FlyBase Gene Report: Dmel\mlt

General Information

Symbol

Dmel\mlt

Species

D. melanogaster

Name

mulet

Annotation Symbol

CG12214

Feature Type

protein_coding_gene

FlyBase ID

FBgn0265512

Gene Model Status

Current

Stock Availability

11 publicly available

Gene Summary

mulet (mlt) encodes a tubulin-specific chaperone E-like protein involved in sperm individualization and regulation of microtubule depolymerization. [Date last reviewed: 2019-09-26] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Interactive Fly

Key Links

Genomic Location

Cytogenetic map

46F3-46F3

Sequence location

2R:10,155,916..10,158,931 [-]

Recombination map

2-62

RefSeq locus

NT_033778 REGION:10155916..10158931

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

Gene Ontology (GO) Annotations (8 terms)

Molecular Function (1 term)

Terms Based on Experimental Evidence (0 terms)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

enables alpha-tubulin binding

inferred from biological aspect of ancestor with PANTHER:PTN001031924

(GO Reference Genome Project, 2011-)

Biological Process (5 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

involved_in positive regulation of microtubule depolymerization

inferred from mutant phenotype

(Fabrizio et al., 2012)

involved_in sperm individualization

inferred from mutant phenotype

(Fabrizio et al., 2012, Nuwal et al., 2012, Fabrizio et al., 1998)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

involved_in microtubule cytoskeleton organization

inferred from biological aspect of ancestor with PANTHER:PTN001031924

(GO Reference Genome Project, 2011-)

involved_in post-chaperonin tubulin folding pathway

inferred from biological aspect of ancestor with PANTHER:PTN001031924

(GO Reference Genome Project, 2011-)

involved_in tubulin complex assembly

inferred from biological aspect of ancestor with PANTHER:PTN001031924

(GO Reference Genome Project, 2011-)

Cellular Component (2 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

located_in axoneme

inferred from direct assay

(Nuwal et al., 2012)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

is_active_in cytoplasm

inferred from biological aspect of ancestor with PANTHER:PTN001031924

(GO Reference Genome Project, 2011-)

Gene Group (FlyBase)

Protein Family (UniProt)

Protein Signatures (InterPro)

Summaries

Gene Snapshot

mulet (mlt) encodes a tubulin-specific chaperone E-like protein involved in sperm individualization and regulation of microtubule depolymerization. [Date last reviewed: 2019-09-26]

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\mlt 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 Q7K549)

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 5.45

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

mlt-RA

FBtr0088370

NM_136718

2195

459

mlt-RB

FBtr0088369

NM_165754

2542

459

mlt-RC

FBtr0331945

NM_001273924

2669

459

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

mlt-PA

FBpp0087458

53.1

459

5.13

FBpp0087457

53.1

459

5.13

FBpp0304278

53.1

459

5.13

Q7K549

NP_001260853

AGB93386

Polypeptides with Identical Sequences

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

459 aa isoforms: mlt-PA, mlt-PB, mlt-PC

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

Nucleotide / Polypeptide Records

UniProt

Q7K549

Mapped Features

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

External Data

Crossreferences

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

mlt_1

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).

4.33

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism

Comment: maternally deposited

(Fisher et al., 2012)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

mass spectroscopy

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage

adult head

• membrane

(Aradska et al., 2015)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in axoneme

inferred from direct assay

(Nuwal et al., 2012)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\mlt in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\mlt

View exonic expression by tissue for Dmel\mlt

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

FBgn0265512

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

36072/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0265512

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0033495

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0265512

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG12214

Images

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 15 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 5 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of mlt

Transgenic constructs containing regulatory region of mlt

Aberrations (Deficiencies and Duplications) ( 4 )

Inferred from experimentation ( 4 )

Gene disrupted in

Df(2R)BSC281

(Fabrizio et al., 2012)

Df(2R)BSC350

(Fabrizio et al., 2012)

Df(2R)stan2

(Fabrizio et al., 2012)

Gene partially disrupted in

Df(2R)CRIMIC-CR71036

(Gene Disruption Project members, 2018-)

Inferred from location ( 5 )

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

viable

viable, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

mlt^GD7531

viable, with Scer\GAL4^Mef2.PR

mlt^GD7531

viable, with Scer\GAL4^pnr-MD237

mlt^GD7531

viable, with Scer\GAL4^tin.CΔ4

mlt^GD7531

viable (with Df(2R)BSC281)

mlt^NP4786

viable | adult stage

mlt⁴²¹⁰

Sterility

Allele

female fertile

female fertile (with Df(2R)BSC281)

mlt^NP4786

fertile

mlt^EY02157

male semi-sterile

mlt^G18151

male semi-sterile (with mlt^e00818)

mlt^G18151

male semi-sterile (with mlt^G18151)

mlt^e00818

male sterile

male sterile (with Df(2R)BSC281)

mlt^NP4786

male sterile | recessive

mlt⁴²¹⁰

Other Phenotypes

Allele

abnormal courtship behavior | male limited

mlt^G18151

Phenotype manifest in

Allele

investment cone

investment cone (with Df(2R)BSC281)

investment cone (with Df(2R)BSC350)

mlt^G18151

investment cone (with Df(2R)stan2)

mlt^EY02157

investment cone (with mlt¹)

investment cone (with mlt²)

mlt^EY02157

investment cone (with mlt⁴²¹⁰)

mlt^EY02157

investment cone (with mlt^EY02157)

investment cone (with mlt^G18151)

primary spermatocyte cyst

mlt⁴²¹⁰

spermatozoon

mlt⁴²¹⁰

sperm individualization complex

sperm individualization complex (with Df(2R)BSC281)

sperm individualization complex (with Df(2R)BSC350)

mlt^G18151

sperm individualization complex (with Df(2R)stan2)

mlt^EY02157

sperm individualization complex (with mlt¹)

sperm individualization complex (with mlt²)

mlt^EY02157

sperm individualization complex (with mlt⁴²¹⁰)

mlt^EY02157

sperm individualization complex (with mlt^EY02157)

sperm individualization complex (with mlt^G18151)

wing blade posterior compartment, with Scer\GAL4^en-e16E

mlt^UY1505

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (25)

Hsap\TBCEL

14 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (23)

Rnor\Tbcel

14 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Rnor\Nisch

1 of 14

Rnor\Podnl1

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Mus musculus (laboratory mouse) (21)

Mmus\Tbcel

13 of 14

Yes

Mmus\Aspn

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (21)

Xtro\tbcel

11 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Xtro\lrrc23

1 of 13

Xtro\lrriq1

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

Yes

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

Danio rerio (Zebrafish) (18)

Drer\tbcelb

13 of 14

Yes

Drer\tbcela

12 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Caenorhabditis elegans (Nematode, roundworm) (6)

Cele\coel-1

13 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Anopheles gambiae (African malaria mosquito) (21)

Agam\AgaP_AGAP011141

12 of 12

Yes

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Yes

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Yes

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Arabidopsis thaliana (thale-cress) (11)

Atha\PFI

3 of 13

Yes

Atha\AT4G36180

1 of 13

Yes

Atha\AT4G39270

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Atha\RLP26

1 of 13

Atha\RLP30

1 of 13

Atha\RLP35

1 of 13

Yes

Saccharomyces cerevisiae (Brewer's yeast) (2)

Scer\PAC2

3 of 13

Yes

Scer\CYR1

1 of 13

Schizosaccharomyces pombe (Fission yeast) (3)

Spom\alp21

2 of 12

Yes

Spom\cyr1

1 of 12

Spom\mex67

1 of 12

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:mlt. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (14)

Tbce

3 of 13

2mit

1 of 13

atk

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Lrt

1 of 13

Sur-8

1 of 13

trn

1 of 13

U2A

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 ( 0 )

Human Ortholog

Disease

Evidence

References

Modifiers Based on Experimental Evidence ( 1 )

Allele

Disease

Interaction

References

mlt^e00818

ameliorates childhood spinal muscular atrophy

modeled by Smn^73Ao

(Chang et al., 2008)

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?

TBCEL; tubulin folding cofactor E like

14 of 14

610451

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

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

Smn

suppressible

mlt

(Chang et al., 2008)

External Data

Linkouts

DroID - A comprehensive database of gene and protein interactions.

FBgn0265512

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

Class of Gene

gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

2-62

Cytogenetic map

46F3-46F3

Sequence location

2R:10,155,916..10,158,931 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

46F3-46F3

Limits computationally determined from genome sequence between P{PZ}Pfk⁰⁶³³⁹&P{PZ}CCS⁰³²²¹ and P{lacW}Hr46^k10308

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

46F-46F

(determined by in situ hybridisation)

(Fabrizio et al., 1998)

Experimentally Determined Recombination Data

Location

2-62

(FlyBase, 2016)

2-60.3

(Comeron et al., 2012)

2-[59]

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (11)

Aberrations including deletions of this gene Aberrations including duplications of this gene

Bloomington

2491

P{A92}mlt¹/CyO; ry⁵⁰⁶

10714

P{ry11}mlt², cn¹/CyO; ry⁵⁰⁶

20138

y¹ w^67c23; P{EPgy2}mlt^EY02157

13767

y¹ w^67c23; P{SUPor-P}mlt^KG03017

27434

y¹ w^*; P{EP}mlt^G18151 KCNQ^G18151

605485

y¹ w^*; Df(2R)CRIMIC-CR71036 TI{KozakGAL4}mlt^{CR71036-KO-kG4}/SM6a

Kyoto

101991

P{A92}mlt¹ / CyO ; ry⁵⁰⁶

104780

w^*; P{GawB}KCNQ^NP4786 / CyO

NIG-Fly

12214R-2

P{NIG.12214R}2

2RG-0694

y² VhaAC39-1^cho-1 v¹; P{U6-mlt.NIG.sgRNA}attP40/CyO

VDRC

v105480

P{KK107715}VIE-260B

Genomic Clones (15)

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

cDNA Clones (68)

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

GH13040

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.

FBgn0265512

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Nuwal et al., 2012)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

Postmeiotic differentiation defect.

(Castrillon et al., 1993)

Relationship to Other Genes

Source for database merge of

Source for merge of: mlt CG12214

(Fabrizio et al., 2012)

Additional comments

Although alleles of the mlt (mulet) complementation group map within both the CG12214 annotation and an intron of KCNQ (nested genes, CG12214 is entirely within a large intron of KCNQ), complementation analysis indicates that the mutant phenotype caused by the "mlt" complementation group (spermatid individualisation defects) is due to an effect on CG12214 and not on KCNQ.

(Fabrizio et al., 2012)

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (11)

Reported As

Symbol Synonym

CG12214

(National Institute of Genetics Fly Stocks, 2022-, Aradska et al., 2015, Sen et al., 2013, Fabrizio et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Nuwal et al., 2012, Pauli et al., 2010, Keleman et al., 2009.8.5, Chang et al., 2008, Wang et al., 2004)

TBCE-like

(Nuwal et al., 2012)

TBCEL

(Nuwal et al., 2012)

Tbcel

(Nuwal et al., 2012)

mlt

(Banho et al., 2021, Meltzer et al., 2019, Croset et al., 2018, Gene Disruption Project members, 2018-, Fabrizio et al., 2012, Nuwal et al., 2012, Rathke et al., 2007, Bazinet and Rollins, 2003)

ms(2)3

ms(2)4210

(Fabrizio et al., 1998)

ms(2)ry-3

ms(2)ry3

mulet

(Fabrizio et al., 2020)

Name Synonyms

mulet

(Fabrizio et al., 2012, Nuwal et al., 2012, Rathke et al., 2007, Bazinet and Rollins, 2003)

tubulin-specific chaperone E-like protein

(Nuwal et al., 2012)

Secondary FlyBase IDs

FBgn0013647
FBgn0011226
FBgn0033495

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 41 )

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.

36072

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.

Q7K549

UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information

Q7K549

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.

Q7K549

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0265512

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

36072/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0265512

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0265512

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

dme:Dmel_CG12214

MARRVEL_MODEL - MARRVEL (model organism gene)

36072

Linkouts

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0265512

DroID - A comprehensive database of gene and protein interactions.

FBgn0265512

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

mlt_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0033495

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0265512

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG12214

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC007700

References (44)

The following external sites may use different assemblies or annotations than FlyBase.

4.33

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