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General Information
Symbol
Dmel\Yp2
Species
D. melanogaster
Name
Yolk protein 2
Annotation Symbol
CG2979
Feature Type
FlyBase ID
FBgn0005391
Gene Model Status
Stock Availability
Enzyme Name (EC)
Carboxylesterase (3.1.1.1)
Gene Snapshot
Also Known As

YP, fs(1)K313, vitellogenin, yolk protein, fs(1)M35

Key Links
Genomic Location
Cytogenetic map
Sequence location
X:10,050,951..10,052,636 [-]
Recombination map

1-30

RefSeq locus
NC_004354 REGION:10050951..10052636
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (7 terms)
Molecular Function (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from electronic annotation with InterPro:IPR000734
(assigned by InterPro )
inferred from biological aspect of ancestor with PANTHER:PTN000176726
(assigned by GO_Central )
Biological Process (2 terms)
Terms Based on Experimental Evidence (0 terms)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000176726
(assigned by GO_Central )
traceable author statement
non-traceable author statement
Cellular Component (3 terms)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
inferred from direct assay
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
traceable author statement
(assigned by UniProt )
inferred from biological aspect of ancestor with PANTHER:PTN000176726
(assigned by GO_Central )
Protein Family (UniProt)
Belongs to the AB hydrolase superfamily. Lipase family. (P02844)
Catalytic Activity (EC)
Experimental Evidence
-
Predictions / Assertions
A carboxylic ester + H(2)O = an alcohol + a carboxylate (3.1.1.1)
Summaries
Gene Group (FlyBase)
OTHER CARBOXYLIC ESTER HYDROLASES -
Other carboxylic ester hydrolases are a collection of hydrolases that do not fall into any other major carboxylic ester hydrolases.
Protein Function (UniProtKB)
Vitellogenin is the major yolk protein of eggs where it is used as a food source during embryogenesis.
(UniProt, P02844)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
Yp2
Structural gene for the yolk protein YP2 found in recently-emerged female flies. Protein migrates at different rates in SDS-polyacrylamide gels when encoded by the electrophoretic variants Yp2F (fast) and Yp2S (slow), alleles that are female fertile and produce normal amounts of YP2. A mutant Yp2M (=Yp212-1245) is female fertile but lays fewer eggs than normal (Mohler, Postlethwait, and Shirk) and does not contain yolk protein in the hemolymph or ovaries.
Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\Yp2 or the JBrowse view of Dmel\Yp2 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 5.49

Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0071424
1553
442
FBtr0339648
1616
442
Additional Transcript Data and Comments
Reported size (kB)

1.67, 1.6 (northern blot)

Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0071359
49.7
442
8.18
FBpp0308710
49.7
442
8.18
Polypeptides with Identical Sequences

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

442 aa isoforms: Yp2-PA, Yp2-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

442 (aa); 49.6 (kD predicted)

Comments
External Data
Post Translational Modification

Tyrosine sulfation occurs in the female only and plays an essential functional role.

(UniProt, P02844)
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\Yp2 using the Feature Mapper tool.

External Data
Crossreferences
Linkouts
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
expression microarray
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from direct assay
Expression Deduced from Reporters
Reporter: P{Yp2.M13.G1}
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Yp2 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
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 6 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 10 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Yp2
Transgenic constructs containing regulatory region of Yp2
Deletions and Duplications ( 2 )
Phenotypes
Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
3 of 15
Yes
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
2 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Rattus norvegicus (Norway rat) (6)
3 of 13
Yes
No
2 of 13
No
No
2 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (4)
2 of 12
Yes
No
1 of 12
No
No
1 of 12
No
No
1 of 12
No
No
Danio rerio (Zebrafish) (5)
2 of 15
Yes
No
2 of 15
Yes
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (0)
No records found.
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091909EV )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091505WQ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Lucilia cuprina
Australian sheep blowfly
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( None identified )
No non-Dipteran orthologies identified
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( None identified )
No non-Insect Arthropod orthologies identified
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (32)
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
3 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v8.0 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    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 see the Physical Interaction reports below for full details
    protein-protein
    Physical Interaction
    Assay
    References
    Summary of Genetic Interactions
    esyN Network Diagram
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    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.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map

    1-30

    Cytogenetic map
    Sequence location
    X:10,050,951..10,052,636 [-]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    9A5-9A5
    Limits computationally determined from genome sequence between P{EP}nejEP1149&P{EP}nejEP1179 and P{EP}α-Man-IEP1628&P{EP}α-Man-IEP1307
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    8F-9A
    (determined by in situ hybridisation)
    8F-9B
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (5)
    Genomic Clones (9)
     

    Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

    cDNA Clones (513)
     

    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 sequenced
    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)
    Other clones
    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
     
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Additional comments
    Other Comments

    The p47 polypeptide seen previously in ACF (ATP-utilizing chromatin assembly and remodeling factor) fractions (FBrf0095281) has been found to be identical to Yp2 and appears to have been a contaminant in these earlier ACF fractions.

    20-Hydroxyecdysone, but not juvenile hormone, regulation of yolk protein gene expression can be mapped to cis-acting DNA sequences. It is not clear whether regulation by 20-hydroxyecdysone is direct or indirect. Methoprene up-regulation is only observed when native yolk protein genes are assayed, suggesting that it may operate through influencing stability of the message.

    Proteins which specifically bind the Yp1/Yp2 ovarian enhancer 1 element have been identified.

    By comparing methylation by Ecol\dam methylase between euchromatic and heterochromatic genes it was determined that the heterochromatic state does not prevent methylase accessibility in vivo.

    srp is expressed in the ovaries of adult flies where it produces an ovary specific protein isoform. The srp protein binds to a 12bp ovarian follicle cell-specific regulatory element located between the divergently transcribed Yp1 and Yp2. The 12bp element activates both in vivo and in vitro transcription of Yp1 and Yp2.

    Insertion of the gypsy\su(Hw)BR into the Yp1 Yp2 intergenic region does not alter temporal or tissue-specific expression of the Adh or Ecol\lacZ reporter genes but does repress the Yp1 Yp2 fat body enhancer elements.

    The yolk protein genes Yp1 and Yp2 are only expressed in the ovary and fat body of females if they are supplied with proteinaceous food. Several regulatory regions have been shown to independently confer nutritional regulation on the expression of Yp1 and Yp2.

    343bp of immediately upstream Yp2 sequences are sufficient for expression in the fat bodies.

    Pattern of osmium zinc iodide staining in the cortical cytoplasm varies in relation to the number of yolk protein structural genes.

    CrebA protein binds to the fat body specific enhancers of Dmul\Adh1, Adh, Yp1 and Yp2 and may be an important component of tissue specific regulation.

    Newly synthesised yolk proteins in normal and mutant strains share secretory vesicles with putative vitelline membrane proteins. Translocation of follicle cell yolk protein is not through the membrane along the interfollicular spaces, but directly through plasmalemma facing the oocyte.

    Evolutionary conservation of specificity of yolk protein uptake by the oocyte is studied throughout Diptera.

    Female flies with varying numbers of Yp1, Yp2 and Yp3 genes have been generated. Each yolk protein gene makes an equivalent contribution to the fecundity and fertility of the female, and they do not individually provide unique functions to the embryo. The number of eggs laid by a female depends on the number of genes encoding yolk proteins present in the genome, and the probability of an egg producing an adult depends on the number of yolk protein genes present in the mother.

    The male and female products of dsx when expressed in E.coli bind specifically to the fat body enhancer (FBE) of Yp1 and Yp2. This demonstrates a direct interaction between the sex determination hierarchy and a target gene.

    Yp1 and Yp2 are transcribed in the same sub-populations of ovarian follicle cells. This expression is directed by two enhancers: ovarian enhancer 1, located in the 1226bp intergenic region, and ovarian enhancer 2 located within the first exon of Yp2.

    Two cis-acting regions influence the transcription of both Yp1 and Yp2 in the ovaries. One is located in the 1224bp intergenic region and determines the stage and cell type specificity of ovarian transcription. The other is in the first intron of Yp2 and acts across the Yp2 promoter region to stimulate Yp1 transcription in ovaries.

    The site of tyrosine sulfation in the Yp2 protein has been identified.

    Yolk proteins share a certain homologous domains with human low-density lipoproteins and human lipoprotein lipase.

    The sex-, time- and tissue-specific expression of the Yp1 and Yp2 genes, divergently transcribed and separated by 1225bp, indicates that there are two tissue-specifying elements acting on each gene. One is necessary for expression in fat body and the other for expression in the ovary.

    Yp2 has been cloned and sequenced, and compared with Yp1. The intergenic region between Yp1 and Yp2 has also been sequenced.

    Yp2 has been cloned.

    The exon-intron structure of Yp2 has been determined.

    The three major yolk proteins (encoded by Yp1, Yp2 and Yp3) have been isolated from mature eggs and characterised.

    Hormonal and genetic regulation of yolk formation has been reviewed.

    Structural gene for the yolk protein YP2 found in recently-emerged female flies. Protein migrates at different rates in SDS-polyacrylamide gels when encoded by the electrophoretic variants Yp2F (fast) and Yp2S (slow), alleles that are female fertile and produce normal amounts of YP2. A mutant Yp2M (=Yp212-1245) is female fertile but lays fewer eggs than normal (Mohler, Postlethwait and Shirk) and does not contain yolk protein in the hemolymph or ovaries.

    Origin and Etymology
    Discoverer

    Postlethwait, Jan. 1979.

    Etymology
    Identification
    External Crossreferences and Linkouts ( 37 )
    Sequence Crossreferences
    NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
    GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
    GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
    RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
    UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
    UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
    Other crossreferences
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
    Flygut - An atlas of the Drosophila adult midgut
    GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
    InterPro - A database of protein families, domains and functional sites
    KEGG Genes - Molecular building blocks of life in the genomic space.
    modMine - A data warehouse for the modENCODE project
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    DRSC - Results frm RNAi screens
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
    FlyMine - An integrated database for Drosophila genomics
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (19)
    Reported As
    Symbol Synonym
    fs(1)A59
    Secondary FlyBase IDs
    • FBgn0000847
    • FBgn0004046
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (246)