FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\shd
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General Information
Symbol
Dmel\shd
Species
D. melanogaster
Name
shade
Annotation Symbol
CG13478
Feature Type
protein_coding_gene
FlyBase ID
FBgn0003388
Gene Model Status
Current
Stock Availability
9 publicly available
Enzyme Name (EC)
ecdysone 20-monooxygenase (1.14.99.22)
Gene Summary
shade (shd) encodes 20-hydroxylase and is responsible for converting Ecdysone into 20-hydroxyecdysone, the active form of the steroid. It is required in all tissues that produce active Ecdysone and thus contributes to larval moulting, metamorphosis, growth, neuroblast diversity and egg chamber maturation. [Date last reviewed: 2018-09-13] (FlyBase Gene Snapshot)
All Summaries
Also Known As

Cyp314a1

Key Links
Genomic Location
Cytogenetic map
70E4-70E4
Sequence location
Recombination map
3-42
RefSeq locus
NT_037436 REGION:14614458..14622205
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (14 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables ecdysone 20-monooxygenase activity
inferred from direct assay
(Petryk et al., 2003)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
enables heme binding
inferred from electronic annotation with InterPro:IPR001128, InterPro:IPR002401, InterPro:IPR036396
(InterPro Project Members, 2004-)
enables iron ion binding
(InterPro Project Members, 2004-)
enables monooxygenase activity
inferred from electronic annotation with InterPro:IPR001128, InterPro:IPR002401, InterPro:IPR036396
(InterPro Project Members, 2004-)
enables oxidoreductase activity, acting on paired donors, with incorporation or reduction of molecular oxygen
(InterPro Project Members, 2004-)
Biological Process (8 terms)
Terms Based on Experimental Evidence (8 terms)
CV Term
Evidence
References
involved_in border follicle cell migration
inferred from mutant phenotype
(Domanitskaya et al., 2014)
involved_in central nervous system development
inferred from mutant phenotype
(Giesen et al., 2003)
involved_in dorsal closure
inferred from mutant phenotype
(Giesen et al., 2003, Petryk et al., 2003)
involved_in ecdysteroid biosynthetic process
inferred from direct assay
(Petryk et al., 2003)
inferred from mutant phenotype
(Petryk et al., 2003)
involved_in growth of a germarium-derived egg chamber
inferred from mutant phenotype
(Petryk et al., 2003)
involved_in head involution
inferred from mutant phenotype
(Giesen et al., 2003)
involved_in midgut development
inferred from mutant phenotype
(Petryk et al., 2003)
involved_in pupation
inferred from mutant phenotype
(Akagi et al., 2016)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in mitochondrion
inferred from direct assay
(Petryk et al., 2003)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the cytochrome P450 family. (Q9VUF8)
Protein Signatures (InterPro)
Catalytic Activity (EC/Rhea)
ecdysone 20-monooxygenase activity
ecdysone + AH2 + O2 = 20-hydroxyecdysone + A + H2O (1.14.99.22)
RHEA 14021:
Summaries
Gene Snapshot
shade (shd) encodes 20-hydroxylase and is responsible for converting Ecdysone into 20-hydroxyecdysone, the active form of the steroid. It is required in all tissues that produce active Ecdysone and thus contributes to larval moulting, metamorphosis, growth, neuroblast diversity and egg chamber maturation. [Date last reviewed: 2018-09-13]
Gene Group (FlyBase)
CYTOCHROME P450 - MITO CLAN -
Cytochrome P450, or CYP genes, constitute one of the largest families of genes, with representatives in virtually all living organisms. P450 enzymes are heme-thiolate proteins best known for their monooxygenase role, catalyzing the transfer of one atom of molecular oxygen to a substrate and reducing the other to water. The MITO clan includes D. melanogaster CYP12, 49, 301, 302, 314 and 315 family genes. Members are localized to mitochondria and at least some are linked to the ecdysteroid metabolism pathway. (Adapted from FBrf0248421.)
Protein Function (UniProtKB)
Required for CNS development; midline glial cells. Involved in the metabolism of insect hormones; responsible for all ecdysone 20-monooxygenase activity during embryonic, larval and adult stages. May be involved in the breakdown of synthetic insecticides.
(UniProt, Q9VUF8)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
shd: shade
Embryonic lethal. No differentiation of cuticle or head skeleton.
Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\shd for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9VUF8)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

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

Gene model reviewed during 5.46

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
shd-RB
FBtr0075676
NM_206352
2730
540
shd-RC
FBtr0304700
NM_140452
1633
433
shd-RD
FBtr0333031
NM_001274914
2985
540
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
shd-PB
FBpp0089324
61.7
540
7.09
Q9VUF8
NP_996074
AAS65010
shd-PC
FBpp0293243
49.0
433
5.02
NP_648709
AAF49727
shd-PD
FBpp0305245
61.7
540
7.09
Q9VUF8
NP_001261843
AGB94536
Polypeptides with Identical Sequences

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

540 aa isoforms: shd-PB, shd-PD
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\shd using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.19

Transcript Expression
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in mitochondrion
inferred from direct assay
(Petryk et al., 2003)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\shd in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 11-12
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 7 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 7 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of shd
Transgenic constructs containing regulatory region of shd
Aberrations (Deficiencies and Duplications) ( 6 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal - all die during embryonic stage
lethal - all die during embryonic stage | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4twi.2PE, shd1/shd2
viable (with shd1), with Scer\GAL4twi.2PE, shdUAS.cPa
viable (with shd2), with Scer\GAL4twi.2PE, shdUAS.cPa
Sterility
Allele
female semi-fertile, with Scer\GAL4GMR47A04
female sterile, with Scer\GAL4twi.2PE, shd1/shd2
female sterile (with shd1), with Scer\GAL4twi.2PE, shdUAS.cPa
female sterile (with shd2), with Scer\GAL4twi.2PE, shdUAS.cPa
Other Phenotypes
Allele
abnormal cell number, with Scer\GAL4phtm.PO, Scer\GAL80spok.1.45, Spyo\Cas9UAS.Tag:NLS(Ude)
abnormal developmental rate | prepupal stage, with Scer\GAL4C855a
abnormal developmental rate | prepupal stage, with Scer\GAL4Cg.PA
abnormal developmental rate | prepupal stage, with Scer\GAL4elav.PLu
abnormal developmental rate | prepupal stage, with Scer\GAL4how-24B
abnormal developmental rate | prepupal stage, with Scer\GAL4Mef2.PR
decreased cell growth rate | adult stage | male, with Scer\GAL4esg.PU
decreased cell number | adult stage, with Scer\GAL4Tk.PS
some die during embryonic stage
some die during embryonic stage | recessive
Phenotype manifest in
Allele
adult anterior midgut, with Scer\GAL4Tk.PS
adult enteroendocrine cell | decreased number, with Scer\GAL4Tk.PS
border follicle cell | somatic clone
cephalopharyngeal skeleton
egg chamber, with Scer\GAL4twi.2PE, shd1/shd2
egg chamber (with shd1), with Scer\GAL4twi.2PE, shdUAS.cPa
egg chamber (with shd2), with Scer\GAL4twi.2PE, shdUAS.cPa
embryo | dorsal closure stage
embryo | embryonic stage 14
embryonic/first instar larval cuticle
embryonic/larval cuticle
embryonic/larval foregut
embryonic/larval hindgut
embryonic/larval midgut
embryonic/larval midgut primordium
embryonic dorsal epidermis
embryonic epidermis
embryonic head
embryonic midgut constriction
embryonic ventral epidermis
enteroendocrine cell | decreased number, with Scer\GAL4phtm.PO, Scer\GAL80spok.1.45, Spyo\Cas9UAS.Tag:NLS(Ude)
epicuticle
glial cell
larval tracheal system
larval ventral nerve cord
lipid droplet | adult stage, with Scer\GAL4Tk.PS
male accessory gland secondary cell, with Scer\GAL4esg.PU
midline glial cell | ectopic
presumptive embryonic/larval central nervous system
presumptive embryonic/larval peripheral nervous system
sensory neuron | embryonic stage
symmetrical commissure
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (29)
Hsap\CYP11A1
3 of 14
Yes
No
Hsap\CYP11B1
3 of 14
Yes
Yes
Hsap\CYP11B2
3 of 14
Yes
Yes
Hsap\CYP24A1
3 of 14
Yes
No
1  
Hsap\CYP27A1
3 of 14
Yes
No
Hsap\CYP27B1
3 of 14
Yes
No
Hsap\CYP27C1
3 of 14
Yes
No
Hsap\CYP2E1
1 of 14
No
No
Hsap\CYP2J2
1 of 14
No
No
Hsap\CYP2U1
1 of 14
No
No
Hsap\CYP2W1
1 of 14
No
No
Hsap\CYP3A4
1 of 14
No
No
1  
Hsap\CYP3A5
1 of 14
No
No
Hsap\CYP3A7-CYP3A51P
1 of 14
No
No
Hsap\CYP3A7
1 of 14
No
No
Hsap\CYP3A43
1 of 14
No
No
Hsap\CYP4A11
1 of 14
No
No
Hsap\CYP4B1
1 of 14
No
No
Hsap\CYP4F3
1 of 14
No
No
Hsap\CYP4F8
1 of 14
No
No
Hsap\CYP4F12
1 of 14
No
No
Hsap\CYP4V2
1 of 14
No
No
Hsap\CYP4X1
1 of 14
No
No
Hsap\CYP4Z1
1 of 14
No
No
Hsap\CYP19A1
1 of 14
No
No
Hsap\CYP21A2
1 of 14
No
No
Hsap\CYP26A1
1 of 14
No
No
Hsap\CYP46A1
1 of 14
No
No
Hsap\CYP51A1
1 of 14
No
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (20)
Rnor\Cyp11a1
3 of 14
Yes
No
Rnor\Cyp11b2
3 of 14
Yes
Yes
Rnor\Cyp11b3
3 of 14
Yes
Yes
Rnor\Cyp24a1
3 of 14
Yes
No
Rnor\Cyp27a1
3 of 14
Yes
No
Rnor\Cyp27b1
3 of 14
Yes
No
Rnor\Cyp2j3
1 of 14
No
No
Rnor\Cyp2j16
1 of 14
No
No
Rnor\Cyp3a2
1 of 14
No
No
Rnor\Cyp3a9
1 of 14
No
No
Rnor\Cyp3a18
1 of 14
No
No
Rnor\Cyp3a23-3a1
1 of 14
No
No
Rnor\Cyp3a62
1 of 14
No
No
Rnor\Cyp4a1
1 of 14
No
No
Rnor\Cyp4b1
1 of 14
No
No
Rnor\Cyp4f5
1 of 14
No
No
Rnor\Cyp4f18
1 of 14
No
No
Rnor\Cyp4v3
1 of 14
No
No
Rnor\Cyp11b1
1 of 14
No
Yes
Rnor\LOC680316
1 of 14
No
Yes
Mus musculus (laboratory mouse) (22)
Mmus\Cyp11a1
3 of 14
Yes
No
Mmus\Cyp11b1
3 of 14
Yes
No
Mmus\Cyp24a1
3 of 14
Yes
No
Mmus\Cyp27b1
3 of 14
Yes
No
Mmus\Cyp11b2
2 of 14
No
No
Mmus\Cyp27a1
2 of 14
No
No
Mmus\Cyp2j7
1 of 14
No
No
Mmus\Cyp2j8
1 of 14
No
No
Mmus\Cyp2j11
1 of 14
No
No
Mmus\Cyp2w1
1 of 14
No
No
Mmus\Cyp3a11
1 of 14
No
No
Mmus\Cyp3a13
1 of 14
No
No
Mmus\Cyp3a16
1 of 14
No
No
Mmus\Cyp3a25
1 of 14
No
No
Mmus\Cyp3a41a
1 of 14
No
No
Mmus\Cyp3a41b
1 of 14
No
No
Mmus\Cyp3a44
1 of 14
No
No
Mmus\Cyp3a57
1 of 14
No
No
Mmus\Cyp3a59
1 of 14
No
No
Mmus\Cyp4f37
1 of 14
No
No
Mmus\Cyp4v3
1 of 14
No
No
Mmus\Cyp17a1
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (30)
Xtro\cyp11a1
2 of 13
Yes
No
Xtro\cyp24a1
2 of 13
Yes
No
Xtro\cyp27b1
2 of 13
Yes
No
Xtro\il17d
2 of 13
Yes
No
Xtro\LOC101731070
2 of 13
Yes
No
Xtro\XB1001094
2 of 13
Yes
No
Xtro\cyp1c1
1 of 13
No
No
Xtro\cyp1d1
1 of 13
No
No
Xtro\cyp2a6.1
1 of 13
No
No
Xtro\cyp2a6.3
1 of 13
No
No
Xtro\cyp2a6.8
1 of 13
No
No
Xtro\cyp2a6.9
1 of 13
No
No
Xtro\cyp2c8l
1 of 13
No
No
Xtro\cyp26a1
1 of 13
No
No
Xtro\cyp27a1.3
1 of 13
No
No
Xtro\cyp27a1
1 of 13
No
No
Xtro\cyp27c1
1 of 13
No
No
Xtro\cyp46a1.3
1 of 13
No
No
Xtro\LOC100491603
1 of 13
No
No
Xtro\LOC100491773
1 of 13
No
No
Xtro\LOC100496967
1 of 13
No
No
Xtro\LOC100497899
1 of 13
No
No
Xtro\LOC100498621
1 of 13
No
No
Xtro\LOC116407844
1 of 13
No
No
Xtro\rem2
1 of 13
No
No
Xtro\XB980013
1 of 13
No
No
Xtro\XB984297
1 of 13
No
No
Xtro\XB5810926
1 of 13
No
No
Xtro\XB5880541
1 of 13
No
No
Xtro\XB5919979
1 of 13
No
No
Danio rerio (Zebrafish) (24)
Drer\cyp11a1
3 of 14
Yes
No
Drer\cyp11a2
3 of 14
Yes
No
Drer\cyp11c1
3 of 14
Yes
Yes
Drer\cyp27a1.2
3 of 14
Yes
No
Drer\cyp27a1.4
3 of 14
Yes
No
Drer\cyp27a7
3 of 14
Yes
No
Drer\cyp27c1
3 of 14
Yes
No
Drer\si:ch211-113j14.1
3 of 14
Yes
No
Drer\si:dkey-91i10.3
3 of 14
Yes
No
Drer\zgc:136333
3 of 14
Yes
No
Drer\cyp24a1
2 of 14
No
No
Drer\cyp27b1
2 of 14
No
No
Drer\cyp2n13
1 of 14
No
No
Drer\cyp2p10
1 of 14
No
No
Drer\cyp2v1
1 of 14
No
No
Drer\cyp3a65
1 of 14
No
No
Drer\cyp3c1
1 of 14
No
No
Drer\cyp3c3
1 of 14
No
No
Drer\cyp4v7
1 of 14
No
No
Drer\cyp4v8
1 of 14
No
No
Drer\cyp46a1.2
1 of 14
No
No
Drer\cyp46a1.4
1 of 14
No
No
Drer\tbxas1
1 of 14
No
No
Drer\zgc:123299
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (32)
Cele\cyp-44A1
3 of 14
Yes
No
Cele\cyp-13A1
1 of 14
No
No
Cele\cyp-13A2
1 of 14
No
No
Cele\cyp-13A3
1 of 14
No
No
Cele\cyp-13A5
1 of 14
No
No
Cele\cyp-13A6
1 of 14
No
No
Cele\cyp-13A10
1 of 14
No
No
Cele\cyp-13A11
1 of 14
No
No
Cele\cyp-13A12
1 of 14
No
No
Cele\cyp-13B1
1 of 14
No
No
Cele\cyp-13B2
1 of 14
No
No
Cele\cyp-25A1
1 of 14
No
No
Cele\cyp-25A2
1 of 14
No
No
Cele\cyp-25A3
1 of 14
No
No
Cele\cyp-25A4
1 of 14
No
No
Cele\cyp-29A2
1 of 14
No
No
Cele\cyp-29A3
1 of 14
No
No
Cele\cyp-29A4
1 of 14
No
No
Cele\cyp-31A2
1 of 14
No
No
Cele\cyp-31A3
1 of 14
No
No
Cele\cyp-32A1
1 of 14
No
No
Cele\cyp-32B1
1 of 14
No
No
Cele\cyp-33C2
1 of 14
No
No
Cele\cyp-33E1
1 of 14
No
No
Cele\cyp-33E2
1 of 14
No
No
Cele\cyp-34A5
1 of 14
No
No
Cele\cyp-34A6
1 of 14
No
No
Cele\cyp-34A7
1 of 14
No
No
Cele\cyp-37A1
1 of 14
No
No
Cele\cyp-37B1
1 of 14
No
No
Cele\cyp-42A1
1 of 14
No
No
Cele\npp-24
1 of 14
No
Yes
Anopheles gambiae (African malaria mosquito) (64)
Agam\AgaP_AGAP002429
12 of 12
Yes
Yes
Agam\AgaP_AGAP005774
4 of 12
No
No
Agam\AgaP_AGAP005992
4 of 12
No
No
Agam\AgaP_AGAP006082
4 of 12
No
No
Agam\AgaP_AGAP000284
3 of 12
No
No
Agam\AgaP_AGAP008018
3 of 12
No
No
Agam\AgaP_AGAP008019
3 of 12
No
No
Agam\AgaP_AGAP008020
3 of 12
No
No
Agam\AgaP_AGAP008022
3 of 12
No
No
Agam\AgaP_AGAP012800
3 of 12
No
No
Agam\AgaP_AGAP000088
1 of 12
No
No
Agam\AgaP_AGAP000193
1 of 12
No
No
Agam\AgaP_AGAP000194
1 of 12
No
No
Agam\AgaP_AGAP001076
1 of 12
No
No
Agam\AgaP_AGAP001443
1 of 12
No
No
Agam\AgaP_AGAP001861
1 of 12
No
No
Agam\AgaP_AGAP001864
1 of 12
No
No
Agam\AgaP_AGAP002195
1 of 12
No
No
Agam\AgaP_AGAP002197
1 of 12
No
No
Agam\AgaP_AGAP002204
1 of 12
No
No
Agam\AgaP_AGAP002205
1 of 12
No
No
Agam\AgaP_AGAP002206
1 of 12
No
No
Agam\AgaP_AGAP002208
1 of 12
No
No
Agam\AgaP_AGAP002209
1 of 12
No
No
Agam\AgaP_AGAP002210
1 of 12
No
No
Agam\AgaP_AGAP002211
1 of 12
No
No
Agam\AgaP_AGAP002416
1 of 12
No
No
Agam\AgaP_AGAP002417
1 of 12
No
No
Agam\AgaP_AGAP002418
1 of 12
No
No
Agam\AgaP_AGAP002419
1 of 12
No
No
Agam\AgaP_AGAP002555
1 of 12
No
No
Agam\AgaP_AGAP002865
1 of 12
No
No
Agam\AgaP_AGAP002866
1 of 12
No
No
Agam\AgaP_AGAP002868
1 of 12
No
No
Agam\AgaP_AGAP002870
1 of 12
No
No
Agam\AgaP_AGAP003343
1 of 12
No
No
Agam\AgaP_AGAP003608
1 of 12
No
No
Agam\AgaP_AGAP006047
1 of 12
No
No
Agam\AgaP_AGAP006049
1 of 12
No
No
Agam\AgaP_AGAP007480
1 of 12
No
No
Agam\AgaP_AGAP008021
1 of 12
No
No
Agam\AgaP_AGAP008204
1 of 12
No
No
Agam\AgaP_AGAP008205
1 of 12
No
No
Agam\AgaP_AGAP008208
1 of 12
No
No
Agam\AgaP_AGAP008209
1 of 12
No
No
Agam\AgaP_AGAP008212
1 of 12
No
No
Agam\AgaP_AGAP008214
1 of 12
No
No
Agam\AgaP_AGAP008217
1 of 12
No
No
Agam\AgaP_AGAP008356
1 of 12
No
No
Agam\AgaP_AGAP008358
1 of 12
No
No
Agam\AgaP_AGAP008552
1 of 12
No
No
Agam\AgaP_AGAP009240
1 of 12
No
No
Agam\AgaP_AGAP009241
1 of 12
No
No
Agam\AgaP_AGAP009246
1 of 12
No
No
Agam\AgaP_AGAP010414
1 of 12
No
No
Agam\AgaP_AGAP012292
1 of 12
No
No
Agam\AgaP_AGAP012957
1 of 12
No
No
Agam\AgaP_AGAP013224
1 of 12
No
No
Agam\AgaP_AGAP013241
1 of 12
No
No
Agam\AgaP_AGAP013305
1 of 12
No
No
Agam\AgaP_AGAP013490
1 of 12
No
No
Agam\CYP6P4
1 of 12
No
No
Agam\CYP6Z2
1 of 12
No
No
Agam\CYP9L1
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (61)
Atha\CYP71B8
2 of 13
Yes
No
Atha\AT2G12190
1 of 13
No
No
Atha\BAS1
1 of 13
No
No
Atha\CYP71A13
1 of 13
No
No
Atha\CYP71A15
1 of 13
No
No
Atha\CYP71A18
1 of 13
No
No
Atha\CYP71A19
1 of 13
No
No
Atha\CYP71A20
1 of 13
No
No
Atha\CYP71A23
1 of 13
No
No
Atha\CYP71A25
1 of 13
No
No
Atha\CYP71A26
1 of 13
No
No
Atha\CYP71B4
1 of 13
No
No
Atha\CYP71B13
1 of 13
No
No
Atha\CYP71B21
1 of 13
No
No
Atha\CYP71B24
1 of 13
No
No
Atha\CYP71B25
1 of 13
No
No
Atha\CYP71B31
1 of 13
No
No
Atha\CYP71B34
1 of 13
No
No
Atha\CYP71B35
1 of 13
No
No
Atha\CYP71B36
1 of 13
No
No
Atha\CYP76C1
1 of 13
No
No
Atha\CYP76C3
1 of 13
No
No
Atha\CYP76C5
1 of 13
No
No
Atha\CYP76C7
1 of 13
No
No
Atha\CYP79A2
1 of 13
No
No
Atha\CYP81D4
1 of 13
No
No
Atha\CYP81D7
1 of 13
No
No
Atha\CYP81D11
1 of 13
No
No
Atha\CYP81F3
1 of 13
No
No
Atha\CYP81F4
1 of 13
No
No
Atha\CYP81H1
1 of 13
No
No
Atha\CYP82C2
1 of 13
No
No
Atha\CYP82C4
1 of 13
No
No
Atha\CYP82G1
1 of 13
No
No
Atha\CYP83A1
1 of 13
No
No
Atha\CYP86B1
1 of 13
No
No
Atha\CYP91A2
1 of 13
No
No
Atha\CYP93D1
1 of 13
No
No
Atha\CYP96A1
1 of 13
No
No
Atha\CYP96A4
1 of 13
No
No
Atha\CYP96A10
1 of 13
No
No
Atha\CYP97A3
1 of 13
No
No
Atha\CYP97B3
1 of 13
No
No
Atha\CYP703A2
1 of 13
No
No
Atha\CYP704A1
1 of 13
No
No
Atha\CYP704A2
1 of 13
No
No
Atha\CYP705A13
1 of 13
No
No
Atha\CYP705A18
1 of 13
No
No
Atha\CYP705A20
1 of 13
No
No
Atha\CYP705A23
1 of 13
No
No
Atha\CYP705A27
1 of 13
No
No
Atha\CYP705A28
1 of 13
No
No
Atha\CYP705A30
1 of 13
No
No
Atha\CYP706A3
1 of 13
No
No
Atha\CYP706A6
1 of 13
No
No
Atha\CYP709B3
1 of 13
No
No
Atha\CYP711A1
1 of 13
No
No
Atha\CYP712A2
1 of 13
No
No
Atha\GA3
1 of 13
No
Yes
Atha\LUT1
1 of 13
No
No
Atha\TT7
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
Scer\DIT2
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:shd. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (57)
Cyp49a1
5 of 13
Cyp301a1
5 of 13
dib
5 of 13
Cyp12a4
4 of 13
Cyp12a5
4 of 13
Cyp12b2
4 of 13
Cyp12c1
4 of 13
Cyp12d1-d
4 of 13
Cyp12d1-p
4 of 13
Cyp12e1
4 of 13
sad
3 of 13
Cyp313a1
2 of 13
Cyp4aa1
1 of 13
Cyp4ac1
1 of 13
Cyp4ac2
1 of 13
Cyp4ac3
1 of 13
Cyp4ad1
1 of 13
Cyp4ae1
1 of 13
Cyp4c3
1 of 13
Cyp4d1
1 of 13
Cyp4d2
1 of 13
Cyp4d8
1 of 13
Cyp4d14
1 of 13
Cyp4d20
1 of 13
Cyp4d21
1 of 13
Cyp4e1
1 of 13
Cyp4e2
1 of 13
Cyp4e3
1 of 13
Cyp4g1
1 of 13
Cyp4g15
1 of 13
Cyp4p1
1 of 13
Cyp4p3
1 of 13
Cyp4s3
1 of 13
Cyp6a2
1 of 13
Cyp6a8
1 of 13
Cyp6a13
1 of 13
Cyp6a14
1 of 13
Cyp6a17
1 of 13
Cyp6a18
1 of 13
Cyp6a19
1 of 13
Cyp6a20
1 of 13
Cyp6a22
1 of 13
Cyp6d4
1 of 13
Cyp6d5
1 of 13
Cyp6g1
1 of 13
Cyp18a1
1 of 13
Cyp28a5
1 of 13
Cyp28d1
1 of 13
Cyp28d2
1 of 13
Cyp311a1
1 of 13
Cyp312a1
1 of 13
Cyp313a2
1 of 13
Cyp313a3
1 of 13
Cyp313a4
1 of 13
Cyp313a5
1 of 13
Cyp313b1
1 of 13
Cyp318a1
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 7 )
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    shdHMC06461
    ameliorates  carcinoma
    modeled by Ras85DV12.UAS, dlg1GD4689
    (Santabárbara-Ruiz and Léopold, 2021)
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    CYP11A1; cytochrome P450 family 11 subfamily A member 1
    3 of 14
    118485
    CYP11B1; cytochrome P450 family 11 subfamily B member 1
    3 of 14
    610613
    CYP11B2; cytochrome P450 family 11 subfamily B member 2
    3 of 14
    124080
    CYP24A1; cytochrome P450 family 24 subfamily A member 1
    3 of 14
    126065
     
    CYP27A1; cytochrome P450 family 27 subfamily A member 1
    3 of 14
    606530
    CYP27B1; cytochrome P450 family 27 subfamily B member 1
    3 of 14
    609506
    CYP27C1; cytochrome P450 family 27 subfamily C member 1
    3 of 14
    620605
        Functional Complementation Data
        Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
        Interactions
        Summary of Physical Interactions
        Interaction Browsers
        View in FlyBase Interactions Browser   View in MIST tool (external link)
        Please see the Physical Interaction reports below for full details
        protein-protein
        Physical Interaction
        Assay
        References
        shd - CCT6
        experimental knowledge based
        (Guruharsha et al., 2011)
        shd - CCT8
        experimental knowledge based
        (Guruharsha et al., 2011)
        Summary of Genetic Interactions
        Interaction Browsers
        View in FlyBase Interactions Browser   View in MIST tool (external link)
        Please look at the allele data for full details of the genetic interactions
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Dp|E2f1
        enhanceable
        shd
        (Sekar et al., 2023)
        External Data
        Linkouts
        DroID - A comprehensive database of gene and protein interactions.
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Pathways
        Signaling Pathways (FlyBase)
        Metabolic Pathways
        FlyBase
        External Links
        KEGG Metabolic Pathways - A collection of manually drawn metabolic pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        External Data
        Linkouts
        KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
        Class of Gene
        Genomic Location and Detailed Mapping Data
        Chromosome (arm)
        3L
        Recombination map
        3-42
        Cytogenetic map
        70E4-70E4
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        70E4-70E4
        Limits computationally determined from genome sequence between P{PZ}Mpcp00564 and P{PZ}l(3)rO220rO220
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        70E4-70E5
        (Dunkov, 2000.6.29)
        Experimentally Determined Recombination Data
        Location

        3-42

        (FlyBase, 2016)

        3-36.3

        (Comeron et al., 2012)

        3-41

        (Tearle and Nusslein-Volhard, 1987)
        Left of (cM)
        Right of (cM)
        Notes
        Stocks and Reagents
        Stocks (9)
        Genomic Clones (19)
         

        Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

        cDNA Clones (10)
         

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

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

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

        cDNA Clones, End Sequenced (ESTs)
        BDGP DGC clones
          Other clones
          RNAi and Array Information
          Linkouts
          DRSC - Results frm RNAi screens
          Antibody Information
          Laboratory Generated Antibodies
           
          Commercially Available Antibodies
           
          Cell Line Information
          Publicly Available Cell Lines
           
            Other Stable Cell Lines
             
              Other Comments

              Four EMS induced alleles have been identified in a screen for mutations affecting commissure formation in the CNS of the embryo.

              (Hummel et al., 1999)

              shd mutants display no differentiation of the cuticle or head skeleton.

              (Jurgens et al., 1984)
              Relationship to Other Genes
              Source for database merge of

              Source for merge of: shd Cyp314a1

              (Petryk, 2002.2.15)
              Additional comments
              Nomenclature History
              Source for database identify of
              Nomenclature comments
              Etymology
              Synonyms and Secondary IDs (12)
              Reported As
              Symbol Synonym
              314a1
              (Nelson, 2001.2.5)
              CG13478
              (Hosono et al., 2015, Keleman et al., 2009.8.5, Perkins et al., 2009)
              CYP314A1
              (Nirusimhan et al., 2022, Pan et al., 2020, Chen and Li, 2007, Rewitz et al., 2007)
              Cyp314a1
              (Scanlan et al., 2020, Good et al., 2014, Niwa et al., 2011, Rewitz et al., 2007, Ranson et al., 2002, Tijet et al., 2001)
              DmCYP314A1
              (Lin et al., 2023)
              Shade
              (Sood et al., 2021)
              Shd
              (Wen et al., 2023, Kamiyama and Niwa, 2022)
              cyp314a1
              (Gangishetti et al., 2009)
              shade
              (Truman and Riddiford, 2023, Meschi and Delanoue, 2021, Jacobs et al., 2020, Gangishetti et al., 2009)
              shd
              (Chen et al., 2025, Peng et al., 2024, Zhang et al., 2024, Suyama et al., 2023, Carlson et al., 2022, Gupta et al., 2022, Keith et al., 2021, Santabárbara-Ruiz and Léopold, 2021, Zhao et al., 2021, Scanlan et al., 2020, Leiblich et al., 2019, Russo et al., 2019, Swevers, 2019, Li et al., 2018, Shimell et al., 2018, Revaitis et al., 2017, Akagi et al., 2016, Borsos et al., 2015, Deady et al., 2015, Xie et al., 2015, Domanitskaya et al., 2014, Good et al., 2014, Deng and Kerppola, 2013, Gangishetti et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Luo et al., 2011, Chavoshi et al., 2010, Pankotai et al., 2010, Xiang et al., 2010, Huang et al., 2008, McBrayer et al., 2007)
              Name Synonyms
              Shade
              (Kannangara et al., 2021, Niwa and Niwa, 2016, Pahi et al., 2015, Heimiller et al., 2014)
              cytochrome P450
              (Chung et al., 2009)
              ecdysone 20-monooxygenase
              (Rewitz et al., 2007)
              shade
              (Wang et al., 2025, Chan et al., 2023, Christensen et al., 2020, Buhler et al., 2018, Akagi et al., 2016, Di Cara and King-Jones, 2016, Saadin and Starz-Gaiano, 2016, Wang et al., 2016, Wieschaus and Nüsslein-Volhard, 2016, Borsos et al., 2015, Domanitskaya et al., 2014, Chavoshi et al., 2010, Pankotai et al., 2010, Xiang et al., 2010, Huang et al., 2008, Rewitz et al., 2007, Hummel et al., 1999)
              Secondary FlyBase IDs
              • FBgn0036432
              Datasets (0)
              Study focus (0)
              Experimental Role
              Project
              Project Type
              Title
              Study result (0)
              Result
              Result Type
              Title
              External Crossreferences and Linkouts ( 44 )
              Sequence Crossreferences
              NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
              GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
              GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
              RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
              UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
              UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
              UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
              Other crossreferences
              AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
              BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
              DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
              EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
              FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
              FlyMine - An integrated database for Drosophila genomics
              InterPro - A database of protein families, domains and functional sites
              KEGG Genes - Molecular building blocks of life in the genomic space.
              MARRVEL_MODEL - MARRVEL (model organism gene)
              Linkouts
              Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
              DroID - A comprehensive database of gene and protein interactions.
              DRSC - Results frm RNAi screens
              Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
              FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
              FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
              Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
              Flygut - An atlas of the Drosophila adult midgut
              iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
              KEGG Metabolic Pathways - A collection of manually drawn metabolic pathway maps representing knowledge of molecular interaction, reaction and relation networks.
              KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
              MIST (protein-protein) - An integrated Molecular Interaction Database
              References (129)