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General Information
Symbol
Dmel\ed
Species
D. melanogaster
Name
echinoid
Annotation Symbol
CG12676
Feature Type
FlyBase ID
FBgn0000547
Gene Model Status
Stock Availability
Gene Snapshot
echinoid (ed) encodes a cell adhesion molecule of adherens junctions that mediate cell adhesion/recognition. It participates in multiple signaling pathways including Egfr, Notch and Hippo during organogenesis. It is also required in multiple steps of dorsal closure during embryogenesis. [Date last reviewed: 2019-03-07]
Also Known As
l(2)k01102
Key Links
Genomic Location
Cytogenetic map
Sequence location
2L:4,031,377..4,115,749 [+]
Recombination map
2-11
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
-
Molecular Function (GO)
[Detailed GO annotations]
Summaries
Pathway (FlyBase)
Positive Regulators of Hippo Signaling Pathway -
The Hippo signaling pathway is an intracellular kinase cascade in which hpo kinase in complex with sav, phosphorylates wts kinase which, in turn, phosphorylates yki transcriptional co-activator leading to its cytosolic retention. Positive regulators of the pathway, enhance the cytosolic retention of yki, negatively regulating tissue growth (Adapted from FBrf0224870).
Negative Regulators of Epidermal Growth Factor Receptor Signaling Pathway -
Negative regulators of Epidermal Growth Factor Receptor signaling down-regulate the pathway, suppressing the activation of ERK kinase (rl) or acting on downstream effectors.
Positive Regulators of Notch Signaling Pathway -
The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. Positive regulators of the pathway enhance the signal from the sending cell or the response in the receiving cell. (Adapted from FBrf0225731 and FBrf0192604).
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
ed: echinoid
Eyes large and rough. Easily classified, although not as extreme as ec. Su(S)/ed is echinoid in phenotype (Gelbart). RK1.
Gene Model and Products
Number of Transcripts
3
Number of Unique Polypeptides
2

Please see the GBrowse view of Dmel\ed or the JBrowse view of Dmel\ed 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 5.47
Sequence Ontology: Class of Gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0077465
5065
1332
FBtr0334265
6378
1332
FBtr0334266
5112
1281
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0077154
146.5
1332
6.51
FBpp0306381
146.5
1332
6.51
FBpp0306382
140.5
1281
6.70
Polypeptides with Identical Sequences

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

1332 aa isoforms: ed-PA, ed-PB
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

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

External Data
Crossreferences
Linkouts
Gene Ontology (32 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (21 terms)
Terms Based on Experimental Evidence (21 terms)
CV Term
Evidence
References
inferred from mutant phenotype
inferred from direct assay
inferred from mutant phenotype
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:fz; FB:FBgn0001085
inferred from genetic interaction with FLYBASE:pnt; FB:FBgn0003118
inferred from genetic interaction with FLYBASE:stan; FB:FBgn0024836
inferred from genetic interaction with FLYBASE:cno; FB:FBgn0259212
inferred from mutant phenotype
inferred from genetic interaction with FLYBASE:yki; FB:FBgn0034970
inferred from genetic interaction with FLYBASE:sav; FB:FBgn0053193
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (8 terms)
Terms Based on Experimental Evidence (8 terms)
CV Term
Evidence
References
inferred from high throughput direct assay
inferred from direct assay
inferred from direct assay
inferred from high throughput direct assay
inferred from direct assay
Terms Based on Predictions or Assertions (0 terms)
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
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
organism

Comment: maternally deposited

Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
ed staining can be found in floor cells, but not in adjacent br-positive roof cells.
During embryogenesis, ed is initially detected uniformly at the apical domain of all epidermal and amnioserosa cells. Just before dorsal closure, levels of ed begin to decrease in the amnioserosa cells. Cells that abut the DME cells are the first to exhibit complete loss of ed. By stage 13 (epithelial sweeping phase), ed is absent from all amnioserosa cells and remains absent during stage 14 (zippering phase) and stage 15 (termination phase).
ed is detected in trachea from embryonic stage 10 until the end of embryogenesis. It is present in the fusion cells from stage 14 on. ed expression levels among the tracheal cells, including the fusion cells, appears uniform during invagination, branching, and fusion of the tracheal repeats into a continuous tubular network.
High levels of ed protein localize at the apical surface of all cells in the MF and through row 1. In rows 3 and 4, ed is reduced specifically within the cells that will soon begin to rotate, making the photoreceptor clusters appear as holes within the imaginal disc epithelium. This staining pattern persists until row 7. High ed levels are observed in photoreceptors R1, R6 and R7 when they are recruited into the growing ommatidium in rows 5/6. A dramatic shift in the relative levels of ed within the ommatidia and in the interommatidial cells occurs when cone cells are recruited. ed becomes prominent in two bands, one at the interface between the cone cells and the photoreceptors and a second at the interface between the cone cells and the interommatidial cells.
Marker for
 
Subcellular Localization
CV Term
Evidence
References
inferred from high throughput direct assay
inferred from direct assay
inferred from direct assay
inferred from high throughput direct assay
inferred from direct assay
Expression Deduced from Reporters
Stage
Tissue/Position (including subcellular localization)
Reference
Stage
Tissue/Position (including subcellular localization)
Reference
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\ed 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
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 51 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 32 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of ed
Transgenic constructs containing regulatory region of ed
Deletions and Duplications ( 52 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
adult thorax & microchaeta | ectopic
chemosensory ventral triple row & microchaeta | ectopic
chemosensory ventral triple row & microchaeta | ectopic (with edk01102)
chemosensory ventral triple row & microchaeta | ectopic (with edlH23)
dorsal triple row & macrochaeta | ectopic
dorsal triple row & microchaeta | ectopic (with edk01102)
dorsal triple row & microchaeta | ectopic (with edlH23)
embryonic leading edge cell & actomyosin
eye (with ed2B8)
sensillum campaniformium & wing | supernumerary | somatic clone
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (5)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
No
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
No
 
1 of 15
Yes
No
 
1 of 15
Yes
No
 
1 of 15
Yes
Yes
Rattus norvegicus (Norway rat) (3)
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
Xenopus tropicalis (Western clawed frog) (0)
No records found.
Danio rerio (Zebrafish) (6)
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (4)
8 of 15
Yes
Yes
1 of 15
No
No
1 of 15
No
Yes
1 of 15
No
No
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) ( EOG091900RH )
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) ( EOG091500TK )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
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
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
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) ( EOG090W00KL )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch 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
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X00AG )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Stegodyphus mimosarum
African social velvet spider
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (17)
6 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 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
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 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
    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
    suppressible
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    suppressible
    suppressible
    suppressible
    suppressible
    suppressible
    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 (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    Positive Regulators of Hippo Signaling Pathway -
    The Hippo signaling pathway is an intracellular kinase cascade in which hpo kinase in complex with sav, phosphorylates wts kinase which, in turn, phosphorylates yki transcriptional co-activator leading to its cytosolic retention. Positive regulators of the pathway, enhance the cytosolic retention of yki, negatively regulating tissue growth (Adapted from FBrf0224870).
    Negative Regulators of Epidermal Growth Factor Receptor Signaling Pathway -
    Negative regulators of Epidermal Growth Factor Receptor signaling down-regulate the pathway, suppressing the activation of ERK kinase (rl) or acting on downstream effectors.
    Positive Regulators of Notch Signaling Pathway -
    The Notch receptor signaling pathway is activated by the binding of the transmembrane receptor Notch (N) to transmembrane ligands, Dl or Ser, presented on adjacent cells. This results in the proteolytic cleavage of N, releasing the intracellular domain (NICD). NICD translocates into the nucleus, interacting with Su(H) and mam to form a transcription complex, which up-regulates transcription of Notch-responsive genes. Positive regulators of the pathway enhance the signal from the sending cell or the response in the receiving cell. (Adapted from FBrf0225731 and FBrf0192604).
    External Data
    Linkouts
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2L
    Recombination map
    2-11
    Cytogenetic map
    Sequence location
    2L:4,031,377..4,115,749 [+]
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    24D4-24D6
    Limits computationally determined from genome sequence between P{EP}EP2595EP2595 and P{lacW}edk01102
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    24D3-24D4
    (determined by in situ hybridisation)
    24D3-24D5
    Cytological location determined using genetic mapping.
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (69)
    Genomic Clones (73)
    cDNA Clones (80)
     

    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
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for database merge of
    Source for merge of: ed l(2)k01102
    Source for merge of: ed l(2)k01102 CG15424 CG12676 E(Elp)24D
    Source for merge of: ed CG16842
    Additional comments
    Source for merge of ed CG16842 was sequence comparison ( date:010202 ).
    Other Comments
    ed and fred are both required in photoreceptors R1, R6 and R7 and in the cone cells for correct ommatidial rotation.
    ed and Grip may form a signaling complex, where competition between the N-terminal and C-terminal PDZ domains of the Grip protein for ed protein binding controls signaling function.
    S2-derived S2-NP cells are treated with dsRNA made from templates generated with primers directed against ed to assess the role of this gene in Stat92E tyrosine phosphorylation.
    ed is required for os-induced Stat92E phosphorylation.
    ed interacts synergistically with the N signaling pathway.
    ed has a role in restricting neurogenic potential during embryonic neurogenesis. ed is also required to suppress sensory bristles and for correct specification of wing veins during adult development.
    Loss of function mutations in ed give rise to the development of extra macrochaetae near the extant ones in the notum and also increase the density of the microchaetae. ed loss of function mutants show synergistic interactions with components of the N pathway.
    Nrg appears to act as a heterophilic ligand and receptor of ed which in turn antagonises Egfr signaling.
    ed mutants show extra photoreceptors and cone cells, while ed overexpression leads to a reduction in the number of photoreceptors. ed acts cell non-autonomously in the generation of supernumerary R7 cells. Epistasis analysis suggests that ed acts parallel to or downstream of phl, Ras85D, pnt, aop and sina but upstream of ttk to specify R7 cell fates.
    The intracellular domain of ed is required for the repression of Egfr signalling.
    ed defines a pathway that antagonizes Egfr signalling by regulating the activity, but not the level, of the ttk TTK88 transcriptional repressor.
    ed protein binds Egfr directly and becomes tyrosine-phosphorylated in response to Egfr activation.
    Developing eye discs mutant for ed display extra and mis-patterned R8 cells; staining with antibodies against phosphorylated MAP kinase suggests that this may be a consequence of increased Egfr activity.
    ed is one of several proteins necessary to regulate Egfr activity during development.
    An EMS-induced mutation that defines a novel negative regulator of Egfr.
    Mutants isolated in a screen of the second chromosome identifying genes affecting disc morphology.
    Origin and Etymology
    Discoverer
    Bridges, 16th Jan. 1931.
    Etymology
    Identification
    External Crossreferences and Linkouts ( 60 )
    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/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
    iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
    KEGG Genes - Molecular building blocks of life in the genomic space.
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    modMine - A data warehouse for the modENCODE project
    Linkouts
    ApoDroso - Functional genomic database for photoreceptor development, survival and function
    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
    FlyMine - An integrated database for Drosophila genomics
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (13)
    Reported As
    Symbol Synonym
    CG15424
    E(Elp)24D
    Secondary FlyBase IDs
    • FBgn0022256
    • FBgn0031582
    • FBgn0031583
    • FBgn0031584
    Datasets (0)
    Study focus (0)
    Experimental Role
    Project
    Project Type
    Title
    References (194)