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

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 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

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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