General Information
Symbol
Dmel\rut
Species
D. melanogaster
Name
rutabaga
Annotation Symbol
CG9533
Feature Type
FlyBase ID
FBgn0003301
Gene Model Status
Stock Availability
Enzyme Name (EC)
Adenylate cyclase (4.6.1.1)
Gene Snapshot
rutabaga (rut) encodes a membrane-bound Ca[2+]/calmodulin-activated adenylyl cyclase that is responsible for synthesis of cAMP. The product of rut plays a key role in regulating behavioral, neuroanatomical, and electrophysiological plasticity. [Date last reviewed: 2018-01-25]
Also Known As
AC
Genomic Location
Cytogenetic map
Sequence location
X:14,787,478..14,826,073 [-]
Recombination map
1-49
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
GO Summary Ribbons
Families, Domains and Molecular Function
Gene Group Membership (FlyBase)
Protein Family (UniProt, Sequence Similarities)
Belongs to the adenylyl cyclase class-4/guanylyl cyclase family. (P32870)
Catalytic Activity (EC)
Experimental Evidence
ATP = 3',5'-cyclic AMP + diphosphate (4.6.1.1)
Predictions / Assertions
ATP = 3',5'-cyclic AMP + diphosphate (4.6.1.1)
Summaries
Gene Group Membership
ADENYLATE CYCLASES -
Adenylate cyclases catalyze the synthesis of cAMP from ATP, yielding diphosphate as a by-product. (Adapted from FBrf0238856).
UniProt Contributed Function Data
This is a membrane-bound, calmodulin-sensitive adenylyl cyclase. Inactivation of this cyclase leads to a learning and memory defect.
(UniProt, P32870)
Phenotypic Description from the Red Book (Lindsley and Zimm 1992)
rut: rutabaga (J.C. Hall)
Mutant males and homozygous females impaired in several types of learning and memory; associative conditioning defective in tests using either reward (Tempel et al., 1983) or aversive unconditioned stimuli (e.g. Dudai, 1983, Proc. Nat. Acad. Sci. USA 80: 5445-48; Dudai, Svi, and Segel, 1984, J. Comp. Physiol. 155: 569-76; Livingstone et al.), including tests of "classical" (e.g. Tully and Quinn, 1985) and "operant" conditioning (Mariath, 1985); able to learn in associative conditioning tests involving visual cues, but at subnormal levels (Folkers, 1982, J. Insect. Physiol. 28: 535-39), and memory appears to be normal. Learning scores subnormal when measured immediately after certain types of training; then either scores decay rapidly with time (Tempel et al., 1983; Tully and Quinn, 1985) or there is no indication of memory (Mariath). Although defective in some aspects of learning, heterozygous females behave essentially normally in shock/odor tests (Dudai et al., 1983). Courtship also defective; unlike wild-type males, rut males court inseminated and virgin females with equal vigor; they may be unable to distinguish them (Gailey et al.). In tests of non-associative conditioning, rut shows aberrant habituation and sensitization to sugar stimuli (Duerr and Quinn); rut males subnormal in learning to avoid courtship of immature males; and homozygous or hemizygous rut females defective in "priming" of mating behavior by prestimulations with artificial courtship songs; effects of such acoustical prestimulations decay more rapidly than normal (Kyriacou and Hall). In either the homozygous or heterozygous condition rut acts as a partial suppressor of the sterility of homozygous dnc females inversely related to degree of rescue, suggesting both a maternal and a zygotic role of rut (Bellen, Gregory, Olsson, and Kiger, 1987, Dev. Biol. 121: 432-44; Bellen and Kiger, 1988, Roux's Arch. Dev. Biol. 197: 258-68). Double mutant females mated to Canton-S males lay many eggs, but most of the eggs fail to hatch. Biochemically, rut influences adenylate cyclase activity (Dudai et al., Livingstone et al.); it seems to abolish a calcium or calmodulin stimulated component of adenylate cyclase activity (Livingstone, Dudai, and Zvi, 1984, Neurosci. Lett. 47: 119-24), while leaving intact a component of activity stimulated by guanyl nucleotides, fluoride, or monoamines, suggesting that rut may directly affect the catalytic subunit of the adenylate cyclase complex (Livingstone et al., Dudai et al., 1984); consistent with this hypothesis is the observation that cyclase activity in rut is lower than normal, even in the presence of forskolin (Dudai et al., 1984; Dudai, Sher, Segal, and Yovell, 1985, J. Neurogenet. 2: 365-80). rut primarily affects total cyclase activity in the adult abdomen, with progressively milder effects on thoracic and head cyclase (Livingstone et al., Dudai and Svi, 1985, J. Neurochem. 45: 355-64); reduction of abdominal adenylate cyclase activity of rut1>rut2>rut3 (Bellen et al.); the majority of adenylate cyclase activity in wild type is in a particulate fraction, and rut lacks up to 35% of total particulate activity (Dudai and Zvi, 1985). That rut may in fact encode a component of the fly's adenylate cyclase catalytic subunit is suggested by altered Km of enzyme activity in mutant flies (e.g. Dudai et al., 1983, 1985) and by the fact that hypoploidy of rut+ in females leads to approximately half normal levels of that cyclase activity specifically affected by the rut mutations (Livingstone et al.), and hyperploidy for the normal allele leads to increased activity (Livingstone). The biochemical results suggest that rut1 could be a null mutation.
Gene Model and Products
Number of Transcripts
7
Number of Unique Polypeptides
6

Please see the GBrowse view of Dmel\rut or the JBrowse view of Dmel\rut for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Comments on Gene Model
Gene model reviewed during 5.50
Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.
Annotated transcripts do not represent all supported alternative splices within 5' UTR.
Gene model reviewed during 5.46
Low-frequency RNA-Seq exon junction(s) not annotated.
Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated.
Gene model reviewed during 5.52
Multiphase exon postulated: exon reading frame differs in alternative transcripts; overlap >20aa.
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0073992
9269
2248
FBtr0305581
8690
2055
FBtr0333654
8963
2146
FBtr0333655
8976
2171
FBtr0333656
7557
1391
FBtr0333657
7905
1507
FBtr0340467
9038
2171
Additional Transcript Data and Comments
Reported size (kB)
9.5, 7.5 (northern blot)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0073809
248.8
2248
6.81
FBpp0294032
228.6
2055
6.66
FBpp0305810
238.4
2146
6.68
FBpp0305811
241.4
2171
6.71
FBpp0305812
153.7
1391
7.88
FBpp0305813
166.5
1507
7.82
FBpp0309409
241.4
2171
6.71
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

2171 aa isoforms: rut-PD, rut-PG
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Linkouts
Sequences Consistent with the Gene Model
Nucleotide / Polypeptide Records
 
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\rut using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (33 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
traceable author statement
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN002899368
(assigned by GO_Central )
Biological Process (29 terms)
Terms Based on Experimental Evidence (22 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
(assigned by UniProt )
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (10 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002899368
(assigned by GO_Central )
traceable author statement
non-traceable author statement
inferred from biological aspect of ancestor with PANTHER:PTN000938101
(assigned by GO_Central )
non-traceable author statement
non-traceable author statement
traceable author statement
traceable author statement
non-traceable author statement
Cellular Component (2 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN002899368
(assigned by GO_Central )
Expression Data
Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
rut transcripts are expressed at high levels in the mushroom body perikarya and at lower levels in most cells of the central brain, the optic lobes, the subesophageal ganglion and the thoracic/abdominal ganglia.
rut transcripts levels are below wild type levels in the rutMB769 mutant.
rut transcripts are reduced to nearly background levels in the rut178 mutant.
rut transcripts are reduced to nearly background levels in the rutMB2769 mutant.
rut transcript levels are nearly wild type in the rutMB1951 mutant.
rut transcripts are reduced to nearly background levels in the rut1084 mutant.
rut transcripts are reduced to nearly background levels in the rut2080 mutant.
A similar pattern of hybridization is observed on northern blots of RNA isolated from rut1 homozygotes and wild type flies.
rut transcripts were detected in adult RNA.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
rut protein is strongly expressed in the mushroom body alpha', beta', alpha, beta and gamma lobes, and spur, and weakly expressed in the mushroom body calyx.
No rut protein immunostaining above background was observed in mushroom bodies of rut178 mutants.
rut protein is strongly expressed in the neuropil of the mushroom bodies along the stalk and in the α, β, and γ lobes, suggesting that it is transported into the axonal projections of the mushroom body neurons. It is expressed at lower levels in the mushroom body calyx and in the neuropil of the central brain, the optic lobes, the subesophageal ganglion, and the thoracic/abdominal ganglia.
rut protein immunostaining was reduced but detectable in mushroom bodies of rutMB769 mutants.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
Expression Deduced from Reporters
Reporter: P{lArB}rut178
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut743
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut1084
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut2080
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut4453
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut5124
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut5249
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rut5399
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rutMB769
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rutMB1951
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{lArB}rutMB2769
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\rut in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, Transgenic Constructs and Phenotypes
Classical and Insertion Alleles ( 30 )
For All Classical and Insertion Alleles Show
 
Allele of rut
Class
Mutagen
Associated Insertion
Stocks
Known lesion
Other relevant insertions
insertion of mobile activating element
Transgenic Constructs ( 9 )
Deletions and Duplications ( 12 )
Summary of Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
anterior fascicle & synapse, with Scer\GAL4elav-C155
axon & motor neuron | conditional ts
cell body & giant fibers
growth cone & embryonic neuron
neuromuscular junction & abdominal 4 ventral longitudinal muscle 3 & larva
neuromuscular junction & abdominal 4 ventral longitudinal muscle 4 & larva
RP3 neuron & synapse
RP3 neuron & synaptic vesicle
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (8)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
14 of 15
Yes
Yes
3 of 15
No
No
 
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
 
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (8)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
13 of 15
Yes
Yes
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
Rattus norvegicus (Norway rat) (9)
8 of 13
Yes
Yes
3 of 13
No
No
3 of 13
No
No
3 of 13
No
No
2 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (5)
7 of 12
Yes
Yes
2 of 12
No
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (12)
9 of 15
Yes
Yes
7 of 15
No
Yes
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
3 of 15
No
No
2 of 15
No
No
2 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (3)
6 of 15
Yes
No
3 of 15
No
No
2 of 15
No
No
Arabidopsis thaliana (thale-cress) (0)
No orthologs reported.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No orthologs reported.
Schizosaccharomyces pombe (Fission yeast) (0)
No orthologs reported.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091900L1 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091500YD )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0232 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X020K )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G05JR )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Strongylocentrotus purpuratus
Purple sea urchin
Ciona intestinalis
Vase tunicate
Ciona intestinalis
Vase tunicate
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Gallus gallus
Domestic chicken
Human Disease Model Data
FlyBase Human Disease Model Reports
Alleles Reported to Model Human Disease (Disease Ontology)
Download
Models ( 0 )
Allele
Disease
Evidence
References
Interactions ( 2 )
Comments ( 0 )
 
Human Orthologs (via DIOPT v7.1)
Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
Functional Complementation Data
Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
Interactions
Summary of Physical Interactions
esyN Network Diagram
Show neighbor-neighbor interactions:
Select Layout:
Legend:
Protein
RNA
Selected Interactor(s)
Interactions Browser

Please look at the Interaction Group reports for full details of the physical interactions
RNA-RNA
Interacting group
Assay
References
Summary of Genetic Interactions
esyN Network Diagram
esyN Network Key:
Suppression
Enhancement

Please look at the allele data for full details of the genetic interactions
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
suppressible
Starting gene(s)
Interaction type
Interacting gene(s)
Reference
External Data
Linkouts
BioGRID - A database of protein and genetic interactions.
DroID - A comprehensive database of gene and protein interactions.
InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
Pathways
Genomic Location and Detailed Mapping Data
Chromosome (arm)
X
Recombination map
1-49
Cytogenetic map
Sequence location
X:14,787,478..14,826,073 [-]
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
12F4-12F5
Limits computationally determined from genome sequence between P{EP}EP1599 and P{EP}EP334&P{EP}EP977
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
12F-13A
(determined by in situ hybridisation)
12F5-12F7
(determined by in situ hybridisation)
Experimentally Determined Recombination Data
Location
Notes
Stocks and Reagents
Stocks (13)
Genomic Clones (25)
cDNA Clones (220)
 

Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see GBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequences
BDGP DGC clones
Other clones
    Drosophila Genomics Resource Center cDNA clones

    For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

    cDNA Clones, End Sequenced (ESTs)
    RNAi and Array Information
    Linkouts
    DRSC - Results frm RNAi screens
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    Antibody Information
    Laboratory Generated Antibodies
     
    polyclonal
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Source for merge of: rut EP1603
    Additional comments
    "l(1)G0344" may affect "rut". "l(1)G0498" may affect "rut". "l(1)G0206" may affect "rut". "l(1)G0325" may affect "rut".
    Other Comments
    RNAi generated by PCR using primers directed to this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R+ cells.
    rut expression in the mushroom bodies of adult flies is both necessary and sufficient to correct the olfactory memory defect in rut flies - ruling out a requirement for rut expression during development of the mushroom bodies.
    The acute role for the rut-encoded type I adenyl cyclase is consistent with rut functioning as a molecular coincidence detector for the convergence of simultaneous signals (e.g. those from a conditioned and an unconditioned stimulus) on a neuron.
    Both Itp-r83A and rut signalling pathways are involved in regulating larval molting.
    Expression of rut in the median bundle appears to be sufficient to provide rut function in learning and memory as tested in a spatial learning paradigm with a heat punishment procedure.
    rut is required exclusively in the Kenyon cells of the mushroom bodies for a component of olfactory short-term memory.
    Activation of rut protein through heterotrimeric guanine nucleotide-binding protein-coupled receptor is regulated by Nf1.
    Subsets of dnc and rut neurons display abnormal spontaneous spikes and altered firing patterns in 'giant' neuron cultures. Abnormal spike activity and reduced K+ current remain in dnc rut double mutants suggesting that the opposite effects on cAMP metabolism by dnc and rut do not counterbalance the mutual functional defects. The aberrant spontaneous activity and altered frequency coding in different stimulus paradigms may present problems in the stability and reliability of neural circuits for information processing during certain behavioural tasks, raising the possibility of modulation in neuronal excitability as a cellular mechanisms underlying learning and memory.
    Nf1P2 eliminates the activation of the rut-encoded adenylyl cyclase by eliminating the pituitary-adenylyl-cyclase-activating polypeptide (PACAP38) response.
    Mutations of rut affect habituation of the electrically stimulated giant fibre response.
    The spatial expression of DopR2 coincides with that of rut, and possibly initiates biochemical cascades underlying learning.
    Phenotypic analysis reveals control of growth cone mobility requires optimal cAMP levels within an operational range.
    Injection of Acp70A into mutant virgin females elicits a small response in behaviour.
    The learning and memory mutant, rut, has impaired adenylyl cyclase activity.
    Synaptic current and modulation of K+ current triggered by Pacap38 are mediated by coactivation of the Ras/Raf (Ras85D/phl) and rut cyclase pathways.
    rut gene product is preferentially expressed in the mushroom bodies. Cell specific ablation of the mushroom bodies, by hydroxyurea, demonstrates they mediate associative odour learning in flies.
    A double mutant of an allele of rut and stnA7 is no more inviable than stnA7 alone.
    The rutabaga gene was cloned from a Ecol\lacZ enhancer trap-induced rut allele: wild type rut expression is elevated in mushroom bodies, suggesting that mushroom bodies are the principal sites mediating olfactory learning and memory.
    The open reading frame is homologous to a major portion of the mammalian adenylyl cyclases. Nucleotide sequence analysis of the open reading frame predicts the encoded protein is 2249 amino acids in length. The amino terminal half appears to share the same overall structure and hydropathy profile as the previously isolated mammalian adenylyl cyclases (Feinstein, 1991, in press, Gao, 1991, in press).
    rut plays a secondary role in the maternal regulation of embryonic cAMP content.
    rut mutant analysis suggests that the cAMP cascade plays a role in the shaping neuronal connectivity. The sensory neuron innervating the antero-notopleural bristle has an abnormally large number of side branches and varicosities in a defined segment of the axon. Ultrastructure studies suggest that the varicosities are potential synaptic sites.
    rut mutants are defective in a step of the cAMP cascade: they show impaired synaptic facilitation and post-tetanic potentiation as well as abnormal responses to direct application of dibutyryl cAMP. Results suggest that the cAMP cascade plays a role in synaptic facilitation and potentiation and indicate that synaptic plasticity is altered in memory mutants.
    Mutations in rut affect the sensory fatigue due to anteronotopleural bristle deflection, not sensory adaptation.
    Mutant analysis provides evidence for the participation of a G0-like protein in learning and memory.
    Mutations in rut significantly reduce the females song memory after prestimulation with courtship hums. This suggests a simple sensitization process may be involved with the female pulse song memory (Kyriacou and Hall, Nature 308: 62).
    Mutant males and homozygous females impaired in several types of learning and memory; associative conditioning defective in tests using either reward (Tempel, Bonini, Dawson and Quinn, 1983) or aversive unconditioned stimuli (e.g. Dudai, 1983; Dudai, Svi and Segel, 1984; Livingstone, Sziber and Quinn, 1984), including tests of 'classical' (e.g. Tully and Quinn, 1985) and 'operant' conditioning (Mariath, 1985); able to learn in associative conditioning tests involving visual cues, but at subnormal levels (Folkers, 1982) and memory appears to be normal. Learning scores subnormal when measured immediately after certain types of training; then either scores decay rapidly with time (Tempel, Bonini, Dawson and Quinn, 1983; Tully and Quinn, 1985) or there is no indication of memory (Mariath, 1985). Although defective in some aspects of learning, heterozygous females behave essentially normally in shock/odor tests (Dudai, Uzzan and Zvi, 1983). Courtship also defective; unlike wild-type males, rut males court inseminated and virgin females with equal vigor; they may be unable to distinguish them (Gailey, Jackson and Siegel, 1984). In tests of non-associative conditioning, rut shows aberrant habituation and sensitization to sugar stimuli (Duerr and Quinn, 1982); rut males subnormal in learning to avoid courtship of immature males; and homozygous or hemizygous rut females defective in 'priming' of mating behavior by prestimulations with artificial courtship songs; effects of such acoustical prestimulations decay more rapidly than normal (Kyriacou and Hall, 1984). In either the homozygous or heterozygous condition rut acts as a partial suppressor of the sterility of homozygous dnc females inversely related to degree of rescue, suggesting both a maternal and a zygotic role of rut (Bellen, Gregory, Olsson and Kiger, 1984; Bellen and Kiger, 1988). Double mutant females mated to Canton-S males lay many eggs, but most of the eggs fail to hatch. Biochemically, rut influences adenylate cyclase activity (Dudai, Svi and Segel, 1984; Livingstone, Dudai and Zvi, 1984); it seems to abolish a calcium or calmodulin stimulated component of adenylate cyclase activity (Livingstone, Dudai and Zvi, 1984), while leaving intact a component of activity stimulated by guanyl nucleotides, fluoride, or monoamines, suggesting that rut may directly affect the catalytic subunit of the adenylate cyclase complex (Livingstone, Dudai and Zvi, 1984; Dudai, Svi and Segel, 1984); consistent with this hypothesis is the observation that cyclase activity in rut is lower than normal, even in the presence of forskolin (Dudai, Svi and Segel, 1984; Dudai, Sher, Segal and Yovell, 1985). rut primarily affects total cyclase activity in the adult abdomen, with progressively milder effects on thoracic and head cyclase (Livingstone, Dudai and Zvi, 1984; Dudai and Svi, 1985); reduction of abdominal adenylate cyclase activity of rut1 > rut2 > rut3 (Bellen et al.); the majority of adenylate cyclase activity in wild type is in a particulate fraction and rut lacks up to 35% of total particulate activity (Dudai and Zvi, 1985). That rut may in fact encode a component of the fly's adenylate cyclase catalytic subunit is suggested by altered Km of enzyme activity in mutant flies (e.g. Dudai, Uzzan and Zvi, 1983; Dudai, Sher, Segal and Yovell 1985) and by the fact that hypoploidy of rut+ in females leads to approximately half normal levels of that cyclase activity specifically affected by the rut mutations (Livingstone et al.) and hyperploidy for the normal allele leads to increased activity (Livingstone). The biochemical results suggest that rut1 could be a null mutation.
    Origin and Etymology
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    Synonyms and Secondary IDs (13)
    Reported As
    Symbol Synonym
    EP1603
    rut
    (Baggett et al., 2018, Louis et al., 2018, Murakami et al., 2017, Park et al., 2017, Ueno et al., 2017, Vonhoff and Keshishian, 2017, Murmu and Martin, 2016, Naganos et al., 2016, Pitmon et al., 2016, Scholz-Kornehl and Schwärzel, 2016, Schwartz et al., 2016, Androschuk et al., 2015, Del Pino et al., 2015, Dissel et al., 2015, Grotewiel and Bettinger, 2015, Ostrowski et al., 2015, Ueda and Wu, 2015, Zhang et al., 2015, Cressy et al., 2014, Del Pino et al., 2014, Xu et al., 2014, Dubnau and Chiang, 2013, Ganguly and Lee, 2013, Ishimoto et al., 2013, Kwon et al., 2013, Neckameyer and Argue, 2013, Perisse et al., 2013, Robinson and Atkinson, 2013, Scheunemann et al., 2013, Terriente-Felix et al., 2013, Vilmos et al., 2013, Xie et al., 2013, Yoon et al., 2013, Zhang and Emery, 2013, Chen and Ganetzky, 2012, Japanese National Institute of Genetics, 2012.5.21, Kikuchi et al., 2012, Koon and Budnik, 2012, Paranjpe et al., 2012, Pezzulo et al., 2012, Rodriguez et al., 2012, Scheunemann et al., 2012, Sudhakaran et al., 2012, Tsai et al., 2012, Ueda and Wu, 2012, Wu et al., 2012, Arthaud et al., 2011, Guan et al., 2011, Guo et al., 2011, Kang et al., 2011, Knapek et al., 2011, Koon et al., 2011, Trannoy et al., 2011, Gervasi et al., 2010, Jumbo-Lucioni et al., 2010, Knapek et al., 2010, Larkin et al., 2010, Shuai et al., 2010, Tan et al., 2010, Zars, 2010, Blum et al., 2009, Claridge-Chang et al., 2009, Donlea et al., 2009, Honjo and Furukubo-Tokunaga, 2009, Iijima-Ando et al., 2009, Koganezawa et al., 2009, Lebreton and Martin, 2009, Li et al., 2009, Liu et al., 2009, Pan et al., 2009, Ruedi and Hughes, 2009, Seugnet et al., 2009, Tomchik and Davis, 2009, Ueda and Wu, 2009, Anaka et al., 2008, Brembs and Plendl, 2008, Hong et al., 2008, Lee et al., 2008, Masek and Heisenberg, 2008, Mensch et al., 2008, Neuser et al., 2008, Shakiryanova and Levitan, 2008, Ueno and Kidokoro, 2008, Acevedo et al., 2007, Asztalos et al., 2007, Balling et al., 2007, Lu et al., 2007, Motosaka et al., 2007, Peng et al., 2007, Perisse et al., 2007, Thum et al., 2007, Tinette et al., 2007, Tong et al., 2007, van Swinderen, 2007, Xia and Tully, 2007, Yamazaki et al., 2007, Akalal et al., 2006, Chabaud et al., 2006, Diegelmann et al., 2006, Ferris et al., 2006, Ganguly-Fitzgerald et al., 2006, Guo and Zhong, 2006, Molnar et al., 2006, Blow et al., 2005, Haddrill et al., 2005, Honjo and Furukubo-Tokunaga, 2005, Yu et al., 2005, Baines, 2004, Cho et al., 2004, Davis, 2004, Gerber et al., 2004, Mee et al., 2004, Zhong and Wu, 2004, Aravamudan and Broadie, 2003, Devaud et al., 2003, Alshuaib and Mathew, 2002, Hall, 2002, Schwaerzel et al., 2002, Hendricks et al., 2001, Kamyshev et al., 2000, Barth and Heisenberg, 1997)
    Name Synonyms
    adenylyl cyclase
    rutabaga
    (Lowenstein and Velazquez-Ulloa, 2018, Romey-Glüsing et al., 2018, Widmann et al., 2018, Murakami et al., 2017, Tomita et al., 2017, Vonhoff and Keshishian, 2017, Crocker et al., 2016, Pitmon et al., 2016, Davis, 2015, Dissel et al., 2015, Lee, 2015, Ueda and Wu, 2015, Singh et al., 2014, Wright, 2014, Dubnau and Chiang, 2013, Duvall and Taghert, 2013, Guichard et al., 2013, Neckameyer and Argue, 2013, Paulk et al., 2013, Perisse et al., 2013, Potdar and Sheeba, 2013, Scheunemann et al., 2013, Vilmos et al., 2013, Chen and Ganetzky, 2012, Duvall and Taghert, 2012, Morozova et al., 2012, Paranjpe et al., 2012, Tsai et al., 2012, Ueda and Wu, 2012, Chow et al., 2011, Guan et al., 2011, Guo et al., 2011, Knapek et al., 2011, Lawlor et al., 2011, Trannoy et al., 2011, Buchanan and Davis, 2010, Gervasi et al., 2010, Knapek et al., 2010, Knight et al., 2010, Larkin et al., 2010, Lee and Wu, 2010, Murakami et al., 2010, Qin and Dubnau, 2010, Tan et al., 2010, Blum et al., 2009, Claridge-Chang et al., 2009, Donlea et al., 2009, Honjo and Furukubo-Tokunaga, 2009, Iijima-Ando et al., 2009, Khurana et al., 2009, Koganezawa et al., 2009, Lebreton and Martin, 2009, Li et al., 2009, Liu et al., 2009, Ruedi and Hughes, 2009, Seugnet et al., 2009, Tomchik and Davis, 2009, Ueda and Wu, 2009, Anaka et al., 2008, Hong et al., 2008, Masek and Heisenberg, 2008, Neuser et al., 2008, Asztalos et al., 2007, Balling et al., 2007, Shadan, 2007, Tinette et al., 2007, Tong et al., 2007, van Swinderen, 2007, Xia and Tully, 2007, Yamazaki et al., 2007, Akalal et al., 2006, Diegelmann et al., 2006, Ferris et al., 2006, Katsov and Clandinin, 2006, Reaume and Sokolowski, 2006, Blow et al., 2005, Haddrill et al., 2005, Honjo and Furukubo-Tokunaga, 2005, Baines, 2004, Davis, 2004, Gerber et al., 2004, Mee et al., 2004, Tinette et al., 2004, Aravamudan and Broadie, 2003, Dubnau et al., 2003, Alshuaib and Mathew, 2002, Hall, 2002, Kitamoto, 2002, Zhang et al., 2002, Hendricks et al., 2001, Kamyshev et al., 2000, Barth and Heisenberg, 1997)
    Secondary FlyBase IDs
    • FBgn0044471
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    References (468)