Drosophila Genes

Drosophila genes listed by biochemical function

Enzymes and cofactors exclusive of transmembrane receptor tyrosine kinases and signaling proteins

abnormal wing discs

functions in endocytosis - required for proper internalization of receptors controlling different signaling

pathways - microtubule-associated nucleoside diphosphate kinase that converts GDP to GTP - tumor suppressor

absent, small, or homeotic discs 1

histone methyl transferase that indirectly prevents gene silencing - provides a specific signal

for establishment of active transcription

Activator of SUMO 1

E1 SUMO-activating complex comprising the Aos1/Uba2 heterodimeric pair - required for SUMOylation - tumor suppressor regulating cell cycle, imaginal disc development, hematopoesis, and CNS development

Acyl-CoA synthetase long-chain

regulates fatty acid metabolism - modifies the composition of fatty acids and membrane lipid - inhibits neuromuscular

junction growth - facilitates Rab11-dependent receptor recycling - regulates axonal transport of synaptic vesicles

Adenosine deaminase-related growth factor A

a secreted enzyme that converts extracellular adenosine into inosine - regulates adenosine levels - opposes the effects

of Hedgehog signaling within the hematopoietic progenitor cells - regulates energy allocation during immune response

ALG3, alpha-1,3- mannosyltransferase

tumor suppressor gene - glycosyltransferase - glycosylation of the fly tumor necrosis factor Grindelwald

alicorn

regulatory subunit of AMP-activated protein kinase - regulation of cellular energy homeostasis - maintainance

of integrity of mature neurons under conditions of increased activity - regulation of developmental and stress-induced autophagy

approximated

DHHC family protein, responsible for the palmitoylation of selected cytoplasmic proteins - acts by controlling the normal subcellular localization

and activity of Dachs in the Fat signaling pathway

airadne 1

E3 ubiquitin-ligase essential for neuronal development - mutants display a lower rate of spontaneous neurotransmitter

release due to failures at the pre-synaptic side - evoked release in Ari-1 mutants was enhanced in a Ca(2+) dependent manner

without modifications in the number of active zones, indicating that the probability of release per synapse is increased

in these mutants - regulates myonuclear organization together with Parkin and is associated with aortic aneurysms

armitage

RNA helicase involved in posttranscriptional gene silencing - mutations disrupt mRNA translational silencing

of oskar in the oocyte and silencing of Stellate in male germ cells

Argonaute 2

endonuclease - an essential component for siRNA-directed RNA interference (RNAi) response -

required for the unwinding of siRNA duplex and in consequence, assembly of siRNA into RISC

Argonaute 3

cleaves transposon derived piwi-interacting RNAs to form an adaptive system for transposon control

atlastin

dynamin-family transmembrane GTPase that drives endoplasmic reticulum membrane fusion in a GTP-dependent fashion - self-associating protein

that is able to form trans-oligomeric complexes - required for normal growth of muscles and synapses at the neuromuscular junction

ATP synthase, subunit d

mitochondrial ATP synthesis coupled proton transport - component of electron transport complex V- regulates aging and stress

auxilin

serine/threonine protein kinase - a J-domain-containing protein that cooperates with Hsc70 in the disassembly of clathrin coats from clathrin-coated vesicles

an integral component of the Notch signaling pathway, participating in the ubiquitin-dependent endocytosis of Delta in the signal receiving cell

ballchen

multifunctional serine/threonine protein kinase - asymmetric cell division pathway - participates in proliferation control and

prevents the differentiation of neuronal stem cells - acts with MASK downstream of obscurin in development of a well defined M-line

and Z-disc of muscle - required for chromosome condensation in oocytes - nucleosomal H2A histone kinase

belle

helicase that unwinds RNA - translational repression - regulates expression of transgenes, essential for male germline stem cell maintenance and

division - suppresses the effects of ectopic Wg signaling, regulates ecdysone-triggered transcriptional cascades - regulates

the temporal pattern of Notch signaling in Drosophila follicle cells, regulates chromosome segregation

bendless

ubiquitin-conjugating enzyme that functions as a developmental switch that permits the transition from axonal growth and incipient synapse

formation to synaptic growth and maturation - required in the IMD immune response signaling pathway for IKK activation leading to NFkappaB activation

Bloom syndrome helicase

RecQ helicase - repairs replication fork damage and double-strand breaks in mitosis, promotes repair through non-crossover

mechanisms - dissolution of double Holliday junctions - promotes meiotic crossover patterning and homolog disjunction

brainiac

glycosyltransferase - directs glycosphingolipid biosynthesis - functions in oogenesis and EGF-R signaling -

helps maintain epithelial structures

Bre1

E3 ubiquitin ligase required for the monoubiquitination of histone H2B and, indirectly, for H3K4 methylation -

directs histone modification that is critical for the transcription of Notch target genes

brummer

lipid storage droplet-associated lypase - a key factor of energy homeostasis control - involved in storage-fat mobilization

capsuléen

protein arginine methyltransferase - together with its partner Valois Capsuléen is involved in protein localization in germ cells -

required for the localization of specific components of the nuage and pole plasm, and in particular of Tudor

cdc6

AAA-superfamily ATP helicase involved in initiation of DNA replication - involved in the formation of the prereplicative complex -

checkpoint protein involved in controlling the G2/M transition

Ceramidase

enzyme that hydrolyzes ceramide to sphingosine, a critical structural components of cell membranes that act as second messengers

in diverse signal transduction cascades - maintains photoreceptor homeostasis - regulates synaptic vesicle exocytosis and trafficking

chameau

histone acetyltransferase - acts as antagonistic cofactor to the histone deacetylase Rpd3 to modulate JNK-dependent

transcription during pupal thorax metamorphosis and JNK-induced apoptosis - recruited by DJun and DFos

Choline acetyltransferase

catalyzes the biosynthesis of the neurotransmitter acetylcholine - serves as specific marker for cholinergic neurons of the

ventral cord and brain - expressed in the mushroom body - transported in bulk in the axons by the heterotrimeric Kinesin-2 motor

Chromodomain-helicase-DNA-binding protein

chromodomain protein in the SNF2-like family of ATPases - functions in assembly of active chromatin

courtless

ubiquitin conjugation enzyme - plays a role in protein degradation - mutation disrupts early steps

of courtship behavior in Drosophila males and the development of their sperm

cryptochrome

potential transcription factor - light response protein - cryptochrome/photolyase

cylindromatosis ortholog (H. sapiens)

deubiquitinating enzyme that deubiquitinates dTRAF2 and prevents dTRAF2 from ubiquitin-mediated

proteolytic degradation - involved in stress response - required for Eiger-triggered JNK activation and cell death

deadhead

ovary-specific thioredoxin - linked to the distinctive redox-state balance set at the oocyte-to-embryo

transition - redox proteome - required to unlock sperm chromatin at fertilization - disulfide bond reduction is required for protamine eviction from sperm chromatin

Death caspase-1

ICE/CED-3 protease involved in apoptosis

Death related ced-3/Nedd2-like protein

ICE/CED-3 protease involved in apoptosis

Defender against apoptotic cell death 1

regulation of N-glycosylation - dolichyl-diphosphooligosaccharide-protein glycotransferase activity - crucial for protein N-glycosylation

in developing tissues - reduction of dDad1 triggers ER stress and activates unfolded protein response (UPR) signaling - loss of

dDad1 function activates JNK signaling and blocking the JNK pathway in dDad1 knock-down tissues suppresses cell apoptosis

degringolade

SUMO-targeted ubiquitin ligase that acts as a negative regulator of the repressor Hairy and its corepressor Groucho

during embryonic segmentation and neurogenesis - acts as a regulator in sex determination by interfering

with the repressive activities of Deadpan and Groucho

Dicer-1

ribonuclease III family, double-stranded RNA domain binding domain, DEAD/DEAH box helicase, PAZ domain -

an enzyme involved in degrading RNA - involved in double-stranded RNA interference (RNAi)

and post-transcriptional gene regulation (PTGS)

Dicer-2

DEAD/DEAH box helicase - mutants are defective in processing small interfering RNAs

dodo

peptidylprolyl isomerase - facilitates the degradation of the transcription factor CF2, which regulates expression

of the rhomboid gene in follicle cells

domino

DNA helicase/adenosine triphosphatase - component of the Tip60 chromatin-remodeling complex

that acetylates nucleosomal phospho-H2Av and exchanges it with an unmodified H2Av

dorado

Cullin-RING E3 ubiquitin ligase - Substrate recognition component of a SCF-like E3 ubiquitin-protein ligase complex that promotes target-directed

microRNA degradation - catalyzes ubiquitination and subsequent degradation of AGO1, thereby exposing miRNAs for degradation

drosha

ribonuclease - acts on primary microRNA transcripts by cleaving it to excise the pre-microRNA polymer - physically interacts with Pasha,

a double-stranded RNA binding protein

Dual oxidase (common alternative name: Curly)

ROS-generating NADPH oxidase - functions to transfer electrons across biological membranes to generate ROS

by transferring electrons from NADPH to oxygen - responsible for the Curly wing phenotype - apoptosis induced proliferation

Dynamin related protein 1

a Dynamin GTPase - regulates mitochondrial fission in normal cells including primary spermatocytes -

mediates mitochondrial fragmentation during programmed cell death

Dynein heavy chain 64C

a molecular motor

easter

required for establishment of embryonic polarity - serine protease, trypsin family - generates processed Späzle, which in turn acts as the Toll ligand

easily shocked

ethanolamine kinase - catalyzes the first step of the synthesis of phosphatidylethanolamine, one of the three major membrane phospholipids - a behavioral mutant

that belongs to a family of bang-sensitive paralytic mutants - required in mushroom body neuroblasts to allow a rapid rate of cell division at metamorphosis

ebony

an β-alanyl-dopamine synthase regulating β-alanyl conjugation of dopamine and histamine, thus 'trapping' these biogenic amines

preventing their further function - regulates pigmentation, photoreceptor activity and behavioral rhythmicity

effete

E2 ubiquitin-conjugating enzyme - mediates anaphase-promoting complex degradation - controls germ-line stem cell maintenance

by targeting Cyclin A - regulates programmed cell death and neuronal process remodeling

egghead

glycosyltransferase - required for glycosphingolipid sugar chain elongation - affects extracellular Gurken gradiant

by controlling Gurken planar transport in the extracellular space

eggless

SET domain hydroxymethyltransferase required for oogenesis - trimethylates histone H3 at its K9 residue

enoki mushroom

acetyltransferase, regulates transposon silencing and piRNA cluster transcription, promotes rhino expression by acetylating H3K23, but also

directly enhances transcription of piRNA clusters by facilitating Rhino recruitment, positively impacts the maintenance of trxG-regulated gene

activation by inhibiting PRC1-mediated transcriptional repression, Enok acetyltransferase complex interacts with Elg1 and negatively

regulates PCNA unloading to promote the G1/S transition, regulates oocyte polarization by promoting expression

of the actin nucleation factor spire, maintains female germline stem cells through regulating Bruno and the niche

eukaryotic translation initiation factor 4A

dead-box RNA helicase - eIF3 complex and the helicase eIF4A are required for dendritic pruning - inactivates TORC1 in response to amino acid starvation -

resistance to eIF4A inhibition in conferred by the insulin receptor cellular IRES - controls germline stem cell self-renewal by directly

inhibiting BAM function in the ovary - negative regulator of Dpp/BMP signalling that mediates SMAD degradation

eyes closed

a fly homolog of p47, a protein co-factor of the p97 ATPase involved in membrane fusion -

mutation results in a lethal failure of nuclear envelope assembly in early zygotic divisions

and results in defective photoreceptor morphogenesis

eyes absent (preferred name: clift)

transcriptional co-activator and protein tyrosine phosphatase - cell survival factor - functions in eye development -

also plays a role in the embryonic determination of somatic gonadal precursor (SGP) cell fate

fat facets

conserved Cys and His domains - ubiquitin-specific protease

Fatty acid synthetase 1

acetylation of FASN1 is involved developmental lipogenesis - acetylation of FASN1 provides a novel mechanism for protein autoacetylation,

through which Drosophila larvae control metabolic homeostasis by linking AcCoA, lysine acetylation, and de novo

lipogenesis - Drosophila STING interacts with lipid synthesizing enzymes acetyl-CoA carboxylase (ACC) and fatty acid synthase. ACC and FASN also

interact with each other, indicating that all three proteins may be components of a large multi-enzyme complex - NAD kinase sustains lipogenesis

by maintaining the pool of NADPH - NADK and fatty acid synthase 1 regulate mitochondrial mass and function by altering the levels of acetyl-CoA and fatty acids

female sterile (1) Yb (common alternative name: Yb)

tudor domain RNA helicase, post-transcriptional gene regulation - regulates transposon silencing via the piRNA pathway -

component of the Yb body, a site for Piwi-associated RNA biogenesis, regulates Piwi transport to the nucleus

four-jointed

transmembrane protein kinase that targets Fat and its transmembrane ligand Dachsous

four wheel drive

a Golgi-localized lipid kinase, the homologue of phosphatidylinositol 4-kinase IIIβ which mediates the phosphorylation of phosphatidylinositol to generate

phosphatidylinositol 4-phosphate, involved in spermatocyte cytokinesis and male fertility, mutants exhibit profound locomotor deficits and shortened lifespan

gastrulation-defective

a protease involved in establishment of dorsal/ventral polarity - targets Snake

GART trifunctional enzyme

trifunctional polypeptide with the activities of phosphoribosylglycinamide formyltransferase, phosphoribosylglycinamide synthetase, and

phosphoribosylaminoimidazole synthetase - expressed in glia, fat body, and gut - positively regulates feeding behavior via cooperation and

coordination - acting in the gut, Gart is crucial for maintaining endogenous feeding rhythms and food intake, while Gart in glia and

fat body regulates energy homeostasis between synthesis and metabolism - directly regulated by the CLOCK/CYCLE heterodimer via canonical E-box

Glucocerebrosidase 1b

Gba1b mutants revealed dysregulation of proteins involved in extracellular vesicle (EV) biology, and altered protein composition of EVs was

found in Gba1b mutants - Gba1b mutant protein aggregation may depend on relative depletion of specific ceramide species often

enriched in EVs - human mutant GCase contributes to accumulation and aggregation of α-synuclein - In the fly, this aggregation leads to development

of more severe parkinsonian signs - mutant flies showed activation of the Unfolded Protein Response (UPR) and

presented inflammation and neuroinflammation that culminated in neuronopathic disease - model for Gaucher's disease

Glucosidase 1

disrupts N-glycosylation of extracellular matrix proteins covering the fat body - invertebrate self-recognition system -

N-glycans attached to extracellular matrix proteins serve as a self signal - activated hemocytes attack tissues lacking this signal

G9a

histone H3 Lys 9-specific histone methyltransferase - regulator of peripheral dendrite development - larval locomotor behavior - non-associative

learning and courtship memory - regulator the expression of insulin-like peptide genes - regulation of starvation-induced autophagy

grappa

a histone methyltransferase that modifies the lysine 79 residue of histone H3 - necessary for the maintenance phase

of Bithorax complex expression - has characteristics of both polycomb- and trithorax-group genes

Guanylyl cyclase at 76C

cGMP production - receptor-type protein possessing a single transmembrane domain - required for Semaphorin-1a-Plexin A directed repulsive axon guidance

HDAC6

a microtubule-associated deacetylase - interacts with polyubiquitinated proteins to stimulate autophagy as a compensatory degradation system

when the ubiquitin-proteasome system is impaired

Hakai

E3 ubiquitin ligase - required for stabilization of core components of the m⁶A mRNA methylation machinery - functions in the sex determination pathway - mediates splicing of Sex lethal

Histone deacetylase 3

chromatin component - deacetylates histone H4 - counteracts the overgrowth induced by InR, PI3K or S6K over-expression -

a target of the PI3K pathway - helps retain chromatin compaction and gene silencing in nuclear lamina-interacting multigenic chromatin domains

Helicase at 25E

DExH/D RNA helicase required in the nucleus for mRNA export - required in the cytoplasm for remodeling

the transacting factors that dictate mRNA cytoplasmic destination

Hen1

RNA methyltransferase - carries out methylation of small interfering RNAs, the final step in assembly of RISC occurring

after the Argonaute-bound siRNA duplex is converted to single-stranded RNA

Hertzog

Protein-serine/threonine phosphatase, Prion-like protein, maternal effect gene required for proper establishment of segment polarity in embryos

HIF prolyl hydroxylase (alternative name: fatiga)

oxygen sensor - a specific prolyl-4-hydroxylases required for hydroxylation of key prolyl residues in the HIFα sequence -

Fe (II) and 2-oxoglutarate dependent dioxygenase superfamily protein that uses O₂ as a co-substrate for catalysis -

involved in tracheal development, oogenesis and drives cell growth downstream of Cyclin D

hiiragi

poly(A) polymerase involved in both nuclear and cytoplasmic polyadenylation of mRNA

HMG Coenzyme A reductase (common alternative name: columbus)

catalyzes the biosynthesis of a mevalonate precursor for isoprenoids - functions in the hedgehog signaling pathway,

implicated in the production of a signal by the somatic gonadal precursor cells that attracts migrating germ cells

hyperplastic discs

a HECT domain E3 ubiquitin ligase that is likely to act by targeting Cubitus interruptus and an unknown activator

of hedgehog expression for proteolysis

Ice

a dowstream effector caspase - activated during spermatogenesis and necessary for the removal

of cytoplasm and the generation of functional sperm

inactivation no afterpotential E

diacylglycerol lipase involved in visual signal transduction and lipid metabolism

Inducer of meiosis 4

methyltransferase - internal modification of mRNA - regulates alternative splicing and Notch

signaling - regulates sex determination and dosage compensation - modulates neural functions

Inositol-requiring enzyme-1

protein kinase, endonuclease, a regulator of the unfolded protein response, leading to the activation of the

transcription factor Xbp1 - regulated Ire1-dependent decay - regulation of RNA splicing - photoreceptor differentiation and rhabdomere morphogenesis

katanin-60

AAA+ family enzyme that suppresses polymerization of microtubule plus-ends - regulates cell migration -

acts in mitosis to stimulates microtubule plus-end depolymerization and Pacman-based chromatid motility

Klp10A

mitotic kinesin required to depolymerize microtubules at their pole-associated minus ends thereby moving chromatids by means of poleward flux

Kinesin heavy chain

plus end directed vesicular transport motor protein

kismet

member of the SWI2/SNF2 family of ATP-dependent chromatin-remodeling factors -

facilitates an early step

in transcriptional elongation by Pol II

krotzkopf verkehrt

a key enzyme that catalyzes the synthesis of chitin, an important component of the Drosophila epidermis, trachea,

and other tissues - development of ball-and-socket joints in the Drosophila leg - wound repair - extracellular matrix

kuzbanian

metalloprotease functioning in notch pathway

lace

encodes Serine Palmitoyl-transferase (SPT), the first enzymatic step for synthesis of sphingolipids - neuromuscular junction, enriched in lipid rafts - facilitates

glial ensheathment of peripheral nerves - suppresses dystrophic muscle phenotypes in a model of Duchenne muscular dystrophy

lazaro

lipid phosphate phosphohydrolase that functions during phototransduction - along with rdgA

regulates amplification and response termination during phototransduction

lesswright (common alternative names: Ubc9 and Semushi)

SUMO E2-conjugating enzyme - stabilizes proteins against ubiquitinization and subsequent degradation -

regulates hematopoiesis - holds the Toll pathway in check

Lipin

phosphatidate phosphatase activity - transcriptional co-activator - downstream effector mediating effects of insulin and TORC1 signaling on lipid metabolism -

influences fat body development and function - affects cellular growth

little imaginal discs

trimethyl H3K4 demethylase - physically interacts with Myc to regulator Myc-induced cell growth

Lysine (K)-specific demethylase 4A

histone demethylase - regulates H3K36me3 levels - regulates heterochromatin organization and function - Jun recruits the HP1a/KDM4A complex to its gene body region upon osmotic stress to

reduce H3K36 methylation levels and disrupt H3K36 methylation-dependent histone deacetylation - along with Kdm4B, Kdm4A is essential for mediating ecdysteroid hormone signaling during larval development

mahjong

E3 ubiquitin ligase - mediates competition of cells mutated for lethal giant larvae (lgl) - triggers cell competition through an Xrp1-dependent

pathway like that in Rp/+ cells, and distinct from cell competition of lgl or scrib clones, which do not express or depend

on Xrp1 function for elimination - proteotoxic is the underlying cause of the loser status for Minute competition and competition induced by mahjong -

forms a protein complex with Warts - promotes the ubiquitination of Wts triggering neural stem cell reactivation by inhibition of Wts

Matrix metalloproteinase 1 and Matrix metalloproteinase 2

enzymes required for several stereotyped motor axon pathfinding decisions and essential for axon fasciculation -

Mmp2 promotes dendrite reshaping through local degradation of the basement membrane - in trachea Mmp2 inhibits FGF

morphogenetic function - Mmp2 is essential for wing imaginal disc:trachea association and air sac tubulogenesis

metro

a MAGUK serving as a major binding partner of perisynaptic Dlg at larval neuromuscular junctions - stabilizes Dlg

in a complex with adaptor protein Lin-7 - controls NMJ expansion and proper establishment of synaptic boutons

MICAL

flavoprotein monooxygenase - large, multidomain protein expressed in axons - interacts with the neuronal Plexin A -

enzymatic function required for Semaphorin 1a/PlexA-mediated repulsive axon guidance

mind-bomb

E3 ubiquitin ligase - regulates endocytosis of Delta and Serrate

Minichromosome maintenance 5

AAA+ ATPase family - required in the Mcm complex for licensing and initiating origins of replication - it acts during elongation

as a helicase at the replication forks - acts in meiosis in maturation of double-strand breaks into crossovers

mummy

UDP-N-acetylglucosamine diphosphorylase - functions in apical extracellular matrix formation by producing

GlcNAc residues< needed for chitin synthesis and protein glycosylation - regulates cuticle production,

tracheal morphogenesis, and axon guidance

mutagen-sensitive 209 (preferred name: Proliferating cell nuclear antigen)

Polymerase-delta/epsilon processivity factor

myosin II (preferred name: zipper)

non-muscle myosin - motor protein - crucial functions in motility, cytokinesis, dorsal closure and cytoplasmic transport

NAD kinase 1a

sustains lipogenesis by maintaining the pool of NADPH - NADPH production rescues the lipid storage defect in the fat body of NADK RNAi

animals -NADK and fatty acid synthase 1 regulate mitochondrial mass and function by altering the levels of acetyl-CoA and fatty acids.

NADPH oxidase

generation of reactive oxygen species, Nox knockdown in mature follicle cells leads to a reduction in superoxide and to defective

ovulation - Nox in required for induction of the secreted murandot family proteins through the action of Unpaired proteins - extracellular

actin detection via Nox dependent Src-family kinase-dependent cascade is means of detecting cell injury - expressed in midgut

Nedd2-like caspase

cysteine protease - a functional homolog of CED-3/caspase-9 - involved in programmed cell death

Nedd4

E3 ubiquitin ligase - negatively regulates the Notch signaling pathway and the genes comm and Amph - contributes to neuromuscular

synaptogenesis, transverse tubule formation in muscles, and muscle function - formation and actin-dependent patterning of the heart

Neprilysin 4

endopeptidase involved in the extra-cellular metabolism of signalling peptides - metallopeptidase - regulates

insulin-like peptide (ILP) expression and food intake - required in the mushroom body for middle-term and long-term memory>

neurotic (preferred name: O-fucosyltransferase 1)

modifies Notch by attaching fucose within EGF domains - positively required for Notch signaling,

essential for the physical interaction of Notch with its ligand Delta and for the ability of Fringe

to modulate this interaction

nicastrin

component of a proteolytic complex, along with Presenilin, that targets Notch and ß amyloid protein -

Zn-dependent exopeptidase

Nicotinamide mononucleotide adenylyltransferase

NAD salvage pathway - catalyzing the last step of NAD synthesis - stress response protein - required for thermotolerance and mitigation of oxidative stress-induced

shortened lifespan - protects against axonal degeneration through chaperone activity - protects against neurodegeneration through a proteasome-mediated

pathway - required for maintaining active zone structural integrity by interacting Bruchpilot - regulated post-transcriptionally by Highwire function

Nitric oxide synthase

Ca++/Calmodulin sensitive enzyme

non-stop

ubiquitin-specific protease expressed in glia - involved in development of laminal glia

no poles

E3 ubiquitin ligase required for the preservation of genomic integrity during early embryogenesis

no receptor potential A

phosphatidylinositol-specific phospholipase C used in visual and odor signal transduction

Notopleural

enzyme - homologous to ST14 transmembrane serine protease matriptase - morphogenesis and remodelling of apical ECM during

tracheal system developmen and essential for maintenance of the transepithelial barrier function - both Np and Tpr degrade

the zona pellucida-domain (ZP-domain) protein Dumpy, a component of the transient tracheal apical ECM.

Notum (common alternative name: wingful)

alpha/ß-hydrolase - Wingless antagonist - acts to induce cleavage of the Dlp glypican at the level of its GPI anchor leading to shedding of Dlp -

spatially regulated modification of Dlp by Notum employs the ligand binding activity of Dlp to promote or inhibit signaling in a context-dependent manner

Nucleolar protein at 60B

pseudouridylate synthase - enzymatically modifies ribosomal RNA -

required for maintenance of germ-line stem cells

nudel

serine protease - docking protein - dorsal group

Nucleosome assembly protein

Histone chaperone - chromatin assembly

O-fucosyltransferase 1 (alternative name: neurotic)

modifies Notch by attaching fucose within EGF domains - positively required for Notch signaling,

essential for the physical interaction of Notch with its ligand Delta and for the ability

of Fringe to modulate this interaction

O-GlcNAc transferase (preferred name: super sex combs)

enzyme and polycomb factor - induces Hipk-mediated tumor-like growth proteasomal degradation - Hipk is

O-GlcNAcylated by OGT - cooperates with N-glycanase to regulate proliferation in intestinal stem cells (ISCs) and apoptosis in

differentiated enterocytes - controls of synaptic size and synaptic bouton number at the neuromuscular junction/a<> - part of a

clock-regulated buffering mechanism that prevent excessive O-GlcNAcylation at non-optimal times of the day-night cycle - O-GlcNAcylation of

TDP-43 suppresses ALS-associated proteinopathies and promotes TDP-43's splicing function - plays a role in

habituation learning - O-GlcNAcylation is needed for Polyhomeotic to form functional, ordered assemblies

okra

DNA helicase involved in DNA repair - required for meiosis

Ornithine decarboxylase antizyme (common alternative name: gutfeeling)

modifies the activity of Ornithine decarboxylase - regulates the nuclear entry and overall levels

of Sex-lethal in early germ cells

OXA1L mitochondrial inner membrane protein

membrane insertase, mediates mitochondrial membrane insertion of cytochrome c oxidase proteins, localizes to cristae and reticular structures

isolated in the matrix in addition to the inner boundary membrane, suggesting that it may participate in the formation of vesicle germination-derived cristae

pacman

exoribonuclease that degrades decapped mRNA - directly interacts with Decapping protein 1 to couple mRNA decapping to 5' exonucleolytic

degradation - required for degradation of mRNAs targeted by NMD and RNAi - regulates polycomb silencing - regulates insulin-like peptide

dilp8 and the neuropeptide-like precursor Nplp2 mRNA levels - affects apoptosis and regulates expression of hid

and reaper - regulates expression of the heat shock protein Hsp67Bc and the microRNA miR-277-3p in wing imaginal discs

pale

tyrosine hydroxylase - catalyzes the first and rate-limiting step in catecholamine biosynthesis - mediates the oxidation of tyrosine to L-DOPA -

DOPA decarboxylase may then metabolize L-DOPA to dopamine

parkin

ubiquitin protein ligase - degrades proteins with aberrant conformations - mutants exhibit muscle degeneration,

male sterility, and defects in brain dopaminergic cells

pavarotti

a kinesin motor protein - required both to establish the structure of the telophase spindle to provide a framework

for assembly of the contractile ring and to mobilize mitotic regulator proteins

Pcaf (common alternative name: Gcn5)

a major histone H3 acetylase in Drosophila that plays a key role in the control of specific morphogenetic cascades

during developmental transitions

Peptidylglycine-alpha-hydroxylating monooxygenase

an enzyme that targets neuropeptides

Peroxinectin-like

cyclooxygenase (Cox-like enzyme) - required for production of prostaglandins - regulation of actin cytoskeleton

during oogenesis - regulation of Fascin to control actin remodeling

persephone

trypsin-like serine protease - specific for fungal detection in a cascade activated by secreted fungal proteases and required

for the sensing of proteases elicited by bacteria in the hemolymph - acts downstream of pattern recognition receptors and upstream of Toll

pch2

conserved AAA+ ATPase that participates in a double strand break independent meiotic prophase checkpoint

Phosphoethanolamine cytidylyltransferase

a critical regulator of hunger-driven feeding - a rate-limiting enzyme in the phosphatidylethanolamine (PE)

biosynthesis pathway - Disrupting Pect activity in Drosophila fat cells causes insulin resistance, dysregulated

lipoprotein delivery to the brain, and a loss of hunger-driven feeding - mutants exhibit alterations in

phospholipid composition, defective phototransduction, and retinal degeneration

Phosphoinositide-dependent kinase 1

a master kinase that is crucial for the activation of AKT (PKB) and many kinases including PKC and S6K - contains pleckstrin homology (PH) domains

that interact with the PtdIns(3,4,5)P₃ second messenger - regulates cell growth - controls synaptic bouton size,

active zone number, and synaptic function - essential for neuronal differentiation Phosphoethanolamine cytidylyltransferase

phyllopod

novel target of ras pathway - protein degradation cofactor

Pi4KIIα

Type II phosphatidylinositol 4-kinase (PI4KII) produces the lipid phosphatidylinositol 4-phosphate, a key regulator

of membrane trafficking - critical for apical membrane transport events required for rhabdomere biogenesis

piRNA methyltransferase

RNA methyltransferase - carries out methylation of small interfering RNAs, the final step in assembly of RISC occurring

after the Argonaute-bound siRNA duplex is converted to single-stranded RNA

pipe

heparan sulfate 2-O-sulfotransferase

PNGase-like

enzyme - deglycosylates the denatured form of N-linked glycoproteins in the cytoplasm and assists their proteasome-mediated

degradation - leaves the beta-aspartyl-glucosamine (GlcNAc) of the glycan and the amide side chain of Asn, converting

Asn to Asp - coordinated regulation between O-GlcNAcylation and PNG1 is critical to balancing proliferation and apoptosis to

maintain gut homeostasis - loss of Pngl results in a severe decrease in the level of Dpp homodimers and abolishes

BMP autoregulation in the visceral mesoderm mediated by Dpp and Tkv homodimers - loss of NGLY1 in the

visceral muscle of the Drosophila larval intestine results in a severe reduction in the level of AMPKalpha, leading to energy metabolism

poly(A) polymerase (preferred name: hiiragi)

involved in both nuclear and cytoplasmic polyadenylation of mRNA

Poly-(ADP-ribose) polymerase

enzyme that attaches ADP-ribose moiety to histone - promotes chromatin silencing by regulating the localization and function of SIR2

Polypeptide N-Acetylgalactosaminyltransferase 4 (common alternative name: Pgant4)

O-glycosyltransferase - initiates the formation of mucin-type O-linked glycans - regulates secretion of components of the peritrophic/mucous membrane in the larval digestive

tract - mutation can be rescued by expression of conserved cargo receptor Tango1 and partially rescued by supplementation with exogenous mucins or treatment with

antibiotics - mutants exhibit disruption of the mucosal barrier, resulting in epithelial expression of the IL-6-like cytokine Upd3, leading to

activation of JAK/STAT signaling, differentiation of cells that form the progenitor cell niche and abnormal proliferation of progenitor cells

porcupine

O-acyltransferase activity - mediates lipidation of Wingless, which targets Wingless to secretory vesicles that deliver the ligand

to specialized microdomains at the cell surface where it can be packaged for secretion

pr-set

Set domain protein - methyltransferase that targets Histone H4 - helps maintain silent chromatin by a histone modification

that precludes neighboring acetylation of the H4 tail

Proliferating cell nuclear antigen (common alternative name: Mutagen-sensitive 209)

Polymerase-delta/epsilon processivity factor

puffyeye

a ubiquitin-specific protease that regulates Myc levels and function - regulates both the ubiquitin ligase Archipelago and its cell cycle substrate Cyclin E

pumpless

glycine cleavage system H protein - involved in appetite control

Putative Achaete Scute Target 1

EF-Hand 1, calcium-binding site - Dynamin superfamily - P-loop containing nucleoside triphosphate hydrolase - plasma membrane protein that genetically interacts with Notch - contributes

to endocytosis and differentiation of photoreceptors R1/R6/R7 and cone cells - required for assembly of the subsynaptic muscle membrane reticulum at the neuromuscular junction

Rad51-like (common alternative name: spindle A)

checkpoint protein essential for recombinational repair of double-stranded

DNA breaks (DSBs) in somatic cells and during meiosis in germ cells

rasp (common alternative name: Sightless)

acetyltransferase acting to acylate Hedgehog or to promote other modifications or trafficking events

necessary for Hedgehog function

recombination-defective

mini-chromosome maintenance (MCM) family protein - facilitates repair synthesis during meiotic recombination

Reptin

member of AAA+ family of ATPases - DNA helicase - participates in epigenetic processes

leading to a repressive chromatin state as part of the TIP60 HAT complex

rhomboid-7

transmembrane mitochondrial protease required for spermatogenesis and muscle maturation

Rm62

DEAD box helicase - involved in generation of precursor-miRNA, double-stranded RNA-mediated silencing, heterochromatin formation,

resilencing of active transcription, maintenance of homeotic genes and antiviral immunity

RNA polymerase III

conserved enzyme complex synthesizing transfer RNAs and other short, non-coding RNAs - inhibition extends lifespan

and reduces gut pathology - the pro-longevity effect of trametinib in ISCs is partially mediated by Maf1,

a repressor of Pol III - Nutrient/TOR-dependent regulation of RNA polymerase II controls tissue and organismal growth

Salsa

a helicase - a component of the NineTeen Complex (NTC), also known as Pre-mRNA-processing factor 19 (Prp19) complex - regulates distinct spliceosome

conformational changes necessary for splicing, rate-limiting for splicing of a subset of small first introns during oogenesis, including the first intron of gurken

scarface

belongs to a subgroup of serine protease homologues with an inactive catalytic site - functions extracellularly during dorsal closure

as an antagonist of the JNK pathway - regulates the polarized localization of laminin A during the development of epithelial polarity

scrawny

protein deubiquitinase required in germline, epithelial, and intestinal stem cells - acts to repress differentiation genes - deubiquitylates histone H2B

and functions in gene silencing - participates in a conserved pathway of chromatin regulation linking H2B ubiquitylation with H3K4me3 methylation

Separase

endopeptidase required for sister chromatid separation during mitosis

serpin-27A

protease inhibitor functioning in the immune response - regulates melanization cascade through specific inhibition

of the terminal protease (prophenoloxidase-activating enzyme)

Serpin77Ba

inhibits a protease cascade, involving the MP1 and MP2 proteases, that activates phenol oxidase, a key enzyme in melanin biosynthesis -

Serpin77Ba inactivation and consequent tracheal melanization induces systemic expression of the antifungal peptide gene Drosomycin

Set1

Histone3 K4 methyltransferase - responsible for the generation of a chromatin structure at active promoters that ensures optimal

Pol II release into productive elongation

Set2

Histone 3 methyltransferase involved in dosage compensation, puparium formation and adult development

Seven in absentia

targets proteins to degradation - C3HC4 RING zinc-finger motif

shaggy (common alternative name: zeste white 3)

glycogen synthase kinase - segment polarity - functions in the Wingless signaling pathway

shutdown

immunophilin - FK506-binding protein domain and a tetratricopeptide repeat - essential for the normal function of the germline stem cells - a conserved cochaperone that collaborates with Hsp90 during piRNA biogenesis, potentially at the loading step of RNAs into Piwi proteins

sightless (preferred name: Rasp)

acetyltransferase acting to acylate Hedgehog or to promote other modifications or trafficking events

necessary for Hedgehog function

Sirtuin 1 (common alternative name: Sir2)

histone deacetylase required for euchromatic and heterochromatic silencing - interacts genetically and

physically with members of the Hairy/Deadpan/E(Spl) family of bHLH euchromatic repressors

smallminded

AAA-protein family - ATPase

Smurf1 (preferred name: lethal with a checkpoint kinase)

ubiquitin-protein ligase that regulates Decapentaplegic pathway signaling by targeting Mothers

against Dpp for degradation

sol narae

a member of A Disintegrin And Metalloprotease with ThromboSpondin motif (ADAMTS) family - positively regulates Wingless

signaling by promoting Wg secretion - activates a subset of Wg signaling whose major function is the regulation

of cell proliferation - a mutually positive genetic interaction exists between ana3 and sona - wing discs

spastin

AAA (ATPase Associated with diverse cellular Activities) family member - regulates microtubule stability -

mutants show defects in synaptic growth and neurotransmission

Stellate

ß subunit of Casein kinase II - a normally inactive gene which, when active, leads to male sterility - a target of

post-transcriptional gene regulation

stem cell tumor (also known as rhomboid-2)

Rhomboid-related transmembrane protein - serine-type protease - activates the oocyte-specific Gurken ligand and

thereby participate in defining posterior cell fates in the early follicular epithelium

Sterile alpha and Armadillo motif

NAD(+) hydrolase activity - required for activation of an injury-induced axon death pathway - Axundead is a mediator of axon death

downstream of Sarm - required cell autonomously for these changes in neuronal cell biology in bystander neurons following axotomy

Structural maintenance of chromosomes 1

ATPase that heterodimerizes with the product of SMC3 to interact with the products of Vtd and Stromalin to form the cohesin ring complex - stabilizes meiotic homolog

pairing - facilitates enhancer-promoter communication and regulates activity of the Polycomb repressive complex 1 at silenced and active genes

stunted

ε-subunit of mitochondrial ATP synthase - required for normal spindle orientation during embryonic divisions -

a circulating insulinotropic peptide produced by fat cells - modulates physiological insulin levels in response to nutrients

super sex combs (common alternative name: O-GlcNAc transferase)

enzyme and polycomb factor - induces Hipk-mediated tumor-like growth proteasomal degradation - Hipk is

O-GlcNAcylated by OGT - cooperates with N-glycanase to regulate proliferation in intestinal stem cells (ISCs) and apoptosis in

differentiated enterocytes - controls of synaptic size and synaptic bouton number at the neuromuscular junction - part of a

clock-regulated buffering mechanism that prevent excessive O-GlcNAcylation at non-optimal times of the day-night cycle - O-GlcNAcylation of

TDP-43 suppresses ALS-associated proteinopathies and promotes TDP-43's splicing function - plays a role in

habituation learning - O-GlcNAcylation is needed for Polyhomeotic to form functional, ordered assemblies

Suppressor of variegation 2-10

SUMO E3 ligase - promotes the efficiency of SUMOylation - links the piRNA-guided target recognition complex to the silencing effector by binding

the piRNA/Piwi complex and inducing SUMO-dependent recruitment of the SetDB1 (Eggless)/Wde histone methyltransferase effector - chromatin modification

Suppressor of variegation 3-3

homolog of the human LSD1 amine oxidase - demethylates H3K4me2 and H3K4me1 and facilitates subsequent H3K9 methylation by SU(VAR)3-9 -

dictates the distinction between euchromatic and heterochromatic domains during early embryogenesis

Suppressor of variegation 3-9

involved in gene silencing - a modifier of position effect variegation - methylates histone H3 thus providing a mark

for the association of HP1 to heterochromatin

Sialyltransferase

mediates sialic acid transfer onto glycan termini - expressed during embryonic development in a tissue- and stage-specific fashion,

with elevated expression in a subset of cells within the central nervous system - mutations result in significantly

decreased life span, locomotor abnormalities, temperature-sensitive paralysis, and defects of neuromuscular junctions.

Tace

an active metalloprotease - causes the shedding of cell surface proteins including the Drosophila TNF homologue Eiger - cleaves Fra to

regulate midline crossing of commissural axons - ADAM17, TNF and the TNF receptor acts in pigmented glial cells of the Drosophila retina - mutation

leads to age-related degeneration of both glia and neurons, preceded by an abnormal accumulation of glial lipid droplets - cleaves Neuroligin3

at its extracellular acetylcholinesterase-like domain to generate the N-terminal fragment essential for maintaining proper locomotor activity - active

in fat body allowing the cleavage and release of adipose Eiger into the hemolymph - in the brain IPCs, Eiger activates its receptor Grindelwald,

leading to JNK-dependent inhibition of insulin production - activates Notch in a ligand-independent manner - targets Delta

Tafazzin

an enzyme in the cardiolipin remodeling pathway that catalyzes the incorporation of an acyl group into lipid acceptors -

modifies lipid composition of mitochondrial inner membranes - regulates spermatogenesis - serves as a Drosophila model for Barth syndrom

tan

hydrolase - abdominal pigmentation - hydrolysis of N-β-alanyl dopamine to dopamine during cuticular melanization -

hydrolysis of carcinine to histamine in the metabolism of photoreceptor neurotransmitter

tankyrase

a positive regulator in the JNK signaling pathway - initiates degradation-independent ubiquitination on two lysine residues of JNK to

promote its kinase activity and in vivo functions - Proteolysis of TNKS substrates is mediated through their ubiquitination by the

poly-ADP-ribose (pADPr)-dependent RING-domain E3 ubiquitin ligase RNF146/Iduna - mediates proteolysis of Axin - downregulation of tankyrase

reduces TDP-43 accumulation in the cytoplasm and potently mitigates neurodegeneration - Wingless pathway activation is promoted

by a critical threshold of Axin maintained by the tumor suppressor APC and the ADP-ribose polymerase Tankyrase

Ter94 (common alternative name: Valosin Containing Protein or VCP)

AAA family ATPase, CDC48 subfamily - modulation of proteolytic degradation - Hedgehog pathway -Wingless pathway - dendritic pruning -

motor neuron degeneration - ER stress response - maintenance of paternal chromosome integrity in the Drosophila zygote

thin (common alternative name: Trim-32)

E3 ligase activity - myofibril assembly and stability, maintenance of glycolytic flux mediated by biochemical interactions with the glycolytic

enzymes - a Drosophila model for Limb Girdle Muscular Dystrophy type 2H required for cell death in the Drosophila abdominal muscles by targeting DIAP1

Tip60

histone acetyltransferase that regulates general metabolism, axon growth and dendritic targeting and sleep-wake cycle -

regulates acetylation of histone H2A and promotes the generation of silent chromatin

Tissue inhibitor of metalloproteases

metalloprotease inhibitor expressed in muscle - neuromuscular junction - limits BMP trans-synaptic

signaling and the downstream synapse-to-nucleus signal transduction

Topoisomerase 2

ATP-dependent homodimeric enzyme that transiently cleaves double stranded DNA, passes a second DNA double helix

through the break and then reseals the break - plays a role in homolog association in meiosis - modulates insulator function

tout-velu

acetylglucosaminyltransferase - involved in heparan sulfate proteoglycan biosynthesis -

affects diffusion of Wingless, Hedgehog and Decapentaplegic

Trap1

mitochondrial chaperone protein of the heat shock protein (HSP90) family - shows an ATPase activity and is involved in neurodegeneration

associated with mitochondrial dysfunction - promotes mitochondrial unfolded protein response - enhances stress resistance, locomotor

activity and fertility - TRAP1 mutation ameliorated oxidative stress sensitivity, mitochondrial dysfunction, and DA neuronal loss in Drosophila PINK1 null mutants

trithorax

histone methyltransferase activity (H3-K4 specific), zinc finger - trithorax group

twin

degrades mRNA poly(A) tails - CCR4 componenet of an enzyme complex catalyzing mRNA deadenylation

twine

cdc25 phosphatase - dephosphorylation of cdc2 by Twine controls entry into meiosis in both males and females

Tyramine β hydroxylase

an enzyme that converts tyramine to octopamine, a neurotransmitter that is the insect equivalent of norepinephrine - octopamine modulates

several physiological functions and behaviors including sleep, aggression and ovulation

Tyrosine decarboxylase 1 and Tyrosine decarboxylase 2

catalyze directly the biosynthesis of the neuromodulator tyramine and indirectly the biosynthesis of octopamine -

Tdc1 acts to synthesize tyramine, a potent diuretic factor in the Malpighian tubule - Tdc2 is required neurally for egg deposition

Ubiquitin activating enzyme 1

Ubiquitin-activating enzyme, E1, activates and transfers ubiquitin to ubiquitin-conjugating enzymes, tumor suppressor

Ubiquitin protein ligase E3A

Ubiquitination - protein degradation - model for Angelman syndrome - neuromuscular junction -

regulation of the actin cytoskeleton, the formation of terminal dendritic branches, and dopamine/serotonin synthesis

Utx

H3K27me3 demethylase - Notch antagonist - suppressor of Rbf-dependent tumors

Vacuolar H+ ATPase subunit 68-2

acidification of the endosomal compartment - required in signal-receiving cells for Notch signaling downstream of ligand activation - promotes degradation of Notch

vihar

E2 ubiquitin-conjugating enzyme - contributes to the spatiotemporal control of Cyclin B degradation

that occurs first during mitosis at the spindle poles

wispy

poly(A) polymerase acting on specific mRNA targets during late oogenesis and early embryogenesis - required at the final stage

of oogenesis for metaphase during meiosis I arrest - interacts with Orb genetically and physically in an ovarian complex.

wunen and wunen-2

lipid phosphate phosphohydrolases - somatic Wun and Wun2 provide a repulsive environment for germ cell migration

by depleting an extracellular, attractive substrate - in germ cells Wun2 competes with somatic Wun and Wun2

for a common lipid phosphate substrate, which is required by germ cells to produce a survival signal

XNP

DEAD-like helicases superfamily - complements HIRA in recognizing exposed DNA and serving as a binding platform

for the efficient recruitment of H3.3 predeposition complexes to chromatin gaps

zipper

Myosin II or non-muscle myosin

zucchini

nuclease that functions in germline RNAi processes, required in dorso-ventral patterning pathway, required early during oogenesis

for the translational silencing of osk mRNA and at later stages for proper expression of the Grk protein

- regulates telomere-specific transposable elements

Drosophila genes listed by biochemical function

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