logo Drosophila genes listed by biochemical function
Secreted proteins and ligands A - N ||P - Z

Accessory gland peptide 26aa (common alternative name: Ovulin)
male ejaculate accessory gland protein - increases ovulation through octopamine based neuronal signaling

Accessory gland peptide 70A (common alternative name: sex-peptide)
male ejaculate accessory gland protein - decreases female receptivity and stimulates egg production in the first mating of virgin females

TGFβ ligand of Baboon - required for tiling of photoreceptor R7 axon terminals

Adipokinetic hormone-like
metabolic neuropeptide, made in the corpora cardiaca, that mediates mobilization of energy substrates from the fat body - regulates starvation-induced foraging behavior

Allatostatin A
neuropeptide secreted by the corpus cardiacum - a modulator of AKH and DILP signaling and feeding choices
between dietary carbohydrates and protein - conveys inhibitory input onto protocerebral dopamine neurons

Allatostatin C
biological clock neurons send inhibitory AstC inputs to the brain insulin-producing cells - orthologous to the vertebrate neuropeptide
somatostatin signaling - clock-neuron-derived AstC mediates evening locomotor activity - AstC-R2 is expressed in LNds, the clock neurons that drive
evening locomotor activity - AstC-R2 inhibits the immune deficiency pathway - AstC is thought to be an immunosuppressive
substance released by nociceptors or Drosophila hemocytes - AstC-R2 also acts to dampen thermal nociception

IG superfamily

pacap-like neuropeptide

secreted glial glycoprotein

argos (common alternative name: giant lens)
ligand for EGF-receptor/Torpedo

beaten path
secreted - Ig superfamily

BM-40-SPARC (common alternative name: Sparc)
basement membrane component required for basal lamina maturation and condensation of the ventral nerve cord

Boss (Preferred name: Bride of sevenless)
transmembrane ligand for sevenless

glycosylphosphatidylinositol anchored membrane protein - essential for tracheal morphogenesis -
contributes to the maintenance of the paracellular barrier and the organisation of the septate junctions -
required for septate junction organisation in chordotonal organ glial cells

FGF homolog - ligand for Breathless

neuropeptide hormone that controls cuticle sclerotization and wing expansion behavior

encodes three neuropeptides, two of the periviscerokinin family (CAPA-1 and CAPA-2) and one pyrokinin
(Pyrokinin-1) - CAPA-1 and CAPA-2 activate GPCRs encoded by CapaR - Pyrokinin-1 activates Pyrokinin 1
receptor (PK1-R) - Capa is expressed in neurosecretory cells innervating the corpora cardiaca
(part of the ring gland) and the abdomen. CAPA-1 and -2 act as diuretic hormones on the Malpighian tubules

Carrier of Wingless
heparin sulfate proteoglycan - secreted from the presynaptic motor neuron - acts to limit synaptic architecture and
neurotransmission strength - acts cooperatively with Notum to coordinate neuromuscular synapse structural and functional
differentiation via negative regulation of Wg trans-synaptic signaling within the extracellular synaptomatrix

a larval fat body and gut produced nutrient-dependent hormone regulator of Drosophila insulin-like peptide -
targets brain insulin-producing cells

Leucine rich repeat cell adhesion protein

Collagen type IV
the main component of basement membranes - synthesized by the fat body, secreted to the hemolymph,
and incorporated into the BM - determines organ shape - regulates BMP signaling

fly ortholog of the vertebrate insulin-like growth factor-binding protein acid-labile subunit (ALS) -
antagonizes Dilp function to control animal growth and carbohydrate and fat metabolism

a secreted neuropeptide produced by dorsolateral peptidergic neurons with axon terminations in the corpora cardiaca, anterior aorta and intestine -
regulates ethanol and trehalose metabolism and male reproductive activity - coordinates increased food intake and diminished energy stores
during stress - coordinates sperm transfer and copulation duration - targets corazonin receptor in salivary glands and adipocytes

twisted gastrulation family member - along with Short gastrulation, Crossveinless promotes BMP signaling
by transporting a Dpp/Gbb heterodimer from the longitudinal veins into the crossvein regions

crossveinless 2
chordin-like cysteine rich domain protein

Crustacean cardioactive peptide (Cardioacceleratory peptide)
a neuropeptide - controls behavior during molting

TGF-ß superfamily ligand - Tolloid-related, by activating Dawdle, provides a permissive signal for axon guidance

ligand - tgf-beta homolog

EGF family - Notch ligand

Diuretic hormone 31
regulation of sleep - regulates a preferred temperature decrease at night-onset - midgut cells that express Dh31 and tachykinin
are activated by the presence of proteins and amino acids - increases the contraction frequency in the anterior midgut

Diuretic hormone 44
hormone produced by brain neurosecretory cells - sensor of nutritive sugar - directs the detection and consumption of nutritive sugar -
controls sperm retention and storage by females - a circadian output hormone that is required for normal rest:activity rhythms

an o-Glycosylated antibacterial peptide with activity against Gram-negative and Gram-positive bacteria - expressed in the fat body
during the systemic immune response and is expressed in various epithelia. The expression of Dro is regulated at the transcriptional level
mostly by the immune deficiency pathway - hemocytes relay an innate immune response to the blood cell reservoir:
through Imd signaling and the ak/Stat pathway ligand Upd3, hemocytes act as sentinels of bacterial infection,
inducing expression of the antimicrobial peptide Drosocin in respiratory epithelia and colocalizing fat body domains

inducible antifungal peptide

neuropeptide - neuromodulator - modulates aggression - controls fighting - regulates sexual arousal - regulates aspects of gut function,
satiety and food ingestion - escape-related locomotion - regulates neuromuscular junction growth through the CREB pathway - nsDSK II, a
non-sulfated version of the hormone, signals through DSK-R2 to influence gut motility and locomotion

Ecdysis triggering hormone
regulates the process of preecdysis and ecdysis behaviors in the process shedding of the old cuticle

Ecdysone-inducible gene L2
a secreted protein that binds Drosophila insulin-like peptide 2 and inhibits growth non-autonomously

Eclosion hormone

TNF superfamily; type II membrane protein - ligand that induces cell death by activating the JNK pathway

functions as a ligand for Eph receptor - expressed on the surface of neuron cell bodies -
acts to confine Eph receptor bearing axons to longitudinal tracts by repulsion

eyes shut
a proteoglycan secreted into the ommatidial apical matrix - critical for the formation of epithelial lumina in the fly retina -
forms an extracellular shield that guards mechanosensory neurons from environmental insult

Fasciclin I
cam - lipid linked to membrane and secreted - novel


folded gastrulation
secreted - novel protein - gastrulation

secreted - novel protein - boundary formation

Fs (common alternative name: Follistatin)
modulates TGF-β ligand signaling - inhibits activin-β signaling but activates Dawdle

Galactose-specific C-type lectin
secreted sugar-binding protein - modulates presynaptic function and neurotransmission strength= - recombinant DL1 protein (Lectin-galC1)
binds to E. coli and E. chrysanthemi - DL1 agglutinates E. coli and markedly intensifies the association of a Drosophila
haemocytes-derived cell line with E. coli, possibly participates in the immune response via the haemocyte-mediated mechanism

glass bottom boat (common alternative name: Transforming growth factor beta at 60A)
TGF-beta superfamily - BMP 7 homolog

Gram-negative bacteria binding protein 3
a pattern recognition receptor that is required for the detection of fungal cell wall components -
required for Toll pathway activation in response to fungal infections

Growth-blocking peptide
a secreted cytokine that mediates acute innate immune reactions through association with IMD - activates Phospholipase C
which promotes increases in the levels of inositol phosphates involved in intracellular signaling - regulates larval growth

TGF alpha homolog - EGF-R ligand

Hedgehog N-terminal signaling domain and C-terminal autoprocessing domain

hikaru genki
synaptic protein - IG superfamily - complement binding domains

controls growth and metabolism - encodes at least two distinct neuropeptides; one, hug-γ, has homology
to an ecdysone triggering hormone and a second, Drm-PK-2, is a pyrokinin with myostimulatory activity

Insulin-like peptide 1
ligand of the insulin/IGF-signaling pathway - expressed in brain neurosecretory cells of larvae, pupae, and adults - Mutation diminishes
organismal weight during pupal development, whereas overexpression increases it - survival during starvation is
strongly diminished in mutants - overexpression decreases survival during starvation in female flies and increases egg laying
and decreases egg to pupal viability - mutants extend lifespan and increase Akh mRNA and protein in a dilp1-dependent manner - expressed
in non-feeding stages and in diapausing flies - is under feedback regulation and appears to play sex-specific functional roles

Insulin-like peptide 6
an insulin-like peptide that is produced by the fat body - induced by metabolic stress -
promotes growth in conditions of nutritional deprivation or following developmentally induced cessation of feeding

Insulin-like peptide 7
Ilp7-producing neurons are wired with Insulin-producing cells - regulation of feeding behavior - The former cells regulate the latter to facilitate larval development at
high temperatures and to regulate systemic Insulin signaling in adults feeding on calorie-rich food lacking dietary yeast - acute release of hub
neuron-derived insulin-like peptide 7 and cognate relaxin family receptor signaling in downstream neurons are required for noxious light
avoidance - knockout of dilp7 causes increase in glycogen levels and simultaneous decrease in triglyceride levels at low protein consumption

Insulin-like peptide 8
ensures developmental stability and robustness - delays metamorphosis by inhibiting ecdysone biosynthesis thus slowing growth
in the imaginal discs - coordinates tgrowth status of tissues with developmental timing

Insulin-related peptide
Insulin-like growth factor that functions as a growth stimulator - regulates of trehalose metabolism

Ion transport peptide
an endocrine regulator of thirst and excretion, which integrates water homeostasis with feeding - regulates sleep through the photoperiod network - expressed in
pars lateralis neurosecretory neurons, three hindgut-innervating neurons in abdominal ganglia, and a stage-specific number of interneurons and peripheral bipolar neurons

jelly belly
a secreted ligand that contains an LDL receptor repeat - required for visceral mesoderm development

ligand for Epidermal growth factor receptor - homolog of Spitz with partially independent functional activity -
provides an autocrine mitogenic stimulus in midgut progenitor cells

Laminin A
laminin - extracellular matrix protein

Laminin B1
extracellular matrix protein - required for normal accumulation of basement membrane components -
involved in morphogenesis of gut, trachea, muscles and nervous system

secreted hormone, regulates insulin pathway, suppresses insulin secretion by insulin-producing cells following starvation,
localized to glucose-sensing, endocrine corpora cardiaca (CC) cells associated with the gut

Lipophorin receptor 1 & Lipophorin receptor 2
a secreted receptor that promotes the extracellular lipolysis of lipophorins - neutral lipid uptake - LpR1 facilitates uptake
of the serpin Necrotic in oocytes and cells of imaginal discs

olfactant-binding-protein, lipocalin family

magu (common alternative name: Pentagone) a secreted protein that acts in a regulatory feedback during establishment and maintenance of BMP/Dpp morphogen signalling during wing
development - modifies the ability of cells to trap and transduce BMP by fine-tuning the levels of the BMP reception system at the plasma
membrane - internalises the Dpp co-receptors Dally and Dally-like protein - required for Wg signalling - spermatogenesis

TGFβ family ligand - concentrated in the foregut - broad expression pattern suggests that signaling could be permissive rather than instructive during
stomatogastric nervous system precursor migration - neuromuscular junction glia release Maverick, which likely activates the muscle activin-type
receptor Punt to potently modulate Gbb-dependent retrograde signaling and synaptic growth

Mesencephalic astrocyte-derived neurotrophic factor
a secreted protein - unfolded protein response - immune modulation - expressed in
glial cells - supports the dopaminergic system in non-cell-autonomous manner

an antifungal peptide that is secreted from the fat body during the systemic immune response\ - expression is regulated at
the transcriptional level by the immune deficiency and/or Toll pathways - targets the iron-sulfur subunit (SdhB)
of succinate-coenzyme Q reductase - interferes with F. graminearum cell wall biosynthesis by targeting the
β(1,3)-glucanosyltransferase Gel1, which is responsible for β(1,3)-glucan chain elongation in the cell wall

mind the gap
secreted carbohydrate-binding protein involved in synapse assembly -
modifier of signaling by the ligand Jelly belly and its receptor Alk

homolog of Collagen XV/XVIII, a matrix protein that can be proteolytically cleaved to release Endostatin, an antiangiogenesis signaling factor -
required for normal calcium channel abundance, presynaptic calcium influx, and neurotransmitter release - functions in motor axon pathfinding

secreted Transforming growth factor-beta family ligand - secreted by glial cells
during larval brain development - for neuronal expression of Ecdysone receptor

Myoinhibiting peptide precursor
ancestral ligand for the Sex peptide receptor - acts directly in the polyamine-detecting olfactory and taste neurons -
regulation of polyamine attraction, stabilization of sleep

Netrin-A and Netrin-B
secreted factor involved in axonogenesis

Neurexin 1
ligand of neuroligins involved in development and function of synaptic architecture in the brain and neuromuscular junction - glutamergic synapse

neuropeptide F
peptide hormone - developmentally programmed to modify foraging and social behavior in Drosophila larvae

Neurotrophin 1
secreted ligand that maintains neuronal survival - expression rescues
naturally occurring cell death and enables targeting by motor neurons

secreted scaffold-like protein - binds chitin - recruits proteins for chitin-matrix growth - modulates secreted scaffold-like protein - binds chitin - recruits proteins for chitin-matrix growth - modulates
localization of proteins at the ECM - controls tracheal tube diameter - affects cuticle
stiffness during wound repair - promotes morphogenesis and axonal growth in the prothoracic gland

chemokine-like protein that binds to phosphatidylserine (PS) - PS exposure on neurons is supplied cell-non-autonomously
to coat dendrites> and to mediate interactions between PS and Draper, thus enabling phagocytosis - involved
in the transformation of glial cells into phagocytes in different developmental neuronal remodeling paradigms - expressed in fat body,
epidermal cells, trachea, hemocytes and glia - necessary for both axonal pruning and removal of axonal debris in the developing mushroom body

outstretched, unpaired 2 and unpaired 3
cytokine-like ligands - activators of JAK/STAT signaling pathway

collagen-like extracellular matrix protein involved in heart development

PDGF- and VEGF-related factor 1
ligands that orchestrate migratory patterns of hemocytes at the embryonic ventral midline - essential cell autonomously
for proper positioning of the salivary glands - border cell migration

PDGF- and VEGF-related factor 2 and PDGF- and VEGF-related factor 3
ligand - orchestrate migratory patterns of hemocytes - essential cell autonomously for proper positioning of the salivary glands

Pigment-dispersing factor

Prothoracicotropic hormone
developmental timing cue involved in promoting larval development and eclosion

pyramus and thisbe
FGFs involved in mesodermal development - ligands for Heartless

secreted - fibrinogen family

TGF-ß superfamily - DPP signaling

Secreted decoy of InR
insulin-binding protein - negative regulator of insulin signaling - regulation of body size - receptor-like decoy molecule

Semaphorin-2a and Semaphorin-2b
secreted - Ig superfamily - axon guidance

transmembrane - EGF homolog - ligand for Notch

sex-peptide (preferred name: Accessory gland peptide 70A)
male ejaculate accessory gland protein - decreases female receptivity and stimulates egg production in the first mating of virgin females

ortholog of the human Wnt inhibitory factor - required for Hh stability and for lipid-modified Hh diffusion -
colocalizes with Hh in the extracellular matrix and interacts with the heparan sulfate proteoglycans

short gastrulation
chordin homolog - segment polarity

short neuropeptide F precursor
neuropeptide that engenders ERK-mediated insulin expression and thus systemically modulates growth, metabolism and lifespan

Ig domain cell adhesion molecule - ligand for Beaten path Ia - axon guidance

extracellular - egf like

secreted - ligand for toll

spatzle 5
neurotrophin - a ligand for the Toll-related receptors - acts as Toll-6 and Toll-7 ligands
in the promotion of motor axon targeting and neuronal survival in CNS - involved in synaptic
targeting in the development of specific neurons at the neuromuscular junction - secreted from
fat body to facilitates the elimination of scrib clones by binding to Toll-6

ligand for Egf-r - egf/tgf alpha homolog

stranded at second
Sas and the receptor-type tyrosine phosphatase PTP10D function as the cell-surface ligand-receptor system that drives tumour-suppressive cell competition - trans-activation
of Sas-PTP10D signalling in loser cells restrains EGFR signalling cell elimination and thereby enables elevated JNK signalling in loser cells, triggering
cell elimination - expression of Sas in neurons and glia and Ptp10D expressed on longitudinal axons are required to prevent axons from abnormally crossing the midline

a circulating insulinotropic peptide produced by fat cells - modulates physiological insulin levels in response to nutrients
ε-subunit of mitochondrial ATP synthase - required for normal spindle orientation during embryonic divisions

a secreted neuropeptide that regulates aggression, response to metabolic stress, and insulin-producing cells

secreted ligand-binding protein

extracellular matrix - neuromuscular junction - synaptic cleft - presynapse - postsynapse - integrin ligand - wing morphogenesis - male genital looping - diameter expansion of the hindgut tube

terribly reduced optic lobes
extracellular matrix component - homolog of perlecan - regulates larval neuroblast division by modulating both FGF and Hedgehog signaling

thisbe and pyramus
FGFs involved in mesodermal development - ligands for Heartless

extracellular matrix ligand produced by tendon cells - essential for the formation of the integrin-mediated myotendinous junction

secreted - extracellular matrix

tolkin common alternative name: tolloid-related-1
BMP-1 homolog - metalloprotease

secreted BMP-1 homolog - metalloprotease

tolloid-related-1 (preferred name: tolkin)
secreted BMP-1 homolog - metalloprotease

Transforming growth factor beta at 60A (preferred name: glass bottom boat)
TGF-beta superfamily - BMP 7 homolog

secreted - ligand for torso

Ig superfamily - ligand or co-receptor - required to restrain dendrite branch formation in neurons with simple arbors,
and to promote dendrite self-avoidance in neurons with complex arbors - regulates the tiling pattern of R7 photoreceptor terminals

twisted gastrulation
secreted growth factor

VAMP-associated protein of 33kDa ortholog A (common alternative name: Vap-33-1)
cleaved secreted ligand for Eph receptors - plays a conserved role in synaptic homeostasis

EGF domain and Ig domain - ligand for EGF-receptor

along with Collagen IV, the main component of basement membranes - synthesized by the fat body,
secreted to the hemolymph, and incorporated into the basement membrane - determines organ shape - regulates BMP signaling
ligand - wnt family - segment polarity

Wnt oncogene analog 6
required for >maxillary palp formation - oogenesis, anterior escort cells are maintained by Wnt6 signaling - required for germline stem cell maintenance

Wnt8 (common alternative name: WntD)
Wnt family ligand - involved in establishment dorsal/ventral patterning and the immune response

Wnt oncogene analog 2
ligand - Wnt family

Wnt oncogene analog 4
ligand - TGF-B homolog - mesodermal

Wnt oncogene analog 5
Wnt superfamily ligand involved in axon guidance and salivary gland morphogenesis

Immunoglobulin domain ligand - present in midline glia - required, along with neuronally-expressed
Neurexin IV, for ensheathment of commissural axons

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Drosophila genes listed by biochemical function

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