logo 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

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

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

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

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

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

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

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

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

glycosyltransferase - directs glycosphingolipid biosynthesis - functions in oogenesis and EGF-R signaling -
helps maintain epithelial structures

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

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

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

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

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

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

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

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

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

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

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)

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

peptidylprolyl isomerase - facilitates the degradation of the transcription factor CF2, which regulates expression
of the rhomboid gene in follicle cells

DNA helicase/adenosine triphosphatase - component of the Tip60 chromatin-remodeling complex
that acetylates nucleosomal phospho-H2Av and exchanges it with an unmodified H2Av

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

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

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

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

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

glycosyltransferase - required for glycosphingolipid sugar chain elongation - affects extracellular Gurken gradiant
by controlling Gurken planar transport in the extracellular space

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

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

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

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

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

a microtubule-associated deacetylase - interacts with polyubiquitinated proteins to stimulate autophagy as a compensatory degradation system
when the ubiquitin-proteasome system is impaired

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

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

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

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

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

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

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

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

metalloprotease functioning in notch pathway

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

ubiquitin protein ligase - degrades proteins with aberrant conformations - mutants exhibit muscle degeneration,
male sterility, and defects in brain dopaminergic cells

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

cyclooxygenase (Cox-like enzyme) - required for production of prostaglandins - regulation of actin cytoskeleton
during oogenesis - regulation of Fascin to control actin remodeling

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

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

novel target of ras pathway - protein degradation cofactor

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

heparan sulfate 2-O-sulfotransferase

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

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

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

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

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

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

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

transmembrane mitochondrial protease required for spermatogenesis and muscle maturation

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

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

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

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

endopeptidase required for sister chromatid separation during mitosis

protease inhibitor functioning in the immune response - regulates melanization cascade through specific inhibition
of the terminal protease (prophenoloxidase-activating enzyme)

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

Histone3 K4 methyltransferase - responsible for the generation of a chromatin structure at active promoters that ensures optimal
Pol II release into productive elongation

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

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

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

AAA (ATPase Associated with diverse cellular Activities) family member - regulates microtubule stability -
mutants show defects in synaptic growth and neurotransmission

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

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

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.

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

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

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

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

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

acetylglucosaminyltransferase - involved in heparan sulfate proteoglycan biosynthesis -
affects diffusion of Wingless, Hedgehog and Decapentaplegic

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

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

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

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

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

E2 ubiquitin-conjugating enzyme - contributes to the spatiotemporal control of Cyclin B degradation
that occurs first during mitosis at the spindle poles

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

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

Myosin II or non-muscle myosin

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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