FlyBase Gene Report: Dmel\milt

General Information

Symbol

Dmel\milt

Species

D. melanogaster

Name

milton

Annotation Symbol

CG43227

Feature Type

protein_coding_gene

FlyBase ID

FBgn0262872

Gene Model Status

Current

Stock Availability

27 publicly available

Gene Summary

Required for kinesin-mediated axonal transport of mitochondria to nerve terminals (PubMed:12495622, PubMed:16717129, PubMed:16887820). The oocyte acquires the majority of its mitochondria by competitive bidirectional transport along microtubules mediated by the Milton adapter. Mitochondria enter the young oocyte en mass from interconnected germ cells to generate the large aggregate known as the Balbiani body (PubMed:16887820). Milt and Miro form an essential protein complex that links Khc to mitochondria for light chain-independent, anterograde transport of mitochondria (PubMed:16717129). (UniProt, Q960V3)

Contribute a Gene Snapshot for this gene.

All Summaries

Alliance
Auto summary
UniProtKB
Interactive Fly

Also Known As

l(2)k04704

Key Links

Genomic Location

Cytogenetic map

27D5-27D7

Sequence location

2L:7,037,823..7,056,010 [+]

Recombination map

2-24

RefSeq locus

NT_033779 REGION:7037823..7056010

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 (13 terms)

Molecular Function (3 terms)

Terms Based on Experimental Evidence (3 terms)

CV Term

Evidence

References

enables myosin binding

inferred from physical interaction with FLYBASE:Myo10A; FB:FBgn0263705

(Liu et al., 2008)

enables protein binding

inferred from physical interaction with UniProtKB:Q8IMX7

(Glater et al., 2006)

enables protein-macromolecule adaptor activity

inferred from mutant phenotype

(Glater et al., 2006)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

enables myosin binding

inferred from biological aspect of ancestor with PANTHER:PTN000405632

(GO Reference Genome Project, 2011-)

Biological Process (8 terms)

Terms Based on Experimental Evidence (6 terms)

CV Term

Evidence

References

involved_in axonal transport of mitochondrion

inferred from direct assay

(Zhang et al., 2025, Glater et al., 2006)

inferred from expression pattern

(Stowers et al., 2002)

inferred from mutant phenotype

(Stowers et al., 2002)

involved_in establishment of mitochondrion localization

inferred from mutant phenotype

(Glater et al., 2006)

involved_in mitochondrion distribution

inferred from mutant phenotype

(Cox and Spradling, 2006)

involved_in Nebenkern assembly

inferred from mutant phenotype

(Aldridge et al., 2007)

involved_in oocyte dorsal/ventral axis specification

inferred from mutant phenotype

(Glater et al., 2006)

involved_in sperm mitochondrion organization

inferred from mutant phenotype

(Aldridge et al., 2007)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

involved_in mitochondrion distribution

inferred from biological aspect of ancestor with PANTHER:PTN000405632

(GO Reference Genome Project, 2011-)

involved_in protein targeting

inferred from biological aspect of ancestor with PANTHER:PTN000405632

(GO Reference Genome Project, 2011-)

involved_in vesicle transport along microtubule

inferred from biological aspect of ancestor with PANTHER:PTN000405632

(GO Reference Genome Project, 2011-)

Cellular Component (2 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

located_in mitochondrion

inferred from direct assay

(Glater et al., 2006)

inferred from direct assay

(Stowers et al., 2002)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

is_active_in cytoplasmic vesicle

inferred from biological aspect of ancestor with PANTHER:PTN000405632

(GO Reference Genome Project, 2011-)

is_active_in mitochondrion

inferred from biological aspect of ancestor with PANTHER:PTN000405632

(GO Reference Genome Project, 2011-)

located_in mitochondrion

inferred from experiment

(Cox and Spradling, 2006)

Gene Group (FlyBase)

Protein Family (UniProt)

Belongs to the milton family. (Q960V3)

Protein Signatures (InterPro)

Summaries

Protein Function (UniProtKB)

(UniProt, Q960V3)

Summary (Interactive Fly)

scaffolding protein that along with Miro forms an essential protein complex that links kinesin heavy chain to mitochondria for light chain-independent, anterograde transport of mitochondria - controls the early acquisition of mitochondria by Drosophila oocytes - contributes to spermatid tail elongation

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\milt 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 Q960V3)

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

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

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

Gene model reviewed during 5.46

Gene model reviewed during 5.52

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

milt-RA

FBtr0306228

NM_164736

4231

1116

milt-RB

FBtr0306230

NM_164738

3873

1122

milt-RC

FBtr0306229

NM_080765

5482

1116

milt-RD

FBtr0306231

NM_001258998

5535

1256

milt-RE

FBtr0306232

NM_001258999

3711

979

milt-RF

FBtr0306233

NM_001259000

5469

1256

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

milt-PA

FBpp0297338

120.2

1116

6.41

FBpp0297340

120.6

1122

5.43

FBpp0297339

120.2

1116

6.41

FBpp0297341

135.3

1256

6.34

FBpp0297342

105.1

979

6.52

FBpp0297343

135.3

1256

6.34

M9NDH7

NP_001245929

AFH03603

Polypeptides with Identical Sequences

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

1116 aa isoforms: milt-PA, milt-PC

1256 aa isoforms: milt-PD, milt-PF

Additional Polypeptide Data and Comments

Reported size (kDa)

Comments

External Data

Subunit Structure (UniProtKB)

Interacts with Miro (PubMed:16717129). Interacts with OGT (PubMed:24995978). Note=Mitochondrial transport by milt and Khc is independent of Klc.

(UniProt, Q960V3)

Crossreferences

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

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\milt 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).

2.66

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 2 -- 5

organism | anterior

Comment: diffuse anterior enrichment

(Lecuyer et al., 2007)

embryonic stage 4 -- 6

anlage in statu nascendi

(Fisher et al., 2012)

central brain anlage in statu nascendi

(Fisher et al., 2012)

mesoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

dorsal head epidermis anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

embryonic stage 7 -- 8

anterior endoderm anlage

(Fisher et al., 2012)

foregut anlage

(Fisher et al., 2012)

head mesoderm

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

posterior endoderm

(Fisher et al., 2012)

trunk mesoderm

(Fisher et al., 2012)

ventral nerve cord primordium

Comment: reported as ventral nerve cord anlage

(Fisher et al., 2012)

embryonic stage 9 -- 10

antennal primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

central brain primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

visual primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

anterior endoderm

(Fisher et al., 2012)

foregut primordium

(Fisher et al., 2012)

head mesoderm

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

posterior endoderm

(Fisher et al., 2012)

trunk mesoderm

(Fisher et al., 2012)

ventral nerve cord primordium

(Fisher et al., 2012)

embryonic stage 11 -- 12

anterior midgut primordium

(Fisher et al., 2012)

central brain primordium

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

posterior midgut primordium

(Fisher et al., 2012)

ventral nerve cord primordium

(Fisher et al., 2012)

embryonic stage 13 -- 16

embryonic brain

(Fisher et al., 2012)

embryonic dorsal apodeme

(Fisher et al., 2012)

embryonic ventral apodeme

(Fisher et al., 2012)

embryonic/larval corpus allatum

(Fisher et al., 2012)

embryonic/larval midgut

(Fisher et al., 2012)

larval nervous system

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

larval ventral nerve cord

(Fisher et al., 2012)

Additional Descriptive Data

milt expression in early embryos is diffusely concentrated at the anterior end of the embryo.

(Lecuyer et al., 2007)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in mitochondrion

inferred from direct assay

(Glater et al., 2006)

inferred from direct assay

(Stowers et al., 2002)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\milt in JBrowse

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

View exonic expression by developmental stage for Dmel\milt

View exonic expression by tissue for Dmel\milt

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

FBgn0262872

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

45683/tissue=All

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

FBgn0262872

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

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

FBgn0262872

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

CG13777

Images

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 29 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 13 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of milt

Transgenic constructs containing regulatory region of milt

Aberrations (Deficiencies and Duplications) ( 8 )

Inferred from experimentation ( 8 )

Gene disrupted in

Df(2L)Dwee-δ5

(Spradling et al., 1999, BDGP Project Members, 1994-1999)

Df(2L)Exel7030

(Ryder, 2004.4.26)

Df(2L)Gpdh1-A

(BDGP Project Members, 1994-1999)

Df(2L)J-H

(Spradling et al., 1999, BDGP Project Members, 1994-1999)

Df(2L)spd^j2

(Spradling et al., 1999)

Df(2L)spd

(BDGP Project Members, 1994-1999)

Gene partially disrupted in

Df(2L)Mnn1^e200

(Busygina et al., 2004)

Df(2L)P556.15

(Busygina et al., 2004)

Inferred from location ( 4 )

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

lethal | recessive

lethal - all die before end of larval stage (with Df(2L)J-H)

lethal - all die before end of larval stage (with milt⁹²)

milt¹⁸⁶

lethal - all die before end of larval stage (with milt¹⁸⁶)

milt⁹²

lethal - all die before end of larval stage | recessive

milt⁹²

partially lethal - majority die

milt^k14514b

partially lethal - majority die, with Scer\GAL4^pnr-MD237

milt^GD10310

viable

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

viable, with Scer\GAL4^Mef2.PR

milt^GD8116

viable, with Scer\GAL4^pnr-MD237

milt^GD8116

viable, with Scer\GAL4^tin.CΔ4

milt^GD8116

Sterility

Allele

female fertile

fertile

Other Phenotypes

Allele

abnormal locomotor behavior

abnormal locomotor behavior, with Scer\GAL4^elav-C155

milt^GD8116

abnormal neuroanatomy

milt¹⁸⁶

abnormal neuroanatomy | adult stage, with Scer\GAL4^GMR.PF

milt^JF03022

abnormal neuroanatomy | adult stage | somatic clone

abnormal neuroanatomy | conditional, with Scer\GAL4^GMR.PF

milt^GD8116

abnormal neurophysiology

abnormal neurophysiology | adult stage

abnormal neurophysiology | adult stage | somatic clone

milt^EY14443

abnormal pain response | adult stage, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

milt^GD8116

abnormal size | third instar larval stage, with Scer\GAL4^Cg.PA

milt^JF03022

decreased efficacy of long-term memory | adult stage, with Scer\GAL4^c739

milt^HMC02365

decreased efficacy of long-term memory | adult stage, with Scer\GAL4^VT030559

decreased rate of adult locomotory behavior, with Scer\GAL4^unspecified

milt^GD8116

flightless, with Scer\GAL4^Mef2.PR

milt^GD10310

some die during first instar larval stage

milt^33-853

some die during larval stage (with Df(2L)J-H)

some die during larval stage (with milt⁹²)

milt¹⁸⁶

some die during larval stage (with milt¹⁸⁶)

milt⁹²

some die during larval stage | recessive

milt⁹²

some die during pupal stage, with Scer\GAL4^pnr-MD237

milt^GD10310

some die during second instar larval stage

milt^33-853

visible

milt^18-872

visible, with Scer\GAL4^pnr-MD237

milt^GD10310

Phenotype manifest in

Allele

adult antennal lobe | somatic clone

milt^33-853

adult brain

milt⁹²

adult olfactory receptor neuron | somatic clone

milt^33-853

adult optic lobe | conditional, with Scer\GAL4^GMR.PF

milt^GD8116

axon

milt⁹²

axon | adult stage, with Scer\GAL4^dpr1-PGaw

milt^HMC02365

axon | adult stage, with Scer\GAL4^GMR15B07

milt^JF03022

axon | adult stage | somatic clone

milt^EY14443

dendrite & dorsal multidendritic neuron ddaC | somatic clone

milt⁹²

egg chamber & mitochondrion | germ-line clone

embryonic/larval fat body | third instar larval stage, with Scer\GAL4^Cg.PA

milt^JF03022

eye

milt^18-872

eye | somatic clone

milt^EY14443

eye photoreceptor cell | P-stage, with Scer\GAL4^GMR.PF

milt^HMC02365

follicle cell & mitochondrion | germ-line clone

milt^k06704

germline cyst & mitochondrion | germ-line clone

gustatory receptor neuron of the wing, with Scer\GAL4^GMR15B07

milt^JF03022

lamina, with Scer\GAL4^GMR.PF

milt^JF03022

larval glial cell, with Scer\GAL4^repo.PL, Scer\GAL4^repo

milt^GD8116

marginal nerve | adult stage, with Scer\GAL4^dpr1-PGaw

milt^HMC02365

mitochondrial cloud | germline clone

mitochondrial derivative | germline clone

milt⁹²

mitochondrion

mitochondrion, with Scer\GAL4^GMR.PF

mitochondrion, with Scer\GAL4^repo.PL, Scer\GAL4^repo

milt^GD8116

mitochondrion & dorsal multidendritic neuron ddaC | somatic clone

milt⁹²

mitochondrion | adult stage, with Scer\GAL4^dpr1-PGaw

milt^HMC02365

mitochondrion | adult stage, with Scer\GAL4^GMR15B07

milt^JF03022

mitochondrion | adult stage | decreased number, with Scer\GAL4^GMR.PF, Scer\GAL4^elav-C155

milt^GD8116

mitochondrion | adult stage | somatic clone

milt^EY14443

mitochondrion | P-stage, with Scer\GAL4^GMR.PF

milt^HMC02365

mitochondrion | somatic clone | third instar larval stage

milt^33-853

mitochondrion | third instar larval stage, with Scer\GAL4^Cg.PA

milt^JF03022

motor neuron | somatic clone | third instar larval stage

milt^33-853

myofibril, with Scer\GAL4^Mef2.PR

milt^GD10310

Nebenkern | germline clone

milt⁹²

neuropil

milt⁹²

oocyte & mitochondrion | germ-line clone

optic cartridge

milt¹⁸⁶

peripheral nervous system & mitochondrion

milt⁹²

photoreceptor

milt⁹²

photoreceptor cell R7

milt^AM472

photoreceptor cell R8

milt^AM472

photoreceptor neuron, with Scer\GAL4^GMR.PF

presynapse | adult stage, with Scer\GAL4^GMR.PF, Scer\GAL4^elav-C155

milt^GD8116

retina | somatic clone

milt^EY14443

sarcomere, with Scer\GAL4^Mef2.PR

milt^GD10310

subesophageal ganglion & mitochondrion

milt⁹²

synapse

milt⁹²

synaptic vesicle

milt¹⁸⁶

synaptic vesicle, with Scer\GAL4^GMR.PF

milt^GD8116

trichogen cell, with Scer\GAL4^pnr-MD237

milt^GD10310

Z disc, with Scer\GAL4^Mef2.PR

milt^GD10310

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) (4)

Hsap\TRAK1

13 of 14

Yes

Hsap\TRAK2

12 of 14

Yes

Hsap\HAP1

4 of 14

Yes

Hsap\BICDL1

1 of 14

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (4)

Rnor\Trak1

13 of 14

Yes

Rnor\Trak2

12 of 14

Yes

Rnor\Hap1

4 of 14

Yes

Rnor\Bicdl1

1 of 14

Mus musculus (laboratory mouse) (4)

Mmus\Trak1

13 of 14

Yes

Mmus\Trak2

11 of 14

Yes

Mmus\Hap1

4 of 14

Yes

Mmus\Bicdl1

1 of 14

Xenopus tropicalis (Western clawed frog) (6)

Xtro\trak2

10 of 13

Yes

Xtro\hap1

7 of 13

Yes

Xtro\trak1

6 of 13

Yes

Xtro\celf5

1 of 13

Xtro\dkk1

1 of 13

Yes

Xtro\f2rl3

1 of 13

Yes

Danio rerio (Zebrafish) (4)

Drer\trak1a

12 of 14

Yes

Drer\trak2

11 of 14

Yes

Drer\hap1

10 of 14

Yes

Drer\trak1b

1 of 14

Yes

Caenorhabditis elegans (Nematode, roundworm) (1)

Cele\trak-1

9 of 14

Yes

Anopheles gambiae (African malaria mosquito) (2)

Agam\AgaP_AGAP008801

6 of 12

Yes

Agam\AgaP_AGAP008800

2 of 12

Yes

Arabidopsis thaliana (thale-cress) (0)

Saccharomyces cerevisiae (Brewer's yeast) (0)

Schizosaccharomyces pombe (Fission yeast) (0)

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:milt. 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) (0)

Human Disease Associations

FlyBase Human Disease Model Reports

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 2 )

Allele

Disease

Evidence

References

milt^HMC02365

model of neurodegenerative disease

CEA

(Vagnoni et al., 2016)

milt^RNAi.UAS.cUa

model of neurodegenerative disease

CEA

(Fang et al., 2012)

Potential Models Based on Orthology ( 1 )

Human Ortholog

Disease

Evidence

References

TRAK1; trafficking kinesin protein 1

model of developmental and epileptic encephalopathy 68

IEA

(FlyBase, 2019-)

Modifiers Based on Experimental Evidence ( 4 )

Allele

Disease

Interaction

References

milt^EY14443

ameliorates tauopathy

modeled by Hsap\MAPT^GMR.Ex.PJ

(Ambegaokar and Jackson, 2011)

milt^GD8116

exacerbates Parkinson's disease

modeled by Hsap\APP^{Aβ42.UAS.Tag:SS(rPENK)}

(Iijima-Ando et al., 2009)

exacerbates tauopathy

modeled by Hsap\MAPT^UAS.cWa

(Iijima-Ando et al., 2012)

exacerbates amyotrophic lateral sclerosis type 8

modeled by Vap33^{P58S.UAS.Tag:FLAG}

(Sanhueza et al., 2015)

milt^JF03022

exacerbates tauopathy

modeled by Hsap\MAPT^UAS.cWa

(Iijima-Ando et al., 2012)

ameliorates Parkinson's disease

modeled by Pink1^B9

(Liu et al., 2012)

milt^LA00951

ameliorates amyotrophic lateral sclerosis type 8

modeled by Vap33^{P58S.UAS.Tag:FLAG}

(Sanhueza et al., 2015)

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?

TRAK1; trafficking kinesin protein 1

13 of 14

608112

DEVELOPMENTAL AND EPILEPTIC ENCEPHALOPATHY 68; DEE68

developmental and epileptic encephalopathy 68

TRAK2; trafficking kinesin protein 2

12 of 14

607334

HAP1; huntingtin associated protein 1

4 of 14

600947

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

milt - Hsap\OGT

anti bait coimmunoprecipitation, western blot

(Pekkurnaz et al., 2014)

milt - Khc

anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation

(Winding et al., 2016, Glater et al., 2006, Stowers et al., 2002)

milt - Miro

anti tag coimmunoprecipitation, anti tag western blot

(Mattedi et al., 2023, Glater et al., 2006)

milt - Myo10A

two hybrid, pull down, autoradiography

(Liu et al., 2008)

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

milt

suppressible

eIF2β

(Shinno et al., 2026)

milt

suppressible

par-1

(Iijima-Ando et al., 2012)

milt

suppressible

tau

(Iijima-Ando et al., 2012)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

Pink1

suppressible

milt

(Liu et al., 2012)

External Data

Subunit Structure (UniProtKB)

Interacts with Miro (PubMed:16717129). Interacts with OGT (PubMed:24995978). Note=Mitochondrial transport by milt and Khc is independent of Klc.

(UniProt, Q960V3 )

Linkouts

DroID - A comprehensive database of gene and protein interactions.

FBgn0262872

MIST (genetic) - An integrated Molecular Interaction Database

45683

MIST (protein-protein) - An integrated Molecular Interaction Database

45683

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

Class of Gene

gene_with_edited_transcript

(Rodriguez et al., 2012, Graveley et al., 2011)

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

2-24

Cytogenetic map

27D5-27D7

Sequence location

2L:7,037,823..7,056,010 [+]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

27D5-27D7

Limits computationally determined from genome sequence between P{PZ}snRNP70K⁰²¹⁰⁷ and P{PZ}wg⁰²⁶⁵⁷

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

27D5-27D6

(determined by in situ hybridisation)

(Spradling et al., 1999)

27C-27D

(Stowers, 1999.4.1)

27D1-27D2

27D5--6

(BDGP Project Members, 1994-1999)

Experimentally Determined Recombination Data

Location

2-24

(FlyBase, 2016)

2-23.2

(Comeron et al., 2012)

(Stowers and Schwarz, 1999)

Left of (cM)

Right of (cM)

Notes

Maps to 2L.

(Stowers and Schwarz, 1999)

Stocks and Reagents

Stocks (27)

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

Bloomington

49171

w¹¹¹⁸; P{GMR26C11-GAL4}attP2

54701

w¹¹¹⁸; P{GMR26C11-lexA}attP40

19198

w¹¹¹⁸; P{XP}milt^d03910

54700

w¹¹¹⁸; P{GMR26C01-lexA}attP40/CyO

28385

y¹ v¹; P{TRiP.JF03022}attP2

43173

y¹ v¹; P{TRiP.GL01515}attP2

44477

y¹ v¹; P{TRiP.HMC02365}attP2

63855

w¹¹¹⁸; PBac{IT.GAL4}milt^0857-G4

NIG-Fly

31630R-3

P{NIG.31630R}3

HMC02365

y¹ v¹; P{TRiP.HMC02365}attP2

VDRC

v110193

P{KK102369}VIE-260B

v41508

w¹¹¹⁸ P{GD8116}v41508

More stocks available...

Genomic Clones (18)

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

cDNA Clones (139)

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

FI19826

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.

FBgn0262872

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

Antibody Information

Laboratory Generated Antibodies

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

milt is required for the proper anchoring of the Nebenkern to the nucleus and the proper elongation of the mitochondrial derivatives during sperm formation.

(Aldridge et al., 2007)

milt has a role in the transport of mitochondria into the oocyte.

(Cox and Spradling, 2006)

Miro binds to milt and can affect its association with mitochondria.

(Glater et al., 2006)

New annotation (CG31630) in release 3 of the genome annotation.

(FlyBase Genome Annotators, 2002-2003)

milt appears to be essential for proper axonal and synaptic localisation of mitochondria but is required neither generally for axonal transport of all kinesin cargos nor for axonal outgrowth.

(Stowers et al., 2002)

Mutants have a defective electroretinogram; there is no on/off transient.

(Stowers and Schwarz, 1999)

Relationship to Other Genes

Source for database merge of

Source for merge of: milt anon-WO0153538.19

(FlyBase, 1996-)

Source for merge of: milton l(2)k04704

(Stowers, 1999.4.1)

Source for merge of: milt CG31630

(FlyBase Genome Annotators, 2011)

Additional comments

Annotations CG13777 and CG31630 merged as CG43227 in release 5.34 of the genome annotation.

(FlyBase Genome Annotators, 2011)

Source for merge of milt anon-WO0153538.19 was sequence comparison ( date:051113 ).

(FlyBase, 1996-)

Source for identity of milton CG13777 was sequence comparison ( date:011219 ).

(FlyBase, 1996-)

Nomenclature History

Source for database identify of

Source for identity of: milton CG13777

(FlyBase, 1996-)

Nomenclature comments

Etymology

Named after the blind British poet Milton.

(Stowers et al., 2002)

Synonyms and Secondary IDs (11)

Reported As

Symbol Synonym

2L6

(Bellen, 2020.5.15)

CG13777

(Keleman et al., 2009.8.5, Perkins et al., 2009, Liu et al., 2008, Papaconstantinou et al., 2005, Herold et al., 2003)

CG31630

(Japanese National Institute of Genetics, 2012.5.21, Ambegaokar and Jackson, 2011)

CG43227

(National Institute of Genetics Fly Stocks, 2022-, Kim et al., 2021, FlyBase Genome Annotators, 2011, Ni et al., 2010.12.1)

Milt

(Du et al., 2021, Gnainsky et al., 2021, Russell et al., 2018, Liu and Lasko, 2015)

Milton

(Mattedi et al., 2023, Du et al., 2021, Ding et al., 2016)

anon-WO0153538.19

dmilton

(Gupta et al., 2022)

l(2)k04704

(Spradling et al., 1999)

milton

(Gonzalez and Wang, 2020, Chen et al., 2016)

Secondary FlyBase IDs

FBgn0022201
FBgn0031884
FBgn0045937
FBgn0051630

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 74 )

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.

45683

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.

Q960V3

UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information

Q960V3

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.

Q960V3

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0262872

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

45683/tissue=All

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

FBgn0262872

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

FBgn0262872

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

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

dme:Dmel_CG43227

MARRVEL_MODEL - MARRVEL (model organism gene)

45683

Linkouts

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

FBgn0262872

DroID - A comprehensive database of gene and protein interactions.

FBgn0262872

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

FBGN0262872

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

CG13777

FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.

FBgn0262872

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

TC011611

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

45683

MIST (protein-protein) - An integrated Molecular Interaction Database

45683

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

References (141)

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

2.66

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