FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Gene: Dmel\mam
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General Information
Symbol
Dmel\mam
Species
D. melanogaster
Name
mastermind
Annotation Symbol
CG8118
Feature Type
protein_coding_gene
FlyBase ID
FBgn0002643
Gene Model Status
Current
Stock Availability
87 publicly available
Gene Summary
mastermind (mam) encodes a transcriptional coactivator that functions in the Notch signaling pathway. It regulates gene expression by interacting with the intracellular domain of the product of N, which is produced upon receptor activation. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
All Summaries
Also Known As

N-2G, l(2)s9998

Key Links
Genomic Location
Cytogenetic map
50C23-50D3
Sequence location
Recombination map
2-68
RefSeq locus
NT_033778 REGION:13991203..14060356
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (24 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables transcription coactivator activity
inferred from direct assay
(Kitagawa et al., 2001)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables transcription coactivator activity
inferred from electronic annotation with InterPro:IPR019082
(InterPro Project Members, 2004-)
Biological Process (16 terms)
Terms Based on Experimental Evidence (15 terms)
CV Term
Evidence
References
involved_in apposition of dorsal and ventral imaginal disc-derived wing surfaces
inferred from mutant phenotype
(Prout et al., 1997)
involved_in asymmetric cell division
inferred from mutant phenotype
(Yedvobnick et al., 2004)
involved_in follicle cell of egg chamber development
inferred from mutant phenotype
(Jia et al., 2015)
involved_in germ-line stem cell population maintenance
inferred from mutant phenotype
(Song et al., 2007)
involved_in heart morphogenesis
inferred from mutant phenotype
(Albrecht et al., 2006)
involved_in imaginal disc-derived wing margin morphogenesis
inferred from mutant phenotype
(Helms et al., 1999)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Dworkin and Gibson, 2006)
involved_in lateral inhibition
inferred from mutant phenotype
(Helms et al., 1999)
involved_in mesodermal cell fate determination
inferred from mutant phenotype
(Corbin et al., 1991)
involved_in neuroblast fate determination
inferred from mutant phenotype
(Bodmer et al., 1993)
involved_in neuron fate determination
inferred from genetic interaction with FLYBASE:N; FB:FBgn0004647
(Schuldt and Brand, 1999)
involved_in Notch signaling pathway
inferred from genetic interaction with FLYBASE:dx; FB:FBgn0000524
(Xu and Artavanis-Tsakonas, 1990)
inferred from direct assay
(Gomez-Lamarca et al., 2018)
inferred from mutant phenotype
(Helms et al., 1999)
inferred from genetic interaction with FLYBASE:Delta; FB:FBgn0000463
(Helms et al., 1999)
inferred from genetic interaction with FLYBASE:N; FB:FBgn0004647
(Helms et al., 1999, Schuldt and Brand, 1999, Xu et al., 1990)
involved_in positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Kitagawa et al., 2001)
involved_in somatic stem cell population maintenance
inferred from mutant phenotype
(Vied and Kalderon, 2009)
involved_in ventral midline development
inferred from mutant phenotype
(Zhang et al., 2011)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
involved_in CENP-A containing chromatin assembly
inferred from biological aspect of ancestor with PANTHER:PTN001046424
(GO Reference Genome Project, 2011-)
involved_in Notch signaling pathway
inferred from electronic annotation with InterPro:IPR019082
(InterPro Project Members, 2004-)
involved_in positive regulation of transcription by RNA polymerase II
inferred from electronic annotation with InterPro:IPR019082
(InterPro Project Members, 2004-)
Cellular Component (7 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
part_of CSL-Notch-Mastermind transcription factor complex
inferred from direct assay
(Gomez-Lamarca et al., 2018, Contreras et al., 2015, Kitagawa et al., 2001)
inferred from physical interaction with FLYBASE:N; FB:FBgn0004647,FLYBASE:Su(H); FB:FBgn0004837
(Maier et al., 2011)
located_in nucleus
inferred from direct assay
(Bettler et al., 1996)
located_in polytene chromosome puff
inferred from direct assay
(Gause et al., 2006)
located_in polytene chromosome
inferred from direct assay
(Gause et al., 2006)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
is_active_in chromatin
inferred from biological aspect of ancestor with PANTHER:PTN001046424
(GO Reference Genome Project, 2011-)
is_active_in chromosome, centromeric region
inferred from biological aspect of ancestor with PANTHER:PTN001046424
(GO Reference Genome Project, 2011-)
located_in nuclear speck
inferred from electronic annotation with InterPro:IPR019082
(InterPro Project Members, 2004-)
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN001046424
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
Belongs to the mastermind family. (P21519)
Protein Signatures (InterPro)
Summaries
Gene Snapshot
mastermind (mam) encodes a transcriptional coactivator that functions in the Notch signaling pathway. It regulates gene expression by interacting with the intracellular domain of the product of N, which is produced upon receptor activation. [Date last reviewed: 2019-03-14]
Pathway (FlyBase)
NOTCH SIGNALING PATHWAY CORE COMPONENTS -
The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. (Adapted from FBrf0225731 and FBrf0192604). Core pathway components are required for signaling from the sending cell and response in the receiving cell.
Gene Group (FlyBase)
CSL-NOTCH-MASTERMIND TRANSCRIPTION FACTOR COMPLEX -
The CSL-Notch-Mastermind transcription factor complex consisting of the CSL protein, Su(H), and mastermind (mam) protein, in complex with Notch intracellular domain (generated by ligand-stimulated N-cleavage), up-regulates transcription of Notch-responsive genes. (Adapted from FBrf0232880).
Protein Function (UniProtKB)
May have a regulatory function possibly in association with the Notch gene product.
(UniProt, P21519)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
mam: master mind (J.C. Hall)
Homozygous embryonic lethal; embryos display neural hyperplasia with compensatory epidermal hypoplasia; caused by failure of most ventral ectodermal cells to differentiate as epidermal cells rather than neuroblasts, as seen in N, amx, bib, neu, Dl, and E(spl); mam tends to have less extreme neural hyperplasia than mutants at the other neurogenic loci. A similar diversion of cells from epidermigenic into neurogenic pathways seen to generate supernumerary peripheral nerve cells (Hartenstein and Campos-Ortega, 1986, Roux's Arch. Dev. Biol. 195: 210-21 (fig.)]. When mam expressed in female germ cells and the ensuing embryos, neural hyperplasia is enhanced, but mam+ embryos from oocytes are normal [Jimenez and Campos Ortega, 1982, Roux's Arch. Dev. Biol. 191: 1901-201 (fig.)]. Homozygous clones in the eye display irregular ommatidial pattern characterized by lack of interommatidial bristles, enlarged facets with supernumerary retinular cells and reduced numbers of pigment cells; in the cuticle, clones homozygous for mam2 are devoid of bristles [Dietrich and Campos-Ortega, 1984, J. Neurogenet. 1: 315-32 (fig.)]. mam1 (formerly N-2G) heterozygotes occasionally exhibit apical wing nicking; not recorded for other alleles. Phenotype of homozygotes for null allele reduced by duplications for normal alleles of other neurogenic loci, N, neu, Dl, E(spl), and H, but not amx or bib (de la Concha, Dietrich, Weigel, and Campos-Ortega, 1988, Genetics 118: 499-508).
Summary (Interactive Fly)

novel protein that physically interacts with Suppressor of Hairless and the intracellular domain of Notch that is produced upon receptor activation - functions as a transcriptional coactivator for Notch signaling

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
2

Please see the JBrowse view of Dmel\mam 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 P21519)

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.

Gene model reviewed during 5.50

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
mam-RA
FBtr0087596
NM_080376
6308
1594
mam-RB
FBtr0087597
NM_166030
7590
1594
mam-RC
FBtr0087598
NM_206109
1941
433
mam-RD
FBtr0339956
NM_001299472
7884
1594
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
mam-PA
FBpp0086722
167.3
1594
8.19
P21519
NP_525115
AAF58299
mam-PB
FBpp0086723
167.3
1594
8.19
P21519
NP_725354
AAF58300
mam-PC
FBpp0086724
44.0
433
5.00
NP_995831
AAS64855
mam-PD
FBpp0308978
167.3
1594
8.19
P21519
NP_001286401
AHN56199
Polypeptides with Identical Sequences

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

1594 aa isoforms: mam-PA, mam-PB, mam-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

1596 (aa); 167 (kD predicted)

(Smoller et al., 1990)
Comments
External Data
Domain

The protein has many AA homomeric domains: 21 poly-Gln runs (from 5 to 16 AA in length), 4 poly-Gly (6 to 10 AA), 3 poly-Asn (3 X 5 AA), 1 poly-Ala (10 AA) and 1 poly-Thr (5 AA) runs.

(UniProt, P21519)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

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

-0.77

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | ubiquitous
(Bettler et al., 1991)
gastrula stage
anterior midgut primordium
(Bettler et al., 1991)
endoderm | presumptive
(Bettler et al., 1991)
mesoderm | presumptive
(Bettler et al., 1991)
cephalic furrow
(Schmid et al., 1996)
ventral furrow
(Schmid et al., 1996)
foregut primordium & inclusive hindgut primordium
(Schmid et al., 1996)
embryonic stage 6
ventral furrow
(Bettler et al., 1991)
embryonic stage 6 -- 8
ventral neurectoderm | restricted

Comment: late gastrula stage

(Bettler et al., 1991)
embryonic stage 8
mesectoderm
(Bettler et al., 1991)
posterior midgut primordium
(Bettler et al., 1991)
ectoderm | ventral
(Bettler et al., 1991)
cephalic furrow
(Bettler et al., 1991)
extended germ band stage
germ band
(Schmid et al., 1996)
embryonic stage 13
embryonic/larval foregut
(Schmid et al., 1996)
ventral midline of embryo
(Schmid et al., 1996)
inclusive hindgut primordium
(Schmid et al., 1996)
larval ventral nerve cord | segmentally repeated
(Schmid et al., 1996)
embryonic stage 13 -- 16
larval midline neuron
(Kearney et al., 2004)
embryonic stage 13 -- 17
larval ventral nerve cord
(Schmid et al., 1996)
larval stage
eye-antennal disc
(Schmid et al., 1996)
leg disc
(Schmid et al., 1996)
eye disc morphogenetic furrow | anterior to
(Schmid et al., 1996)
wing disc
(Schmid et al., 1996)
pharate adult stage
central nervous system | restricted
(Schmid et al., 1996)
adult stage & oogenesis stage S10
nurse cell
(Schmid et al., 1996)
oocyte
(Schmid et al., 1996)
follicle cell
(Schmid et al., 1996)
oogenesis stage S1 -- S6
nurse cell
(Schmid et al., 1996)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 4

Comment: reference states 0-2 hr AEL

(Smoller et al., 1990)
embryonic stage 5 -- 16

Comment: reference states 2-16 hr AEL

(Smoller et al., 1990)
embryonic stage 12 -- 16

Comment: reference states 8-16 hr AEL

(Smoller et al., 1990)
third instar larval stage
organism
(Smoller et al., 1990)
prepupal stage P1 -- P2
organism
(Smoller et al., 1990)
radioisotope in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 10
organism | ubiquitous

Comment: reference states 5 hr AEL

(Smoller et al., 1990)
embryonic stage 16
larval brain

Comment: reference states 16 hr AEL

(Smoller et al., 1990)
larval ventral nerve cord

Comment: reference states 16 hr AEL

(Smoller et al., 1990)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
blastoderm stage
organism | ubiquitous
(Bettler et al., 1996)
gastrula stage
cephalic furrow
(Bettler et al., 1991)
embryonic stage 7 -- 8
ventral midline of embryo
(Bettler et al., 1996)
posterior midgut primordium
(Bettler et al., 1996)
epidermis | dorsal | lateral
(Bettler et al., 1996)
anterior midgut primordium
(Bettler et al., 1996)
ventral neurectoderm
(Bettler et al., 1996)
embryonic/larval hindgut | presumptive
(Bettler et al., 1996)
ectoderm
(Bettler et al., 1996)
cephalic furrow
(Bettler et al., 1996)
embryonic stage 8
mesoderm
(Bettler et al., 1991)
embryonic stage 9 -- 10
neuroblast MP2
(Bettler et al., 1996)
embryonic stage 9 -- 17
ventral neurectoderm
(Bettler et al., 1991)
mesectoderm
(Bettler et al., 1991)
embryonic stage 11
Malpighian tubule
(Bettler et al., 1996)
epidermis
(Bettler et al., 1996)
embryonic/larval hindgut
(Bettler et al., 1996)
embryonic stage 14 -- 15
embryonic/larval proventriculus
(Bettler et al., 1996)
embryonic/larval esophagus
(Bettler et al., 1996)
neuroblast MP2
(Bettler et al., 1996)
gnathal ganglion
(Bettler et al., 1996)
inclusive hindgut primordium
(Bettler et al., 1996)
embryonic/larval pharynx
(Bettler et al., 1996)
embryonic/larval salivary gland
(Bettler et al., 1996)
embryonic/larval midgut
(Bettler et al., 1996)
neuroblast
(Bettler et al., 1996)
embryonic stage 14 -- 17
larval ventral nerve cord
(Bettler et al., 1996)
embryonic stage 17
embryonic/larval hindgut

Comment: early stage 17

(Bettler et al., 1996)
Malpighian tubule

Comment: early stage 17

(Bettler et al., 1996)
embryonic/larval foregut

Comment: early stage 17

(Bettler et al., 1996)
central nervous system
(Bettler et al., 1996)
larval antenno-maxillary sensory complex | restricted
(Bettler et al., 1996)
embryonic/larval midgut

Comment: early stage 17

(Bettler et al., 1996)
third instar larval stage
wing disc
(Bettler et al., 1996)
leg disc
(Bettler et al., 1996)
eye-antennal disc
(Bettler et al., 1996)
larval ventral nerve cord | restricted
(Bettler et al., 1996)
larval brain
(Bettler et al., 1996)
neuroblast
(Bettler et al., 1996)
adult optic lobe
(Bettler et al., 1996)
adult stage & oogenesis
follicle cell
(Bettler et al., 1996)
Additional Descriptive Data

The nuclear localization of mam protein is consistent with its proposed role in gene regulation. Early in embryogenesis, mam protein shows nuclear, punctate localization. Immunohistochemical studies show that mam protein binds to more than 100 locations on polytene chromosomes, some of which colocalize with a subset of RNA polymerase and gro protein binding sites.

(Bettler et al., 1996)

The pattern of mam protein localization is comparable to that of mam transcript. During gastrulation, the highest levels of mam protein are found in the ventral furrow, although expression is also seen in the cephalic furrow. Late in gastrulation, mam protein accumulates in the mesoderm and in the midgut primordia. At the completion of gastrulation, there are high levels of mam protein in the mesectoderm, the neurectoderm, as well as in the midgut primordia and the cephalic furrow.

(Bettler et al., 1991, Smoller et al., 1990)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
part_of CSL-Notch-Mastermind transcription factor complex
inferred from direct assay
(Gomez-Lamarca et al., 2018, Contreras et al., 2015, Kitagawa et al., 2001)
inferred from physical interaction with FLYBASE:N; FB:FBgn0004647,FLYBASE:Su(H); FB:FBgn0004837
(Maier et al., 2011)
located_in nucleus
inferred from direct assay
(Bettler et al., 1996)
located_in polytene chromosome puff
inferred from direct assay
(Gause et al., 2006)
located_in polytene chromosome
inferred from direct assay
(Gause et al., 2006)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\mam in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
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
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 194 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 18 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of mam
Transgenic constructs containing regulatory region of mam
Aberrations (Deficiencies and Duplications) ( 51 )
Inferred from experimentation ( 51 )
Gene disrupted in
Df(2R)50C-16
(Preston et al., 1996)
Df(2R)50C-36
(Preston et al., 1996)
Df(2R)50C-38
(Spradling et al., 1999, Preston et al., 1996, BDGP Project Members, 1994-1999)
Df(2R)50C-40
(Preston et al., 1996)
Df(2R)50C-101
(Spradling et al., 1999, Preston et al., 1996, BDGP Project Members, 1994-1999)
Df(2R)50C-128
(Preston et al., 1996)
Df(2R)BSC18
(Albrecht et al., 2006)
Df(2R)BSC383
(Christensen et al., 2007.10.29)
Df(2R)CX1
(Albrecht et al., 2006, Schuldt and Brand, 1999, Spradling et al., 1999, Prout et al., 1997, Hu et al., 1995, BDGP Project Members, 1994-1999)
Df(2R)Exel7129
(Ryder, 2004.4.26)
Df(2R)mam04615
(Preston et al., 1996)
In(2R)G
(Tearle and Nusslein-Volhard, 1987)
In(2R)vgU
(Simmonds et al., 1997, Williams et al., 1990)
In(2R)vgUR2
(Williams et al., 1990)
In(2R)vgURK1
(Simmonds et al., 1997)
In(2R)vgURK2
(Simmonds et al., 1997)
In(2R)vgURK3
(Simmonds et al., 1997)
In(2R)vgURK4
(Simmonds et al., 1997)
In(2R)vgURK5
(Simmonds et al., 1997)
In(2R)vgURK6
(Simmonds et al., 1997)
In(2R)vgURK7
(Simmonds et al., 1997)
In(2R)vgURK9
(Simmonds et al., 1997)
In(2R)vgURK11
(Simmonds et al., 1997)
In(2R)vgURK12
(Simmonds et al., 1997)
In(2R)vgURK13
(Simmonds et al., 1997)
In(2R)vgURK14
(Simmonds et al., 1997)
In(2R)vgURK15
(Simmonds et al., 1997)
In(2R)vgURK16
(Simmonds et al., 1997)
In(2R)vgURK17
(Simmonds et al., 1997)
Gene partially disrupted in
Df(2R)8328
(Wu et al., 2017)
Df(2R)MK1
(Schuldt and Brand, 1999)
Gene not disrupted in
Df(2R)50C-21
(Preston et al., 1996)
Df(2R)50C-43
(Preston et al., 1996)
Df(2R)50C-45
(Preston et al., 1996)
Df(2R)50C-49
(Preston et al., 1996)
Df(2R)50C-68
(Preston et al., 1996)
Df(2R)50C-69
(Preston et al., 1996)
Df(2R)50C-71
(Preston et al., 1996)
Df(2R)50C-81
(Preston et al., 1996)
Df(2R)50C-85
(Preston et al., 1996)
Df(2R)50C-89
(Preston et al., 1996)
Df(2R)50C-102
(Preston et al., 1996)
Df(2R)50C-107
(Preston et al., 1996)
Df(2R)50C-111
(Preston et al., 1996)
Df(2R)50C-116
(Preston et al., 1996)
Df(2R)50C-129
(Preston et al., 1996)
Df(2R)50C-131
(Preston et al., 1996)
Df(2R)50C-132
(Preston et al., 1996)
Df(2R)50C-133
(Preston et al., 1996)
Df(2R)vg135
(BDGP Project Members, 1994-1999)
Df(2R)vg-B
(BDGP Project Members, 1994-1999)
Inferred from location ( 7 )
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 (with Df(2R)BSC383)
lethal | recessive
lethal - all die during embryonic stage
lethal - all die during embryonic stage | neurogenic phenotype | recessive
lethal - all die during embryonic stage | recessive
partially lethal - majority die
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
Sterility
Allele
female sterile
fertile
male sterile
Other Phenotypes
Allele
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4arm.PS
abnormal neuroanatomy | adult stage, with Scer\GAL4ato.3.6
abnormal neuroanatomy | embryonic stage 13
abnormal neuroanatomy | embryonic stage 16
abnormal neuroanatomy | embryonic stage 16 (with mam8)
abnormal neuroanatomy | embryonic stage 16 (with mamΔC)
abnormal neuroanatomy | embryonic stage, with Scer\GAL4elav-C155
abnormal neuroanatomy | heat sensitive, with Scer\GAL4hs.PH
abnormal neuroanatomy | larval stage, with Scer\GAL4elav-C155
abnormal neuroanatomy | somatic clone
abnormal size | third instar larval stage, with Scer\GAL4en.PU
chemical sensitive, with Scer\GAL4elav-C155
increased cell number | embryonic stage
neurogenic phenotype
some die during embryonic stage
some die during embryonic stage | recessive
visible
visible, with Scer\GAL4309
visible, with Scer\GAL4ap-md544
visible, with Scer\GAL4bbg-C96
visible, with Scer\GAL4bi-omb-Gal4
visible, with Scer\GAL4Bx-MS1096-KE
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4GMR.PF
visible, with Scer\GAL4hs.2sev
visible, with Scer\GAL4pnr-MD237
visible | adult stage, with Scer\GAL4309
visible | adult stage, with Scer\GAL4309, mamEP2022.A
visible | adult stage, with Scer\GAL4309, mamH.UAS
visible | adult stage, with Scer\GAL4bbg-C96
visible | adult stage, with Scer\GAL4bbg-C96, mamEP2022.A
visible | adult stage, with Scer\GAL4bbg-C96, mamH.UAS
visible | adult stage, with Scer\GAL4Dll.PU
visible | adult stage, with Scer\GAL4pnr-MD237
visible | adult stage, with Scer\GAL4pnr-MD237, mamEP2022.A
visible | adult stage, with Scer\GAL4pnr-MD237, mamN.UAS
visible | dominant
visible | heat sensitive, with Scer\GAL4hs.PH
visible | homeotic, with Scer\GAL4ey.PB
visible | recessive | somatic clone
visible | somatic clone
Phenotype manifest in
Allele
adult olfactory receptor neuron Or49a/85f
antenna | ectopic, with Scer\GAL4ey.PB
antennal lobe glomerulus DA1
antennal lobe glomerulus DA3
antennal lobe glomerulus DA4l
antennal lobe glomerulus DC1
antennal lobe glomerulus DL1
antennal lobe glomerulus DL4
antennal lobe glomerulus DM1
antennal lobe glomerulus DM2
antennal lobe glomerulus DM3
antennal lobe glomerulus DM4
antennal lobe glomerulus V
antennal lobe glomerulus VA4
antennal lobe glomerulus VC1
antennal lobe glomerulus VC2
antennal lobe glomerulus VC3
antennal lobe glomerulus VC4
antennal lobe glomerulus VL1
antennal lobe glomerulus VM1
antennal lobe glomerulus VM4
anterior fascicle & synapse | supernumerary, with Scer\GAL4elav-C155
apterous-expressing neuron of the lateral cluster of the thorax, with Scer\GAL4elav-C155
cone cell | P-stage, with Scer\GAL4pros.PSG
cuticle & macrochaeta | somatic clone
dMP2 neuron
ectoderm
ectoderm | heat sensitive, with Scer\GAL4hs.PH
egg chamber | somatic clone
egg chamber | somatic clone, with Scer\GAL4Act.PU, Scer\GAL4Tub.PU
embryo | germline clone | maternal effect
embryonic/larval cuticle
embryonic/larval cuticle | ventral
embryonic/larval dorsal vessel | precursor
embryonic cardioblast | increased number
embryonic epidermis
embryonic epidermis | lateral
embryonic epidermis | ventral
embryonic heart cardioblast | increased number
embryonic heart ostial cardioblast | increased number
embryonic hypopharynx & epidermis
embryonic labial segment & embryonic epidermis
embryonic labral segment & embryonic epidermis
embryonic maxillary segment & embryonic epidermis
embryonic myoblast
embryonic pericardial cell
embryonic posterior midline glial cell | embryonic stage 13
external sensory organ precursor cell | embryonic stage
eye, with Scer\GAL4dpp.blk1
eye, with Scer\GAL4ey.PB
eye, with Scer\GAL4ey.PH
eye, with Scer\GAL4GMR.PF
eye, with Scer\GAL4hs.2sev
eye | heat sensitive, with Scer\GAL4hs.PH
eye | somatic clone
eye disc morphogenetic furrow, with Scer\GAL4dpp.blk1
eye photoreceptor cell | somatic clone
female germline stem cell, with Scer\GAL4C587
follicle stem cell | somatic clone
germarium cap cell, with Scer\GAL4C587
germline cell | male, with Scer\GAL4ptc-559.1
heart primordium
interommatidial bristle | somatic clone
larval A-subperineurial glial cell
larval B-subperineurial glial cell
larval central nervous system
larval DA1 motor neuron
larval H-cell | embryonic stage 16 | increased number
larval H-cell sib | embryonic stage 16
larval head
larval intersegmental nerve, with Scer\GAL4elav-C155
larval neuron | ectopic | heat sensitive, with Scer\GAL4hs.PH
larval RP2 motor neuron
larval RP2 motor neuron | ectopic
larval RP2 motor neuron | ectopic | heat sensitive, with Scer\GAL4hs.PH
larval segmental nerve | embryonic stage
larval somatic muscle cell
larval ventral nerve cord
larval ventral nerve cord | embryonic stage
larval VUM interneuron | embryonic stage 16
larval VUM motor neuron | embryonic stage 16 | increased number
lateral ventral subperineurial glial cell
leg, with Scer\GAL4Dll.PU
lobula columnar neuron LC14, with Scer\GAL4ato.3.6
macrochaeta, with Scer\GAL4309
macrochaeta, with Scer\GAL4pnr-MD237
macrochaeta | ectopic, with Scer\GAL4309
macrochaeta | ectopic, with Scer\GAL4dpp.blk1
macrochaeta | ectopic, with Scer\GAL4pnr-MD237
macrochaeta | increased number | heat sensitive, with Scer\GAL4hs.PH
macrochaeta | somatic clone
medulla, with Scer\GAL4ato.3.6
mesonotum, with Scer\GAL4pnr-MD237
mesonotum, with Scer\GAL4pnr-MD237, mamEP2022.A
mesonotum, with Scer\GAL4pnr-MD237, mamN.UAS
microchaeta, with Scer\GAL4pnr-MD237
microchaeta, with Scer\GAL4pnr-MD237, mamEP2022.A
microchaeta, with Scer\GAL4pnr-MD237, mamN.UAS
microchaeta | ectopic, with Scer\GAL4pnr-MD237
microchaeta | somatic clone
midline glial cell | embryonic stage 16
midline glial cell | embryonic stage 16 (with mam8)
midline glial cell | embryonic stage 16 (with mamΔC)
midline glial cell | embryonic stage 16 | increased number
multidendritic neuron | ectopic
nervous system
neuroblast
neuroblast MNB | embryonic stage 16
notopleural bristle, with Scer\GAL4309
olfactory receptor neuron
ommatidium, with Scer\GAL4dpp.blk1
ommatidium | somatic clone
ovary
pCC neuron
pigment cell | somatic clone
posterior notopleural bristle, with Scer\GAL4309
posterior notopleural bristle, with Scer\GAL4309, mamEP2022.A
posterior notopleural bristle, with Scer\GAL4309, mamH.UAS
posterior notopleural bristle | ectopic, with Scer\GAL4309
presumptive embryonic/larval central nervous system
presumptive embryonic/larval central nervous system, with Scer\GAL4arm.PS
presumptive embryonic/larval nervous system
presumptive embryonic/larval peripheral nervous system, with Scer\GAL4arm.PS
RP2sib neuron
RP2sib neuron | heat sensitive, with Scer\GAL4hs.PH
scutellar bristle
scutellar bristle, with Scer\GAL4309
scutellar bristle | ectopic, with Scer\GAL4309
sensory mother cell, with Scer\GAL4pnr-MD237
stomatogastric nervous system primordium
tendon cell
thecogen cell | somatic clone
tormogen cell | somatic clone
trichogen cell | somatic clone
U neuron
Usib neuron
ventral denticle belt
vMP2 neuron
wing
wing, with Scer\GAL4bbg-C96
wing, with Scer\GAL4bi-omb-Gal4
wing, with Scer\GAL4Bx-MS1096-KE
wing, with Scer\GAL4dpp.blk1
wing | anterior
wing | heat sensitive, with Scer\GAL4hs.PH
wing | somatic clone
wing disc posterior compartment | third instar larval stage, with Scer\GAL4en.PU
wing margin
wing margin, with Scer\GAL4bbg-C96
wing margin, with Scer\GAL4bbg-C96, mamEP2022.A
wing margin, with Scer\GAL4bbg-C96, mamH.UAS
wing margin | somatic clone
wing margin bristle, with Scer\GAL4bbg-C96
wing vein
wing vein, with Scer\GAL4Bx-MS1096-KE
wing vein | somatic clone
wing vein L2
wing vein L3
wing vein L3, with Scer\GAL4ap-md544
wing vein L3, with Scer\GAL4dpp.blk1
wing vein L4
wing vein L5, with Scer\GAL4ap-md544
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (3)
Hsap\MAML1
1 of 14
Yes
Yes
Hsap\MAML2
1 of 14
Yes
Yes
Hsap\MAML3
1 of 14
Yes
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (2)
Rnor\Maml1
1 of 14
Yes
Yes
Rnor\Maml2
1 of 14
Yes
Yes
Mus musculus (laboratory mouse) (3)
Mmus\Maml1
1 of 14
Yes
Yes
Mmus\Maml2
1 of 14
Yes
Yes
Mmus\Maml3
1 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (4)
Xtro\b3gnt3.1
1 of 13
Yes
No
Xtro\LOC100494334
1 of 13
Yes
No
Xtro\maml2
1 of 13
Yes
No
Xtro\maml3
1 of 13
Yes
Yes
Danio rerio (Zebrafish) (3)
Drer\maml1
2 of 14
Yes
Yes
Drer\maml3
2 of 14
Yes
Yes
Drer\zmp:0000001236
1 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (2)
Agam\AgaP_AGAP010410
7 of 12
Yes
Yes
Agam\AgaP_AGAP006460
1 of 12
No
No
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:mam. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
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 ( 4 )
    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
    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
    mam
    enhanceable
    Ada2b
    (Gause et al., 2006)
    mam
    enhanceable
    arm
    (Helms et al., 1999)
    mam
    suppressible
    arm
    (Helms et al., 1999)
    mam
    suppressible
    Atg1
    (Kwon et al., 2013)
    mam
    suppressible
    Awh
    (Singh et al., 2025)
    mam
    enhanceable
    CG14767
    (Kankel et al., 2007)
    mam
    enhanceable
    ct
    (Helms et al., 1999)
    mam
    enhanceable
    Delta
    (Alexander et al., 2006, Helms et al., 1999)
    mam
    suppressible
    Delta
    (Helms et al., 1999)
    mam
    enhanceable
    dom
    (Gause et al., 2006)
    mam
    suppressible
    dsh
    (Helms et al., 1999)
    mam
    enhanceable
    dx
    (Helms et al., 1999)
    mam
    enhanceable
    EcR
    (Kwon et al., 2013)
    mam
    suppressible
    emc
    (Kwon et al., 2013)
    mam
    suppressible
    f06222
    (Kankel et al., 2007)
    mam
    enhanceable
    fz2
    (Helms et al., 1999)
    mam
    suppressible
    GPHR
    (Kankel et al., 2007)
    mam
    enhanceable
    H
    (Helms et al., 1999)
    mam
    suppressible
    H
    (Helms et al., 1999)
    mam
    enhanceable
    klu
    (Kankel et al., 2007)
    mam
    enhanceable
    lilli
    (Kwon et al., 2013)
    mam
    enhanceable
    Lk6
    (Kwon et al., 2013)
    mam
    suppressible
    Lk6
    (Kwon et al., 2013)
    mam
    enhanceable
    lola
    (Kwon et al., 2013)
    mam
    enhanceable
    mahe
    (Surabhi et al., 2015)
    mam
    suppressible
    Mrp4
    (Kankel et al., 2007)
    mam
    suppressible
    mts
    (Kankel et al., 2007)
    mam
    enhanceable
    N
    (Zhang et al., 2011, Helms et al., 1999)
    mam
    suppressible
    N
    (Helms et al., 1999)
    mam
    enhanceable
    Nipped-A
    (Gause et al., 2006, Rollins et al., 1999)
    mam
    enhanceable
    Nipped-B
    (Rollins et al., 1999)
    mam
    enhanceable
    Pabp2
    (Kwon et al., 2013)
    mam
    enhanceable
    Psn
    (Yedvobnick et al., 2001)
    mam
    enhanceable
    pum
    (Kwon et al., 2013)
    mam
    suppressible
    RpL13A
    (Alexander et al., 2006)
    mam
    enhanceable
    RpL13A
    (Alexander et al., 2006)
    mam
    enhanceable
    RpL38
    (Rollins et al., 1999)
    mam
    enhanceable
    sca
    (Kankel et al., 2007)
    mam
    suppressible
    Sema2b
    (Joo et al., 2013)
    mam
    enhanceable
    Ser
    (Helms et al., 1999)
    mam
    suppressible
    Ser
    (Helms et al., 1999)
    mam
    enhanceable
    sgg
    (Helms et al., 1999)
    mam
    enhanceable
    Sin3A
    (Kankel et al., 2007)
    mam
    enhanceable
    sno
    (Yedvobnick et al., 2001)
    mam
    enhanceable
    Su(H)
    (Helms et al., 1999, Fortini and Artavanis-Tsakonas, 1994)
    mam
    suppressible
    tara
    (Kwon et al., 2013)
    mam
    suppressible
    wdb
    (Kwon et al., 2013)
    mam
    enhanceable
    wg
    (Helms et al., 1999)
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    arm
    enhanceable
    mam
    (Kankel et al., 2007)
    Caf1-105
    enhanceable
    mam
    (Yu et al., 2013)
    crb
    suppressible
    mam
    (Richardson and Pichaud, 2010)
    ct
    enhanceable
    mam
    (Morcillo et al., 1996)
    dom
    enhanceable
    mam
    (Kwon et al., 2013)
    Dsor1
    enhanceable
    mam
    (Karim et al., 1996)
    EndoGI
    suppressible
    mam
    (O'Keefe et al., 2011)
    Eogt|Dcr-2
    suppressible
    mam
    (Müller et al., 2013)
    H
    suppressible
    mam
    (Johannes and Preiss, 2002)
    hh
    suppressible
    mam
    (Vied and Kalderon, 2009)
    l(1)sc
    enhanceable
    mam
    (Rottgen et al., 1998)
    mys
    enhanceable
    mam
    (Prout et al., 1997)
    N
    enhanceable
    mam
    (Chang et al., 2021, Verheyen et al., 1996)
    N
    suppressible
    mam
    (Verheyen et al., 1996, Xu et al., 1990)
    noc|elB
    suppressible
    mam
    (Luque and Milan, 2007)
    numb
    suppressible
    mam
    (Rajan et al., 2023)
    numb|fru
    suppressible
    mam
    (Rajan et al., 2023)
    Raf
    enhanceable
    mam
    (Karim et al., 1996)
    Ras85D
    suppressible
    mam
    (Karim et al., 1996)
    Rbf
    enhanceable
    mam
    (Kwon et al., 2013)
    Ser
    suppressible
    mam
    (Helms et al., 1999)
    sev|Ras85D
    suppressible
    mam
    (Maixner et al., 1998)
    Su(H)
    enhanceable
    mam
    (Verheyen et al., 1996)
    Su(H)
    suppressible
    mam
    (Helms et al., 1999)
    su(Hw)|ct
    enhanceable
    mam
    (Morcillo et al., 1996)
    Uvrag
    suppressible
    mam
    (Lee et al., 2011)
    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
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-68
    Cytogenetic map
    50C23-50D3
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    50C23-50D3
    Limits computationally determined from genome sequence between P{lacW}Cp1k15819&P{EP}CG6701EP2054 and P{lacW}mamk05821
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    50D1-50D2
    (determined by in situ hybridisation)
    (Spradling et al., 1999)
    50C20-50C23
    (determined by in situ hybridisation)
    (Preston et al., 1996, Lozovskaya et al., 1993)
    50C20-50C21
    50C22--23
    (BDGP Project Members, 1994-1999)
    50C-50D
    (determined by in situ hybridisation)
    (Smoller et al., 1990, Yedvobnick et al., 1988)
    50C11-50D4
    (determined by in situ hybridisation)
    (Weigel et al., 1987)
    Experimentally Determined Recombination Data
    Location

    2-68

    (FlyBase, 2016)

    2-73.8

    (Comeron et al., 2012)

    2-71

    (Tearle and Nusslein-Volhard, 1987)

    2-70.3

    Left of (cM)
    (Preston and Engels, 1996)
    Right of (cM)
    Notes

    Determined by deletion mapping.

    (Preston and Engels, 1996)
    Stocks and Reagents
    Stocks (87)
    Genomic Clones (58)
    cDNA Clones (56)
     

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

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

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

    cDNA Clones, End Sequenced (ESTs)
    BDGP DGC clones
    Other clones
    RNAi and Array Information
    Linkouts
    DRSC - Results frm RNAi screens
    Antibody Information
    Laboratory Generated Antibodies
     

    monoclonal

    (Bettler et al., 1996)

    polyclonal

    (Smoller et al., 1990)
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       
        Other Comments

        mam is required for normal follicle stem cell maintenance in the ovary, but this function does not act through its known role in the N signalling pathway.

        (Vied and Kalderon, 2009)

        The N signaling pathway is important for the formation and maintenance of the germline stem cell niche in the ovary.

        (Song et al., 2007)

        RNAi generated by PCR using primers directed to this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R+ cells.

        (Boutros et al., 2004)

        mam acts downstream of N to specify neuronal cell fates in the embryonic central nervous system.

        (Schuldt and Brand, 1999)

        Five EMS-induced alleles have been isolated that suppress the macrochaetae, microchaetae and wing vein phenotypes of NAx-16, and the lethality of NAx-9/NAx-E2.

        (Go and Artavanis-Tsakonas, 1998)

        2 alleles of mam have been isolated in a genetic screen for autosomal mutations that produce blisters in somatic wing clones.

        (Prout et al., 1997)

        mam protein is widely expressed during embryonic and postembryonic development. Within salivary glands mam protein binds to over 100 polytene chromosome sites. Comparison of mams basic domain sequence to known proteins suggest that it may be distantly related to a subset of the leucine zipper class of transcription factors.

        (Bettler et al., 1996)

        Chi, but not mam, modulates expression of a gypsy insertion into Ubx. Chi and mam regulate ct alleles lacking gypsy insertions. sd, mam and Chi cooperate synergistically to regulate ct expression.

        (Morcillo et al., 1996)

        mam transcription is highly conserved in D.melanogaster and D.virilis. Results also identify an unusual perinuclear distribution of RNA.

        (Schmid et al., 1996)

        Mutations (unspecified) enhance the rough eye phenotype of SerTM.sev.T:VSV\G and DlTM.sev flies.

        (Sun and Artavanis-Tsakonas, 1996)

        Variation of a microsatellite within the mam locus has been studied in North American populations of D.melanogaster.

        (Goldstein and Clark, 1995)

        Mutations at fs(1)Yb show genetic interactions with the N group of neurogenic genes.

        (Johnson et al., 1995)

        Mutations can act as dominant modifiers of the activated N eye phenotype (FBrf0064452).

        (Verheyen et al., 1995)

        Su(H) shows allele specific interactions with N, Dl, dx and mam.

        (Fortini and Artavanis-Tsakonas, 1994)

        mam is a neurogenic gene required initially to ensure the correct number of PNS precursors. mam continues to be required in the peripheral ectoderm, possibly for maintenance of the commitment to an epidermal fate.

        (Bodmer et al., 1993)

        NM1 defines a new class of Notch allele: similarity with and lack of specificity of interaction of N- and NM1 with H, mam, gro and E(spl) suggest that the NM1 effect is due to modification in the intracellular signalling of the activated N receptor.

        (de Celis et al., 1993)

        The embryonic phenotype of neurogenic mutations was examined in most tissues using Ecol\lacZ enhancer trap lines. All alleles examined show defects in many organs from all three germ layers. At least for ectodermally and endodermally derived tissues, neurogenic gene function is primarily involved in interactions among cells that need to acquire or maintain an epithelial phenotype.

        (Hartenstein et al., 1992)

        The expression of mam RNA and protein during embryogenesis (up to the completion of gastrulation) has been studied.

        (Bettler et al., 1991)

        mam is needed for proper mesoderm differentiation prior to the onset of nau expression: mutant alleles cause hypertrophy in nau expressing cells.

        (Corbin et al., 1991)

        The genomic organisation of Dvir\mam and D.melanogaster mam is similar.

        (Newfeld et al., 1991)

        Mutations of mam, bib and neur in an heterozygous condition had no effect on the expression of NAx-59d or NAx-59b except when coupled in cis with Nfa-g. The neurogenic mutations suppress the wing venation phenotype of N.

        (Portin and Rantanen, 1991)

        mam alleles act as enhancers of spl alleles of N.

        (Brand and Campos-Ortega, 1990)

        mam has been sequenced and its expression pattern analysed.

        (Smoller et al., 1990)

        An extra wild type copy of mam, in combination with dxENU, causes some pupal lethality, escapers have small eyes.

        (Xu and Artavanis-Tsakonas, 1990)

        Analysis of N and mam mutant combinations reveals that reduction of the wild type number of mam was capable of interferring with the mechanism underlying negative complementation in a manner that was not restricted to specific Abruptex combinations.

        (Xu et al., 1990)

        Both neur and mam serve as hotspots for insertion of P{lacW} enhancer trap elements.

        (Bier et al., 1989)

        Neural hyperplasia, caused by mutations in mam, can be prevented by the presence of another neurogenic mutation.

        (Brand and Campos-Ortega, 1988)

        H2 causes reduction of neural hyperplasia due to homozygosity for mam- alleles.

        (de la Concha et al., 1988)

        Regions of mam cross-hybridize to the opa sequence.

        (Knust et al., 1987)

        mam has been molecularly cloned.

        (Weigel et al., 1987)

        Homozygous embryonic lethal; embryos display neural hyperplasia with compensatory epidermal hypoplasia; caused by failure of most ventral ectodermal cells to differentiate as epidermal cells rather than neuroblasts, as seen in N, amx, bib, neu, Dl, and E(spl); mam tends to have less extreme neural hyperplasia than mutants at the other neurogenic loci. A similar diversion of cells from epidermigenic into neurogenic pathways seen to generate supernumerary peripheral nerve cells (Hartenstein and Campos-Ortega, 1986). When mam expressed in female germ cells and the ensuing embryos, neural hyperplasia is enhanced, but mam+ embryos from oocytes are normal (Jimenez and Campos Ortega, 1982). Homozygous clones in the eye display irregular ommatidial pattern characterized by lack of interommatidial bristles, enlarged facets with supernumerary retinular cells and reduced numbers of pigment cells; in the cuticle, clones homozygous for mam2 are devoid of bristles (Dietrich and Campos-Ortega, 1984). mam1 (formerly N-2G) heterozygotes occasionally exhibit apical wing nicking; not recorded for other alleles. Phenotype of homozygotes for null allele reduced by duplications for normal alleles of other neurogenic loci, N, neu, Dl, E(spl) and H, but not amx or bib (de la Concha et al., 1988).

        Relationship to Other Genes
        Source for database merge of

        Source for merge of: mam l(2)s9998

        (Chang et al., 2021)
        Additional comments
        Nomenclature History
        Source for database identify of
        Nomenclature comments
        Etymology
        Synonyms and Secondary IDs (18)
        Reported As
        Symbol Synonym
        CG8118
        (National Institute of Genetics Fly Stocks, 2022-, Peterson et al., 2017, Aerts et al., 2010, Kim et al., 2010, Keleman et al., 2009.8.5, Perkins et al., 2009, Molnar et al., 2006, Kim et al., 2004, Kraut et al., 2001, Kraut et al., 2001)
        DMam
        (Kitagawa, 2016)
        Dm0064
        (Teeter et al., 2000)
        MAM
        (Contreras et al., 2015, Lee et al., 2013, Perdigoto and Bardin, 2013, Guruharsha et al., 2012)
        Mam
        (Sachan et al., 2024, Mishra et al., 2021, Bahrampour and Thor, 2020, Díaz-de-la-Peña et al., 2020, Nagel et al., 2017, Yuan et al., 2016, Auer et al., 2015, Baëza et al., 2015, Shimizu et al., 2014, Zacharioudaki and Bray, 2014, Langen et al., 2013, Mbodj et al., 2013, Ulvklo et al., 2012, Maier et al., 2011, Schaaf et al., 2009, Cave and Caudy, 2008, Yamada et al., 2008, Zhong et al., 2007, Veraksa et al., 2005, Hall et al., 2004, Hall et al., 2003, Lopez et al., 2002, Kitagawa et al., 2001)
        MamH
        (Mukherjee et al., 2016)
        N-2G
        (Jackson, 1983)
        SR2-4
        (Chang et al., 1994)
        dMam
        (Petcherski and Kimble, 2000)
        l(2)03505
        l(2)04615
        l(2)s9998
        (Chang et al., 2021)
        mam
        (Barbaste et al., 2023, Bharti et al., 2023, Candia et al., 2023, Chen et al., 2023, Rajan et al., 2023, Ding et al., 2022, Jullien et al., 2022, Ray and Li, 2022, Chang et al., 2021, McLaughlin et al., 2021, Feuillette et al., 2020, Gahr et al., 2019, Koca et al., 2019, Bischof et al., 2018, Gomez-Lamarca et al., 2018, Komori et al., 2018, Lee et al., 2018, Portela et al., 2018, Salazar and Yamamoto, 2018, Schwarz et al., 2018, Bhattacharya et al., 2017, Liu and Jin, 2017, Morimoto et al., 2016, Xu et al., 2016, Arya et al., 2015, Bivik et al., 2015, Gene Disruption Project members, 2015-, Jia et al., 2015, Kidd et al., 2015, Portela et al., 2015, Sachan et al., 2015, Spratford and Kumar, 2015, Surabhi et al., 2015, Ashwal-Fluss et al., 2014, Galindo et al., 2014, Huang and Kalderon, 2014, Parsons et al., 2014, Rhee et al., 2014, Singari et al., 2014, de Celis et al., 2013.9.11, Ghezzi et al., 2013, Joo et al., 2013, Kwon et al., 2013, Müller et al., 2013, Perea et al., 2013, Terriente-Felix et al., 2013, Webber et al., 2013, Yu et al., 2013, Japanese National Institute of Genetics, 2012.5.21, Murray et al., 2012, Paik et al., 2012, Zacharioudaki et al., 2012, Bhattacharya and Baker, 2011, Carreira et al., 2011, Dworkin et al., 2011, O'Keefe et al., 2011, Zhang et al., 2011, Aerts et al., 2010, Cunha et al., 2010, Houalla et al., 2010, Kong et al., 2010, Monastirioti et al., 2010, Mourikis et al., 2010, Richardson and Pichaud, 2010, Ayroles et al., 2009, Bejarano and Milán, 2009, Bhattacharya and Baker, 2009, Debat et al., 2009, Parks and Muskavitch, 2009.2.3, Sellin et al., 2009, Southall and Brand, 2009, Vied and Kalderon, 2009, Chen et al., 2008, Herranz et al., 2008, Mensch et al., 2008, Pierre et al., 2008, Spresser et al., 2008, Vied and Kalderon, 2008, Bejarano et al., 2007, Beltran et al., 2007, Christensen et al., 2007.10.29, Endo et al., 2007, Junion et al., 2007, Kankel et al., 2007, Luque and Milan, 2007, Minidorff et al., 2007, Minidorff et al., 2007, Reiter et al., 2007, Song et al., 2007, Tao et al., 2007, Vied and Kalderon, 2007, Wang and Lin, 2007, Zeitlinger et al., 2007, Zeitouni et al., 2007, Albrecht et al., 2006, Alexander et al., 2006, Choksi et al., 2006, Dworkin and Gibson, 2006, Dworkin and Gibson., 2006, Garces and Thor, 2006, Gause et al., 2006, Herranz et al., 2006, Molnar et al., 2006, Sandmann et al., 2006, Macdonald and Long, 2005, Lindsley and Zimm, 1992)
        mam(CAG)7
        (Flores and Engels, 1999)
        Name Synonyms
        Mastermind
        (Pinot and Le Borgne, 2024, Papagiannouli, 2022, Revici et al., 2022, Harnish et al., 2021, Xu et al., 2016, Zacharioudaki and Bray, 2014, Lee et al., 2011, Maier et al., 2011, Monastirioti et al., 2010, Quijano et al., 2010, Bhattacharya and Baker, 2009, Schaaf et al., 2009, Herranz et al., 2008, Yamada et al., 2008, Bejarano et al., 2007, Zhong et al., 2007, Herranz et al., 2006, Hall et al., 2004, Voas and Rebay, 2004, Kitagawa et al., 2001)
        lethal (2) s9998
        master mind
        (Lindsley and Zimm, 1992)
        mastermind
        (Josserand et al., 2023, Ng et al., 2019, Salazar and Yamamoto, 2018, Wieschaus and Nüsslein-Volhard, 2016, Baëza et al., 2015, Galindo et al., 2014, Parsons et al., 2014, Rhee et al., 2014, Singari et al., 2014, Joo et al., 2013, Pool et al., 2012, Bhattacharya and Baker, 2011, Meyer et al., 2011, Zhang et al., 2011, Houalla et al., 2010, Mensch et al., 2010, Monastirioti et al., 2010, Mourikis et al., 2010, Richardson and Pichaud, 2010, Vied and Kalderon, 2009, Cave and Caudy, 2008, Mensch et al., 2008, Spresser et al., 2008, Vied and Kalderon, 2008, Endo et al., 2007, Kankel et al., 2007, Song et al., 2007, Tao et al., 2007, Wang and Lin, 2007, Zeitouni et al., 2007, Albrecht et al., 2006, Alexander et al., 2006, Choksi et al., 2006, Dworkin and Gibson, 2006, Gause et al., 2006, Johnson-Schlitz and Engels, 2006, Burnette et al., 2005)
        Secondary FlyBase IDs
        • FBgn0010581
        • FBgn0026797
        • FBgn0028593
        Datasets (0)
        Study focus (0)
        Experimental Role
        Project
        Project Type
        Title
        Study result (0)
        Result
        Result Type
        Title
        External Crossreferences and Linkouts ( 103 )
        Sequence Crossreferences
        NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
        GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
        GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
        RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
        UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
        UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
        UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
        Other crossreferences
        AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
        DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
        EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
        FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
        FlyMine - An integrated database for Drosophila genomics
        InterPro - A database of protein families, domains and functional sites
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        Linkouts
        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
        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
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        References (443)