Drosophila Genes

Drosophila genes listed by biochemical function

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

absent, small, or homeotic discs 1

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

for establishment of active transcription

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

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

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

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

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

Chromodomain-helicase-DNA-binding protein

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

CYLD

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

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

Death caspase-1

ICE/CED-3 protease involved in apoptosis

Death related ced-3/Nedd2-like protein

ICE/CED-3 protease involved in 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

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

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

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

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

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

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

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

four-jointed

transmembrane protein kinase that targets Fat and its transmembrane ligand Dachsous

gastrulation-defective

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

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

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

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

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

kuzbanian

metalloprotease functioning in notch pathway

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

little imaginal discs

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

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 (common alternative name: Proliferating cell nuclear antigen)

Polymerase-delta/epsilon processivity factor

myosin II(preferred name: zipper)

non-muscle myosin

Nedd2-like caspase

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

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

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

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

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

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

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

phyllopod

novel target of ras pathway - protein degradation cofactor

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

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

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 (preferred name: Mutagen-sensitive 209)

Polymerase-delta/epsilon processivity factor

pch2

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

pumpless

glycine cleavage system H protein - involved in appetite control

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

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

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

sightless (preferred name: Rasp)

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

necessary for Hedgehog function

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

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

against Dpp for degradation

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

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

tout-velu

acetylglucosaminyltransferase - involved in heparan sulfate proteoglycan biosynthesis -

affects diffusion of Wingless, Hedgehog and Decapentaplegic

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

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

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.

vihar

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

that occurs first during mitosis at the spindle poles

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

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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