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Gene Dmel\hh

General Information
Symbol	Dmel\hh	Species	D. melanogaster
Name	hedgehog	Annotation symbol	CG4637
Feature type	protein_coding_gene	FlyBase ID	FBgn0004644
Gene Model Status	Current	Stock availability	31 publicly available
Also Known As	bar-3, l(3)neo56
Genomic Location
Chromosome (arm)	3R	Recombination map	3-81.2
Cytogenetic map	94E1-94E1	Sequence location	3R:18,953,425..18,967,881 [-]
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 hedgehog is referred to in FlyBase by the symbol Dmel\hh (CG4637, FBgn0004644). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: cell surface binding; protein binding. There is experimental evidence for 25 unique biological process terms, many of which group under: anatomical structure development; biological regulation; localization; organ morphogenesis; sensory organ development; cellular component organization or biogenesis; cellular process involved in reproduction; multicellular organismal reproductive process; organ development; locomotion. 178 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; adult segment; primordium; late extended germ band embryo; adult mesothoracic segment; ventral nerve cord primordium; organ system; abdominal ventral denticle belt; imaginal precursor; germarium. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Hedgehog, N-terminal signaling domain; Hedgehog/DD-peptidase; Hedgehog/intein hint domain, C-terminal; Hedgehog/intein hint, N-terminal; Intein splice site; Peptidase C46, hedgehog protein; Peptidase C46, hedgehog protein, hint region. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of very low expression. Peak expression observed within 00-12 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult crop, larval midgut, adult hindgut. Comments on Affy2 ProbeSet: ProbeSet 1626527_at completely aligns to an exonic region common to each of the 2 FlyBase-annotated transcript isoforms of hh. Gene sequence location is 3R:18953425..18967881.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	hh: hedgehog A segment polarity type of embryonic lethal. Homozygous embryos have the posterior naked portion of the ventral surface of each segment deleted and replaced by a mirror image of the anterior denticle belts. Embryos appear to lack segmental boundaries. In strong alleles, there is no obvious segmentation; the larvae are approximately 40% the length of the wild-type larvae, and there is a lawn of denticles arranged in a number of whorls on the ventral surface as a result of loss of naked cuticle. In intermediate alleles, naked cuticle is also lost from the ventral region, but the lawn of denticles is arranged in segmental arrays in mirror-image symmetry. The weak alleles show fusions that delete the naked cuticle usually between abdominal segments 1 and 2 and 6, 7, and 8 (Mohler, 1988). Temperature shift experiments with a temperature-sensitive allele (viable and normal at 18, and mutant at 25) indicate two phases of hh activity at 25, the first during early embryogenesis (3-6 hr of development) and the second during the late larval and early pupal stages (4-7 days of development). hh1 A weak hypomorphic allele that is not complemented by other hh alleles. Eye of homozygote small and narrow with about 150 facets. Eye disc size reduced; deep cleft at anterior edge cell; clusters at cleft look mature (Renfranz and Benzer, 1989, Dev. Biol. 136: 411-29). Mir: Mirabile (M. Muskavitch) Mirror-image duplication of tergite structure. Microchaetae are eliminated from the anterior portion of the tergite and replaced by a duplication consisting of an anteriorly oriented row of macrochaetae and the darkly pigmented cuticle normally found in the posterior portion of the tergite. Fat body and oenocytes underneath the tergite are also duplicated with mirror-image symmetry (Madhavan and Madhavan). Mrt: Moonrat (J.A. Kennison) Heterozygote shows partial transformation of anterior wing to posterior (triple row bristles replaced by double row bristles in patches). A network of extra veins appears in the anterior compartment, beginning at the distal edge in the least affected flies, and covering the entire anterior compartment in the more extreme cases. Wing blade expanded anteriorly at the distal edge. Wing blade expansion and extra veins resemble phenotypes seen in en1 homozygotes in the presence of Minute mutations. Bubbles often form in the wing blade. More rarely, a mirror-image outgrowth from the anterior edge is present. Mirror-image duplications sometimes appear in halteres. Legs sometimes appear deformed (similar to phenotype of enlethal clones induced in the larva). Dominant phenotypes strongly temperature-sensitive. Penetrance greater than 99% at 18 (with strong expressivity) but only 30-40% at 29 (with very weak expressivity). Shows paternal effect. Penetrance greater when mutant allele inherited from father than when inherited from mother. Mrt/+/+ indistinguishable from Mrt/+.
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	References González et al., 2008, Bioinformatics 24(16): i234--i240 Barrett et al., 2008, Dev. Biol. 317(1): 234--245 Babcock et al., 2011, Curr. Biol. 21(18): 1525--1533 Wojcinski et al., 2011, Dev. Biol. 358(1): 168--180 Alleles hh[Scer\UAS.T:Arus\HRP] Transgenic Constructs P{UAS-hh.HRP}
FB2012_01	References Wang et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(27): 11139--11144 Casso et al., 2011, Genetics 187(2): 485--499 Ayers et al., 2010, Dev. Cell 18(4): 605--620 Nahmad, 2011, J. R. Soc. Interface 8(63): 1429--1439 Sequence features BKN45504 BKN28916 HFA16965
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 94E1-94E1   Limits computationally determined from genome sequence between P{lacW}GclmL0580 and P{EP}hhEP3521
Experimentally Determined Cytological Location
Cytogenetic map Notes References 94E1-94E3   (determined by in situ hybridisation)   (Ranz et al., 2001) 94E2-94E2   (determined by in situ hybridisation)   (Abdelilah-Seyfried et al., 2000) 94E1-94E4   (determined by in situ hybridisation)   (Spradling et al., 1999) 94E-94E     (Gellon et al., 1997) 94D-94E   (determined by in situ hybridisation)   (Beachy, 1992) 94D1-94E13   94E1--4   (BDGP Project Members, 1994-1999) 94D10-94E5   (determined by in situ hybridisation)   (Mohler and Vani, 1992) 94D10-94D13   (determined by in situ hybridisation)   (Mohler et al., 1991) 94D-94E   (determined by in situ hybridisation)   (Cooley et al., 1988) 94E-94E   On the basis of meiotic mapping.   (Karim et al., 1996)
Experimentally Determined Recombination Data
Location	3-81 (Tearle and Nusslein-Volhard, 1987) 3-81.2   3-79.1 (Gonzalez et al., 1989, Atkinson et al., 1991, Mohler et al., 1992)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\hh 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) hh-RA FBtr0084404  NM_079735   3089   471 hh-RB FBtr0100506  NM_001038976   2370   471
Additional Transcript Data & Comments
Reported size (kB)	2.4 (northern blot) (Lee et al., 1992, Tabata et al., 1992) 2.3 (northern blot) (Tashiro et al., 1993)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein hh-PA   FBpp0083796   52.1   471   8.23     NP_524459   AAF56102 hh-PB   FBpp0099945   52.1   471   8.23     NP_001034065   ABC66186
Additional Polypeptide Data & Comments
Reported size (kDa)	471 (aa); 52 (kD predicted) (Tabata et al., 1992) 471 (aa); 52 (kD) (Lee et al., 1992) 471 (aa) (Tashiro et al., 1993)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Hedgehog, N-terminal signaling domain (IPR000320) Peptidase C46, hedgehog protein (IPR001657) Peptidase C46, hedgehog protein, hint region (IPR001767) Hedgehog/intein hint domain, C-terminal (IPR003586) Hedgehog/intein hint, N-terminal (IPR003587) Intein splice site (IPR006141) Hedgehog/DD-peptidase (IPR009045) MEROPS - An information resource for peptidases and the proteins that inhibit them • C46.001 PDB - Protein Data Bank. An information portal to biological macromolecular structures • 1AT0
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA201458   AA263416   AE014297 AAF56102   AE014297 ABC66186   AI519048   AQ026358   AQ034105   AQ034132   AQ254728   AX146557   AX282805 CAD10621   AX354872 CAD22996   AX417094 CAD35282   BG635849   BI142134   BI164506   BI214827   BI565435   BI586434   BI589144   BI622694   BI625170   BI632636   BI641128   CO266552   CO273477   CO303982   CO311934   CS126494 CAJ18813   EC060492   EC061587   EC087454   EC202430   EC236939   L02793 AAA16458   L05404 AAA28604   L05405   S66384 AAB28646   Z11840   Z83558
UniProtKB/Swiss-Prot	Q02936
UniProtKB/TrEMBL
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
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference embryonic stage   labral segment   (Lee et al., 1992) proctodeum   (Lee et al., 1992) labial segment   (Lee et al., 1992) organism | segment polarity expression pattern   (Lee et al., 1992) embryonic/larval foregut   (Tashiro et al., 1993) embryonic stage | mid   embryonic/larval posterior spiracle   (Lee et al., 1992) anal pad   (Lee et al., 1992) in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage   embryonic/larval esophagus   (Mohler and Vani, 1992, Tabata et al., 1992) embryonic/larval foregut   (Mohler and Vani, 1992, Tabata et al., 1992) procephalic segment   (Mohler and Vani, 1992) antennal lobe   (Mohler and Vani, 1992) mandibular segment   (Mohler and Vani, 1992, Lee et al., 1992) embryonic/larval salivary gland   (Tabata et al., 1992) maxillary segment   (Mohler and Vani, 1992) intercalary segment   (Mohler and Vani, 1992, Lee et al., 1992) embryonic/larval pharynx   (Mohler and Vani, 1992, Tabata et al., 1992) embryonic/larval hindgut   (Mohler and Vani, 1992, Tabata et al., 1992, Tashiro et al., 1993) labial segment   (Mohler and Vani, 1992) antennal segment   (Lee et al., 1992) organism | segment polarity expression pattern   (Lee et al., 1992, Tashiro et al., 1993) embryonic stage | mid   embryonic/larval anterior spiracle   (Lee et al., 1992) embryonic stage 5   organism | 10% egg length   (Mohler and Vani, 1992) Comment:5% egg length organism | 10-70% egg length   (Mohler and Vani, 1992) organism | 90% egg length   (Mohler and Vani, 1992) Comment:97% egg length maxillary segment   (Lee et al., 1992) embryonic stage 5 -- 17   organism | segment polarity expression pattern   (Mohler and Vani, 1992) embryonic stage 6 -- 17   organism | segment polarity expression pattern   (Tabata et al., 1992) hindgut anlage   (Lee et al., 1992) embryonic stage 10 -- 11   esophagus | posterior   (De Velasco et al., 2004) epipharynx   (De Velasco et al., 2004) embryonic stage 13   small intestine primordium   (Iwaki et al., 2001) rectum primordium   (Iwaki et al., 2001) larval stage   wing disc | posterior   (Tabata et al., 1992) Kenyon cell   (Noveen et al., 2000)
Additional Descriptive Data
	hh transcripts are expressed in embryos, larvae, pupae and adults with peaks of expression in 6-12hr embryos and early pupae. Transcripts are first detected in embryonic stage 5 in a few stripes at the anterior and posterior ends of the embryo. The number of stripes gradually increases to 17. The terminal stripes are 2-3 cells wide and the internal stripes are 1 cell wide. Expression is stronger in every second stripe and is stronger laterally than in dorsal and ventral regions. The expression in stripes reaches a maximum at stages 8-11. By the end of germband retraction, the stripes are situated in the posterior compartments of the lateral ectoderm. (Tashiro et al., 1993) hh transcripts are first detected at the cellular blastoderm stage in 17 segmental stripes. 14 of the stripes are one cell wide and extend from 10-70% egg length. There are two 3-cell-wide stripes at 5% and 75% egg length and a dorsal anterior spot at 97% egg length. The stripes appear asynchronously. Even parasegmentally-numbered stripes precede odd-numbered stripes and anterior stripes precede more posterior stripes. The stripes are activated around the entire circumference of the embryo but disappear from the amnioserosa and mesoderm after gastrulation. At stage 11, the stripes are located just posterior to the parasegmental furrow and are spaced one cell anterior to the tracheal pit in each segment. Stripes persist after germ band retraction and are located in the posteriormost portion of the lateral ectoderm of each segment. hh transcripts are also expressed in the fore- and hindguts following gastrulation and germ band extension as well as in the cephalic region of the embryo. Up to stage 10, the intensity of staining is heavier in the nucleus than in the cytoplasm. After stage 10, RNA staining is predominantly cytoplasmic. (Mohler and Vani, 1992) At the cellular blastoderm stage, hh transcripts are located predominantly in a single stripe at 75% egg length with additional transcripts at the anterior tip and along the ventral side. At gastrulation, hh is expressed in 14 single-cell-wide stripes between 30% and 65% egg length. The stripes are coincident with en expression and occur in the cells of the posterior compartments. They appear in a characteristic order, even-numbered ones before odd-numbered ones and anterior ones before posterior ones. hh is also expressed in a block of cells at the anterior end and in wide stripes at 10% and 75% egg length. hh transcripts are expressed later in the foregut, pharynx, esophagus, hindgut, and salivary glands. hh transcripts are exressed in imaginal discs where they are localized to the posterior compartments. The differences between hh and en expression are noted. (Tabata et al., 1992) Peaks of hh expression are observedin 2-10hr embryos and in pupae. In embryos, hh transcripts are firstexpressed in a discrete pattern in the maxillary segment followed by apattern of 14 parasegmental stripes. At germ band extension, a 15th stripeis seen. hh expression in the metameric portion of the embryo closelyresembles en expression. Expression is also described in a variety ofsites in the nonmetameric portion of the embryo including the intercalaryand antennal segments, the procephalon, the gnathal segments, and portionsof the hindgut. Expression in imaginal discs is described for theassociated Ecol\lacZ insertion. hh expression is pair-rule dependent. Inftz mutants, expression in the even-numbered parasegments is missing.wg mutations caused diminished expression and ptc mutants causeexpression in an ectopic stripe in each segment. nkd mutations causebroadening of the stripes. (Lee et al., 1992)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference wandering third instar larval stage   eye-antennal disc | restricted   (Shyamala and Bhat, 2002)
Additional Descriptive Data
	Expression of hh in the eye-antennal disc is more restricted than the expression of ptc. hh protein is present in cells adjacent to ptc-expressing cells in only certain regions. (Shyamala and Bhat, 2002)
Marker for
	embryonic rectum
Subcellular Localization
CV Term
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 FBgn0004644    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0004644 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 very low expression. Peak expression observed within 00-12 hour embryonic 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 FBgn0004644 expression level Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    525 embryo 04-06hr    1929 embryo 06-08hr    1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr  (525) embryo 04-06hr  (1929) embryo 06-08hr  (1487) embryo 08-10hr  (1185) embryo 10-12hr  (1253) embryo 12-14hr  (794) embryo 14-16hr  (385) embryo 16-18hr  (183) embryo 18-20hr  (257) embryo 20-22hr  (182) embryo 22-24hr  (173) larva L1  (145) larva L2  (119) larva L3 12hr old    80 larva L3 puffstage 1-2  (296) larva L3 puffstage 3-6  (456) larva L3 puffstage 7-9  (697) white prepupae new  (801) white prepupae 12hr  (818) white prepupae 24hr  (1194) pupae 2d postWPP  (742) pupae 3d postWPP  (158) pupae 4d postWPP  (143) adult male 01day  (155) adult male 05day  (198) adult male 30day  (218) adult female 01day  (110) adult female 05day    37 adult female 30day    54 Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    525 embryo 04-06hr  (1929) embryo 06-08hr  1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    525 embryo 04-06hr    1929 embryo 06-08hr    1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 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    40 embryo 02-04hr    525 embryo 04-06hr    1929 embryo 06-08hr    1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0004644 expression level Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    525 embryo 04-06hr    1929 embryo 06-08hr    1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 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    40 embryo 02-04hr  (525) embryo 04-06hr  (1929) embryo 06-08hr  (1487) embryo 08-10hr  (1185) embryo 10-12hr  (1253) embryo 12-14hr  (794) embryo 14-16hr  (385) embryo 16-18hr  (183) embryo 18-20hr  (257) embryo 20-22hr  (182) embryo 22-24hr  (173) larva L1  (145) larva L2  119 larva L3 12hr old    80 larva L3 puffstage 1-2  (296) larva L3 puffstage 3-6  (456) larva L3 puffstage 7-9  (697) white prepupae new  (801) white prepupae 12hr  (818) white prepupae 24hr  (1194) pupae 2d postWPP  (742) pupae 3d postWPP  (158) pupae 4d postWPP  143 adult male 01day  (155) adult male 05day  (198) adult male 30day  (218) adult female 01day  110 adult female 05day    37 adult female 30day    54 Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    525 embryo 04-06hr  1929 embryo 06-08hr  1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    40 embryo 02-04hr    525 embryo 04-06hr    1929 embryo 06-08hr    1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 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    40 embryo 02-04hr    525 embryo 04-06hr    1929 embryo 06-08hr    1487 embryo 08-10hr    1185 embryo 10-12hr    1253 embryo 12-14hr    794 embryo 14-16hr    385 embryo 16-18hr    183 embryo 18-20hr    257 embryo 20-22hr    182 embryo 22-24hr    173 larva L1    145 larva L2    119 larva L3 12hr old    80 larva L3 puffstage 1-2    296 larva L3 puffstage 3-6    456 larva L3 puffstage 7-9    697 white prepupae new    801 white prepupae 12hr    818 white prepupae 24hr    1194 pupae 2d postWPP    742 pupae 3d postWPP    158 pupae 4d postWPP    143 adult male 01day    155 adult male 05day    198 adult male 30day    218 adult female 01day    110 adult female 05day    37 adult female 30day    54 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 FBgn0004644    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0004644 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 crop, larval midgut, adult hindgut. [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 FBgn0004644 expression level Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0004644 expression level Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    3.45 Larval Midgut    108.8 Larval Hindgut    58.5 Larval Malpighian Tubules    0.7 Larval Fat Body    9.3 Larval Salivary Gland    2.5 Larval Trachea    3.2 Larval Carcass    13.75 Adult Head    9.1 Adult Eye    1.925 Adult Brain    0.7 Adult Thoracic-Abdominal Ganglion    2.1 Adult Crop    181.5 Adult Midgut    88.9 Adult Hindgut    169.8 Adult Malpighian Tubules    2.4 Adult Fat Body    5.5 Adult Salivary Gland    6.3 Adult Heart    3.325 Adult VirginFemale Spermatheca    61.4 Adult InseminatedFemale Spermatheca    77.7 Adult Ovary    1.3 Adult Testis    45.1 Adult Male Accessory Gland    1.8 Adult Carcass    6.1 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     Adult Fat Body     Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    9.1   NA  Eye    1.925   NA  Brain    0.7   3.45  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    2.1   NA  Crop    181.5   108.8  Midgut    88.9   58.5  Hindgut    169.8   0.7  Malpighian Tubules    2.4   9.3  Fat Body    5.5   2.5  Salivary Gland    6.3   NA  Heart    3.325   3.2  Trachea    NA   NA  VirginFemale Spermatheca    61.4   NA  InseminatedFemale Spermatheca    77.7   NA  Ovary    1.3   NA  Testis    45.1   NA  Male Accessory Gland    1.8   13.75  Carcass    6.1 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_11
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004644 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG4637-RA
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal hh8 hhAC hhMrtr1 hhMrtr2 lethal, with Scer\GAL4sca-4512 hhEP3521 lethal | dominant, with Scer\GAL4bi-omb-Gal4 hhEP3521 lethal | dominant, with Scer\GAL4dpp.blk1 hhEP3521 lethal | embryonic stage hhhs.PI lethal | embryonic stage, with Scer\GAL4en-e16E hhN.Scer\UAS lethal | embryonic stage | heat sensitive hh4 lethal | embryonic stage | recessive hh3 hh8 hh11 hh21 hhA hhAC hhAE hhAM hhMrtr1 hhMrtr2 lethal | embryonic stage | recessive | segment polarity expression pattern hh2 hh5 hh8 hh9 hh10 hh11 hh15 hh21 hh22 hh737 hh1582 hhunspecified lethal | embryonic stage | recessive | segment polarity expression pattern | heat sensitive hh3 hh4 lethal | heat sensitive hh4 hh5 hhts2 lethal | pupal stage, with Scer\GAL430A hhScer\UAS.cIa lethal | pupal stage | recessive hhMrt lethal | recessive hh21 hhA hhAC hhAE hhAM hhneo56 hhneo57 hhQ50 hhrJ413 hhSC1 hhSC2 hhSC3 hhSC4 hhSC5 hhSC6 hhSC7 hhSC8 hhSC9 hhSC10 hhSC11 hhSC12 hhSC13 hhSC14 hhSC15 hhSC16 hhSC17 hhSC18 hhSC19 hhSC20 hhSC21 hhSCG1 hhSCG2 hhSCG3 hhXE-2134 hhXE-3161 lethal | recessive | heat sensitive hhts2 semi-lethal | dominant, with Scer\GAL4hs.2sev hhEP3521 viable hhbar3 hhfse hhh9D hhMrt hhts2 viable, with Scer\GAL4pnr-MD237 hhGD193 hhGD6242 Sterility Allele fertile hhh9D Other Phenotypes Allele increased cell death, with Scer\GAL4sim.PS hhScer\UAS.cIa increased cell death | embryonic stage hh3 mitotic cell cycle defective | somatic clone hhts2 neuroanatomy defective, with Scer\GAL4sim.P3.7 hhScer\UAS.cAa planar polarity defective | recessive hhbar3 size defective, with Scer\GAL4MD-638 hhNIG.4637R visible hhMrt hhts2 hhαTub84B.PB visible, with Scer\GAL4bi-omb-Gal4 hhC84S.Scer\UAS hhScer\UAS.cLa visible, with Scer\GAL4dpp.3KK hhScer\UAS.cIa visible, with Scer\GAL4en-e16E hhC84S.Scer\UAS hhScer\UAS.cCa hhScer\UAS.cIa hhScer\UAS.cLa visible, with Scer\GAL4en-e16E, hhC84S.Scer\UAS hhts2 visible, with Scer\GAL4en-e16E, hhts2 hhC84S.Scer\UAS visible, with Scer\GAL4en-GAL4-33 hhScer\UAS.cUa visible, with Scer\GAL4ey.PH hhC84S.Scer\UAS hhEP3521 hhScer\UAS.cCa visible, with Scer\GAL4MD-638 hhNIG.4637R visible, with Scer\GAL4ptc-559.1 hhC84S.Scer\UAS visible, with Scer\GAL4sca-537.4 hhEP3521 visible, with Scer\GAL4T155 hhScer\UAS.cPb visible (with hh8) hhbar3 visible (with hhbar3) hh8 visible | dominant hhMir hhMrt visible | dominant, with Scer\GAL4hs.2sev hhEP3521 visible | dominant | heat sensitive hhhs.PI visible | dominant | paternal effect | cold sensitive hhMrt visible | heat sensitive hhts2 visible | recessive hh4 hhbar3 hhlacZ visible | recessive | heat sensitive hhts2 visible | recessive | somatic clone hhAC hhSH2 hhSH3 hhSH7 hhSH9 hhSH17 visible | semidominant hhh9D visible | somatic clone, with Scer\GAL4Ubx.PdC hhN.Scer\UAS.T:Avic\GFP wild-type hhP30 Phenotype manifest in Allele 1st posterior cell hhbar3 1st posterior cell, with Scer\GAL4bi-omb-Gal4 hhC84S.Scer\UAS hhScer\UAS.cLa 1st posterior cell, with Scer\GAL4en-e16E hhC84S.Scer\UAS hhScer\UAS.cCa hhScer\UAS.cIa hhScer\UAS.cLa 1st posterior cell, with Scer\GAL4en-e16E, hhC84S.Scer\UAS hhts2 1st posterior cell, with Scer\GAL4en-e16E, hhts2 hhC84S.Scer\UAS 1st posterior cell, with Scer\GAL4MD-638 hhNIG.4637R 2nd posterior cell hh2 hhbar3 abdominal 1 ventral denticle belt hh11 hh3 hh8 abdominal 2 ventral denticle belt hh11 hh3 hh8 abdominal 3 ventral denticle belt hh11 hh3 hh8 abdominal 4 ventral denticle belt hh11 hh3 hh8 abdominal 5 ventral denticle belt hh11 hh3 hh8 hhhs.PI abdominal 6 ventral denticle belt hh11 hh3 hh8 hhhs.PI abdominal 7 ventral denticle belt hh11 hh3 hh8 hhhs.PI abdominal 8 ventral denticle belt hh11 hh3 hh8 abdominal lateral oblique muscle 1 founder cell hh21 abdominal segment hhMir abdominal sternite bristle | heat sensitive hhts2 abdominal tergite & macrochaeta | conditional ts hhts2 abdominal tergite & microchaeta hhMirRevBx1 abdominal tergite & microchaeta | conditional ts hhhs.PI abdominal tergite | anterior & macrochaeta hhαTub84B.PB abdominal tergite | anterior & microchaeta hhαTub84B.PB abdominal tergite | anterior & trichome hhαTub84B.PB abdominal tracheal pit 1 hh8 abdominal tracheal pit 2 hh8 abdominal tracheal pit 6 hh8 abdominal tracheal pit 7 hh8 abdominal ventral acute muscle 1 founder cell hh21 abdominal ventral acute muscle 2 founder cell hh21 abdominal ventral acute muscle 3 founder cell hh21 abdominal ventral transverse muscle 1 founder cell hh21 adult abdomen & cuticle | anterior compartment hhαTub84B.PB adult abdomen & macrochaeta | anterior compartment hhαTub84B.PB adult abdomen & microchaeta | anterior compartment hhαTub84B.PB adult cuticle & head capsule | dorsal | conditional ts hhts2 adult eye primordium hh21 anal plate hhαTub84B.PB hh4 hhlacZ antenna | distal hhMrt antenna | heat sensitive hhts2 antennal sense organ hh8 anterior commissure, with Scer\GAL4sca-537.4 hhScer\UAS.cIa anterior crossvein, with Scer\GAL4MD-638 hhNIG.4637R anterior crossvein, with Scer\GAL4ptc-559.1 hhC84S.Scer\UAS anterior scutellar bristle hh4 basal apodeme of penis hh4 hhlacZ basal stalk hhαTub84B.PB hhhs.PI Bolwig organ hh21 hh8 Bolwig organ, with Scer\GAL4da.G32 hhScer\UAS.cKa Bolwig organ, with Scer\GAL4h-1J3 hhScer\UAS.cKa Bolwig organ | ectopic | heat sensitive hhhs.PI Bolwig organ | heat sensitive hhhs.PI hhts2 cardial valve | precursor hh21 cardioblast hh8 cephalopharyngeal skeleton, with hha.cBa hhcBa cephalopharyngeal skeleton, with hhcBa hha.cBa chemosensory ventral triple row bristle, with Scer\GAL430A hhScer\UAS.cIa commissure hhunspecified costal cell | somatic clone, with Scer\GAL4Ubx.PdC hhN.Scer\UAS.T:Avic\GFP cuticle hh21 hh3 hh8 hhAC hhMrtr1 hhMrtr2 cytoneme & dorsal mesothoracic disc | somatic clone hhts2 denticle hhdsRNA.cRa denticle | ectopic hhAC denticle belt hh10 hh11 hh15 hh2 hh21 hh22 hh3 hh4 hh5 hh5A7 hh8 hh9 hhA hhAC hhAE hhAM denticle belt, with hha.cBa hhcBa denticle belt, with hhcBa hha.cBa denticle belt, with Scer\GAL4en-e16E hhN.Scer\UAS hhScer\UAS.cIa denticle belt, with Scer\GAL4ptc-559.1 hhΔCT.Scer\UAS hhScer\UAS.cIa denticle belt | supernumerary hhAC denticle row 1 hhAC denticle row 1 | ectopic, with Scer\GAL469B hhScer\UAS.cIa denticle row 2 hhAC denticle row 2 | ectopic, with Scer\GAL469B hhScer\UAS.cIa denticle row 3 hhAC denticle row 3, with Scer\GAL469B hhScer\UAS.cIa denticle row 4 hhAC denticle row 4, with Scer\GAL469B hhScer\UAS.cIa denticle row 5 hhAC denticle row 5, with Scer\GAL469B hhScer\UAS.cIa denticle row 5 | ectopic hhAC denticle row 6 hhAC denticle row 6, with Scer\GAL469B hhScer\UAS.cIa denticle row 6, with Scer\GAL4en-e16E hhΔCT.Scer\UAS hhScer\UAS.cIa denticle row 6, with Scer\GAL4hs.PB hhΔCT.Scer\UAS hhScer\UAS.cIa dorsal denticle hh4 hh8 hhts2 dorsal double row hhh9D dorsal double row, with Scer\GAL4en-e16E hhScer\UAS.cIa dorsal double row | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa dorsal double row | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa dorsal fine hair | ectopic hhts2 dorsal thick hair hh4 hh8 hhts2 dorsal thick hair, with Scer\GAL4en-e16E hhN.Scer\UAS dorsal thick hair, with Scer\GAL4hs.PB hhΔCT.Scer\UAS hhScer\UAS.cIa dorsal triple row hhh9D dorsal triple row, with Scer\GAL430A hhScer\UAS.cIa dorsal triple row | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa dorsal triple row | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa ectoderm hh8 egg chamber hhαTub84B.PB hh11 hh4 hhhs.PI hhts2 egg chamber (with hhAC) hhts2 egg chamber (with hhts2) hhAC ejaculatory bulb apodeme hhαTub84B.PB embryo | anterior | ectopic, with Scer\GAL469B hhScer\UAS.cIa embryo | segment polarity expression pattern hhAC embryonic/first instar larval cuticle hh21 hh8 hhAC embryonic/first instar larval cuticle, with Scer\GAL4en-e16E, hhAC hhC85S.Scer\UAS.cGa embryonic/first instar larval cuticle, with Scer\GAL4en-e16E, hhC85S.Scer\UAS.cGa hhAC hhSF.Scer\UAS.T:Rnor\CD2 embryonic/first instar larval cuticle, with Scer\GAL4en-e16E, hhSF.Scer\UAS.T:Rnor\CD2 hhC85S.Scer\UAS.cGa embryonic/first instar larval cuticle, with Scer\GAL4Scer\FRT.Rnor\CD2.Act5C, hhAC hhSF.Scer\UAS.T:Rnor\CD2 embryonic/first instar larval cuticle, with Scer\GAL4Scer\FRT.Rnor\CD2.Act5C, hhSF.Scer\UAS.T:Rnor\CD2 hhAC embryonic/first instar larval cuticle | dorsal hh4 hhhs.PI embryonic/first instar larval cuticle | germline clone hhAC embryonic/first instar larval cuticle | segment polarity expression pattern hhAC embryonic/first instar larval cuticle | segment polarity expression pattern (with hhAC) hhts2 embryonic/first instar larval cuticle | segment polarity expression pattern (with hhts2) hhAC embryonic/first instar larval cuticle | ventral hh068 hh20-107 hh278 hh279 hh281 hh287 hh302 hh4 hh5 hh8 hh9 hhAC hhts2 hhunspecified embryonic/larval cuticle, with Scer\GAL4en-e16E hhN.Scer\UAS embryonic/larval dorsal branch hh8 embryonic/larval dorsal branch, with Scer\GAL4btl.PS hhScer\UAS.cIa embryonic/larval dorsal branch, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 hhScer\UAS.cIa embryonic/larval dorsal trunk hh8 embryonic/larval dorsal vessel hh8 hhhs.PI embryonic/larval epidermis hhAC embryonic/larval epidermis, with Scer\GAL4en-e16E, hhAC hhC85S.Scer\UAS.cGa embryonic/larval epidermis, with Scer\GAL4en-e16E, hhC85S.Scer\UAS.cGa hhAC embryonic/larval epidermis, with Scer\GAL4Scer\FRT.Rnor\CD2.Act5C, hhAC hhSF.Scer\UAS.T:Rnor\CD2 embryonic/larval epidermis, with Scer\GAL4Scer\FRT.Rnor\CD2.Act5C, hhSF.Scer\UAS.T:Rnor\CD2 hhAC embryonic/larval fat body hh21 embryonic/larval foregut | embryonic stage hh21 embryonic/larval optic stalk hhbar3 hhts2 embryonic/larval posterior spiracle hh21 embryonic/larval tracheal system hh8 embryonic/larval tracheal system, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 hhScer\UAS.cIa embryonic/larval visceral branch hh8 embryonic abdomen, with Scer\GAL469B hhScer\UAS.cIa embryonic abdominal segment 1 & cuticle, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 1 & denticle belt, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 3 & denticle belt, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 3 & denticle belt | supernumerary, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 5 & denticle belt, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 5 & denticle belt | supernumerary, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 7 & denticle belt, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic abdominal segment 7 & denticle belt | supernumerary, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic brain hh21 embryonic dorsal epidermis hh6L93 embryonic dorsal epidermis, with Scer\GAL4en-e16E hhN.Scer\UAS hhScer\UAS.cAa embryonic epidermis hh10 hh11 hh15 hh1582 hh2 hh21 hh22 hh3 hh4 hh5 hh737 hh8 hh9 hhAC hhdsRNA.cRa embryonic epidermis, with Scer\GAL4en-e16E hhScer\UAS.cIa embryonic epidermis | dorsal hh4 hhts2 embryonic gnathal segment hhAC embryonic head hh5A7 hh6L93 hhAC hhAE embryonic head epidermis hh21 embryonic head epidermis | heat sensitive hhhs.PI embryonic hindgut hh21 embryonic large intestine hh21 embryonic optic lobe hh21 embryonic optic lobe | ectopic | heat sensitive hhhs.PI embryonic optic lobe | heat sensitive hhhs.PI embryonic rectum hh21 embryonic segment | heat sensitive hhts2 embryonic small intestine hh21 embryonic thoracic segment & cuticle, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic thoracic segment & denticle belt, with Scer\GAL4prd.RG1 hhScer\UAS.cIa embryonic ventral epidermis, with Scer\GAL4en-e16E hhN.Scer\UAS hhScer\UAS.cAa epidermis hh21 eye hhAC hhMrt hhbar3 hhfse hhts2 eye, with Scer\GAL4ey.PH hhEP3521 hhScer\UAS.cCa eye, with Scer\GAL4hs.2sev hhEP3521 eye (with hh8) hhbar3 eye (with hhbar3) hh8 eye & ommatidium hhαTub84B.PB eye | anterior hhbar3 eye | cell non-autonomous | somatic clone hhαTub84B.PB eye | somatic clone hhSH17 hhSH2 hhSH3 hhSH7 hhSH9 eye-antennal disc hhts2 eye-antennal disc, with Scer\GAL4dpp.blk1 hhScer\UAS.cIa eye disc hhts2 eye disc | somatic clone hhrJ413 hhts2 eye disc | somatic clone, with Scer\GAL4Act5C.PI hhScer\UAS.cSa eye equator | ectopic hhαTub84B.PB fat body/gonad primordium hhunspecified female genital disc, with Scer\GAL4cad-em459 hhScer\UAS.cCa female genitalia | supernumerary, with Scer\GAL4cad-em459 hhScer\UAS.cCa follicle cell hh11 hh4 hhhs.PI follicle cell, with Scer\GAL4hs.PB hhN.Scer\UAS follicle stem cell hhts2 frons hhαTub84B.PB fronto-orbital plate hhαTub84B.PB fusome hhhs.PI gastric caecum primordium hh8 gastric caecum primordium | heat sensitive (with hh4) hh8 gastric caecum primordium | heat sensitive (with hh8) hh4 genital arch hh4 hhlacZ genital disc hhts2 genital disc primordium hhunspecified germarium hhhs.PI hhts2 germarium region 2a hh11 hh4 hhhs.PI germarium region 2a, with Scer\GAL4hs.PB hhN.Scer\UAS germarium region 2b hh11 hh4 hhhs.PI germarium region 2b, with Scer\GAL4hs.PB hhN.Scer\UAS germarium region 3 hh11 hh4 germline cell (with hh4) hh11 germline cell (with hh11) hh4 germline cell | embryonic stage, with Scer\GAL4elav.PLu hhScer\UAS.cIa germline cell | embryonic stage, with Scer\GAL4h-1J3 hhScer\UAS.cIa germline cell | embryonic stage, with Scer\GAL4how-24B hhScer\UAS.cIa germline cell | embryonic stage, with Scer\GAL4twi.PG hhScer\UAS.cIa glial cell hhbar3 hhts2 glial cell & eye disc | somatic clone | cell non-autonomous, with Scer\GAL4Act5C.PP hhScer\UAS.cAa gonad hh18 gonadal sheath proper primordium hhunspecified gonopod long bristle hhαTub84B.PB hh4 hhlacZ gonopod long bristle | somatic clone hhAC gonopod thorn bristle hhαTub84B.PB hh4 hhlacZ gonopod thorn bristle | somatic clone hhAC haltere | distal hhMrt head capsule hhhs.PI head capsule | dorsal | heat sensitive hhts2 hypandrial process hh4 hhlacZ imaginal disc hh1582 hh737 interfollicle cell | ectopic hhαTub84B.PB interfollicle cell | supernumerary, with Scer\GAL4hs.PB hhScer\UAS.cAa interocellar bristle hhts2 interocellar bristle | heat sensitive hhts2 interommatidial bristle, with Scer\GAL4lz-gal4 hhScer\UAS.cIa lamina hhbar3 hhts2 lamina & neuron hhbar3 hhts2 lamina & neuron | precursor hhts2 lamina anlage glial cell | ectopic hhts2 lamina forming neuroblast hhαTub84B.PB hhbar3 hhts2 lamina intrinsic neuron hhbar3 lamina tangential neuron hhbar3 lateral ocellus hhts2 lateral trunk anterior branch hh8 lateral trunk posterior branch, with Scer\GAL4btl.PS hhScer\UAS.cIa leg hhαTub84B.PB leg | anterior compartment hhαTub84B.PB leg | distal hhMrt leg | heat sensitive hhts2 macrochaeta & tarsal segment 5 | distal hh4 macrochaeta | ectopic hhαTub84B.PB macrochaeta | ectopic, with Scer\GAL4C-734 hhScer\UAS.cCa macrochaeta | ectopic | somatic clone, with Scer\GAL4Ubx.PdC hhN.Scer\UAS.T:Avic\GFP male genital disc, with Scer\GAL4cad-em459 hhScer\UAS.cCa male genitalia hhαTub84B.PB hh4 hhlacZ male genitalia | somatic clone hhAC male genitalia | supernumerary, with Scer\GAL4cad-em459 hhScer\UAS.cCa male terminalia sensillum hhαTub84B.PB hh4 hhlacZ mechanosensory ventral triple row bristle, with Scer\GAL430A hhScer\UAS.cIa medial ocellus hhts2 medial triple row hhh9D medial triple row, with Scer\GAL430A hhScer\UAS.cIa medial triple row | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa medial triple row | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa mesothoracic tarsal bristle longitudinal row 1 hhbar3 mesothoracic tarsal bristle longitudinal row 2 hhbar3 mesothoracic tarsal bristle longitudinal row 2.5 hhbar3 mesothoracic tarsal bristle longitudinal row 3 hhbar3 mesothoracic tarsal bristle longitudinal row 3.5 hhbar3 mesothoracic tarsal bristle longitudinal row 4 hhbar3 mesothoracic tarsal bristle longitudinal row 5 hhbar3 mesothoracic tarsal bristle longitudinal row 5.5 hhbar3 mesothoracic tarsal bristle longitudinal row 6 hhbar3 mesothoracic tarsal bristle longitudinal row 6.5 hhbar3 mesothoracic tarsal bristle longitudinal row 7 hhbar3 mesothoracic tarsal bristle longitudinal row 8 hhbar3 mesothoracic tergum | somatic clone, with Scer\GAL4Ubx.PdC hhN.Scer\UAS.T:Avic\GFP metathoracic tracheal pit hh8 microchaeta & tarsal segment 1 hh4 microchaeta & tarsal segment 2 hh4 microchaeta & tarsal segment 3 hh4 microchaeta & tarsal segment 4 hh4 microchaeta & tarsal segment 5 hh4 microchaeta & wing | anterior | proximal hhh9D midline glial cell hhunspecified midline glial cell, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 hhScer\UAS.cIa midline glial cell, with Scer\GAL4sca-537.4 hhScer\UAS.cIa midline glial cell, with Scer\GAL4sim.P3.7 hhScer\UAS.cAa morphogenetic furrow hhbar3 hhfse hhts2 morphogenetic furrow, with Scer\GAL4upd-E132 hhScer\UAS.cPb morphogenetic furrow | ectopic | somatic clone, with Scer\GAL4Act5C.PI hhScer\UAS.cSa MP1 neuron, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 hhScer\UAS.cIa MP1 neuron, with Scer\GAL4sca-537.4 hhScer\UAS.cIa muscle attachment site hhMir neuroblast | larval stage | supernumerary hh21 neuroblast | larval stage | supernumerary | heat sensitive hhts2 neuroblast NB2-1 hh19 neuroblast NB2-2 hh19 neuroblast NB2-3 hh19 neuroblast NB2-4 hh19 neuroblast NB3-3 hh19 neuroblast NB5-1 hh19 neuroblast NB5-4 hh19 neuroblast NB5-5 hh19 neuroblast NB6-1 hh19 neuroblast NB6-2 hh19 neuroblast NB6-4 hh19 neuroblast NB7-3 hh19 hhAC ocellar bristle hhts2 ocellar bristle | heat sensitive hhts2 ocellus, with Scer\GAL4ey.PH hhC84S.Scer\UAS ocellus | heat sensitive hhts2 oenocyte hhMir ommatidial precursor cluster hhAC hhbar3 hhfse ommatidial precursor cluster, with Scer\GAL4lz-gal4 hhScer\UAS.cIa ommatidial precursor cluster (with hhAC) hhbar3 ommatidial precursor cluster (with hhbar3) hhAC ommatidium hhAC hhMrt hhbar3 hhts2 ommatidium (with hh8) hhbar3 ommatidium (with hhbar3) hh8 ommatidium | ectopic hhαTub84B.PB oocyte hhαTub84B.PB hhhs.PI oocyte, with Scer\GAL4hs.PB hhN.Scer\UAS oocyte & microtubule hhαTub84B.PB oocyte nucleus hhαTub84B.PB ovariole hhhs.PI ovariole, with Scer\GAL4hs.PB hhN.Scer\UAS ovariole (with hh4) hh11 ovariole (with hh11) hh4 ovariole (with hhAC) hhts2 ovariole (with hhts2) hhAC penis hhαTub84B.PB hh4 hhlacZ penis | somatic clone hhAC peripodial epithelium, with Scer\GAL4en-e16E hhN.Scer\UAS.cGa peripodial epithelium | heat sensitive hhts2 peripodial epithelium | somatic clone hhts2 photoreceptor cell hhts2 photoreceptor cell, with Scer\GAL4ey.PH hhN.Scer\UAS.T:Ivir\HA1 photoreceptor cell & axon hhbar3 hhts2 photoreceptor cell | somatic clone hhrJ413 photoreceptor cell R7 & axon hhbar3 photoreceptor cell R8 & axon hhbar3 polar follicle cell hhhs.PI polar follicle cell, with Scer\GAL4hs.PB hhN.Scer\UAS polar follicle cell | ectopic hhαTub84B.PB hhhs.PI pole cell | dorsal closure stage hhMrt hhunspecified pole cell | dorsal closure stage (with hhMrt) hhunspecified pole cell | dorsal closure stage (with hhunspecified) hhMrt posterior scutellar bristle hh4 postvertical bristle hhts2 postvertical bristle | heat sensitive hhts2 prescutum, with Scer\GAL4pnr-MD237 hhScer\UAS.cCa prescutum | ectopic hhαTub84B.PB presumptive embryonic/larval central nervous system hhunspecified prothoracic leg disc hhts2 proventriculus outer layer hh21 pseudotracheal sensillum | somatic clone hhAC retina, with Scer\GAL4lz-gal4 hhScer\UAS.cIa scutellar bristle, with Scer\GAL4en-e16E hhN.Scer\UAS hhScer\UAS.cAa scutellar bristle | ectopic, with Scer\GAL4sca-537.4 hhEP3521 scutellum hhαTub84B.PB scutellum, with Scer\GAL4C-734 hhScer\UAS.cCa scutellum, with Scer\GAL4en-e16E hhScer\UAS.cCa scutellum, with Scer\GAL4pnr-MD237 hhScer\UAS.cCa scutellum & macrochaeta hhαTub84B.PB scutellum & macrochaeta, with Scer\GAL4C-734 hhScer\UAS.cCa scutellum & macrochaeta, with Scer\GAL4en-e16E hhScer\UAS.cCa scutum hhαTub84B.PB scutum, with Scer\GAL430A hhScer\UAS.cIa scutum, with Scer\GAL4pnr-MD237 hhScer\UAS.cCa sensillum campaniformium of dorsal radius, with Scer\GAL4en-e16E hhScer\UAS.cCa sensory mother cell, with Scer\GAL4C-734 hhScer\UAS.cCa sensory mother cell, with Scer\GAL4dpp.blk1 hhScer\UAS.cCa sensory mother cell, with Scer\GAL4en-e16E hhScer\UAS.cCa spectrosome hhhs.PI sternite | heat sensitive hhhs.PI hhts2 subretinal glial cell | ectopic hhts2 tergite hhMirRevBx1 tergite, with Scer\GAL4T155 hhScer\UAS.cPb tergite | anterior hhαTub84B.PB hhMir tergite | anterior | heat sensitive hhhs.PI hhts2 tergite | ectopic | posterior, with Scer\GAL4T155 hhScer\UAS.cPb tergite | ectopic | posterior compartment hhαTub84B.PB tergite | heat sensitive hhhs.PI tergite | posterior hhαTub84B.PB tergite | posterior | heat sensitive hhts2 tracheal system hh520 hh688 tracheal tip cell | embryonic stage | supernumerary, with Scer\GAL4NP5133 hhScer\UAS.cKb trichogen cell, with Scer\GAL4sca-537.4 hhEP3521 unguis hhts2 ventral double row hhh9D ventral double row, with Scer\GAL4en-e16E hhScer\UAS.cIa ventral double row | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa ventral double row | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa ventral midline hh3 ventral midline, with Scer\GAL4sim.PS hhScer\UAS.cIa ventral nerve cord commissure, with Scer\GAL4sca-4512 hhEP3521 ventral thoracic disc | heat sensitive hhts2 ventral triple row hhh9D ventral triple row, with Scer\GAL430A hhScer\UAS.cIa ventral triple row | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa ventral triple row | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa visceral mesoderm hh8 VUM neuron, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 hhScer\UAS.cIa wing hhαTub84B.PB hhMrt wing, with Scer\GAL4en-e16E hhScer\UAS.cCa wing, with Scer\GAL4en-GAL4-33 hhScer\UAS.cUa wing | anterior hhMrt wing | anterior, with Scer\GAL430A hhScer\UAS.cCa wing | anterior, with Scer\GAL4Bx-MS1096 hhScer\UAS.cCa wing | anterior, with Scer\GAL4C-765 hhScer\UAS.cCa wing | anterior, with Scer\GAL4unspecified hhN.Scer\UAS wing | anterior/posterior compartment boundary, with Scer\GAL4C-734 hhScer\UAS.cCa wing | anterior/posterior compartment boundary, with Scer\GAL4dpp.blk1 hhScer\UAS.cCa wing | anterior | distal hhMrt wing | anterior | distal | cold sensitive hhMrt wing | anterior | proximal hhh9D wing | anterior | proximal, with Scer\GAL430A hhScer\UAS.cIa wing | anterior | proximal, with Scer\GAL434B hhScer\UAS.cIa wing | anterior compartment hhαTub84B.PB hhMrt hhhs.PI wing | anterior compartment, with Scer\GAL434B hhScer\UAS.cIa wing | anterior compartment | pharate adult stage, with Scer\GAL4Bx-MS1096 hhScer\UAS.cCa wing | ectopic hhαTub84B.PB wing | heat sensitive hhts2 wing | pharate adult stage, with Scer\GAL4MS941 hhScer\UAS.cCa wing | somatic clone hhAC hhSH17 hhSH2 hhSH3 hhSH7 hhSH9 wing blade, with Scer\GAL4MD-638 hhNIG.4637R wing disc hhMrt hhts2 wing disc, with Scer\GAL430A hhScer\UAS.cIa wing disc, with Scer\GAL4en-e16E hhN.Scer\UAS hhN.Scer\UAS.cGa wing disc | anterior compartment hhts2 wing disc | anterior compartment, with Scer\GAL430A hhΔCT.Scer\UAS wing disc | anterior compartment, with Scer\GAL4en-e16E hhN.Scer\UAS.T:Ivir\HA1 wing disc | anterior compartment, with Scer\GAL4en-e16E, hhN.Scer\UAS.T:Ivir\HA1 hhts2 wing disc | anterior compartment, with Scer\GAL4en-e16E, hhts2 hhN.Scer\UAS.T:Ivir\HA1 wing disc | cold sensitive hhMrt wing disc | heat sensitive hhts2 wing margin hhMrt wing vein hhαTub84B.PB hhMrt hhhs.PI wing vein, with Scer\GAL471B hhScer\UAS.cCa wing vein, with Scer\GAL4en-e16E hhN.Scer\UAS hhScer\UAS.cAa wing vein, with Scer\GAL4unspecified hhN.Scer\UAS wing vein | ectopic hhMrt wing vein | ectopic | somatic clone, with Scer\GAL4Ubx.PdC hhN.Scer\UAS.T:Avic\GFP wing vein | supernumerary hhMrt wing vein L1 hhh9D wing vein L1 | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa wing vein L1 | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa wing vein L2 hhMrt hhh9D wing vein L2 | cold sensitive hhMrt wing vein L2 | pharate adult stage, with Scer\GAL430A hhScer\UAS.cCa wing vein L2 | pharate adult stage, with Scer\GAL471B hhScer\UAS.cCa wing vein L2 | pharate adult stage, with Scer\GAL4zfh2-MS209 hhScer\UAS.cCa wing vein L3 hhMrt wing vein L3, with Scer\GAL4dpp.3KK hhScer\UAS.cIa wing vein L3, with Scer\GAL4en-e16E hhScer\UAS.cCa hhScer\UAS.cIa wing vein L3, with Scer\GAL4MD-638 hhScer\UAS.cCa hhNIG.4637R wing vein L3, with Scer\GAL4ptc-559.1 hhC84S.Scer\UAS wing vein L3, with Scer\GAL4unspecified hhN.Scer\UAS wing vein L3 | cold sensitive hhMrt wing vein L3 | pharate adult stage, with Scer\GAL471B hhScer\UAS.cCa wing vein L3 | pharate adult stage, with Scer\GAL4MS941 hhScer\UAS.cCa wing vein L4, with Scer\GAL4MD-638 hhNIG.4637R wing vein L4, with Scer\GAL4ptc-559.1 hhC84S.Scer\UAS wing vein L4, with Scer\GAL4unspecified hhN.Scer\UAS wing vein L4 | somatic clone hhAC
Classical Alleles ( 106 )
For All Classical Alleles Show
Allele of hh Class Mutagen Stocks Known lesion hhbar3 hypomorphic allele - genetic evidence spontaneous 6 Yes hhAC amorphic allele - genetic evidence, loss of function allele Delta2-3 2 Yes hh2 ethyl methanesulfonate 2 -- hhMir gain of function allele X ray 2 -- hhts2 hypomorphic allele - genetic evidence gamma ray 2 Yes hh21 loss of function allele ethyl methanesulfonate 1 Yes hhEP3521 P-element activity 1 Yes hhfse gamma ray 1 Yes hhH90 P-element activity 1 -- hhIJ32 1 -- hhMrt neomorphic allele - genetic evidence, gain of function allele spontaneous ethyl methanesulfonate 1 -- hhneo56 P-element activity 1 -- hhP30 P-element activity 1 Yes hhPyR215 P-element activity 1 Yes hhrJ413 P-element activity 1 -- hh10 amorphic allele - genetic evidence gamma ray 0 -- hh11 amorphic allele - genetic evidence gamma ray 0 -- hh3 loss of function allele, amorphic allele - genetic evidence ethyl methanesulfonate 0 -- hh8 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes hhAE amorphic allele - genetic evidence Delta2-3 0 Yes hh068 0 Yes hh10E 0 -- hh12 gamma ray 0 -- hh13 gamma ray 0 -- hh14 gamma ray 0 -- hh1582 0 -- hh15 PM hybrid dysgenesis 0 -- hh16 PM hybrid dysgenesis P-element activity 0 Yes hh17 PM hybrid dysgenesis P-element activity 0 Yes hh18 ethyl methanesulfonate 0 -- hh19 ethyl methanesulfonate 0 -- hh20-107 0 Yes hh20 ethyl methanesulfonate 0 -- hh22 triethylenemelamine 0 -- hh278 0 Yes hh279 0 Yes hh281 0 Yes hh287 0 Yes hh302 0 Yes hh4 ethyl methanesulfonate 0 Yes hh520 ethyl methanesulfonate 0 -- hh5 ethyl methanesulfonate 0 Yes hh5A7 ethyl methanesulfonate 0 -- hh688 ethyl methanesulfonate 0 -- hh6 ethyl methanesulfonate 0 -- hh6L93 ethyl methanesulfonate 0 -- hh6n16e 0 -- hh737 0 -- hh9 ethyl methanesulfonate 0 Yes hh9D94 0 -- hhA loss of function allele Delta2-3 0 Yes hhAM loss of function allele Delta2-3 0 Yes hhG31 0 -- hhGal4 P-element activity 0 -- hhh9D P-element activity 0 Yes hhlacZ P-element activity 0 -- hhMir-rv1 X ray 0 -- hhMir-rv2 X ray 0 -- hhMir-rv3 X ray 0 -- hhMir-rv4 X ray 0 -- hhMir-rv5 X ray 0 -- hhMir-rv6 X ray 0 -- hhMir-rv7 X ray 0 -- hhMirRevBx1 hypomorphic allele - genetic evidence 0 -- hhMrtr1 X ray ethyl methanesulfonate 0 Yes hhMrtr2 X ray ethyl methanesulfonate 0 Yes hhneo57 P-element activity 0 -- hhPIE-7 ethyl methanesulfonate 0 Yes hhQ50 hypomorphic allele - genetic evidence P-element activity 0 Yes hhSC10 ethyl methanesulfonate 0 -- hhSC11 ethyl methanesulfonate 0 -- hhSC12 ethyl methanesulfonate 0 -- hhSC13 ethyl methanesulfonate 0 -- hhSC14 ethyl methanesulfonate 0 -- hhSC15 ethyl methanesulfonate 0 -- hhSC16 ethyl methanesulfonate 0 -- hhSC17 ethyl methanesulfonate 0 -- hhSC18 ethyl methanesulfonate 0 -- hhSC19 ethyl methanesulfonate 0 -- hhSC1 ethyl methanesulfonate 0 -- hhSC20 ethyl methanesulfonate 0 -- hhSC21 ethyl methanesulfonate 0 -- hhSC2 ethyl methanesulfonate 0 -- hhSC3 ethyl methanesulfonate 0 -- hhSC4 ethyl methanesulfonate 0 -- hhSC5 ethyl methanesulfonate 0 -- hhSC6 ethyl methanesulfonate 0 -- hhSC7 ethyl methanesulfonate 0 -- hhSC8 ethyl methanesulfonate 0 -- hhSC9 ethyl methanesulfonate 0 -- hhSCE-2 ethyl methanesulfonate 0 -- hhSCG1 gamma ray 0 -- hhSCG2 gamma ray 0 -- hhSCG3 gamma ray 0 -- hhSFE-4 ethyl methanesulfonate 0 Yes hhSGE-2 ethyl methanesulfonate 0 Yes hhSH17 ethyl methanesulfonate 0 -- hhSH2 ethyl methanesulfonate 0 -- hhSH3 ethyl methanesulfonate 0 -- hhSH7 ethyl methanesulfonate 0 -- hhSH9 ethyl methanesulfonate 0 -- hhSHE-2 ethyl methanesulfonate 0 Yes hhSLE-1 ethyl methanesulfonate 0 Yes hhunspecified ethyl methanesulfonate 0 -- hhXE-2134 X ray 0 -- hhXE-3161 X ray 0 --
Alleles Carried on Transgenic Constructs ( 72 )
For All Alleles Carried on Transgenic Constructs Show
Allele of hh Class Mutagen Stocks Known lesion hhGD193 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes hhGD6242 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hhHMS00492 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hhJF01270 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hhJF01488 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hhJF01804 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hh90-471 in vitro construct 0 Yes hha.cBa in vitro construct - RNAi in vitro construct 0 Yes hhAct5C.PL in vitro construct - regulatory fusion 0 Yes hhAct5C.PN in vitro construct - regulatory fusion 0 Yes hhAct5C.PT in vitro construct - regulatory fusion 0 Yes hhAt1 in vitro construct - deletion 0 Yes hhbar3.T:Ecol\lacZ 0 Yes hhbar3L.T:Ecol\lacZ 0 Yes hhbar3L1.T:Ecol\lacZ 0 Yes hhbar3L2.T:Ecol\lacZ 0 Yes hhbar3L2.ΔEts.T:Ecol\lacZ 0 Yes hhbar3R.T:Ecol\lacZ 0 Yes hhC.Act5C in vitro construct 0 Yes hhC.hs in vitro construct - deletion 0 Yes hhC84S.Scer\UAS in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes hhC85S.N.Scer\UAS.T:Avic\GFP in vitro construct - regulatory fusion 0 Yes hhC85S.N.Scer\UAS in vitro construct - deletion in vitro construct - site directed mutagenesis 0 Yes hhC85S.N.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes hhC85S.Scer\UAS.cGa in vitro construct - regulatory fusion in vitro construct - site directed mutagenesis 0 Yes hhC85S.Scer\UAS.T:Avic\GFP in vitro construct - site directed mutagenesis 0 Yes hhC85S.Scer\UAS in vitro construct - regulatory fusion in vitro construct - amino acid replacement 0 Yes hhC85S.T:Ivir\HA1 in vitro construct - amino acid replacement 0 Yes hhcBa in vitro construct - RNAi in vitro construct 0 Yes hhdsRNA.cRa in vitro construct - RNAi 0 Yes hhF1 in vitro construct 0 Yes hhfse.T:Ecol\lacZ 0 Yes hhH329A.hs in vitro construct - regulatory fusion 0 Yes hhhs.PC in vitro construct - regulatory fusion 0 Yes hhhs.PI in vitro construct - regulatory fusion 0 Yes hhhs.PK in vitro construct - regulatory fusion 0 Yes hhhs.PP in vitro construct - genomic fragment 0 Yes hhKK108916 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes hhMtnA.PL in vitro construct - regulatory fusion 0 Yes hhMtnA.PvO in vitro construct - regulatory fusion 0 Yes hhN.Act5C.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes hhN.MtnA in vitro construct - regulatory fusion 0 Yes hhN.Scer\UAS.cGa in vitro construct - regulatory fusion in vitro construct - deletion in vitro construct - site directed mutagenesis 0 Yes hhN.Scer\UAS.T:Avic\GFP in vitro construct - site directed mutagenesis 0 Yes hhN.Scer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes hhN.Scer\UAS in vitro construct - deletion in vitro construct - regulatory fusion in vitro construct 0 Yes hhN.T:Ivir\HA1 in vitro construct - coding region fusion 0 Yes hhNIG.4637R in vitro construct - RNAi 0 Yes hhScer\FRT.Rnor\CD2.smo.αTub84B in vitro construct - regulatory fusion 0 Yes hhScer\FRT.Rnor\CD2.αTub84B in vitro construct - regulatory fusion 0 Yes hhScer\FRT.αTub84B in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cAa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cCa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cIa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cKa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cKb in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cLa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cPb in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cSa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.cUa in vitro construct - regulatory fusion 0 Yes hhScer\UAS.T:Arus\HRP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes hhScer\UAS.T:Avic\GFP-EGFP in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes hhScer\UAS.T:Ivir\HA1 in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes hhSF.Scer\UAS.T:Rnor\CD2 in vitro construct - coding region fusion in vitro construct - deletion 0 Yes hht14 in vitro construct - genomic fragment 0 Yes hht8.5 in vitro construct - genomic fragment 0 Yes hhT:Ivir\HA1 in vitro construct - coding region fusion 0 Yes hhU-CA.hs in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes hhU.Scer\UAS in vitro construct - regulatory fusion 0 -- hhαTub84B.PB FLPase 0 Yes hhΔCT.Scer\UAS in vitro construct - deletion 0 Yes hhΔSC.Scer\UAS in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(3R)B204 (Jurgens et al., 1984) Df(3R)BSC618 (Christensen et al., 2008.9.29) Df(3R)BSC619 (Christensen et al., 2008.9.29) Df(3R)BSC803 (Christensen et al., 2009.5.6) Df(3R)EB6 (Mohler et al., 1995, Therond et al., 1996) Df(3R)ED6096 (Ryder and Roote, 2004.11.10) Df(3R)ED6103 (Ryder and Roote, 2004.11.10) Df(3R)Exel6193 (Ryder, 2004.4.26) Df(3R)hhE23 (Spradling et al., 1999, BDGP Project Members, 1994-1999) Df(3R)r90b (Mohler et al., 1992) Df(3R)r94a (Mohler et al., 1992) Df(3R)S485 (Muller et al., 1999) T(1;3)Mir (Kopp et al., 1997) Ts(YLt;3Lt)B27 (Jurgens et al., 1984, Tearle and Nusslein-Volhard, 1987) Ts(YLt;3Lt)B172 (Jurgens et al., 1984)
Not disrupted in	Df(3R)e-N19 (Spradling et al., 1999, BDGP Project Members, 1994-1999) Ts(YLt;3Lt)D100+Ts(YLt;3Rt)H173 (Jones, 1971) Ts(YLt;3Lt)D100+Ts(YSt;3Rt)B27 (Jones, 1971) Ts(YSt;3Lt)R13 (Tearle and Nusslein-Volhard, 1987)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD193} NA P{GD6242} NA P{hhCD2} NA P{KK108916} NA P{TRiP.HMS00492} NA P{TRiP.JF01270} NA P{TRiP.JF01488} NA P{TRiP.JF01804} NA P{UAS-hh.A} NA P{UAS-hh.C84S} NA P{UAS-hh.C85S.G} NA P{UAS-hh.C85S.GFP} NA P{UAS-hh.C85S.N.GFP} NA P{UAS-hh.C85S.N} NA P{UAS-hh.C85S} NA P{UAS-hh.C} NA P{UAS-hh.F.HA} NA P{UAS-hh.GFP} NA P{UAS-hh.HRP} NA P{UAS-hh.I} NA P{UAS-hh.K} NA P{UAS-hh.Kb} NA P{UAS-hh.L} NA P{UAS-hh.N.G} NA P{UAS-hh.N.GFP} NA P{UAS-hh.N.HA} NA P{UAS-hh.N} NA P{UAS-hh.P} NA P{UAS-hh.S} NA P{UAS-hh.U} NA P{UAS-hh.Unspecified} NA P{UAS-hh.ΔCT} NA P{UAS-hh.ΔSC} NA reporter construct P{bar3-hs-lacZ} No P{bar3L2.ΔEts-hs-lacZ} No P{bar3L2-hs-lacZ} No P{bar3L-hs-lacZ} No P{bar3R-hs-lacZ} No P{barL1-hs-lacZ} No P{fse-hs-lacZ} No P{hh-lacZ.VM} No heat-shock construct P{HShh.I} NA P{hsHh} NA P{hshh} NA P{hsHh-C} NA P{hshhH329A} NA P{hsHh-UCA} NA P{hsp-hh.K} NA characterization construct P{hh.8.5-w} NA P{hh.14-w} NA P{hh.P1-w} NA P{hh.P2-lacZ} NA P{NIG.4637R} NA P{UAS-hh::Fas1.GPI} NA P{UAS-hh::Fas1.ΔGPI} NA P{αTub84B(FRT.y+)hh.B} NA P{αTub84B(FRT.y+CD2)hh.Z} NA P{αTub84B(-FRT)hh.B} NA P{αTub84B(-FRT)hh.D} NA P{αTub84B(-FRT)hh.Z} NA
Insertions
	Type of insertions Name Expression data insertion of enhancer trap P{lacW}A937A No P{lacZ}hhlacZ No P{lwB}hhH90 No P{PyR}hhPyR215 No P{PZ}hhP30 No P{PZ}hhQ50 No P{PZ}hhrJ413 No P{PZ}P2023-44 No miscellaneous insertions P{?'PZ}hhA NA P{?'PZ}hhAC NA P{?'PZ}hhAE NA P{?'PZ}hhAM NA P{hsneo}hhneo56 NA P{hsneo}hhneo57 NA P{hsp-hh.K}hhh9D NA insertion of mobile activating element P{EP}hhEP3521 NA P{GSV1}C279C279 NA insertion of enhancer trap binary system P{GAL4}hhGal4 No
Gene Ontology: Function, Process & Cellular Component ( 74 unique terms )
Terms Based on Experimental Evidence ( 31 terms )
Molecular Function
CV term Evidence References
cell surface binding inferred from direct assay (Yao et al., 2006, McLellan et al., 2006) protein binding inferred from physical interaction with ihog (McLellan et al., 2006)
Biological Process
CV term Evidence References
anterior commissure morphogenesis inferred from mutant phenotype (Bossing and Brand, 2006) Bolwig's organ morphogenesis inferred from mutant phenotype (Suzuki and Saigo, 2000) cytoneme assembly inferred from mutant phenotype (Ramirez-Weber and Kornberg, 1999) epithelial cell migration, open tracheal system inferred from mutant phenotype (Glazer and Shilo, 2001) eye morphogenesis inferred from mutant phenotype (Kent et al., 2006) genital disc anterior/posterior pattern formation inferred from expression pattern (Freeland and Kuhn, 1996) genital disc development inferred from mutant phenotype (Chen et al., 2005) germ cell migration inferred from mutant phenotype (Moore et al., 1998) glial cell migration inferred from mutant phenotype (Hummel et al., 2002) gonadal mesoderm development inferred from mutant phenotype (Moore et al., 1998) heart formation inferred from mutant phenotype (Park et al., 1996) hindgut morphogenesis inferred from mutant phenotype (Takashima and Murakami, 2001) labial disc development inferred from mutant phenotype (Joulia et al., 2005) morphogenesis of larval imaginal disc epithelium inferred from mutant phenotype (McClure and Schubiger, 2005) pole cell migration inferred from genetic interaction with disp (Deshpande and Schedl, 2005) inferred from genetic interaction with Hmgcr (Deshpande and Schedl, 2005) positive regulation of hh target transcription factor activity inferred from genetic interaction with Su(fu) AND inferred from genetic interaction with fu (Ohlmeyer and Kalderon, 1998) positive regulation of transcription factor import into nucleus inferred from mutant phenotype (Wang et al., 2000) progression of morphogenetic furrow involved in compound eye morphogenesis inferred from expression pattern AND inferred from genetic interaction with dpp AND inferred from genetic interaction with h AND inferred from genetic interaction with sca AND inferred from mutant phenotype (Ma et al., 1993) protein autoprocessing inferred from direct assay AND inferred from mutant phenotype (Hall et al., 1997) regulation of mitotic cell cycle inferred from mutant phenotype (Zhang and Kalderon, 2001) regulation of protein import into nucleus inferred from mutant phenotype (Horabin et al., 2003) segment polarity determination inferred from expression pattern (assigned by UniProtKB) (Mohler and Vani, 1992) inferred from mutant phenotype (Merabet et al., 2005) smoothened signaling pathway inferred from genetic interaction with smo (van den Heuvel and Ingham, 1996) spiracle morphogenesis, open tracheal system inferred from mutant phenotype (Merabet et al., 2005) ventral midline development inferred from mutant phenotype (Bossing and Brand, 2006)
Cellular Component
CV term Evidence References
cytoplasm inferred from direct assay (assigned by UniProtKB) (Mohler and Vani, 1992) cytoplasmic membrane-bounded vesicle inferred from direct assay (Gallet et al., 2003) endocytic vesicle inferred from direct assay (Callejo et al., 2008) nucleus inferred from direct assay (assigned by UniProtKB) (Mohler and Vani, 1992)
Terms Based on Predictions or Assertions ( 51 terms )
Molecular Function
CV term Evidence References
cysteine-type endopeptidase activity inferred from sequence or structural similarity (Coates, 1999.11) endopeptidase activity non-traceable author statement (Swiss-Prot Project Members, 1994.2.1) morphogen activity traceable author statement (Sanchez and Guerrero, 2001) patched binding non-traceable author statement (Swiss-Prot Project Members, 1994.2.1) traceable author statement (Chen and Struhl, 1998, Marsh and Theisen, 1999, Ogden et al., 2004) peptidase activity inferred from electronic annotation with InterPro:IPR001767 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
analia development traceable author statement (Sanchez and Guerrero, 2001) anterior/posterior axis specification, embryo non-traceable author statement (Taylor, 2002) anterior/posterior lineage restriction, imaginal disc traceable author statement (Milan and Cohen, 2000) anterior head segmentation traceable author statement (Peel, 2004) cell-cell signaling inferred from electronic annotation with InterPro:IPR000320 (FlyBase Curators et al., 2004-) compartment pattern specification traceable author statement (McNeill, 2000) compound eye morphogenesis traceable author statement (Suzuki and Saigo, 2000) compound eye photoreceptor cell differentiation traceable author statement (Treisman, 2004) developmental pigmentation traceable author statement (Wittkopp et al., 2003) ectoderm development non-traceable author statement (Chen and Fishman, 2000) epidermis development traceable author statement (Hatini et al., 2000) foregut morphogenesis traceable author statement (Murakami et al., 1999) genital disc development traceable author statement (Sanchez and Guerrero, 2001) germ cell attraction traceable author statement (Coffman, 2003) germ-line stem cell division traceable author statement (Deng and Lin, 2001, Lin, 2002) growth non-traceable author statement (Oldham et al., 2000) heart development non-traceable author statement (Cripps and Olson, 2002) traceable author statement (Chen and Fishman, 2000) hindgut morphogenesis traceable author statement (Murakami et al., 1999) imaginal disc-derived wing vein specification traceable author statement (Milan and Cohen, 2000) imaginal disc growth traceable author statement (Hafen and Stocker, 2003) imaginal disc pattern formation non-traceable author statement (Potter and Xu, 2001, Swiss-Prot Project Members, 1994.2.1) intein-mediated protein splicing inferred from electronic annotation with InterPro:IPR006141 (FlyBase Curators et al., 2004-) leg disc morphogenesis traceable author statement (Sanchez and Guerrero, 2001) mesoderm development non-traceable author statement (Chen and Fishman, 2000) open tracheal system development traceable author statement (Ghabrial et al., 2003) ovarian follicle cell development traceable author statement (Dobens and Raftery, 2000) ovarian follicle cell stalk formation traceable author statement (Dobens and Raftery, 2000) positive regulation of smoothened signaling pathway traceable author statement (Leahy, 1999, Marsh and Theisen, 1999) posterior head segmentation traceable author statement (Peel, 2004) progression of morphogenetic furrow involved in compound eye morphogenesis traceable author statement (Peters, 2002) protein autoprocessing non-traceable author statement (Swiss-Prot Project Members, 1994.2.1) traceable author statement (Leahy, 1999) protein splicing traceable author statement (Leahy, 1999) proteolysis inferred from electronic annotation with InterPro:IPR001767 (FlyBase Curators et al., 2004-) R8 cell fate specification non-traceable author statement (Frankfort and Mardon, 2002) regulation of cell proliferation non-traceable author statement (Sanchez and Guerrero, 2001) regulation of organ growth traceable author statement (Potter and Xu, 2001) regulation of transcription from RNA polymerase II promoter traceable author statement (Sanson, 2001) segment polarity determination non-traceable author statement (Sanson, 2001, Swiss-Prot Project Members, 1994.2.1) traceable author statement (Peel, 2004) smoothened signaling pathway non-traceable author statement (McNeill, 2000, Swiss-Prot Project Members, 1994.2.1) traceable author statement (Farese and Herz, 1998, Ogden et al., 2004) somatic stem cell division traceable author statement (Lin, 2002) trunk segmentation traceable author statement (Peel, 2004) wing disc anterior/posterior pattern formation traceable author statement (Milan and Cohen, 2000, McNeill, 2000, de Celis, 2003, Ogden et al., 2004, Crozatier et al., 2004) wing disc proximal/distal pattern formation traceable author statement (de Celis, 2003)
Cellular Component
CV term Evidence References
cytoplasm non-traceable author statement (Swiss-Prot Project Members, 1994.2.1) extracellular region non-traceable author statement (Swiss-Prot Project Members, 1994.2.1) traceable author statement (Chen and Struhl, 1998, Fuller, 1998, Leahy, 1999) nucleus non-traceable author statement (Swiss-Prot Project Members, 1994.2.1)
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 hh allele Gene References hh2 sgl (Benevolenskaya et al., 1998) hh4 Wnt4 (Buratovich et al., 2000) hh5A7 Dfd (Harding et al., 1995) hh8 amos (Chanut et al., 2002) ro (Chanut et al., 2000) trr (Sedkov et al., 2003) wg (Cox and Baylies, 2005) hh18 CycE (Brumby et al., 2004) hh21 ci (Chen et al., 1999) CycE (Brumby et al., 2004) os (Bach et al., 2003) hh737 ro (Chanut et al., 2000) hh1582 ro (Chanut et al., 2000) hhAC ato (White and Jarman, 2000) ci (Gallet et al., 2000) CycE (Brumby et al., 2004) sgl (Benevolenskaya et al., 1998) wg (Deshpande et al., 2001, Cox and Baylies, 2005) hhbar3 amos (Heberlein et al., 1993) ey (Kango-Singh et al., 2003) oro (Epps et al., 1997) sca (Chou and Chien, 2002) trr (Sedkov et al., 2003) hhG31 os (Bach et al., 2003) hhh9D ptc (Kojima et al., 1994) hhhs.PI fs(1)Yb (King et al., 2001) ph-p (Maschat et al., 1998) piwi (King et al., 2001) hhMir bi (Kopp and Duncan, 1997) hhMrt Asx (Randsholt et al., 2000) brm (Brizuela et al., 1994, Felsenfeld and Kennison, 1995) dally (Jackson et al., 1997) fu (Alves et al., 1998) gpp (Shanower et al., 2005) mxc (Randsholt et al., 2000) ph-d (Randsholt et al., 2000) pho (Randsholt et al., 2000) Psc (Randsholt et al., 2000) ptc (Johnson et al., 1995) Su(hh)I (Haines and van Den Heuvel, 2000) Su(hh)II (Haines and van Den Heuvel, 2000) Su(hh)III (Haines and van Den Heuvel, 2000) Su(hh)IV (Haines and van Den Heuvel, 2000) Su(hh)V (Haines and van Den Heuvel, 2000) Su(hh)VI (Haines and van Den Heuvel, 2000) Su(hh)VII (Haines and van Den Heuvel, 2000) vtd (Schulze et al., 2001) hhN.Scer\UAS cos (Lum, 2003) hhrJ413 bun (Treisman et al., 1995) dac (Mardon et al., 1994) hyd (Lee et al., 2002) hhSC10 gl (Ma et al., 1996) hhSC11 gl (Ma et al., 1996) hhSC12 gl (Ma et al., 1996) hhSC13 gl (Ma et al., 1996) hhScer\UAS.cAa ci (Kango-Singh et al., 2003) dpp (Kango-Singh et al., 2003) ey (Kango-Singh et al., 2003) hhScer\UAS.cCa kn (Vervoort et al., 1999) ptc (Mullor and Guerrero, 2000) ro (Chanut et al., 2000) hhScer\UAS.cIa ci (Gallet et al., 2000) fu (Alves et al., 1998) tsh (Gallet et al., 2000) ttv (Gallet et al., 2003) hhScer\UAS.cKa ci (Suzuki and Saigo, 2000) fu (Suzuki and Saigo, 2000) hhScer\UAS.cPb gish (Hummel et al., 2002) hhScer\UAS.cSa ato (White and Jarman, 2000) hhts2 dpp (Borod and Heberlein, 1998) kn (Vervoort et al., 1999) ph-p (Maschat et al., 1998) Pka-C1 (Strutt et al., 1995, Zhang and Kalderon, 2000) ptc (Ma et al., 1996, Zhang and Kalderon, 2000) wg (Borod and Heberlein, 1998) hhunspecified Dfd (Gellon et al., 1997) oro (Tanda et al., 1996) hhXE-2134 Dsor1 (Karim et al., 1996) phl (Karim et al., 1996) phl::tor (Karim et al., 1996) Ras85D (Karim et al., 1996) hhXE-3161 Dsor1 (Karim et al., 1996) phl (Karim et al., 1996) phl::tor (Karim et al., 1996) Ras85D (Karim et al., 1996) unspecified ci (Pankratz and Hoch, 1995) cos (Ruel et al., 2003) dpp (McClure and Schubiger, 2005) Egfr (Read et al., 2005) en (Maschat et al., 1998) fu (Sanchez-Herrero et al., 1996, Ohlmeyer and Kalderon, 1998) gl (Ma et al., 1996) L (Singh et al., 2005) lin (Bokor and DiNardo, 1996) pb (Joulia et al., 2005) Psn (Mahoney et al., 2006) ptc (Bokor and DiNardo, 1996, Epps et al., 1997, Ruel et al., 2003) Ret (Read et al., 2005) rho (Sturtevant and Bier, 1995) smo (Ruel et al., 2003) snama (Jones et al., 2006) Su(fu) (Ohlmeyer and Kalderon, 1998) Su(hh)2A (Reifegerste and Moses, 1998) Su(hh)3A (Reifegerste and Moses, 1998)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 67682 DroID - A comprehensive database of gene and protein interactions. • FBgn0004644 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 42737
Orthologs
Genome-wide drosophilid orthologs	Dana\GF18029 Dere\GG12458 Dgri\GH23852 Dmoj\GI24573 Dper\GL23598 Dpse\GA18321 Dsec\GM23589 Dsim\GD18403 Dvir\GJ22641 Dwil\GK12833 Dyak\GE23980
Curated drosophilid orthologs	Dhyd\hh   Dmau\hh   Dpse\GA18321   Drep\hh (Ranz et al., 2001, Ranz et al., 2001) Dsim\hh (Coyne, 1997.1.2)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL006708-PA Aedes aegypti (mosquito) AGAP001412-PA Anopheles gambiae (mosquito) XP_001943707.1 Acyrthosiphon pisum (pea aphid) 84974 Branchiostoma floridae (lancelet) BGIBMGA012535-PA Bombyx mori (silk worm) ENSBTAP00000000135 Bos taurus (bovine) ENSCAFP00000012759 Canis familiaris (dog) ENSCINP00000013864 Ciona intestinalis (sea squirt) CPIJ003906 Culex pipens (mosquito) ENSCPOP00000002048 Cavia porcellus (guinea pig) RP19440 Caenorhabditis remanei (nematode) ENSCSAVP00000017051 Ciona savignyi (sedentary tunicate) 129621 Capitella spl (polychaete worm) 290571 Daphnia pulex (daphnia) ENSDARP00000056746 Danio rerio (zebra fish) ENSECAP00000008540 Equus caballus (horse) ENSGACP00000005121 Gymnopilus aculeatus (mushroom) ENSGALP00000010292 Gallus gallus (chicken) 154346 Hypsipyla robusta (cedar shoot borer) ENSP00000266991 Homo sapiens (human) 121665 Lottia gigantea (owl limpet) ENSMODP00000019154 Monodelphis domestica (gray opossum) ENSMMUP00000001060 Macaca mulatta (rhesus monkey) ENSMUSP00000023737 Mus musculus (house mouse) 241466 Nematostella vectensis (starlet anemone) XP_001605475.1 Nasonia vitripennis (wasp) ENSOANP00000000534 Ornithorynchus anatinus (platypus) ENSORLP00000013115 Oryzias latipes (japanese killifish) PHUM002563-PA Pediculus humanus (human body louse) ENSPPYP00000005114 Pongo pygmaeus (orangutan) ENSPTRP00000022106 Pan troglodytes (chimpanzee) ENSRNOP00000020563 Rattus norvegicus (norway rat) NP_001012720.1 Strongylocentrotus purpuratus (purple sea urchin) NP_001107837.1 Tribolium castaneum (red flour beetle) ENSTNIP00000015753 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000045862 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000024437 Xenopus tropicalis (frog) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB13786 NV16490 LH13486 ISCW009092 ISCW007045 FBgn0241122 FBgn0214842 FBgn0209745 FBgn0189928 FBgn0178454 FBgn0250482 JGI_V11_290571 FBgn0078326 FBgn0161188 FBgn0147295 FBgn0131309 FBgn0104746 FBgn0095047 CPIJ003906 BGIBMGA012535 ACYPI27225 ACYPI006525 GB30509 GB14574 AGAP001412 AAEL006708 PHUM018090 GLEAN_01364
Stocks & Reagents
Stocks Listed in FlyBase ( 31 )
Bloomington	1376 hhbar3 tx1 35562 w1118; hhfse 26166 hhMrt/TM3, Sb1 5530 ry506 P{PZ}hhP30 5338 P{ftz/lacC}4; hh21/TM3, Sb1
Kyoto	105915 hhbar3 106293 hhbar3 tx1 122063 w1118; P{EP}hhEP3521 108431 T(1;3)Mir, hhMir/TM6B, Tb1
VDRC	v1402 w1118; P{GD193}v1402 v1403 w1118; P{GD193}v1403 v43255 w1118; P{GD6242}v43255
	More stocks available...
Genomic Clones ( 2 )
	BACR03N12 BACR17P04 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 22 )
	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
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD04474 LD07204 LD38166 RE04204 RE20958 RH26379 RH30894 RH54606 RH63210 RH65176 SD16140 SD24332 SD26463
Other clones	11123 35991 97298 104024 EK111112 EK146832 EK227247 EK243330 SD13349
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0004644 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 42737
Antibody Information
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: hh anon-WO0134654.19 (FlyBase, 1996-) Source for merge of: hh l(3)neo56 (Spradling et al., 1999)
Additional comments	Source for merge of hh anon-WO0134654.19 was sequence comparison (date:051113). (FlyBase, 1996-)
Other Comments
	Cholesterol modification of hh is necessary for hh-dependent graded cell fate specification in the dorsal epidermis. (Gallet et al., 2006) One of 42 Drosophila genes identified as being most likely to reveal molecular and cellular mechanisms of nervous system development or plasticity relevant to human Mental Retardation disorders. (Inlow and Restifo, 2004) Cholesterol modification of hh protein is necessary (in the embryonic epidermis) for its assembly in large punctate subcellular structures and apical sorting through the activity of the disp protein. Movement of these specialized structures containing hh protein through the cellular field is contingent upon the activity of ttv. (Gallet et al., 2003) hh is responsible for maintaining Con expression and its own expression in the embryonic trunk mesoderm during gut and tracheal development. (Hosono et al., 2003) gish, so, ey and hh act in the posterior region of the eye disc to prevent precocious glial cell migration. (Hummel et al., 2002) The expression of hh in the wing disc, once activated, is dependent on ph-p, trx and brm. This may be due to an element upstream of the hh transcriptional start site (hh-CMM) that can bind Pc protein and is able to act as a cellular memory module (CMM) when placed upstream of a UAS sequence in reporter constructs. (Maurange and Paro, 2002) Misexpression of hh in the soma induces germ cells to migrate to inappropriate locations in the developing embryo. (Deshpande et al., 2001) hh may act as an attractive guidance cue in germ cell migration in the embryo. (Deshpande et al., 2001) Migration of all tracheal branches is absent or stalled in hh-mutant embryos. (Glazer and Shilo, 2001) hh expression in the prospective rectum is necessary for the expression of dpp at the posterior end of the adjacent large intestine. hh expression is also required for the development of the rectum. (Takashima and Murakami, 2001) hh acts as a somatic stem cell factor in the ovary. (Zhang and Kalderon, 2001) Clones of hh mutants in the peripodial membrane disrupt disc growth. (Cho et al., 2000) The secreted proteins encoded by hh, wg and dpp are expressed in the peripodial membrane yet they control the expression of Dl and Ser in the disc proper. (Cho et al., 2000) hh signalling from the peripodial membrane, but not from the disc proper, is required for eye disc patterning and growth. (Cho et al., 2000) ptc protein destabilises smo protein in the absence of hh protein. (Denef et al., 2000) Bolwig's organ formation is governed by ato, the expression of which is under the control of hh, eya and so. (Suzuki and Saigo, 2000) hh activates vn expression. This activation is mediated through the gene ci. (Amin et al., 1999) hh induces Egfr signalling during head development. (Amin et al., 1999) hh is required in the developing eye both for the induction of ato expression that prefigures the position of the R8 cells, and for the repression of ato expression between the nascent proneural clusters. Both effects are due to the direct stimulation of responding cells by the hh gene product itself. (Dominguez, 1999) hh plays a role in ommatidial development by regulating ato expression both positively and negatively. (Dominguez, 1999) In the absence of hh activity, prothoracic leg disc fragments fail to undergo anterior/posterior conversion, but can still regenerate missing anterior pattern elements. hh-independent regeneration (integration) may be mediated by the wg and dpp positional cues. (Gibson and Schubiger, 1999) hh is required for activation of en during regeneration of fragmented imaginal discs. (Gibson and Schubiger, 1999) The en/hh interface in the embryonic epidermis imposes asymmetry on wg signaling. (Gritzan et al., 1999) Three EMS induced alleles were identified in a screen for mutations affecting commissure formation in the CNS of the embryo. (Hummel et al., 1999) hh signalling, coming from the adjacent P compartments across both Anterior/Posterior and Posterior/Anterior boundaries in the abdomen, organizes the pattern of all the Anterior cells. (Lawrence et al., 1999) Cell affinities in the adult abdomen depend on hh: cells of the A compartment show two gradients of affinity, both of which depend on direct readouts of the level of hh function. (Lawrence et al., 1999) cad acts in combination with the hh pathway to specify the different components of the analia. (Moreno and Morata, 1999) hh plays a role in the formation of the posterior barrier to wg movement at the presumptive embryonic segment border. Increased hh signalling decreases the domain of wg action in the anterior direction. (Sanson et al., 1999) fu is required autonomously in anterior cells neighboring hh to maintain ptc and wg expression. wg is in turn maintaining en and hh expression. The hh signalling components smo and ci are required in cells posterior to hh to maintain ptc expression, whereas fu is not necessary in these cells. (Therond et al., 1999) The levels of glycosaminoglycans (in which sgl plays a role) are rate limiting for cell-cell signalling pathways such as those of wg and hh, which mediate changes in gene expression. (Benevolenskaya et al., 1998) hh is required at the posterior margin of the eye disc to maintain expression of dpp and ato. (Borod and Heberlein, 1998) ptc protein normally binds hh gene product without any help of the smo gene product, though smo is also a part of the receptor complex that binds hh and transduces the hh signal. The mechanism of signal transduction may involve hh binding specifically to ptc and inducing a conformational change leading to the release of latent smo activity. (Chen and Struhl, 1998) The division of the limb into two antagonistic domains, as defined by exd function and hh signaling, may be a general feature of limb development. (Gonzalez-Crespo et al., 1998) hh and spi bring about the concerted assembly of ommatidial and synaptic cartridge units, imposing the "neurocrystalline" order of the compound eye onto the post-synaptic target field. hh encodes an inductive signal that is transported along retinal axons from the developing eye, and induces the expression of Egfr in post-synaptic precursor cells. (Huang et al., 1998) In the absence of hh signalling results propose that Su(fu) inhibits ci by binding to it and that, upon reception of the hh signal, fu is activated and counteracts Su(fu), leading to the activation of ci. (Monnier et al., 1998) Mutants are isolated in an EMS mutagenesis screen to identify zygotic mutations affecting germ cell migration at discrete points during embryogenesis: mutants exhibit segment polarity pattern defects. (Moore et al., 1998) The hh product stimulates maturation of ci into a labile transcriptional activator. (Ohlmeyer and Kalderon, 1998) CrebA hh double mutant phenotype confirms that CrebA is not involved in segment polarity. (Andrew et al., 1997) Processing of the full length ci protein is inhibited by hh, an observation that represents the first direct evidence that ci transduces the hh signal. (Aza-Blanc et al., 1997) Each primordia of the genital disc (female genital, male genital and anal primordia) is divided into anterior and posterior compartments. Clonal phenotype of genes known to play compartment specific functions demonstrate the anterior/posterior patterning functions of these genes are conserved in the genital disc. (Chen and Baker, 1997) Genetic combinations with mutants of nub cause additive phenotypes. (Cifuentes and Garcia-Bellido, 1997) Clonal analysis demonstrates hh has two distinct functions: expression is required in the photoreceptor cells to drive the morphogenetic furrow and in addition hh secreted from cells at the posterior disc margin is absolutely required for the initiation of patterning and predisposes ommatidial precursor cells to enter ommatidial assembly later. (Dominguez and Hafen, 1997) hh induces ommatidial development in the absence of its secondary signals wg and dpp. Regulatory relationships between hh, dpp and wg in the eye are similar to those found in other imaginal discs, such as the leg. (Dominguez and Hafen, 1997) Cross-regulatory relationships among hh, wg and en, as well as their initial mode of activation, in the anterior head are significantly different from those in the trunk. (Gallitano-Mendel and Finkelstein, 1997) Identified in a screen for modifiers of the Dfd13/Dfd3 mutant phenotype. Shows no interaction with the Pc mutant phenotype. (Gellon et al., 1997) ci forms a negative feedback loop with ptc that is regulatd by hh signal transduction. (Hepker et al., 1997) bi is the primary target of hh signaling in the adult abdomen, mediating both the morphogenetic and polarity-reversal functions of hh. (Kopp and Duncan, 1997) hh protein secreted by posterior compartment cells plays a key role in patterning the posterior portion of the anterior compartment in adult abdominal segments. (Kopp et al., 1997) dpp specifies the position of most of the sensory organ precursors (SOPs) in the notum and some of them in the wing. Close to the A/P compartment border of the wing, however, SOPs are specified by hh rather than by dpp alone. (Mullor et al., 1997) dpp only mediates a subset of hh functions in the morphogenetic furrow. (Penton et al., 1997) dpp does not appear to be the principal mediator of hh function in the eye. (Pignoni and Zipursky, 1997) Loss of smo function causes a hh-like phenotype. smo activity is required for transduction of hh but not wg. smo acts downstream from ptc to transduce the hh signal. (Quirk et al., 1997) 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) The affinity boundary that segregates A and P cells into adjacent but immiscible cell populations is to a large extent a consequence of local hh signalling, rather than a reflection of an intrinsic affinity difference between A and P cells. (Rodriguez and Basler, 1997) cos encodes a kinesin-related protein that accumulates preferentially in cells capable of responding to hh signal. (Sisson et al., 1997) Comparing the biological activities of secreted and membrane-tethered hh protein provides evidence that hh forms a local concentration gradient and functions as a concentration-dependent morphogen in the wing. (Strigini and Cohen, 1997) The pattern of expression of hh in the larval and adult abdomen has been analysed. (Struhl et al., 1997) The function of hh in morphogenetic furrow progression is indirect. Cells that cannot receive/transduce the hh signal (as in smo clones) are still capable of entering a furrow fate and differentiating normally. However hh is required to promote furrow progression and regulate its rate of movement across the disc, since the furrow is delayed in smo clones. (Strutt and Mlodzik, 1997) hh and ptc can regulate transcription from a wg enhancer element containing ci protein binding sites by modulating the activity of ci protein. (Von Ohlen and Hooper, 1997) smo encodes a seven-pass membrane protein, a putative receptor of the hh signal. (Alcedo et al., 1996) Elevated levels of ci are sufficient to activate hh target genes, even in the absence of hh activity. ci activates transcription in yeast by a GLI consensus-binding site and the zinc finger domain is sufficient for its target specificity. Results strongly support a role for ci as the transcriptional activator that mediates hh signaling. (Alexandre et al., 1996) hh is required for the normal activation of bap and srp in anterior portions of each parasegment, whereas wg is required to suppress bap and srp expression in posterior portions. hh and wg play opposing roles in mesoderm segmentation. (Azpiazu et al., 1996) wg and hh signaling account for all cell types across the dorsal epidermis. dpp does not appear to mediate this hh dorsal epidermis signaling. hh antagonizes the activity of ptc in the specification of primary and secondary but not tertiary cell types. hh also antagonizes lin function. (Bokor and DiNardo, 1996) Distinction between dorsal and ventral fates is maintained through mutual repression by dpp and wg. Expression of wg and dpp in their normal domains depends on the hh signal. Cells that are not likely to be within range of the wg or dpp signals have a different capacity to respond to hh. (Brook and Cohen, 1996) Loss of da disrupts the progression of the morphogenetic furrow and this effect is mediated by the loss of both hh and dpp. (Brown et al., 1996) smo activity is required in wing anterior cells along the A/P boundary for these cells both to transduce hh and to limit its further movement into the anterior compartment. ptc regulates smo activity in response to hh signalling. (Chen and Struhl, 1996) Ectodermal and mesodermal Dr expression depend on wg and hh. (D'Alessio and Frasch, 1996) Cells in anterior compartments lacking ci express hh and adopt a posterior fate without expressing en. Increased levels of ci can induce the expression of dpp independent of hh. Expression of ci in anterior cells controls limb development by restricting hh secretion to posterior cells and by conferring competence to respond to hh by mediating transduction of the hh signal. (Dominguez et al., 1996) hh is required for the proliferation and specification of ovarian somatic cells prior to egg chamber formation. hh signalling during egg chamber assembly appears to be closley related to, or part of pathways involving the neurogenic genes. (Forbes et al., 1996) ptc and ci are expressed in a pattern complementary to hh and en in adult ovaries. Ectopic expression of hh results in the ectopic expression of ptc. hh directly effects region 2 somatic cells of the germarium via a signalling pathway which includes ptc and ci, but not wg or dpp. (Forbes et al., 1996) The expression pattern of a number of genes in the larval genital discs, including a hh-Ecol\lacZ reporter gene, has been studied to determine the segment-parasegment organisation of the genital discs. (Freeland and Kuhn, 1996) ara-caup expression at patches on the wing, located one at each side of the DV compartment border, is mediated by the hh signal through its induction of high levels of ci protein in anterior cells near to the AP compartment border. (Gomez-Skarmeta and Modolell, 1996) exd is expressed in a normal pattern in the absence of hh function. (Gonzalez-Crespo and Morata, 1996) hh, wg and dpp are required for the establishment of signaling centres that coordinate morphogenesis in the hindgut epithelium. Activation of these genes in the developing hindgut and foregut requires fkh. hh and wg activities in the gut epithelial cells are required for the expression of the homeobox gene bap in the ensheathing visceral mesoderm. (Hoch and Pankratz, 1996) The secreted hh product regulates the temporal assembly of photoreceptor precursor cells into ommatidia in the eye and is transmitted along the retinal axons to serve as the inductive signal in the brain, triggering neurogenesis in the developing visual centers. hh acts in the first of two retinal axon-mediated steps in the assembly of lamina synaptic cartridges. (Huang and Kunes, 1996) A combination of hh and wg is required to specify the most posterior fates of the A compartment. (Lawrence et al., 1996) Four segment polarity genes, hh, wg, gsb and en all function in concert to determine the formation and specifications of three hh-dependent eg-neuroblasts (6-4, 7-3 and 2-4). (Matsuzaki and Saigo, 1996) hh is required in the early gastrula for heart development, overexpression of hh increases the amount of heart formation. Overexpression of wg restores the heart deficit of hh mutant embryos. (Park et al., 1996) The hh autoprocessing reaction proceeds via an internal thioester intermediate and results in a covalent modification that increases the hydrophobic character of the signalling domain and influences its spatial and subcellular distribution. Truncated, unprocessed amino terminal protein causes embryonic mispatterning, suggesting a role for autoprocessing in spatial regulation of hh signalling. (Porter et al., 1996) Cholesterol is the lipophilic moiety covalently attached to the amino-terminal signalling domain during autoprocessing. The carboxy-terminal domain acts as an intramolecular cholesterol transferase. (Porter et al., 1996) hh and wg specify the identities of specific regions of the head capsule. During eye-antennal disc development hh and wg expression initially overlap, but subsequently segregate. This regional segregation is critical to head specification and is regulated by oc. oc is a candidate hh target gene during early eye-antennal disc development. (Royet and Finkelstein, 1996) In competition binding, cross-linking and co-immunoprecipitation experiments no binding of tagged hh protein to smo protein or its rat homolog could be detected, although hh protein can bind to the protein encoded by the mouse homolog of ptc. (Stone et al., 1996) en is not required for hh activation or maintenance in the eye imaginal disc. (Strutt and Mlodzik, 1996) fu protein is phosphorylated during embryogenesis as a result of hh activity. Results from cell culture studies suggest that fu and Pka-C1 function downstream of hh but in parallel pathways that eventually converge distal to fu. (Therond et al., 1996) Segment polarity gene smo is required for the response of cells to hh signalling during the development of both the embryonic segments and imaginal discs. Structure of the smo protein suggests it may act as a receptor for the hh ligand. (van den Heuvel and Ingham, 1996) 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) gro and hh regulate en expression in the anterior compartment of the wing. (de Celis and Ruiz-Gomez, 1995) hh protein acts in the wing as a signal to instruct neighbouring cells to adopt fates appropriate to the region of the wing just anterior to the compartmental boundary. Some members of the trx group genes are involved in the transcriptional regulation of genes in the hh signalling pathway during imaginal development, Pc group genes are not involved in this regulation pathway. (Felsenfeld and Kennison, 1995) Ectopic expression of the amino-terminal half of the hh protein results in effects similar to those induced by the wild-type protein, altering the identity of cells of both the dorsal and ventral ectoderm of the developing embryo and of cells of the anterior compartment of the imaginal discs. Ectopic expression of a form of the protein in which the signal cleavage sequence is mutated has no effect on larval or adult development. Results suggest that all signaling activity of the hh protein is most likely to reside in the amino terminal fragment generated by autoproteolysis. (Fietz et al., 1995) Mutations of hh interact with Dfd to reduce the viability of the Dfd3/Dfd13 combination. (Harding et al., 1995) Ectopic expression of hh produces ectopic furrows in the anterior eye disc. In addition to changes in cell shape the ectopic furrows are associated cell proliferation, cell cycle synchronisation and pattern formation, events that parallel normal furrow progression. Results propose that the morphogenetic furrow coincides with a transient boundary that coordinates growth and differentiation of the eye disc and hh is necessary and sufficient to propagate this boundary across the epithelium. (Heberlein et al., 1995) Ectopic hh causes respecification of the wing anterior compartment. Reorganisation of the anterior wing is presaged by ectopic expression of dpp and ptc. (Ingham and Fietz, 1995) Pka-C1 and hh have antagonistic effects on a common substrate which regulates transcription of dpp and wg. (Jiang and Struhl, 1995) Pka-C1 is essential during limb development to prevent inappropriate dpp and wg expression. A constitutively active form of Mmus\Pkaca, can prevent inappropriate dpp and wg expression but does not interfere with their normal induction by hh. The basal activity of Pka-C1 imposes a block on the transcription of dpp and wg and hh exerts its organizing influence by alleviating the block. (Jiang and Struhl, 1995) Pka-C1 activity is not regulated by ptc but may be regulated by hh. (Li et al., 1995) The effects of mutations in the anterior gap genes hkb, tll, oc, ems and btd on the spatial expression of hh and wg during embryogenesis have been investigated. (Mohler, 1995) fu and hh modulate the post-transcriptional regulation of ci protein. (Motzny and Holmgren, 1995) Pka-C1 is a component of the signalling pathway that represses dpp expression in the anterior compartment in appendage imaginal discs and anterior to the morphogenetic furrow in eye discs. (Pan and Rubin, 1995) hh, wg and mys are required for epithelial morphogenesis during proventriculus organ development. The morphogenetic process is suppressed by dpp. These results identify a novel cell signalling centre in the foregut that operates through a distinct genetic circuitry in the midgut to direct the formation of a multiply folded organ from a simple epithelial tube. (Pankratz and Hoch, 1995) The en-hh-ptc regulatory loop that is responsible for segmental expression of wg in the embryo is reused in imaginal disks to create a stripe of dpp expression along the A/P compartment boundary. (Sanicola et al., 1995) Both ptc and Pka-C1 act downstream of hh in the developing eye. (Strutt et al., 1995) hh pathway mutants induce ectopic morphogenetic furrows. Results show that ommatidial clusters are self-organising units whose polarity in one axis is determined by the direction of furrow progression and which can independently define the position of an equator without reference to the global coordinates of the eye disc. (Strutt and Mlodzik, 1995) en governs growth and patterning in both anterior and posterior wing compartments by controlling the expression of the hh and dpp products as well as the response of the cells to them. en activity programs wing cells to express hh whereas the absence of en activity programs them to respond to hh by expressing dpp. Consequently, posterior cells secrete hh product and induce a stripe of neighboring anterior cells across the compartment boundary to secrete dpp. dpp may exert its organizing influence by acting as a gradient morphogen in contrast to hh which appears to act principally as a short range inducer of dpp. (Zecca et al., 1995) Ectopic expression of hh can induce ectopic wg and dpp expression in anterior cells and reorganise the anterior compartment pattern. Loss of endogenous hh blocks wg and dpp expression along the compartment boundary and impedes growth and patterning in both compartments. (Basler and Struhl, 1994) Ectopic expression of hh in the anterior compartment of the wing disc causes overgrowth and pattern duplications in both anterior and posterior compartments of the wing disc, similar to alterations seen with ectopic dpp expression. These results indicate that hh is acting as a regulator of dpp expression and dpp acts as an organising molecule controlling growth and patterning in the wing imaginal disc. (Capdevila and Guerrero, 1994) The maintenance of wg expression by the hh signal is limited to early development. In a later role, hh organizes segmental pattern acting in a concentration-dependent manner, as a morphogen. (Heemskerk and DiNardo, 1994) hh mediates the interaction between anterior dpp-expressing cells and posterior en-expressing cells. (Hidalgo, 1994) Direct wg autoregulation differs from wg signalling to adjacent cells in the importance of fu, smo and ci relative to sgg and arm. Early wg autoregulation during the hh-dependent stage differs from later wg autoregulation. (Hooper, 1994) Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events. (Patel, 1994) hh gene product is involved in regulating ptc expression in both embryo and discs, through its role in regulating gene expression along the anterior-posterior compartment border. hh function establishes the proximodistal axis in discs. hh protein is secreted and can cross embryo parasegment borders and the anterior-posterior compartment border of imaginal discs to neighbouring cells that express neither en nor hh. In the embryo hh regulation of ptc apparently facilitates ptc and wg expression. In the discs hh regulation of ptc and other genes in the anterior compartment helps to establish the proximodistal axis. Cell-cell communication mediated by hh links the special properties of compartment borders with specification of the proximodistal axis in imaginal development. (Tabata and Kornberg, 1994) Wild type activity of five segment polarity genes, wg, ptc, en, nkd and hh, can account for most of the ventral pattern elements in the embryo. wg is required for naked cuticle and en is required for the first row of denticles in each abdominal denticle belt. Remaining cell types are produced by different combinations of the five gene activities. wg generates the diversity of cell types within the segment but each specific cell identity depends on the activity of ptc, en, nkd and hh. hh and en contribute to the pattern independently. hh and ptc show mutual suppression through opposing effects on wg expression. hh alters the competence of cells to respond to wg signal. (Bejsovec and Wieschaus, 1993) Transcriptional control of both ptc and wg by hh is mediated by the same signal transduction pathway. (Forbes et al., 1993) Developing retinal cells drive the progression of morphogenesis using the products of the hh and dpp genes. Clonal analysis suggests that gene products act as diffusible signals. hh induces the expression of dpp, the primary mediator of furrow movement. (Heberlein et al., 1993) Segment polarity mutations cause stripes of abnormal patterning within sectors of the leg disc, which may be mediated by regional perturbations in growth. (Held, 1993) The ptc and hh genes encode components of a signal transduction pathway that regulate the expression of wg transcription following its activation by pair rule genes, but most other aspects of wg expression are independent of ptc and hh. Maintenance of wg expression depends upon the activity of hh, which acts only on neighboring cells to maintain wg expression. Expression of wg in the absence of ptc depends on hh. (Ingham and Hidalgo, 1993) Competence of cells to express wg is independent of their ability to receive the hh signal. wg activation requires the function of fu, this suggests that the putative hh signal is transduced by the serine/threonine kinase that fu encodes. (Ingham, 1993) The role of hh in the regulation of run mRNA expression in the early embryo has been investigated. (Klingler and Gergen, 1993) A hh-related gene family has been identified in the zebrafish. Over-expression of one member, sonic hedgehog, in fly embryos, can activate the hh-dependent pathway. (Krauss et al., 1993) Many alleles of hh act as dominant alleles of gl. (Ma et al., 1993) hh expression posterior to the morphogenetic furrow in the developing eye disc is continuously required for its progression. The forward diffusion of hh protein induces anterior cells to enter the furrow. hh acts upstream of gl, sca, h and dpp in the developing eye. (Ma et al., 1993) Although hh is essential for wg function in segmentation, wg appears to be still capable of some action in hh's absence. (Sampedro et al., 1993) Probably encodes a secreted or transmembrane protein. (Tashiro et al., 1993) The pattern of hh protein expression during embryonic development has been analysed. (Taylor 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) hh has been isolated and characterised. (Lee et al., 1992) hh gene cloned by plasmid rescue, the encoded protein is targetted to the secretory pathway (consistant with the non-cell autonomous requirement for hedgehog in cuticular patterning) and is expressed coincidentally with engrailed in embryos and imaginal discs. Maintainance of hh expression is dependent upon other segment polarity genes including engrailed and wingless. The amino acid sequence shows no similarity to any known protein but hydropathy analysis highlights a prominent hydrophobic region. (Lee et al., 1992) Sequence analysis of hh indicates that the gene product contains a putative transmembrane domain which suggests that it may be localized at the cell surface and be involved in cell-cell communication. (Mohler and Vani, 1992) hh gene cloned and the sequence suggests a membrane associated protein. Expression pattern analysed and found to coincide with that of en in the epidermis. Though initially independent of en, hh expression later becomes en-dependent. (Tabata et al., 1992) hh cannot completely rescue the ptc phenotype when in double mutant combinations. (Hidalgo, 1991) The role of ptc in positional signalling is permissive rather than instructive, its activity is required to suppress wg transcription in cells predisposed to express wg. These cells receive an extrinsic signal, encoded by hh, that antagonises the repressive activity of ptc. Results suggest that ptc protein may be the receptor for the hh signal, implying that this is an usual mechanism of ligand-dependent receptor inactivation. (Ingham et al., 1991) hh is essential for maintaining the normal pattern of ptc expression. (Hidalgo and Ingham, 1990) Role of hh in neurogenesis has been studied. (Patel et al., 1989) Adult eyes are small, narrow with about 150 facets, eye disc is small due to precursor cell defects. (Renfranz and Benzer, 1989) The mutational effects of hh on larval and adult cuticular patterns has been studied. Defects in the distal portions of the legs and antenna occur in association with homozygous hh clones in the posterior compartments of the structures. (Mohler, 1988) Genetic mosaics were used to determine that hh is not autonomous at the level of the single cell. (Wieschaus and Riggleman, 1987) hh mutants display segment polarity segmentation defects. (Jurgens et al., 1984) A segment polarity type of embryonic lethal. Homozygous embryos have the posterior naked portion of the ventral surface of each segment deleted and replaced by a mirror image of the anterior denticle belts. Embryos appear to lack segmental boundaries. In strong alleles, there is no obvious segmentation; the larvae are approximately 40% the length of the wild-type larvae and there is a lawn of denticles arranged in a number of whorls on the ventral surface as a result of loss of naked cuticle. In intermediate alleles, naked cuticle is also lost from the ventral region, but the lawn of denticles is arranged in segmental arrays in mirror-image symmetry. The weak alleles show fusions that delete the naked cuticle usually between abdominal segments 1 and 2 and 6, 7 and 8 (Mohler, 1988). Temperature shift experiments with a temperature-sensitive allele (viable and normal at 18oC, and mutant at 25oC) indicate two phases of hh activity at 25oC, the first during early embryogenesis (3-6 hr of development) and the second during the late larval and early pupal stages (4-7 days of development).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_079735 NM_001038976 RefSeq (Proteins) • NP_524459 NP_001034065 DDBJ / EMBL / GenBank • AAA16458 AAA28604 AAB28646 AAF56102 AAF56102.1 ABC66186 AQ026358 AQ034105 AQ034132 AQ254728 AX146557 AX282805 AX354872 AX417094 CAD10621 CAD22996 CAD35282 CAJ18813 CS126494 L02793 L05404 L05405 S66384 Z11840 Z83558 Entrez Gene - A searchable database of RefSeq genes. • 42737 UniProtKB/Swiss-Prot • Q02936
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Hedgehog, N-terminal signaling domain (IPR000320) Peptidase C46, hedgehog protein (IPR001657) Peptidase C46, hedgehog protein, hint region (IPR001767) Hedgehog/intein hint domain, C-terminal (IPR003586) Hedgehog/intein hint, N-terminal (IPR003587) Intein splice site (IPR006141) Hedgehog/DD-peptidase (IPR009045) MEROPS - An information resource for peptidases and the proteins that inhibit them • C46.001 PDB - Protein Data Bank. An information portal to biological macromolecular structures • 1AT0
Linkouts
	BioGRID - A database of protein and genetic interactions • 67682 DroID - A comprehensive database of gene and protein interactions. • FBgn0004644 DRSC - Results from RNAi screens. • FBgn0004644 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004644 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG4637-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0004644 FlyReactome - A curated knowledgebase of Drosophila melanogaster pathways • 92192 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 42737 InParanoid A subset of ortholog calls from InParanoid. • AAEL006708-PA Aedes aegypti (mosquito) AGAP001412-PA Anopheles gambiae (mosquito) XP_001943707.1 Acyrthosiphon pisum (pea aphid) 84974 Branchiostoma floridae (lancelet) BGIBMGA012535-PA Bombyx mori (silk worm) ENSBTAP00000000135 Bos taurus (bovine) ENSCAFP00000012759 Canis familiaris (dog) ENSCINP00000013864 Ciona intestinalis (sea squirt) CPIJ003906 Culex pipens (mosquito) ENSCPOP00000002048 Cavia porcellus (guinea pig) RP19440 Caenorhabditis remanei (nematode) ENSCSAVP00000017051 Ciona savignyi (sedentary tunicate) 129621 Capitella spl (polychaete worm) 290571 Daphnia pulex (daphnia) ENSDARP00000056746 Danio rerio (zebra fish) ENSECAP00000008540 Equus caballus (horse) ENSGACP00000005121 Gymnopilus aculeatus (mushroom) ENSGALP00000010292 Gallus gallus (chicken) 154346 Hypsipyla robusta (cedar shoot borer) ENSP00000266991 Homo sapiens (human) 121665 Lottia gigantea (owl limpet) ENSMODP00000019154 Monodelphis domestica (gray opossum) ENSMMUP00000001060 Macaca mulatta (rhesus monkey) ENSMUSP00000023737 Mus musculus (house mouse) 241466 Nematostella vectensis (starlet anemone) XP_001605475.1 Nasonia vitripennis (wasp) ENSOANP00000000534 Ornithorynchus anatinus (platypus) ENSORLP00000013115 Oryzias latipes (japanese killifish) PHUM002563-PA Pediculus humanus (human body louse) ENSPPYP00000005114 Pongo pygmaeus (orangutan) ENSPTRP00000022106 Pan troglodytes (chimpanzee) ENSRNOP00000020563 Rattus norvegicus (norway rat) NP_001012720.1 Strongylocentrotus purpuratus (purple sea urchin) NP_001107837.1 Tribolium castaneum (red flour beetle) ENSTNIP00000015753 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000045862 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000024437 Xenopus tropicalis (frog) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /segment/hedghog1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 42737 modMine - Data generated by the modENCODE project. • FBgn0004644 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB13786 NV16490 LH13486 ISCW009092 ISCW007045 FBgn0241122 FBgn0214842 FBgn0209745 FBgn0189928 FBgn0178454 FBgn0250482 JGI_V11_290571 FBgn0078326 FBgn0161188 FBgn0147295 FBgn0131309 FBgn0104746 FBgn0095047 CPIJ003906 BGIBMGA012535 ACYPI27225 ACYPI006525 GB30509 GB14574 AGAP001412 AAEL006708 PHUM018090 GLEAN_01364 REDfly - A database of transcriptional regulatory elements. • FBgn0004644
Synonyms & Secondary IDs ( 21 )
Reported As
Symbol Synonym	anon-WO0134654.19   anon-WO0182946.19 (Reilly, 2002.2.6) bar (Gonzalez et al., 1989) bar-3 (Lloyd et al., 1997, Renfranz and Benzer, 1989, ) bar3 (Atkinson et al., 1991) CG4637 (Molnar et al., 2006, Perkins et al., 1.15.2009, Ni et al., 2008, Ni et al., 2010.12.1, Keleman et al., 2009.8.5) hg (Fini et al., 1997) hh (Casso et al., 2008, Peel et al., 2005, Chanut-Delalande et al., 2006, Ramos and Mohler, 2006, Yasunaga et al., 2006, Chanas and Maschat, 2005, Ishii, 2005, Briscoe and Therond, 2005, Gorfinkiel et al., 2005, Jones et al., 2006, Lu et al., 2006, Lindner et al., 2007, Casso et al., 2007, Bejarano et al., 2007, Bras-Pereira and Casares, 2007, Smelkinson and Kalderon, 2006, Fraser, 2006, Umetsu et al., 2006, Chotard et al., 2005, Nystul and Spradling, 2006, Freeland and Kuhn, 1996, Chien-Hsiang et al., 2007, Wheeler et al., 2006, Rogers et al., 2005, Kent et al., 2006, Yao et al., 2006, Mahoney et al., 2006, Wang and Struhl, 2004, Roederer et al., 2005, de Velasco et al., 2006, Ntini E and Wimmer, 2007, Pichaud et al., 2007, Mandal et al., 2007, Sandmann et al., 2007, Tountas and Fortini, 2007, Pfleger et al., 2007, Friedrich, 2006, Cheesman et al., 2004, Bras-Pereira et al., 2006, D'Costa et al., 2006, Williams et al., 2010, Akimoto et al., 2005, Besse et al., 2005, Colosimo and Tolwinski, 2006, Chanana et al., 2007, Deshpande and Schedl, 2005, Molnar et al., 2011, Yuva-Aydemir et al., 2011, Martin-Lanneree et al., 2006, Deshpande and Schedl, 2005, Song et al., 2007, Deshpande et al., 2007, Casso et al., 2008, Chen et al., 2008, Maeda et al., 2007, Molnar et al., 2006, Seong et al., 2010, Torroja et al., 2005, Liu et al., 2007, Christensen et al., 2008.9.29, Christensen et al., 2008.9.29, Birdsall et al., 2000, Wang and Price, 2008, Callejo et al., 2006, Chu et al., 2006, Smelkinson et al., 2007, Fan and Bergmann, 2008, Gallet et al., 2008, Takashima et al., 2008, McLellan et al., 2008, Bornemann et al., 2008, Ou et al., 2007, Sakurai et al., 2007, Lechner et al., 2007, Price et al., 2006, Farzan et al., 2008, Walthall et al., 2007, Melicharek et al., 2008, Schuettengruber et al., 2009, Vincent et al., 2008, Larsen et al., 2008, Ogden et al., 2008, Jia et al., 2009, Cheng et al., 2010, Monnier et al., 2002, Dawber et al., 2005, Lindner et al., 2007, Foronda et al., 2009, Levine et al., 1997, Xie et al., 2005, Bejarano and Milán, 2009, Hallson et al., 2008, Christensen et al., 2009.5.6, Vrailas and Moses, 2006, Blanco et al., 2009, Renault et al., 2009, Vied and Kalderon, 2009, Bejarano et al., 2007, Lim et al., 2008, Orme and Leevers, 2005, Khaliullina et al., 2009, Baker et al., 2009, de Velasco et al., 2007, Glise et al., 2005, Zhang et al., 2005, Bantignies et al., 2011, Mulinari and Häcker, 2009, Gazi et al., 2009, Zhao et al., 2008, Wang and Huang, 2009, Sato et al., 2008, Schlichting and Dahmann, 2008, Brás-Pereira and Casares, 2008, Landsberg et al., 2009, Venken et al., 2009, Eivers et al., 2009, Wang et al., 2008, Nahmad and Stathopoulos, 2009, Guichard et al., 2006, DasGupta et al., 2007, Magalhaes et al., 2007, Gutierrez-Aviño et al., 2009, Glazov et al., 2005, Klein et al., 2010, Wang et al., 2010, Lopes and Casares, 2010, Irons et al., 2010, Maurel-Zaffran et al., 2010, Zheng et al., 2010, Salzer and Kumar, 2010, Baig et al., 2010, Molnar et al., 2007, Tokhunts et al., 2010, Terriente-Félix et al., 2010, Yavari et al., 2010, Bergantiños et al., 2010, Dilks and DiNardo, 2010, Schwartz et al., 2010, Pospisilik et al., 2010, Liu et al., 2010, Beltran et al., 2007, Subramanian and Gadgil, 2010, Suh et al., 2006, Terriente-Félix et al., 2011, Watson et al., 2011, Schilling et al., 2011, Chang et al., 2010, Ayers et al., 2010) Hh (Friggi-Grelin et al., 2008, Ma, 2005, Holmgren et al., 2005, Gonzalez-Gaitan, 2003, Ma and Beachy, 2002, Lee and Treisman, 2002, Merabet et al., 2001, Selleck et al., 2000, Zhang and Kalderon, 2000, Ziegenhorn et al., 2006, Sisson et al., 2006, Jia and Jiang, 2006, Chanana et al., 2006, Osterlund and Kogerman, 2006, Goodman et al., 2006, Zhou et al., 2006, Wang et al., 2007, Sun and Deng, 2007, Farzan et al., 2007, Vied and Kalderon, 2007, Firth and Baker, 2007, Bejarano et al., 2007, Plessis et al., 2007, Callejo et al., 2007, Smelkinson and Kalderon, 2006, Horabin, 2005, Lander, 2007, Liu et al., 2006, Croker et al., 2006, Joshi et al., 2006, Kirkbride, 2005, Chien-Hsiang et al., 2007, Fisher and Howie, 2006, McLellan et al., 2006, Beenken and Mohammadi, 2006, Ueyama et al., 2008, Gallet et al., 2006, Giuliani et al., 2007, Wojcinski et al., 2007, Molnar et al., 2007, Gallet et al., 2007, Aikin et al., 2007, Scholler Joulie et al., 2007, Kugler and Nagel, 2007, Bangi et al., 2007, Ogden et al., 2006, Zhang et al., 2006, Voas and Rebay, 2004, Nybakken et al., 2005, Eldar and Barkai, 2005, Maricich and Zoghbi, 2006, Finan et al., 2011, Perrimon and Mathey-Prevot, 2007, Wilson and Chuang, 2006, Wendler et al., 2006, Fan and Bergmann, 2008, Bornemann et al., 2008, Wojcinski et al., 2008, Liu et al., 2008, Zhao and Jiang, 2008, Williams et al., 2008, Vied and Kalderon, 2008, Su et al., 2008, De Rivoyre et al., 2006, Linder and Deschenes, 2004, Zhao et al., 2007, Bovolenta and Marti, 2005, Mehlen et al., 2005, Reig et al., 2007, Firth and Baker, 2005, Katanaev et al., 2008, Eugster et al., 2007, Umemori et al., 2007, Callejo et al., 2008, Escudero and Freeman, 2007, Su et al., 2007, Benítez et al., 2009, Franch-Marro et al., 2008, Makhijani et al., 2007, Wang et al., 2000, Sisson et al., 2006, Gazi et al., 2009, Gim et al., 2001, Li et al., 2010, González et al., 2008, Raisin et al., 2010, Yan et al., 2010, Farzan et al., 2009, Wang et al., 2011, Silver et al., 2007, Chou et al., 2010, Zhou and Kalderon, 2010, Jia et al., 2010, Wang et al., 2011) HH (Roy and Ingham, 2002, Scadden, 2006, Sanial and Plessis, 2007, Plessis et al., 2007, Legent et al., 2008, Zhao et al., 2007, Claret et al., 2007) l(3)hh (Perrimon and Mahowald, 1987) l(3)neo56 (BDGP Project Members, 1994-1999, Cooley et al., 1988, ) l(3)neo57   Mir   Mrt (Cifuentes and Garcia-Bellido, 1997, )
Name Synonym	bar-on-3   hedgehog (Gellon et al., 1997, Mulinari et al., 2007, Bras-Pereira and Casares, 2007, Fraser, 2006, Lander, 2007, Kruger, 2005, Thomas, 2005, Nystul and Spradling, 2006, Wheeler et al., 2006, Rogers et al., 2005, Roederer et al., 2005, Thompson et al., 2007, Tountas and Fortini, 2007, de Velasco et al., 2006, Williams et al., 2010, Akimoto et al., 2005, Besse et al., 2005, Chanana et al., 2007, Finan et al., 2011, Song et al., 2007, Muller and Kassis, 2006, Mulinari et al., 2008, Vrailas and Moses, 2006, Maeda et al., 2007, Gonzalez-Reyes, 2003, Hashimoto and Yamaguchi, 2006, Hime et al., 2004, Birdsall et al., 2000, Takashima et al., 2008, McLellan et al., 2008, Ou et al., 2007, Lechner et al., 2007, Price et al., 2006, Melicharek et al., 2008, Dansereau and Lasko, 2008, Jia et al., 2009, Cheng et al., 2010, Bejarano et al., 2007, Orme and Leevers, 2005, Mulinari and Häcker, 2009, Schlichting and Dahmann, 2008, Eivers et al., 2009, Martinez et al., 2009, DasGupta et al., 2007, Gutierrez-Aviño et al., 2009, Glazov et al., 2005, Lopes and Casares, 2010, Salzer and Kumar, 2010, Baig et al., 2010, Yavari et al., 2010, Dilks and DiNardo, 2010, Schwartz et al., 2010, Liu et al., 2010, Terriente-Félix et al., 2011) Hedgehog (Schwartz and Pirrotta, 2007, Coudreuse and Korswagen, 2007, Goodman et al., 2006, Payre et al., 2007, Mandal et al., 2007, Bulanin and Orenic, 2007, Sanial and Plessis, 2007, Lindner et al., 2007, Dahmann and Schlichting, 2007, Bejarano et al., 2007, Plessis et al., 2007, Plessis et al., 2007, Beach et al., 2006, Panakova and Eaton, 2006, Vrailas et al., 2006, Horabin, 2005, Maurange et al., 2006, Chien-Hsiang et al., 2007, Bossing and Brand, 2006, McLellan et al., 2006, Gallet et al., 2006, Schlichting and Dahmann, 2007, Giuliani et al., 2007, Wojcinski et al., 2007, Molnar et al., 2007, Escudero et al., 2007, Ogden et al., 2006, Niki et al., 2006, Friedrich, 2006, Colosimo and Tolwinski, 2006, Pocha et al., 2011, Voas and Rebay, 2004, Molnar et al., 2011, Arbouzova and Zeidler, 2006, Casso et al., 2008, Fan and Bergmann, 2008, Bornemann et al., 2008, Lorigan et al., 2008, Liu et al., 2008, Legent et al., 2008, Zhao and Jiang, 2008, Williams et al., 2008, Ming et al., 2008, Vied and Kalderon, 2008, Lawrence, 2004, Linder and Deschenes, 2004, Strigini, 2005, Torroja et al., 2005, Zhao et al., 2007, Reig et al., 2007, Kerszberg and Wolpert, 2007, Liu et al., 2007, Vied et al., 2007, Chu et al., 2006, Eugster et al., 2007, Gallet et al., 2008, Claret et al., 2007, Pallavi and Shashidhara, 2005, Franch-Marro et al., 2008, Farzan et al., 2008, Walthall et al., 2007, Monnier et al., 2002, Dawber et al., 2005, Escudero and Freeman, 2007, Su et al., 2007, Buchon et al., 2009, Renault et al., 2009, Lim et al., 2008, Khaliullina et al., 2009, Baker et al., 2009, Tolhuis et al., 2006, Sisson et al., 2006, Gazi et al., 2009, Gim et al., 2001, Irons et al., 2010, Maurel-Zaffran et al., 2010, Zheng et al., 2010, Terriente-Félix et al., 2010, Raisin et al., 2010, Yan et al., 2010, Farzan et al., 2009, Schilling et al., 2011, Chou et al., 2010, Zhou and Kalderon, 2010, Jia et al., 2010, Chang et al., 2010, Wang et al., 2011) Hedghog (Landsberg et al., 2009) Mirabile   Moonrat
Secondary FlyBase IDs
	FBgn0000159 FBgn0001191 FBgn0002748 FBgn0002793 FBgn0011486 FBgn0011487 FBgn0044801
References ( 1521 )
Generate a list of	All references relating to this gene ( 1521 )
List References by type	Research paper  ( 652 ) Supplementary material  ( 6 ) Review  ( 408 ) Erratum  ( 3 ) Personal communication to FlyBase  ( 14 ) Abstract  ( 391 ) FlyBase analysis  ( 2 ) DNA/RNA sequence record  ( 3 ) Computer file  ( 3 ) Meeting report  ( 1 ) Conference report  ( 11 ) Stock list  ( 2 ) Patent  ( 4 ) Letter  ( 12 ) Curated genome annotation  ( 1 ) Protein sequence record  ( 1 ) Automatic genome annotation  ( 1 ) Interview  ( 2 ) Book  ( 1 ) Biography  ( 1 ) Book review  ( 1 ) Poem  ( 1 )
Recent research papers ( 49 )
	Babcock et al., 2011, Curr. Biol. 21(18): 1525--1533 Hedgehog signaling regulates nociceptive sensitization. [FBrf0216292] Bantignies et al., 2011, Cell 144(2): 214--226 Polycomb-Dependent Regulatory Contacts between Distant Hox Loci in Drosophila. [FBrf0212775] Casso et al., 2011, Genetics 187(2): 485--499 A Novel Interaction Between hedgehog and Notch Promotes Proliferation at the Anterior-Posterior Organizer of the Drosophila Wing. [FBrf0212986] Eliazer et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(17): 7064--7069 Loss of lysine-specific demethylase 1 nonautonomously causes stem cell tumors in the Drosophila ovary. [FBrf0213593] Finan et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(14): 5566--5571 From the Cover: Proteomics approach to study the functions of Drosophila myosin VI through identification of multiple cargo-binding proteins. [FBrf0213355] Gabilondo et al., 2011, Mech. Dev. 128(3-4): 208--221 A targeted genetic screen identifies crucial players in the specification of the Drosophila abdominal Capaergic neurons. [FBrf0213290] Molnar et al., 2011, PLoS Genet. 7(3): e1001335 Role of the Drosophila non-visual ß-arrestin kurtz in hedgehog signalling. [FBrf0213301] Nahmad, 2011, J. R. Soc. Interface 8(63): 1429--1439 Steady-state invariant genetics: probing the role of morphogen gradient dynamics in developmental patterning. [FBrf0214797] Parker et al., 2011, Sci. Signal. 4(176): ra38 The cis-Regulatory Logic of Hedgehog Gradient Responses: Key Roles for Gli Binding Affinity, Competition, and Cooperativity. [FBrf0213871] Pocha et al., 2011, Curr. Biol. 21(13): 1111--1117 Retromer controls epithelial cell polarity by trafficking the apical determinant crumbs. [FBrf0214250] Schilling et al., 2011, PLoS Comput. Biol. 7(4): e1002025 Cell-sorting at the a/p boundary in the Drosophila wing primordium: a computational model to consolidate observed non-local effects of hh signaling. [FBrf0213439] Swarup and Verheyen, 2011, Proc. Natl. Acad. Sci. U.S.A. 108(24): 9887--9892 Drosophila homeodomain-interacting protein kinase inhibits the Skp1-Cul1-F-box E3 ligase complex to dually promote Wingless and Hedgehog signaling. [FBrf0214241] Terriente-Félix et al., 2011, Dev. Biol. 350(2): 382--392 A conserved function of the chromatin ATPase Kismet in the regulation of hedgehog expression. [FBrf0212899] Wang et al., 2011, Infect. Immun. 79(2): 606--616 Host and Pathogen Glycosaminoglycan-Binding Proteins Modulate Antimicrobial Peptide Responses in Drosophila melanogaster. [FBrf0212806] Wang et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(27): 11139--11144 Sexually dimorphic regulation of the Wingless morphogen controls sex-specific segment number in Drosophila. [FBrf0214239] Watson et al., 2011, Development 138(7): 1285--1295 Drosophila hedgehog signaling and engrailed-runt mutual repression direct midline glia to alternative ensheathing and non-ensheathing fates. [FBrf0213237] Wojcinski et al., 2011, Dev. Biol. 358(1): 168--180 DSulfatase-1 fine-tunes Hedgehog patterning activity through a novel regulatory feedback loop. [FBrf0215253] Xie et al., 2011, Genetics 188(4): 823--834 Inhibition of RNA interference and modulation of transposable element expression by cell death in Drosophila. [FBrf0214636] Yorimitsu et al., 2011, Dev. Biol. 356(2): 598--607 Defective proventriculus specifies the ocellar region in the Drosophila head. [FBrf0214458] Yuva-Aydemir et al., 2011, J. Neurosci. 31(19): 7005--7015 Spinster Controls Dpp Signaling during Glial Migration in the Drosophila Eye. [FBrf0213705] Ayers et al., 2010, Dev. Cell 18(4): 605--620 The long-range activity of Hedgehog is regulated in the apical extracellular space by the glypican Dally and the hydrolase Notum. [FBrf0214999] Baig et al., 2010, Genetics 184(3): 731--744 The chromatin-remodeling protein Osa interacts with CyclinE in Drosophila eye imaginal discs. [FBrf0210399] Bergantiños et al., 2010, Development 137(7): 1169--1179 Cell death-induced regeneration in wing imaginal discs requires JNK signalling. [FBrf0210204] Chang et al., 2010, Brain Res. 1324: 1--13 Hedgehog-dependent down-regulation of the tumor suppressor, vitamin D3 up-regulated protein 1 (VDUP1), precedes lamina development in Drosophila. [FBrf0210439] Cheng et al., 2010, Dev. Biol. 337(1): 99--109 Regulation of smoothened by Drosophila G-protein-coupled receptor kinases. [FBrf0209432] Chou et al., 2010, Cell 142(6): 954--966 Patterning axon targeting of olfactory receptor neurons by coupled hedgehog signaling at two distinct steps. [FBrf0211838] Dilks and DiNardo, 2010, Development 137(8): 1395--1404 Non-cell-autonomous control of denticle diversity in the Drosophila embryo. [FBrf0210398] Fabre et al., 2010, Development 137(17): 2885--2894 Mechanosensilla in the adult abdomen of Drosophila: engrailed and slit help to corral the peripheral sensory axons into segmental bundles. [FBrf0211534] Irons et al., 2010, Dev. Biol. 342(2): 180--193 Robustness of positional specification by the Hedgehog morphogen gradient. [FBrf0210817] Jia et al., 2010, J. Biol. Chem. 285(48): 37218--37226 Casein kinase 2 promotes Hedgehog signaling by regulating both smoothened and Cubitus interruptus. [FBrf0214177] Klein et al., 2010, Dev. Biol. 337(2): 458--470 The proprioceptive and contractile systems in Drosophila are both patterned by the EGR family transcription factor Stripe. [FBrf0209777] Li et al., 2010, Genes Dev. 24(9): 933--946 Polycomb group genes Psc and Su(z)2 restrict follicle stem cell self-renewal and extrusion by controlling canonical and noncanonical Wnt signaling. [FBrf0210675] Liu et al., 2010, Sci. Signal. 3(132): ra57 The Drosophila Female Germline Stem Cell Lineage Acts to Spatially Restrict DPP Function Within the Niche. [FBrf0211426] Lopes and Casares, 2010, Dev. Biol. 339(1): 78--88 hth maintains the pool of eye progenitors and its downregulation by Dpp and Hh couples retinal fate acquisition with cell cycle exit. [FBrf0210055] Maurel-Zaffran et al., 2010, Dev. Biol. 343(1-2): 18--27 Reiterative use of signalling pathways controls multiple cellular events during Drosophila posterior spiracle organogenesis. [FBrf0211024] Pospisilik et al., 2010, Cell 140(1): 148--160 Drosophila genome-wide obesity screen reveals hedgehog as a determinant of brown versus white adipose cell fate. [FBrf0212676] 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] Salzer and Kumar, 2010, PLoS ONE 5(1): e8510 Identification of retinal transformation hot spots in developing Drosophila epithelia. [FBrf0209699] Schwartz et al., 2010, PLoS Genet. 6(1): e1000805 Alternative epigenetic chromatin states of polycomb target genes. [FBrf0209708] Seong et al., 2010, PLoS ONE 5(12): e15365 Inhibition of the nuclear import of cubitus interruptus by roadkill in the presence of strong hedgehog signal. [FBrf0212658] Subramanian and Gadgil, 2010, IET Syst. Biol. 4(2): 169 Robustness of the Drosophila segment polarity network to transient perturbations. [FBrf0210287] Terriente-Félix et al., 2010, Genetics 185(2): 671--684 Identification of genes affecting wing patterning through a loss-of-function mutagenesis screen and characterization of med15 function during wing development. [FBrf0211230] Tokhunts et al., 2010, J. Biol. Chem. 285(4): 2562--2568 The full-length unprocessed hedgehog protein is an active signaling molecule. [FBrf0209740] Wang et al., 2010, Dev. Biol. 337(2): 246--258 The role of eyg Pax gene in the development of the head vertex in Drosophila. [FBrf0209801] Williams et al., 2010, Proc. Natl. Acad. Sci. U.S.A. 107(13): 5869--5874 Dally-like core protein and its mammalian homologues mediate stimulatory and inhibitory effects on Hedgehog signal response. [FBrf0210451] Yan et al., 2010, Development 137(12): 2033--2044 The cell-surface proteins Dally-like and Ihog differentially regulate Hedgehog signaling strength and range during development. [FBrf0210906] Yavari et al., 2010, Dev. Cell 19(1): 54--65 Role of Lipid Metabolism in Smoothened Derepression in Hedgehog Signaling. [FBrf0211286] Zheng et al., 2010, Genes Dev. 24(1): 57--71 Genetic and biochemical definition of the Hedgehog receptor. [FBrf0209607] 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 ...
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