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General Information
Symbol
Dmel\trol
Species
D. melanogaster
Name
terribly reduced optic lobes
Annotation Symbol
CG33950
Feature Type
FlyBase ID
FBgn0284408
Gene Model Status
Stock Availability
Gene Snapshot
terribly reduced optic lobes (trol) encodes the extracellular matrix component Perlecan, a secreted heparan sulfate proteoglycan. It regulates cell-signaling by multiple growth factors including those in the hedgehog, wingless/WNT, FGF, TGFbeta, EGF, and VEGF families, thus playing a role in many developmental patterning decisions. [Date last reviewed: 2019-06-27]
Also Known As
Perlecan, zw1, Pcan, l(1)zw1, zw-1
Key Links
Genomic Location
Cytogenetic map
Sequence location
X:2,470,460..2,545,427 [-]
Recombination map
1-1
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
-
Predictions / Assertions
-
Summaries
Pathway (FlyBase)
Negative Regulators of Fibroblast Growth Factor Receptor Signaling Pathway -
Negative regulators of Fibroblast Growth Factor Receptor signaling down-regulate the pathway, suppressing the activation of ERK kinase (rl) or acting on other effectors.
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
l(1)3Ac
Tight alleles [l(1)3Ac9, l(1)3Ac12, l(1)3Ac23, l(1)3Ac33] are polyphasic, dying during L2, L3, and post puparial stages; cell viable. Semilethal alleles [l(1)3Ac2 and l(1)3Ac6] die as pupae; escapers have rough eyes, shriveled or blistered wings, and crippled metathoracic legs; phenotype of X0 more extreme that that of XY males. XY escapers fertile but females sterile [l(1)3Ac2/l(1)3Ac2 and l(1)3Ac2/l(1)3Ac6] or nearly so [l(1)3Ac6/l(1)3A6]. The latter females sterile in crosses to l(1)3Ac6 males, but produce heterozygous daughters when outcrossed (Shannon, Kaufman, Shen, and Judd, 1975, Genetics 72: 615-38). In line freed of l(1)3Ba4 (Robbins), l(1)3Ac14 survives as epidermal clones and as oogenic clones, which produce lethal zygotes (Garcia-Bellido and Robbins, 1983, Genetics 103: 235-47).
Summary (Interactive Fly)
extracellular matrix component - homolog of perlecan - regulates larval neuroblast division by modulating both FGF and Hedgehog signaling
Gene Model and Products
Number of Transcripts
22
Number of Unique Polypeptides
22

Please see the GBrowse view of Dmel\trol or the JBrowse view of Dmel\trol for information on other features

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

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model
Gene model reviewed during 6.03
Gene model reviewed during 5.39
Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.
Low-frequency RNA-Seq exon junction(s) not annotated.
Gene model reviewed during 5.51
Gene model reviewed during 6.06
Gene model reviewed during 6.29
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0342806
14181
4434
FBtr0342807
10611
3105
FBtr0342808
13564
4411
FBtr0342809
12022
3897
FBtr0342810
12866
4176
FBtr0342811
11515
3728
FBtr0342812
12232
3967
FBtr0342813
12679
4150
FBtr0342814
12540
3887
FBtr0342815
11749
3806
FBtr0342816
13776
4299
FBtr0342817
12781
4150
FBtr0342818
12294
3805
FBtr0342819
13666
4479
FBtr0342820
14232
4451
FBtr0342821
12798
3973
FBtr0342822
12900
4007
FBtr0342823
13213
2853
FBtr0342824
12378
3833
FBtr0342825
12846
3989
FBtr0342826
13488
4203
FBtr0475303
13023
4178
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
RefSeq ID
GenBank
FBpp0309642
491.7
4434
4.47
FBpp0309643
345.1
3105
4.49
FBpp0309644
489.1
4411
4.47
FBpp0309645
431.6
3897
4.63
FBpp0309646
461.1
4176
4.57
FBpp0309647
412.8
3728
4.54
FBpp0309648
439.5
3967
4.52
FBpp0309649
460.1
4150
4.52
FBpp0309650
430.3
3887
4.53
FBpp0309651
421.5
3806
4.53
FBpp0309652
476.6
4299
4.49
FBpp0309653
460.0
4150
4.53
FBpp0309654
421.0
3805
4.53
FBpp0309655
496.9
4479
4.48
FBpp0309656
493.8
4451
4.47
FBpp0309657
440.4
3973
4.55
FBpp0309658
444.2
4007
4.52
FBpp0309659
316.8
2853
4.49
FBpp0309660
424.6
3833
4.52
FBpp0309661
442.2
3989
4.55
FBpp0309662
465.2
4203
4.52
FBpp0423412
461.9
4178
4.56
Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
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\trol using the Feature Mapper tool.

External Data
Crossreferences
Linkouts
Gene Ontology (10 terms)
Molecular Function (0 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (9 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002576803
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN002471709
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN002471709
(assigned by GO_Central )
inferred from biological aspect of ancestor with PANTHER:PTN002576803
(assigned by GO_Central )
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002471709
(assigned by GO_Central )
inferred from sequence or structural similarity
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic/larval brain | restricted

Comment: two clusters of cells at inner side of brain hemispheres.

follicle cell | subset

Comment: dorsal follicle cells overlying the oocyte nucleus

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
cultured cell-line

Comment: reference states Kc[[1]] cells

Additional Descriptive Data
trol-protein is expressed in a subset of neurons in the embryonic ventral nerve cord, some of which co-express the motor neuron marker Scer\GAL4exex-Gal4. Strong staining is observed in abdominal apodemes and at branch points of the intersegmental nerve ISNb and the segmental nerve SNa. A fainter expression is detected in several regions of ventral and lateral hypodermal muscles with an accumulation along motor axon trajectories close to the ISNb pathway.
In stage 16 embryos, particular depositions of trol protein are observed in the basement membrane of cardiac cells, the dorsal vessel, muscle attachment sites of dorsal muscles, the central nervous system, CNS channels, and dorsal median cells of the CNS.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{unk}trolGFP311
Stage
Tissue/Position (including subcellular localization)
Reference
Stage
Tissue/Position (including subcellular localization)
Reference
adult brain

Comment: unstructured neuropil throughout adult brain

Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\trol in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 187 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 9 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of trol
Transgenic constructs containing regulatory region of trol
Deletions and Duplications ( 70 )
Disrupted in
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
larval brain & neuroblast
neuroblast & larval brain
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
9 of 15
Yes
Yes
2 of 15
No
No
 
1 of 15
No
No
1 of 15
No
No
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (3)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
6 of 15
Yes
Yes
2 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (1)
1 of 13
Yes
No
Xenopus tropicalis (Western clawed frog) (3)
5 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
Yes
Danio rerio (Zebrafish) (7)
2 of 15
Yes
No
2 of 15
Yes
Yes
2 of 15
Yes
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (4)
11 of 15
Yes
Yes
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190027 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150029 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0034 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X002T )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G048M )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (13)
3 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 0 )
Allele
Disease
Evidence
References
Potential Models Based on Orthology ( 2 )
Modifiers Based on Experimental Evidence ( 2 )
Allele
Disease
Interaction
References
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
 
Disease Associations of Human Orthologs (via DIOPT v7.1 and OMIM)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Functional Complementation Data
Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
Interactions
Summary of Physical Interactions
esyN Network Diagram
Show neighbor-neighbor interactions:
Select Layout:
Legend:
Protein
RNA
Selected Interactor(s)
Interactions Browser

Please see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
RNA-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement

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
External Data
Linkouts
Pathways
Genomic Location and Detailed Mapping Data
Chromosome (arm)
X
Recombination map
1-1
Cytogenetic map
Sequence location
X:2,470,460..2,545,427 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
3A1-3A4
Left limit from in situ hybridisation (FBrf0108529) Right limit from in situ hybridisation (FBrf0108529); Left limit from in situ hybridisation (FBrf0109189) Right limit from in situ hybridisation (FBrf0109189)
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
3A1-3A4
(determined by in situ hybridisation)
3A3-3A4
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (37)
Genomic Clones (37)
cDNA Clones (107)
 

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

cDNA clones, fully sequences
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)
    RNAi and Array Information
    Linkouts
    Antibody Information
    Laboratory Generated Antibodies
     
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Source for merge of: trol pcan
    Source for merge of: trol l(1)G0023 l(1)G0271 l(1)G0374
    Source for merge of: trol anon-WO0153538.72
    Source for merge of: pcan EG:BACR25B3.11
    Source for merge of: trol BcDNA:GM02481
    Source for merge of: EG:BACR25B3.11 CG7981
    Source for merge of: trol EG:BACR25B3.1 EG:BACR25B3.10
    Source for merge of: EG:BACR25B3.2 CG12497
    Source for merge of: trol MRE7
    Source for merge of: trol l(1)G0021 l(1)G0181 l(1)G0412 l(1)G0019 l(1)G0211
    Additional comments
    MRE7 corresponds to a portion of the 3' UTR of a short isoform of trol.
    Source for merge of EG:BACR25B3.2 CG12497 was sequence comparison ( date:000425 ).
    "l(1)G0019" may affect "trol". "l(1)G0021" may affect "trol". "l(1)G0073" may affect "trol". "l(1)G0023" may affect "trol". "l(1)G0374" may affect "trol". "l(1)G0181" may affect "trol". "l(1)G0211" may affect "trol". "l(1)G0377" may affect "trol". "l(1)G0271" may affect "trol". "l(1)G0412" may affect "trol". "l(1)G0362" may affect "trol".
    "l(1)G0021" may affect "trol".
    "l(1)G0181" may affect "trol".
    "l(1)G0412" may affect "trol".
    "l(1)G0019" may affect "trol".
    "l(1)G0211" may affect "trol".
    "l(1)G0023" may affect "trol".
    "l(1)G0271" may affect "trol".
    "l(1)G0374" may affect "trol".
    " EG:BACR25B3.2 " may correspond to "Scim4". Sequence analysis off ends of P{SUPor-P} in Scim insertion mutant places "Scim4" near/in " EG:BACR25B3.2 " and/or " EG:BACR25B3.3 ".
    Source for merge of pcan EG:BACR25B3.11 was sequence comparison ( date:000228 ).
    Source for merge of trol BcDNA:GM02481 was a shared cDNA ( date:030728 ).
    Source for merge of trol anon-WO0153538.72 was sequence comparison ( date:051113 ).
    Source for merge of EG:BACR25B3.11 CG7981 was sequence comparison ( date:000425 ).
    Other Comments
    trol does not have an essential role in apical-basal follicle cell polarity.
    trol is required for Sema-1a-plexA-mediated repulsive axon guidance.
    Loss of trol increases basement membrane tension in a Collagen IV-dependent manner.
    trol gene product is required for the activation of proliferation in quiescent optic lobe and thoracic neuroblasts.
    trol acts downstream of ana, it is possible that trol may function to inactivate or bypass ana repression to activate cell proliferation. trol is required specifically to regulate initiation of proliferation and may be part of a switch that controls the transition from quiescence to active proliferation.
    Mutations in trol cause optic lobe defects.
    Partial maternal insufficiency of trol does not cause embryonic lethality.
    Tight alleles <up>trol9, trol12, trol23, trol33</up> are polyphasic, dying during L2, L3 and post-puparial stages; cell viable. Semilethal alleles <up>trol2 and trol6</up> die as pupae; escapers have rough eyes, shriveled or blistered wings and crippled metathoracic legs; phenotype of X0 more extreme than that of XY males. XY escapers fertile but females sterile <up>trol2/trol2 and trol2/trol6</up> or nearly so <up>trol6/trol6</up>. The latter females sterile in crosses to trol6 males, but produce heterozygous daughters when outcrossed (Shannon, Kaufman, Shen and Judd, 1975). In line freed of sgg4 (Robbins), trol14 survives as epidermal clones and as oogenic clones, which produce lethal zygotes (Garcia-Bellido and Robbins, 1983).
    Origin and Etymology
    Discoverer
    Etymology
    Identification
    External Crossreferences and Linkouts ( 172 )
    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 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.
    Synonyms and Secondary IDs (55)
    Reported As
    Symbol Synonym
    BcDNA:GM02481
    CG33675
    EG:BACR25B3.2
    anon-WO0153538.72
    l(1)9-96
    l(1)G0019
    l(1)G0021
    l(1)G0023
    l(1)G0181
    l(1)G0211
    l(1)G0271
    l(1)G0374
    l(1)G0412
    l(1)VA51
    troll
    Secondary FlyBase IDs
    • FBgn0267911
    • FBgn0261451
    • FBgn0001402
    • FBgn0027946
    • FBgn0029620
    • FBgn0029621
    • FBgn0040379
    • FBgn0040380
    • FBgn0040381
    • FBgn0045878
    • FBgn0053675
    • FBgn0053950
    • FBgn0063205
    • FBgn0026710
    • FBgn0027241
    • FBgn0028307
    • FBgn0086539
    • FBgn0067759
    • FBgn0028348
    • FBgn0028284
    • FBgn0067760
    • FBgn0027269
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (266)