inferred from direct assay (Sisson et al., 1997) protein binding inferred from physical interaction with fu (Fukumoto et al., 2001) protein kinase binding inferred from physical interaction with fu (Ascano et al., 2002) smoothened binding inferred from physical interaction with smo (Jia et al., 2003, Ogden et al., 2003) transcription factor binding inferred from physical interaction with ci (Wang et al., 2000)
Biological Process
CV term Evidence References
cytoplasmic sequestering of transcription factor inferred from mutant phenotype (Wang et al., 2000) imaginal disc-derived wing morphogenesis inferred from mutant phenotype (Dworkin and Gibson, 2006) negative regulation of sequence-specific DNA binding transcription factor activity inferred from mutant phenotype (Wang et al., 2000) negative regulation of transcription factor import into nucleus inferred from mutant phenotype (Wang and Holmgren, 2000) positive regulation of hh target transcription factor activity inferred from mutant phenotype (Wang and Holmgren, 2000) regulation of protein stability inferred from direct assay (Liu et al., 2007) smoothened signaling pathway inferred from mutant phenotype (Wang et al., 2000) inferred from physical interaction with smo (Jia et al., 2003, Ogden et al., 2003)
Cellular Component
CV term Evidence References
cytoplasm inferred from direct assay (Sisson et al., 1997) Hedgehog signaling complex inferred from physical interaction with ci AND inferred from physical interaction with fu (Stegman et al., 2000) microtubule associated complex inferred from physical interaction with ci (Sisson et al., 1997) plasma membrane inferred from direct assay (Stegman et al., 2004) vesicle membrane inferred from direct assay (Stegman et al., 2004)
Terms Based on Predictions or Assertions ( 11 terms )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR001752 (FlyBase Curators et al., 2004-) microtubule motor activity inferred from electronic annotation with InterPro:IPR001752 (FlyBase Curators et al., 2004-) inferred from sequence or structural similarity (Goldstein and Gunawardena, 2000) motor activity inferred from sequence or structural similarity with Khc (Sisson et al., 1997)
Biological Process
CV term Evidence References
microtubule-based movement inferred from electronic annotation with InterPro:IPR001752 (FlyBase Curators et al., 2004-) inferred from sequence or structural similarity (Goldstein and Gunawardena, 2000) negative regulation of smoothened signaling pathway traceable author statement (Ogden et al., 2004) ovarian follicle cell development traceable author statement (Dobens and Raftery, 2000) positive regulation of smoothened signaling pathway traceable author statement (Ogden et al., 2004) regulation of proteolysis non-traceable author statement (Ou et al., 2003) smoothened signaling pathway traceable author statement (Leahy, 1999)
Cellular Component
CV term Evidence References
cytoplasm traceable author statement (Leahy, 1999) kinesin complex inferred from sequence or structural similarity (Goldstein and Gunawardena, 2000) inferred from sequence or structural similarity with Khc (Sisson et al., 1997) inferred from sequence or structural similarity with Saccharomyces KIP3 (FlyBase, 1992-)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions cos allele Gene References cos1 fu (Besse et al., 2002) cos2 hh (Lum, 2003) cos+t6.1 smo (Hooper, 2003) cosV1 Pka-C1 (Grau and Simpson, 1987, Capdevila et al., 1994, Sanchez-Herrero et al., 1996) Pka-R2 (Grau and Simpson, 1987, Collier et al., 2004) cosV2 Pka-C1 (Grau and Simpson, 1987) Pka-R2 (Grau and Simpson, 1987) cosV3 Pka-C1 (Grau and Simpson, 1987) Pka-R2 (Grau and Simpson, 1987) cosV4 Pka-C1 (Grau and Simpson, 1987) Pka-R2 (Grau and Simpson, 1987) cosV5 Pka-C1 (Sanchez-Herrero et al., 1996) unspecified fu (Preat et al., 1993) hh (Ruel et al., 2003) ptc (Ruel et al., 2003) smo (Ogden et al., 2006, Ruel et al., 2003) Su(fu) (Preat et al., 1993)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 61534 DroID - A comprehensive database of gene and protein interactions. • FBgn0000352 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35653
Orthologs
Genome-wide drosophilid orthologs	Dana\GF12234 Dere\GG10758 Dgri\GH20742 Dmoj\GI20825 Dper\GL10940 Dpse\GA14314 Dsec\GM20803 Dsim\GD10265 Dvir\GJ20559 Dwil\GK22018 Dyak\GE24103
Curated drosophilid orthologs	Dafs\cos (Bartolome and Charlesworth, 2006) Dmir\cos (Bartolome and Charlesworth, 2006, Bartolome and Charlesworth, 2006) Dpse\GA14314   Dpse\cos (Lum, 2003)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL010619-PA Aedes aegypti (mosquito) AGAP010883-PA Anopheles gambiae (mosquito) XP_624554.1 Apis mellifera (honey bee) BGIBMGA010383-PA Bombyx mori (silk worm) CPIJ006179 Culex pipens (mosquito) ENSORLP00000011669 Oryzias latipes (japanese killifish) PHUM010096-PA Pediculus humanus (human body louse) XP_973391.1 Tribolium castaneum (red flour beetle) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • LH23857 FBgn0241241 FBgn0224003 FBgn0207699 FBgn0182039 FBgn0175684 JGI_V11_319054 FBgn0074342 FBgn0148550 FBgn0143560 FBgn0128205 FBgn0103063 FBgn0089273 CPIJ006179 BGIBMGA010383 ACYPI003112 AGAP010883 AAEL010619 PB22256 PHUM473100 GLEAN_07589
Stocks & Reagents
Stocks Listed in FlyBase ( 6 )
Bloomington	11156 y1 w67c23; P{lacW}cosk16101/CyO 19759 y1 w67c23; P{EPgy2}cosEY08735/CyO 3919 cos5 cn1 bw1 sp1/CyO, Df(2R)cn-S6
Kyoto	107266 cos5 cn1 bw1 sp1/CyO, Df(2R)cn-S6 111351 yd2 w1118 P{ey-FLP.N}2 P{GMR-lacZ.C(38.1)}TPN1; P{neoFRT}42D P{lacW}cosk16101 /CyO y+ 101353 cos5 cn1 bw1 sp1 / In(2LR)O, Df(2R)cn-S6, Cy1 dplv1 pr1 cn-
Genomic Clones ( 1 )
	BACR26B18 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 30 )
	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 Sequenced
BDGP DGC clones	LD35834 LD42253
Other clones	FI03725
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD01518 LD15871 LD41509 LD47057 RE60632 RE62187 RH45322 RH64529 RH74783
Other clones	32381 60828 70810 72785 99035 106319 173067 194287 CK01.123C.C9 EK093911 EK109308 EK120308 EK141912 EK186955 EK189901 EK201630 EK232065 EK250109
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0000352 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35653
Antibody Information
	polyclonal (Sisson et al., 1997)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: cos CG1708 (Gabrielian, 1999.11)
Source for database merge of
Additional comments
Other Comments
	cos blocks smo phosphorylation through binding with smo. (Liu et al., 2007) Identified as a component of the hh signalling pathway in a genome-wide RNAi screen. dsRNA made from templates generated with primers directed affects the extent of expression of a hh signaling reporter construct in Clone 8 cells. (Nybakken et al., 2005) dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology. (Kiger et al., 2003) dsRNA made from templates generated with primers against this gene tested in an RNAi assay in S2 cells. (Lum, 2003) Epistatic analysis places cos function downstream of ptc and smo. (Ruel et al., 2003) The last 301 amino acids of the cos COOH terminus are sufficient to provide full association with smo. (Ruel et al., 2003) 1 allele of cos been recovered in a screen for mutations with mutant phenotypes in clones in the wing. (Vegh and Basler, 2003) fu protein phosphorylates cos protein. The primary site of phosphorylation is Ser-572, with Ser-931 being phosphorylated to a lesser extent. (Nybakken et al., 2002) Mutants are isolated in a screen of the second chromosome isolating disc morphology defects. (Roch et al., 1998) hh elicits signal transduction via a complex that includes the products of the fu, ci and cos genes. The complex binds with high affinity to microtubules in the absence of hh protein, but not when hh is present. The complex may facilitate signalling from hh by governing access of the ci product to the nucleus. (Robbins et al., 1997) cos encodes a kinesin-related protein that accumulates preferentially in cells capable of responding to hh signal. The product is cytoplasmic and binds microtubules. The ci protein associates with the cos product in a large protein complex, suggesting that cos directly controls the activity of ci. (Sisson et al., 1997) The small lobe of fu may play a role in generating the neomorphic cos phenotype displayed by an unregulated fu protein in a Su(fu)- background. (Alves et al., 1995) cos function is required maternally for the patterning of the central portion of each segment, in the absence of cos activity the central portion is replaced by a mirror image duplication of the anterior partion. The cos protein may be involved in localisation or transport of other segment polarity gene products within the cell, facilitating cell polarisation. (Sisson et al., 1995) Viable mutations in the segmentation genes cos cause specific alterations in dpp expression within the anterior-posterior compartment boundary of the wing disc. (Capdevila et al., 1994) Mutants display hyperplastic phenotype, imaginal disc overgrowth. (Mechler, 1994) cos negatively regulates ptc and wg transcription. (Forbes et al., 1993) Genetic and developmental characteristics of the imaginal disc overgrowth mutant have been determined. (Torok et al., 1993) wg and en expression patterns are studied in all known segment polarity mutants to investigate the requirement of other segment polarity genes in mediating the maintenance of wg and en. (van den Heuvel et al., 1993) cos suppresses the segment polarity and zygotic phenotype of one class of fu alleles and causes a lethal interaction with the other class of fu alleles. (Preat et al., 1989) cos falls into the segment polarity class of genes, and has both a maternal and a zygotic action. (Grau and Simpson, 1987) Leaky alleles of cos display wing duplications that are part of a larger syndrome of polarity defects that affect the embryonic and imaginal segment. The dominant phenotype seen in flies heterozygous for the neomorphic "Cos" mutants is the result of a reduced level of function at the cos locus. Haploidy for cos enhances, whereas triploidy for cos suppresses the phenotype of "Cos" heterozygotes. (Grau and Simpson, 1987) Homozygotes, hemizygotes and heteroallelic heterozygotes for class I alleles die as embryos with a normal cuticular phenotype. When derived from homozygous ovarian clones, however, abnormal deletion-duplication patterns in each segment are observed; posterior rows of abdominal denticles are replaced by mirror-image duplications of the anterior rows, including the segmental boundary, which is thereby duplicated; thoracic denticle belts missing; cos1 especially severely affected in this regard, including loss of entire segments. Heterozygotes derived from homozygous ovarian clones show slightly abnormal segmental patterning. Flies heterozygous for class I alleles or cos deficiencies and at the same time heterozygous for Cos alleles display pattern duplications of wings and halteres. Class II alleles are hemizygous lethal but homozygous viable. They display wing and haltere duplications indistinguishable from those found for cos1/Pka-C1Cos-1 heterozygotes as described by Whittle. Flies heterozygous for class III alleles and Cos show moderately reduced viability and occasional wing duplications; flies homozygous for class III alleles show reduced viability and have wild-type phenotype, but enhance the pattern duplications associated with Cos/+; hemizygotes for class III alleles or heterozygotes with class I alleles are lethal or nearly so in the presence of Cos, the survivors invariably exhibiting pattern duplications of wings and halteres.
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057744 RefSeq (Proteins) • NP_477092 DDBJ / EMBL / GenBank • AA441088 AAB66813 AAF59270 AAN71259 AAT94488 AC005452 AF019250 AI456015 AQ034038 AW943304 BT001504 BT015259 CL610224 Entrez Gene - A searchable database of RefSeq genes. • 35653 UniProtKB/TrEMBL • O16844
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Kinesin, motor domain (IPR001752)
Linkouts
	BioGRID - A database of protein and genetic interactions • 61534 DroID - A comprehensive database of gene and protein interactions. • FBgn0000352 DRSC - Results from RNAi screens. • FBgn0000352 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0000352 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG1708-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0000352 FlyReactome - A curated knowledgebase of Drosophila melanogaster pathways • 205651 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35653 InParanoid A subset of ortholog calls from InParanoid. • AAEL010619-PA Aedes aegypti (mosquito) AGAP010883-PA Anopheles gambiae (mosquito) XP_624554.1 Apis mellifera (honey bee) BGIBMGA010383-PA Bombyx mori (silk worm) CPIJ006179 Culex pipens (mosquito) ENSORLP00000011669 Oryzias latipes (japanese killifish) PHUM010096-PA Pediculus humanus (human body louse) XP_973391.1 Tribolium castaneum (red flour beetle) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /segment/cosl2-1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35653 modMine - Data generated by the modENCODE project. • FBgn0000352 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • LH23857 FBgn0241241 FBgn0224003 FBgn0207699 FBgn0182039 FBgn0175684 JGI_V11_319054 FBgn0074342 FBgn0148550 FBgn0143560 FBgn0128205 FBgn0103063 FBgn0089273 CPIJ006179 BGIBMGA010383 ACYPI003112 AGAP010883 AAEL010619 PB22256 PHUM473100 GLEAN_07589
Synonyms & Secondary IDs ( 22 )
Reported As
Symbol Synonym	CG1708 (Goldstein and Gunawardena, 2000, Nybakken et al., 2005, Molnar et al., 2006, Keleman et al., 2009.8.5) cos (Dworkin and Gibson., 2006, Dworkin and Gibson, 2006, Bartolome and Charlesworth, 2006, Nybakken et al., 2005, Molnar et al., 2006, Raisin et al., 2010) cos2 (Friggi-Grelin et al., 2008, Collier et al., 2004, Nakano et al., 2004, Ogden et al., 2004, Horabin et al., 2003, Lum et al., 2003, Vegh and Basler, 2003, Bai and Montell, 2002, Besse et al., 2002, Jia et al., 2002, Collier et al., 2002, Vied and Horabin, 2001, Lefers et al., 2001, Wang and Holmgren, 2000, Zhang and Kalderon, 2000, Vied and Horabin, 2000, Wang and Holmgren, 1999, Wang and Holmgren, 1999, Kalderon, 1997, Biesecker, 1997, Srivastava, 1997, Sanchez et al., 1997, Sisson et al., 1997, Ruiz i Altaba, 1997, Robbins et al., 1997, Therond et al., 1996, Sisson and Scott, 1996, Alves et al., 1995, Pham et al., 1995, Capdevila and Guerrero, 1994, Capdevila et al., 1994, Martinez Arias, 1993, Yao et al., 2006, Molnar et al., 2011, Malpel et al., 2007, Smelkinson et al., 2007, Ruel et al., 2007, Claret et al., 2007, Farzan et al., 2008, Ayers et al., 2009, Mosimann et al., 2009, Sisson et al., 2006, Zhang et al., 2005) Cos2 (Huangfu and Anderson, 2006, Ho et al., 2005, Ruel et al., 2003, Lum, 2003, Zhang, 2004, Lum et al., 2003, Collins and Cohen, 2003, Beachy et al., 2004, Lum and Beachy, 2004, Kalderon, 2002, Zhang et al., 2005, Walthall et al., 2005, Nybakken et al., 2005, Ruel et al., 2003, Bijlsma et al., 2004, Ogden et al., 2003, Stegman et al., 2004, Kalderon, 2004, Ogden et al., 2004, Lum et al., 2004, Jia et al., 2004, Jia et al., 2003, Ou et al., 2003, Nybakken and Perrimon, 2002, Horabin, 2003, Lefers and Holmgren, 2003, Jiang, 2002, Lefers et al., 2002, Nybakken et al., 2002, Taipale and Beachy, 2001, Wang et al., 2001, Ascano et al., 2001, Lefers et al., 2001, Vied et al., 2001, Matise and Joyner, 1999, Ramirez-Weber et al., 2001, Stegman et al., 2000, Stegman et al., 2000, Marsh and Theisen, 1999, Chen et al., 1999, Maniatis, 1999, Lee, 1998, Cadigan and Nusse, 1997, Perrimon, 1996, Ruel et al., 2005, Ogden et al., 2006, Casso et al., 2006, Osterlund and Kogerman, 2006, Ishii, 2005, Briscoe and Therond, 2005, Singla, 2006, Ogden et al., 2006, Fouix et al., 2003, Ascano et al., 2002, Liu et al., 2008, Zhao and Jiang, 2008, De Rivoyre et al., 2006, Varjosalo et al., 2006, Torroja et al., 2005, Zhao et al., 2007, Liu et al., 2007, Wang and Price, 2008, Wang and Jiang, 2004, Walthall et al., 2007, Ogden et al., 2008, Jia et al., 2009, Monnier et al., 2002, Wang et al., 2000, Raisin et al., 2010, Farzan et al., 2009, Zhou and Kalderon, 2010) Cos-2 (Price, 2006, van den Heuvel, 2003, Greaves, 2001, Zhang et al., 2003, Zhang et al., 2002, Ingham and McMahon, 2001, Murone et al., 1999, Aza-Blanc and Kornberg, 1999, Wang and Holmgren, 1999, Monnier et al., 1999, Johnson and Scott, 1997, Robbins et al., 1997, Tabin and McMahon, 1997, Ingham, 1997, Nusse, 1996) cos-2 (Wicking et al., 1999, Luschnig et al., 2004, Dobens and Raftery, 2000, Liu and Montell, 1999, Theil et al., 1999, Goodrich and Scott, 1998, Ingham, 1998, Roch et al., 1998, Aza-Blanc et al., 1997, Dean, 1996, Hogan, 1996, Sanchez-Herrero et al., 1996, Johnson et al., 1995, Gateff, 1994, Sisson et al., 1995, Klingensmith and Nusse, 1994, DiNardo et al., 1994, Forbes et al., 1993, Preat et al., 1989, Preat et al., 1993, Peifer and Bejsovec, 1992, Woods and Bryant, 1992, Garcia-Bellido and de Celis, 1992, Wilkins and Gubb, 1991, Grau and Simpson, 1987) COS2 (Sanial and Plessis, 2007, Malpel et al., 2007) Cos2 (Perrimon, 1995, Perrimon, 1994) COS2 (Hammerschmidt et al., 1997) l(2)161/28 (Watson et al., 1994, Mechler, 1994, Gateff, 1994, Torok et al., 1993) l(2)cos2 (Whittle, 1976) l(2)k16101   l(2)k16128
Name Synonym	costa   costal2 (Hyman and Scott, 2005, Dubnau et al., 2003, Ho et al., 2000, Therond, 1998, Ho and Scott, 1998, Vied and Horabin, 2000) costal-2 (Janody et al., 2004, Sisson, 1997.8.13, Vied and Horabin, 1999, Cohen, 1993, van den Heuvel et al., 1993, Ingham, 1991, Ingham, 1991, Dworkin and Gibson, 2006) Costal2 (Plessis et al., 2004, Zhu et al., 2003, Zhu et al., 2002, McNeill, 2000, Zhu et al., 2001, Goldstein and Gunawardena, 2000, Ho and Scott, 1999, Farese and Herz, 1998, Ho et al., 1997, Ho et al., 1997, Goshima and Vale, 2003, Bijlsma et al., 2004, Ogden et al., 2006, Ascano et al., 2002, Liu et al., 2008, Zhao and Jiang, 2008, Torroja et al., 2005, Liu et al., 2007, Ruel et al., 2007, Claret et al., 2007, Farzan et al., 2008, Ogden et al., 2008, Monnier et al., 2002, Raisin et al., 2010) Costal-2 (Ruiz i Altaba, 1999, Vied and Horabin, 2002, Dussillol et al., 2000, Dussillol et al., 1999, Leahy, 1999, The et al., 1998, Orsulic and Peifer, 1996, Lum et al., 2004, Zhang et al., 2002, Marsh and Theisen, 1999, Torroja et al., 2005, Nusse, 1996, Walthall et al., 2007, Raisin et al., 2010, Farzan et al., 2009) Costal 2 (Gundersen and Cook, 1999, Sanial and Plessis, 2007, Zhao et al., 2007, Jia et al., 2009) costal 2 (Liu and Montell, 1999, Johnston and Scott, 1998) COSTAL2 (Goldstein and Gunawardena, 2000) Enhancer of Epaulette
Secondary FlyBase IDs
	FBgn0014275 FBgn0021829
References ( 243 )
Generate a list of	All references relating to this gene ( 243 )
List References by type	Research paper  ( 87 ) Supplementary material  ( 11 ) Review  ( 72 ) Erratum  ( 1 ) Personal communication to FlyBase  ( 5 ) Abstract  ( 57 ) FlyBase analysis  ( 2 ) DNA/RNA sequence record  ( 1 ) Meeting report  ( 1 ) Curated genome annotation  ( 2 ) Computer file  ( 3 ) Automatic genome annotation  ( 1 )
Recent research papers ( 3 )
	Molnar et al., 2011, PLoS Genet. 7(3): e1001335 Role of the Drosophila non-visual ß-arrestin kurtz in hedgehog signalling. [FBrf0213301] Raisin et al., 2010, Dev. Biol. 344(1): 119--128 Dynamic phosphorylation of the kinesin Costal-2 in vivo reveals requirement of fused kinase activity for all levels of hedgehog signalling. [FBrf0211312] Zhou and Kalderon, 2010, Dev. Biol. 348(1): 47--57 Costal 2 interactions with Cubitus interruptus (Ci) underlying Hedgehog-regulated Ci processing. [FBrf0212239] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...