FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\lwr
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General Information
Symbol
Dmel\lwr
Species
D. melanogaster
Name
lesswright
Annotation Symbol
CG3018
Feature Type
FlyBase ID
FBgn0010602
Gene Model Status
Stock Availability
Gene Summary
lesswright (lwr) encodes Ubc9, a SUMO conjugating enzyme that accepts SUMO from the SUMO activating enzyme and hands it off to the SUMO conjugation target. It has documented biological functions in innate immunity, meiosis, and anterior patterning of the embryo. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

Ubc9, DmUbc9, semi, dUbc9, semushi

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
2-1.5
RefSeq locus
NT_033779 REGION:540599..542580
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (25 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000629675
Biological Process (19 terms)
Terms Based on Experimental Evidence (19 terms)
CV Term
Evidence
References
involved_in dorsal closure
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:Axs; FB:FBgn0000152
inferred from genetic interaction with FLYBASE:mei-218; FB:FBgn0002709
inferred from genetic interaction with FLYBASE:ncd; FB:FBgn0002924
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:smo; FB:FBgn0003444
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:ptc; FB:FBgn0003892
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000629675
inferred from sequence or structural similarity with SGD:S000002222
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000629675
Gene Group (FlyBase)
Protein Family (UniProt)
-
Summaries
Gene Snapshot
lesswright (lwr) encodes Ubc9, a SUMO conjugating enzyme that accepts SUMO from the SUMO activating enzyme and hands it off to the SUMO conjugation target. It has documented biological functions in innate immunity, meiosis, and anterior patterning of the embryo. [Date last reviewed: 2019-03-14]
Pathway (FlyBase)
POSITIVE REGULATORS OF HEDGEHOG SIGNALING PATHWAY -
Positive regulators of hedgehog signaling up-regulate the pathway, resulting in the activation of transcription of hh-responsive genes.
NEGATIVE REGULATORS OF TOLL-NF-kappaB SIGNALING PATHWAY -
In Drosophila, the canonical Toll signaling pathway is initiated by the binding of a spatzle ligand to Toll (Tl) or a Toll-like receptor leading to the nuclear localization of the NF-κB (dl or Dif) transcription factor. Negative regulators of Toll-NF-κB Signaling lead to the increased cytosolic retention of the NF-κB proteins dl or Dif and/or repression of target gene transcription. (Adapted from FBrf0091014 and FBrf0223077).
NEGATIVE REGULATORS OF IMD SIGNALING PATHWAY -
Negative regulators of the immune deficiency (Imd) pathway result in the decreased activity of the NFκB-like transcription factor Rel in the nucleus. Negative regulators are important in preventing damage to the host from over-activation of the pathway; preventing inappropriate triggering or terminating the response. (Adapted from FBrf0224587 and FBrf0238555.)
Gene Group (FlyBase)
SUMO TRANSFERASES -
SUMO transferases catalyze the transfer of SUMO from either E2 or E3 enzyme to a substrate.
Summary (Interactive Fly)

SUMO E2-conjugating enzyme - stabilizes proteins against ubiquitinization and subsequent degradation - regulates hematopoiesis - holds the Toll pathway in check

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\lwr 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 Q7KNM2)

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

Gene model reviewed during 5.42

Gene model reviewed during 5.46

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0078081
1109
159
FBtr0078082
1294
159
FBtr0310028
1363
159
FBtr0310029
1105
159
Additional Transcript Data and Comments
Reported size (kB)

1.2 (northern blot)

0.736 (longest cDNA)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0077741
18.1
159
8.61
FBpp0077742
18.1
159
8.61
FBpp0301731
18.1
159
8.61
FBpp0301732
18.1
159
8.61
Polypeptides with Identical Sequences

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

159 aa isoforms: lwr-PA, lwr-PB, lwr-PC, lwr-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

159 (aa); 18 (kD predicted)

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\lwr 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).

-0.03

Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

Maternal lwr transcripts are distributed throughout the embryo until stage 10 and then decrease gradually. From stage 14 until the end of embryogenesis, transcripts are observed in the embryonic gonads in the pole cells. Within ovaries, transcripts are weakly detected in the germarium, are strongly expressed in stage S1-10 nurse cells and are weakly detected in the oocyte up until stage S10. lwr transcripts are weakly expressed in spermatogonial cells at the tip region and remain detectable in the spermatocytes and spermatids.

lwr transcripts are provided maternally and are expressed throughout embryogenesis.

lwr transcript accumulates at the anterior end of syncytial blastoderm embryos.

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
Additional Descriptive Data

lwr protein is detected in the nucleus of S2 cells.

lwr protein is detected at high levels in 0-24 hr embryos, and in adult females. When the Western blot is overexposed, lwr protein is detected in all developmental stages studied.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
located_in nucleus
inferred from direct assay
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\lwr in JBrowse
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
Linkouts
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
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 19 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 14 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of lwr
Transgenic constructs containing regulatory region of lwr
Aberrations (Deficiencies and Duplications) ( 10 )
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
Other Phenotypes
Allele
Phenotype manifest in
Allele
hemocyte & nucleus | supernumerary (with lwr4-3)
hemocyte & nucleus | supernumerary (with lwr14)
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (25)
11 of 14
Yes
Yes
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
1  
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (8)
12 of 14
Yes
Yes
9 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (8)
11 of 14
Yes
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (8)
11 of 13
Yes
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (10)
12 of 14
Yes
Yes
12 of 14
Yes
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (12)
12 of 14
Yes
Yes
3 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (13)
12 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (28)
12 of 13
Yes
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (9)
12 of 13
Yes
Yes
1 of 13
No
No
1  
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Schizosaccharomyces pombe (Fission yeast) (5)
11 of 12
Yes
Yes
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:lwr. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (26)
3 of 13
3 of 13
3 of 13
3 of 13
3 of 13
2 of 13
2 of 13
2 of 13
2 of 13
2 of 13
2 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 2 )
Potential Models Based on Orthology ( 0 )
Human Ortholog
Disease
Evidence
References
Modifiers Based on Experimental Evidence ( 2 )
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?
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

Please see the Physical Interaction reports below for full details
protein-protein
Physical Interaction
Assay
References
Summary of Genetic Interactions
External Data
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
MIST (genetic) - An integrated Molecular Interaction Database
MIST (protein-protein) - An integrated Molecular Interaction Database
Pathways
Class of Gene
Genomic Location and Detailed Mapping Data
Chromosome (arm)
2L
Recombination map
2-1.5
Cytogenetic map
Sequence location
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
21E2-21E2
Limits computationally determined from genome sequence between P{lacW}l(2)k09610k09610&P{lacW}Sk09538a and P{PZ}l(2)1068510685&P{lacW}Tango14k00619
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
21C6-21C7
(determined by in situ hybridisation)
21C-21D
(determined by in situ hybridisation)
21D-21D
(determined by in situ hybridisation)
Located at the doublet of 21D.
Experimentally Determined Recombination Data
Left of (cM)
Right of (cM)
Notes
Stocks and Reagents
Stocks (17)
Genomic Clones (13)
 

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

cDNA Clones (62)
 

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)
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

    Expression is enriched in embryonic gonads.

    lwr inhibits dl and Dif activities.

    lwr may play a novel role as a gap gene, more specifically lwr function may be responsible for the degradation or modification of Gap genes or other factors which negatively regulate these genes.

    lwr is involved in the regulation of bcd target genes by bcd, being required for nuclear import of bcd protein.

    Mutation of lwr disrupts anterior segmentation in embryogenesis, by interfering with the regulation of bcd target genes by bcd protein.

    Identification: Yeast two hybrid screen for interaction with Hsp23. Co-immunoprecipitation with an antibody raised against lwr confirms interaction with Hsp23 and Hsp26 and preferentially with Hsp27.

    Identification: Yeast two hybrid study using full length sina to screen a SL2 cDNA library.

    Mutants suppress the nodDTW phenotype.

    Relationship to Other Genes
    Source for database merge of

    Source for merge of: semi lwr Ubc9

    Additional comments

    Genetic complementation analysis reveals lwr functionally rescues a temperature-sensitive Scer\UBC9 mutant.

    Nomenclature History
    Source for database identify of

    Source for identity of: lwr CG3018

    Nomenclature comments
    Etymology

    'semushi' means 'hunchback' in Japanese.

    Synonyms and Secondary IDs (33)
    Reported As
    Symbol Synonym
    FBgn0010602
    l(2)01519
    l(2)02858
    l(2)05486
    l(2)05487
    Secondary FlyBase IDs
    • FBgn0010523
    • FBgn0015755
    • FBgn0022761
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    Study result (0)
    Result
    Result Type
    Title
    External Crossreferences and Linkouts ( 83 )
    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.
    RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
    UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
    UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
    Other crossreferences
    AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
    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
    KEGG Genes - Molecular building blocks of life in the genomic space.
    MARRVEL_MODEL - MARRVEL (model organism gene)
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    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
    FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
    iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Reactome Metabolic Pathways - An open-source, open access, manually curated and peer-reviewed metabolic pathway database.
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    References (204)