FB2024_02 , released April 23, 2024
Gene: Dmel\zld
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General Information
Symbol
Dmel\zld
Species
D. melanogaster
Name
zelda
Annotation Symbol
CG12701
Feature Type
protein_coding_gene
FlyBase ID
FBgn0259789
Gene Model Status
Current
Stock Availability
15 publicly available
Gene Summary
zelda (zld) encodes a zinc finger transcription factor that functions in early blastoderm development. It works as a zygotic genome activator because it has a global role in regulating early expressed genes, such as genes essential for cellularization, sex determination and pattern formation. It is also required at later times in development. [Date last reviewed: 2019-03-21] (FlyBase Gene Snapshot)
All Summaries
Also Known As

vfl, vielfaltig, EP1344

Key Links
Genomic Location
Cytogenetic map
18F1-18F2
Sequence location
Recombination map
1-64
RefSeq locus
NC_004354 REGION:19766465..19783548
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (25 terms)
Molecular Function (8 terms)
Terms Based on Experimental Evidence (7 terms)
CV Term
Evidence
References
enables chromatin loop anchoring activity
inferred from mutant phenotype
(Ogiyama et al., 2018)
enables DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from direct assay
(Brennan et al., 2023, Fernandes et al., 2022, Duan et al., 2021, Larson et al., 2021, Yamada et al., 2019, Hamm et al., 2015, Schulz et al., 2015, Foo et al., 2014, Xu et al., 2014, Harrison et al., 2011, Harrison et al., 2010)
enables nucleosome binding
inferred from mutant phenotype
(McDaniel et al., 2019)
enables protein binding
inferred from physical interaction with FLYBASE:bi; FB:FBgn0000179
(Giannios and Tsitilou, 2013)
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding
inferred from direct assay
(Brennan et al., 2023, Larson et al., 2021, Yamada et al., 2019, Dufourt et al., 2018, Hamm et al., 2015, Sun et al., 2015, Xu et al., 2014, Harrison et al., 2011, Nien et al., 2011)
enables sequence-specific DNA binding
inferred from mutant phenotype
(McDaniel et al., 2019)
inferred from direct assay
(Harrison et al., 2010)
enables transcription cis-regulatory region binding
inferred from direct assay
(Liang et al., 2008)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables DNA-binding transcription activator activity, RNA polymerase II-specific
inferred from biological aspect of ancestor with PANTHER:PTN002811836
(GO Reference Genome Project, 2011-)
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding
inferred from biological aspect of ancestor with PANTHER:PTN002811836
(GO Reference Genome Project, 2011-)
Biological Process (14 terms)
Terms Based on Experimental Evidence (13 terms)
CV Term
Evidence
References
involved_in cellularization
inferred from mutant phenotype
(Liang et al., 2008)
involved_in chromatin remodeling
inferred from direct assay
(Brennan et al., 2023, Duan et al., 2021, Larson et al., 2021, Dufourt et al., 2018, Schulz et al., 2015, Sun et al., 2015, Foo et al., 2014)
involved_in maternal mRNA clearance
inferred from direct assay
(Hamm et al., 2017, Fu et al., 2014, Liang et al., 2008)
involved_in maternal-to-zygotic transition of gene expression
inferred from direct assay
(Brennan et al., 2023, Fernandes et al., 2022, Duan et al., 2021, Yamada et al., 2019, Dufourt et al., 2018, Hamm et al., 2017, Schulz et al., 2015, Sun et al., 2015, Harrison et al., 2011, Nien et al., 2011, Liang et al., 2008)
involved_in miRNA transcription
inferred from direct assay
(Fu et al., 2014)
involved_in mitotic cell cycle
inferred from mutant phenotype
(Staudt et al., 2006)
involved_in neuroblast fate commitment
inferred from direct assay
(Larson et al., 2021)
involved_in nucleosome organization
inferred from direct assay
(Sun et al., 2015)
involved_in positive regulation of DNA-templated transcription
inferred from direct assay
(Hamm et al., 2017, Liang et al., 2008)
involved_in positive regulation of gastrulation
inferred from mutant phenotype
(McDaniel et al., 2019)
involved_in positive regulation of gene expression
inferred from mutant phenotype
(McDaniel et al., 2019)
involved_in positive regulation of neuroblast proliferation
inferred from mutant phenotype
(Reichardt et al., 2018)
involved_in wing disc development
inferred from mutant phenotype
(Giannios and Tsitilou, 2013)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN002811836
(GO Reference Genome Project, 2011-)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
part_of chromatin
inferred from direct assay
(Staudt et al., 2006)
located_in nucleoplasm
inferred from direct assay
(Staudt et al., 2006)
is_active_in nucleus
inferred from direct assay
(Dufourt et al., 2018, Nien et al., 2011)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN002811836
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
zelda (zld) encodes a zinc finger transcription factor that functions in early blastoderm development. It works as a zygotic genome activator because it has a global role in regulating early expressed genes, such as genes essential for cellularization, sex determination and pattern formation. It is also required at later times in development. [Date last reviewed: 2019-03-21]
Gene Group (FlyBase)
C2H2 ZINC FINGER TRANSCRIPTION FACTORS -
Zinc finger C2H2 transcription factors are sequence-specific DNA binding proteins that regulate transcription. They possess DNA-binding domains that are formed from repeated Cys2His2 zinc finger motifs. (Adapted from PMID:1835093, FBrf0220103 and FBrf0155739).
Protein Function (UniProtKB)
Transcription factor required for zygotic genome activation (ZGA), a critical event in early embryonic development during which the developmental control passes from maternally provided mRNAs to the expression of the zygotic genome after fertilization (PubMed:18931655, PubMed:20599892, PubMed:22028662, PubMed:22028675, PubMed:24909324, PubMed:24637116, PubMed:26335633, PubMed:26335634, PubMed:25538246, PubMed:29261646, PubMed:30518940, PubMed:30982648, PubMed:30797686, PubMed:34342574, PubMed:35363606, PubMed:37557175). Binds to regulatory DNA sequences containing a 5'-CAGGTAG-3' sequence motif, which are highly enriched among developmental enhancers (PubMed:18931655, PubMed:22028662, PubMed:22028675, PubMed:35363606, PubMed:37557175). Within 1 hour into development, or by the embryo's 8th nuclear cycle, binds the majority of its motifs genome-wide (PubMed:22028662, PubMed:22028675). Zelda-binding promotes nucleosome depletion and chromatin accessibility, thereby facilitating the binding of patterning transcription factors, including the binding of the dorsoventral patterning transcription factors dorsal (dl) and twist (twi), and the anteroposterior patterning transcription factors bicoid (bcd) and caudal (cad) (PubMed:24909324, PubMed:24637116, PubMed:26335633, PubMed:26335634, PubMed:30518940, PubMed:30982648, PubMed:30797686, PubMed:34342574, PubMed:35363606, PubMed:37557175). Promotes the activity of patterning transcription factors, such as bcd and dl, by lowering the concentration threshold required for transcriptional activation (PubMed:24637116, PubMed:35363606). Required both for the earliest (minor) and major waves of transcription during ZGA (PubMed:22028662). Also involved in maternal mRNA clearance during the maternal-to-zygote transition by promoting expression of microRNAs (miRNAs), such as miR-1, miR-9a and miR-309, which mediate degradation of maternally-loaded RNAs (PubMed:18931655, PubMed:24764079). Also involved in post-blastoderm development: nvolved in nervous system development by maintaining neuroblasts in an undifferentiated state and equired for wing disk development (PubMed:23891688, PubMed:34887421).
(UniProt, Q9VWC6)
Summary (Interactive Fly)

zinc finger transcription factor - functions in the cellular blastoderm to activate many genes essential for cellularization, sex determination and pattern formation

Gene Model and Products
Number of Transcripts
5
Number of Unique Polypeptides
3

Please see the JBrowse view of Dmel\zld 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
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q9VWC6)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.55

Gene model reviewed during 5.45

Gene model reviewed during 5.48

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.56

Sequence Ontology: Class of Gene
gene
gene_with_edited_transcript
(Graveley et al., 2011)
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
zld-RA
FBtr0074808
NM_134512
7562
1596
zld-RB
FBtr0074807
NM_167685
7217
1596
zld-RD
FBtr0307537
NM_001258840
8011
1373
zld-RF
FBtr0344733
NM_001258839
5919
1367
zld-RG
FBtr0345014
NM_001298537
7157
1596
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
UniProt
RefSeq ID
GenBank
zld-PA
FBpp0074577
170.3
1596
6.55
Q9VWC6
NP_608356
AAF49020
zld-PB
FBpp0074576
170.3
1596
6.55
Q9VWC6
NP_728304
AAN09519
zld-PD
FBpp0300192
146.1
1373
6.90
M9NES1
NP_001245769
AFH07481
zld-PF
FBpp0311065
145.3
1367
6.79
X2BZL0
NP_001245768
AFH07480
zld-PG
FBpp0311265
170.3
1596
6.55
Q9VWC6
NP_001285466
AHN59936
Polypeptides with Identical Sequences

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

1596 aa isoforms: zld-PA, zld-PB, zld-PG
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Domain

C2H2-type zinc fingers mediate DNA-binding (PubMed:25538246). C2H2-type zinc finger 2 serves as an inhibitory domain, and is essential for reprogramming the early embryonic genome, but not for its functions later in development (PubMed:29261646).

(UniProt, Q9VWC6)
Crossreferences
InterPro - A database of protein families, domains and functional sites
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\zld using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.43

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 3
organism | ubiquitous

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 1 -- 5
organism
(Liang et al., 2008)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 9 -- 10
dorsal ectoderm
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 11 -- 12
anterior midgut primordium
(Fisher et al., 2012)
central brain primordium
(Fisher et al., 2012)
dorsal epidermis primordium
(Fisher et al., 2012)
embryonic central brain glial cell
(Fisher et al., 2012)
embryonic central brain neuron
(Fisher et al., 2012)
fat body/gonad primordium
(Fisher et al., 2012)
foregut primordium
(Fisher et al., 2012)
hindgut proper primordium
(Fisher et al., 2012)
lateral cord glial cell
(Fisher et al., 2012)
lateral cord neuron
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
somatic muscle primordium
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ectoderm
(Pearson et al., 2012)
embryonic stage 11 -- 16
midline glial cell
(Pearson et al., 2012)
embryonic posterior midline glial cell
(Pearson et al., 2012)
embryonic stage 13 -- 16
dorsal sensory complex proper primordium

Comment: reported as dorsal/lateral sensory complexes

(Fisher et al., 2012)
embryonic brain
(Fisher et al., 2012)
embryonic dorsal epidermis
(Fisher et al., 2012)
embryonic hypopharynx
(Fisher et al., 2012)
embryonic ventral epidermis
(Fisher et al., 2012)
embryonic/larval fat body
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
embryonic/larval nervous system
(Fisher et al., 2012)
embryonic/larval somatic muscle cell
(Fisher et al., 2012)
embryonic/larval visceral muscle cell
(Fisher et al., 2012)
sensory nervous system primordium
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
ventral sensory complex primordium
(Fisher et al., 2012)
embryonic stage 15 -- 16
larval VUM interneuron
(Pearson et al., 2012)
larval VUM motor neuron
(Pearson et al., 2012)
lineage MNB primary interneuron
(Pearson et al., 2012)
embryonic stage 16
embryonic brain
(Pearson et al., 2012)
imaginal disc primordium
(Pearson et al., 2012)
embryonic stage 16 -- 17
embryonic/larval central nervous system
(Liang et al., 2008)
third instar larval stage
optic lobe neuroepithelial cell
(Wang et al., 2013)
wandering third instar larval stage
wing pouch | restricted
(Molnar et al., 2012)
wing hinge primordium | restricted
(Molnar et al., 2012)
adult stage | female
ovary
(Liang et al., 2008)
egg chamber
(Liang et al., 2008)
nurse cell
(Liang et al., 2008)
egg stage
unfertilized egg
(Liang et al., 2008)
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
larval stage
wing disc

Comment: isoforms zld-RD, zld-RB, zld-RF

(Giannios and Tsitilou, 2013)
organism | restricted

Comment: excludes wing discs; isoforms zld-RD, zld-RB, zld-RF

(Giannios and Tsitilou, 2013)
Additional Descriptive Data

zld isoforms zld-RB, zld-RD, and zld-RF can be detected by RT-PCR in wing discs, and in larval tissues other than wing discs; zld-RA is not detected at this stage.

(Giannios and Tsitilou, 2013)

The CNS and brain expression resemble nuclear dots, consistent with a probe detection of pre-spliced nuclear RNA corresponding to zld transcript zld-RD (FBtr0307537). In contrast, the imaginal disc expression appears to be cytoplasmic, likely corresponding to transcript zld-RB (FBtr0074807).

(Pearson et al., 2012)

During stage 12, both anterior midline glial cells and posterior midline glial cells continue to express zld transcript zld-RB, while two midline neurons express zld-RD. At stages 15 and 16, all zld-expressing cells express zld-RD, including iVUM4 (VUM interneuron), mVUM4 (VUM motor neuron), and median neuroblast MNB interneurons (MNB progeny).

(Pearson et al., 2012)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 4
organism
(Nien et al., 2011)
embryonic stage 5
organism | ubiquitous
(Pearson et al., 2012)
embryonic stage 11
embryonic brain
(Pearson et al., 2012)
embryonic/larval central nervous system
(Pearson et al., 2012)
embryonic stage 12
embryonic/larval central nervous system | medial
(Pearson et al., 2012)
embryonic/larval tracheal section
(Pearson et al., 2012)
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 6
organism
(Harrison et al., 2010)
larval stage
wing disc
(Giannios and Tsitilou, 2013)
organism | restricted

Comment: excludes wing discs

(Giannios and Tsitilou, 2013)
Additional Descriptive Data

zld protein is detected in embryos as early as nuclear cycle 2, and appears to accumulate to higher levels in blastoderm embryos. Protein levels appears to increase substantially during the second hour of development, coincident with the activation of the zygotic genome.

(Nien et al., 2011)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
part_of chromatin
inferred from direct assay
(Staudt et al., 2006)
located_in nucleoplasm
inferred from direct assay
(Staudt et al., 2006)
is_active_in nucleus
inferred from direct assay
(Dufourt et al., 2018, Nien et al., 2011)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\zld in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
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
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 25 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 27 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of zld
Transgenic constructs containing regulatory region of zld
Aberrations (Deficiencies and Duplications) ( 1 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal, with Scer\GAL4pnr-MD237
lethal | embryonic stage
lethal | embryonic stage | recessive
lethal | larval stage
lethal | larval stage | recessive
lethal | male
lethal | maternal effect
lethal | pharate adult stage
lethal | pupal stage
lethal | pupal stage, with Scer\GAL4pnr-MD237
lethal | recessive
lethal - all die before end of pupal stage, with Scer\GAL4bi-md653
lethal - all die during embryonic stage | germline clone
partially lethal - majority die
partially lethal - majority die (with zld294)
partially lethal - majority die (with zldJAZ)
partially lethal - majority die (with zldZnF2)
partially lethal - majority die | maternal effect
partially lethal - majority die | maternal effect | recessive | conditional
partially lethal - majority die | recessive
partially lethal - majority live
partially lethal - majority live (with zld294)
partially lethal - majority live (with zldEDD)
partially lethal - majority live (with zldZnF1)
viable
Sterility
Allele
female sterile
fertile
Other Phenotypes
Allele
abnormal mitotic cell cycle
abnormal mitotic cell cycle | embryonic stage, with Scer\GAL4VP16.mat.αTub67C
abnormal neuroanatomy
abnormal neuroanatomy, with Scer\GAL4ftz.ng
abnormal size, with Scer\GAL4en-e16E
abnormal size | adult stage
decreased occurrence of cell division | larval stage, with Scer\GAL4ase.PU
decreased occurrence of cell division | larval stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
majority die during embryonic stage | maternal effect | recessive | conditional
some die during embryonic stage | germline clone
visible, with Scer\GAL4ftz.ng
visible, with Scer\GAL4sca-537.4
visible | adult stage
Phenotype manifest in
Allele
axon, with Scer\GAL4ftz.ng
central nervous system
dorsocentral bristle, with Scer\GAL4sca-537.4
embryo | blastoderm stage
embryo | conditional
embryonic/first instar larval cuticle
embryonic/larval hemocyte | third instar larval stage, with Scer\GAL4Pxn.PS
embryonic/larval neuron
embryonic/larval neuron, with Scer\GAL4ftz.ng
embryonic head
eye
furrow canal | embryonic stage | germline clone
intermediate neuronal progenitor | larval stage, with Scer\GAL4ase.PU
intermediate neuronal progenitor | larval stage, with Scer\GAL4wor.PA, Scer\GAL80ase.PN
nucleus | embryonic stage | germline clone
nucleus | pre-blastoderm stage
presumptive embryonic/larval central nervous system
presumptive embryonic/larval muscle system
presumptive embryonic/larval peripheral nervous system
scutellar bristle, with Scer\GAL4sca-537.4
tormogen cell, with Scer\GAL4sca-537.4
wing, with Scer\GAL4bi-md653
wing | posterior, with Scer\GAL4en-e16E
wing blade, with Scer\GAL4ftz.ng
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (36)
Hsap\BCL6
1 of 14
Yes
Yes
0  
Hsap\FEZF1
1 of 14
Yes
No
Hsap\GFI1B
1 of 14
Yes
No
1  
Hsap\HIC2
1 of 14
Yes
No
Hsap\MYNN
1 of 14
Yes
No
Hsap\OSR1
1 of 14
Yes
No
Hsap\REST
1 of 14
Yes
No
Hsap\SALL1
1 of 14
Yes
No
Hsap\ZBTB14
1 of 14
Yes
No
Hsap\ZBTB47
1 of 14
Yes
No
Hsap\ZBTB48
1 of 14
Yes
No
Hsap\ZFP3
1 of 14
Yes
No
Hsap\ZKSCAN5
1 of 14
Yes
No
Hsap\ZKSCAN8
1 of 14
Yes
No
Hsap\ZNF26
1 of 14
Yes
No
Hsap\ZNF41
1 of 14
Yes
No
1  
Hsap\ZNF48
1 of 14
Yes
No
Hsap\ZNF69
1 of 14
Yes
No
Hsap\ZNF136
1 of 14
Yes
No
Hsap\ZNF175
1 of 14
Yes
No
Hsap\ZNF180
1 of 14
Yes
No
Hsap\ZNF251
1 of 14
Yes
No
Hsap\ZNF266
1 of 14
Yes
Yes
Hsap\ZNF350
1 of 14
Yes
No
Hsap\ZNF423
1 of 14
Yes
No
1  
Hsap\ZNF436
1 of 14
Yes
No
Hsap\ZNF506
1 of 14
Yes
No
Hsap\ZNF541
1 of 14
Yes
No
Hsap\ZNF549
1 of 14
Yes
No
Hsap\ZNF555
1 of 14
Yes
No
Hsap\ZNF559
1 of 14
Yes
No
Hsap\ZNF561
1 of 14
Yes
Yes
Hsap\ZNF580
1 of 14
Yes
Yes
Hsap\ZNF696
1 of 14
Yes
Yes
Hsap\ZSCAN10
1 of 14
Yes
No
Hsap\ZSCAN22
1 of 14
Yes
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (5)
Rnor\Gfi1b
1 of 14
Yes
No
Rnor\Sall1
1 of 14
Yes
No
Rnor\Zbtb33
1 of 14
Yes
Yes
Rnor\Zfp541
1 of 14
Yes
No
Rnor\Zfp580
1 of 14
Yes
Yes
Mus musculus (laboratory mouse) (4)
Mmus\Gfi1b
1 of 14
Yes
No
Mmus\Sall1
1 of 14
Yes
No
Mmus\Zfp541
1 of 14
Yes
No
Mmus\Zfp580
1 of 14
Yes
Yes
Xenopus tropicalis (Western clawed frog) (8)
Xtro\gfi1b
1 of 13
Yes
No
Xtro\LOC100489584
1 of 13
Yes
No
Xtro\LOC100498204
1 of 13
Yes
No
Xtro\sall1
1 of 13
Yes
No
Xtro\siah3
1 of 13
Yes
No
Xtro\zbtb38
1 of 13
Yes
Yes
Xtro\znf420
1 of 13
Yes
No
Xtro\znf853
1 of 13
Yes
No
Danio rerio (Zebrafish) (4)
Drer\sall1b
1 of 14
Yes
No
Drer\si:ch73-40a17.4
1 of 14
Yes
No
Drer\si:ch211-212k18.5
1 of 14
Yes
No
Drer\zbtb26
1 of 14
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (8)
Cele\zfp-2
2 of 14
Yes
Yes
Cele\blmp-1
1 of 14
No
No
Cele\klu-2
1 of 14
No
No
Cele\lsy-2
1 of 14
No
Yes
Cele\Y56A3A.28
1 of 14
No
Yes
Cele\zag-1
1 of 14
No
No
Cele\znf-782
1 of 14
No
No
Cele\ztf-14
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (22)
Agam\AgaP_AGAP002702
8 of 12
Yes
Yes
Agam\AgaP_AGAP001781
2 of 12
No
Yes
Agam\AgaP_AGAP001073
1 of 12
No
No
Agam\AgaP_AGAP001083
1 of 12
No
No
Agam\AgaP_AGAP002163
1 of 12
No
No
Agam\AgaP_AGAP002164
1 of 12
No
Yes
Agam\AgaP_AGAP002165
1 of 12
No
No
Agam\AgaP_AGAP002918
1 of 12
No
No
Agam\AgaP_AGAP005135
1 of 12
No
No
Agam\AgaP_AGAP005390
1 of 12
No
No
Agam\AgaP_AGAP007389
1 of 12
No
No
Agam\AgaP_AGAP007514
1 of 12
No
No
Agam\AgaP_AGAP008032
1 of 12
No
No
Agam\AgaP_AGAP008232
1 of 12
No
No
Agam\AgaP_AGAP008984
1 of 12
No
No
Agam\AgaP_AGAP009010
1 of 12
No
No
Agam\AgaP_AGAP009130
1 of 12
No
No
Agam\AgaP_AGAP009566
1 of 12
No
No
Agam\AgaP_AGAP010500
1 of 12
No
Yes
Agam\AgaP_AGAP010926
1 of 12
No
No
Agam\AgaP_AGAP013246
1 of 12
No
Yes
Agam\AgaP_AGAP013275
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (0)
Saccharomyces cerevisiae (Brewer's yeast) (2)
Scer\ADR1
1 of 13
Yes
Yes
Scer\MOT3
1 of 13
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (2)
Spom\prz1
1 of 12
Yes
Yes
Spom\sfc2
1 of 12
Yes
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:zld. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (49)
ham
2 of 13
ab
1 of 13
Blimp-1
1 of 13
br
1 of 13
Cf2
1 of 13
CG1529
1 of 13
CG1602
1 of 13
CG4282
1 of 13
CG4374
1 of 13
CG4730
1 of 13
CG4820
1 of 13
CG6813
1 of 13
CG7691
1 of 13
CG10348
1 of 13
CG10431
1 of 13
CG11247
1 of 13
CG11902
1 of 13
CG11906
1 of 13
CG12071
1 of 13
CG12236
1 of 13
CG12391
1 of 13
CG12605
1 of 13
CG17328
1 of 13
CG17806
1 of 13
CG32767
1 of 13
CG42726
1 of 13
CG43347
1 of 13
CG44002
1 of 13
Clamp
1 of 13
cliff
1 of 13
erm
1 of 13
esg
1 of 13
hb
1 of 13
ich
1 of 13
Kah
1 of 13
MESR4
1 of 13
MTF-1
1 of 13
Paris
1 of 13
rgr
1 of 13
Sag1
1 of 13
salr
1 of 13
scrt
1 of 13
sens
1 of 13
sna
1 of 13
sqz
1 of 13
wdn
1 of 13
wor
1 of 13
Zaf1
1 of 13
ZIPIC
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    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 ( 1 )
    Allele
    Disease
    Interaction
    References
    zldGFP.FPTB
    ameliorates  Huntington's disease
    modeled by Hsap\HTT128Q.1-231.UAS
    (Onur et al., 2021)
    Disease Associations of Human Orthologs (via DIOPT v9.1 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:
    Show/hide secondary interactors 
    (data from AllianceMine provided by esyN)
    Select Layout:
    Legend:
    Protein
    RNA
    Selected Interactor(s)
    Interactions Browser
    View in Interactions Browser
    Please see the Physical Interaction reports below for full details
    MIST Molecular Interaction Search Tool
    View in MIST tool (external link)
    Please see the Physical Interaction reports below for full details
    RNA-protein
    Physical Interaction
    Assay
    References
    zld - brat
    anti bait coimmunoprecipitation, quantitative reverse transcription pcr, electrophoretic mobility shift assay, autoradiography
    (Reichardt et al., 2018)
    zld - stau
    anti bait coimmunoprecipitation, partial DNA sequence identification by hybridization
    (Laver et al., 2013)
    protein-protein
    Physical Interaction
    Assay
    References
    zld - Kah
    two hybrid array
    (Shokri et al., 2019)
    zld - Ubx
    anti tag coimmunoprecipitation, western blot, proximity-dependent biotin identification, Identification by mass spectrometry, anti tag western blot
    (Carnesecchi et al., 2020)
    zld - mirr
    two hybrid array
    (Shokri et al., 2019)
    zld - pho
    two hybrid array
    (Shokri et al., 2019)
    Summary of Genetic Interactions
    esyN Network Diagram
    Show/hide secondary interactors 
    (data from AllianceMine provided by esyN)
    esyN Network Key:
    Suppression
    Enhancement
    View in Interactions Browser   View in MIST tool (external link)
    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    brat
    suppressible
    zld
    (Reichardt et al., 2018)
    External Data
    Linkouts
    BioGRID - A database of protein and genetic interactions.
    DroID - A comprehensive database of gene and protein interactions.
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    External Data
    Linkouts
    Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-64
    Cytogenetic map
    18F1-18F2
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    18F1-18F2
    Left limit from in situ hybridisation (FBrf0128367) Right limit from in situ hybridisation (FBrf0128367)
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    18F1-18F
    (determined by in situ hybridisation)
    (Peter et al., 2002)
    18F1-18F2
    (determined by in situ hybridisation)
    (Abdelilah-Seyfried et al., 2000, Schäfer et al., 1999.8.12)
    18F-18F
    (determined by in situ hybridisation)
    (Schäfer et al., 1999.6.15, Schäfer et al., 1999.12.2)
    Experimentally Determined Recombination Data
    Location

    1-64

    (FlyBase, 2016)

    1-62.7

    (Comeron et al., 2012)
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (15)
    Genomic Clones (19)
     

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

    cDNA Clones (87)
     

    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 JBrowse 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)
    BDGP DGC clones
    Other clones
    RNAi and Array Information
    Linkouts
    DRSC - Results frm RNAi screens
    Antibody Information
    Laboratory Generated Antibodies
     

    polyclonal

    (Brennan et al., 2023, Koromila et al., 2020, Giannios and Tsitilou, 2013, Nien et al., 2011, Harrison et al., 2010)
    Commercially Available Antibodies
     
    Cell Line Information
    Publicly Available Cell Lines
     
      Other Stable Cell Lines
       

      • New stable cell line derived from S2-unspecified : Stable S2 cell lines containing inducible cDNAs for zld (isoform zld-RB), grh (isoform grh-RH) or twi (isoform twi-RA) were created. Stable S2 cell containing the DNA-binding domains of zld (amino acids 1114-1487) or grh (amino acids 1114-1487) were also created.

        (Gibson et al., 2024)
      Other Comments
      Relationship to Other Genes
      Source for database merge of

      Source for merge of: vfl l(1)G0353 l(1)G0225 l(1)G0427 EP1344

      (Staudt et al., 2006)
      Additional comments

      The vfl gene may have been derived from the CG11902 gene by retroposition.

      (Pan and Zhang, 2009)

      FlyBase curator comment: Correspondence with authors Liang and Rushlow revealed that 'zld' = 'CG12701'.

      (Liang et al., 2008)

      "l(1)G0353" may affect "CG12701".

      (Peter et al., 2002)

      "l(1)G0062" may affect "CG12701". "l(1)G0120" may affect "CG12701". "l(1)G0150" may affect "CG12701". "l(1)G0209" may affect "CG12701". "l(1)G0223" may affect "CG12701". "l(1)G0353" may affect "CG12701". "l(1)G0225" may affect "CG12701".

      (Peter et al., 2002)

      "l(1)G0225" may affect "CG12701".

      (Peter et al., 2002)

      "l(1)G0427" may affect "CG12701".

      (Peter et al., 2002)
      Nomenclature History
      Source for database identify of

      Source for identity of: vfl CG12701

      (Staudt et al., 2006)

      Source for identity of: zld vfl

      (FlyBase, 2009-)
      Nomenclature comments

      Renamed from 'vfl' ('vielfaltig') to 'zld' ('zelda') to reflect clear usage preference in the published literature.

      (FlyBase, 2009-)
      Etymology

      The fullname 'zelda' is itself an acronym for 'zinc finger early drosophila activator'.

      (FlyBase, 2009-)

      For reasons apparent from the mutant phenotype, the gene is named "vielfaltig" (German for versatile or manifold).

      (Staudt et al., 2006)
      Synonyms and Secondary IDs (21)
      Reported As
      Symbol Synonym
      CG12701
      (Sarov et al., 2016, Japanese National Institute of Genetics, 2012.5.21, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Pan and Zhang, 2009, Stofanko et al., 2008, Staudt et al., 2006)
      EP1344
      VFL
      (Blatti et al., 2015, Cheng et al., 2013)
      Vfl
      (Hartmann et al., 2013, Kvon et al., 2012)
      ZLD
      (McDaniel et al., 2019, Lefebvre and Lécuyer, 2018, Li et al., 2014, Giannios and Tsitilou, 2013, Samee and Sinha, 2013)
      Zelda
      (Messina et al., 2023, Stathopoulos and Newcomb, 2020, Tsai et al., 2020, He et al., 2019, McCarthy et al., 2015, Sun et al., 2015, Levine, 2011)
      Zld
      (Hamamoto et al., 2023, Kyrchanova et al., 2023, Pérez-Mojica et al., 2023, Batut et al., 2022, DeAngelis et al., 2021, Espinola et al., 2021, Irizarry and Stathopoulos, 2021, Carnesecchi et al., 2020, Koromila et al., 2020, Schnepf et al., 2020, Small and Arnosti, 2020, Basu et al., 2018, Chambers et al., 2017, Koromila and Stathopoulos, 2017, Samee et al., 2015, Shin and Hong, 2015, Mannervik, 2014, Martinez et al., 2014, Xu et al., 2014, Giannios and Tsitilou, 2013, Saunders et al., 2013, Kvon et al., 2012, Nien et al., 2011, Tsurumi et al., 2011)
      l(1)G0225
      l(1)G0353
      l(1)G0427
      vfl
      (Nadimpalli et al., 2015, dos Santos, 2014, Jiang and Singh, 2014, Rogers et al., 2014, Djiane et al., 2013, Kwon et al., 2013, Spokony, 2013.8.25, Sung et al., 2013, Wang et al., 2013, Yáñez-Cuna et al., 2012, Staudt et al., 2006)
      zelda
      (Connacher and Goldstrohm, 2021, Llorens-Giralt et al., 2021, Parniewska and Stocker, 2020, Eichhorn et al., 2016)
      zld
      (Cho and O'Farrell, 2023, Colonnetta et al., 2023, Beaver et al., 2022, Falo-Sanjuan and Bray, 2022, Velten et al., 2022, Chetverina et al., 2021, Duan et al., 2021, Huang et al., 2021, Onur et al., 2021, Peterson et al., 2021, Winbush and Singh, 2021, Bandodkar et al., 2020, Li and Hidalgo, 2020, Lv et al., 2020, Mahmud et al., 2020, Port et al., 2020, Schloop et al., 2020, Erceg et al., 2019, He et al., 2019, Kwasnieski et al., 2019, Matthews and White, 2019, Shokri et al., 2019, Tiwari et al., 2019, Yamada et al., 2019, Dufourt et al., 2018, Haines and Eisen, 2018, Khajouei and Sinha, 2018, Lucas et al., 2018, Reichardt et al., 2018, Xie and Blankenship, 2018, Barr et al., 2017, Hamm et al., 2017, Hannon et al., 2017, Koenecke et al., 2017, Ribeiro et al., 2017, Levario et al., 2016, Sandler and Stathopoulos, 2016, Sandler and Stathopoulos, 2016, Zhang et al., 2016, Blythe and Wieschaus, 2015, Foo et al., 2014, Darbo et al., 2013, Kanodia et al., 2012, Pearson et al., 2012, Rushlow and Shvartsman, 2012, Harrison et al., 2011, Nien et al., 2011, Struffi et al., 2011, Tsurumi et al., 2011, Harrison et al., 2010, Liang et al., 2008, Liang et al., 2008)
      Name Synonyms
      CG12701
      EP1344
      Vielfaltig
      (Kvon et al., 2012, Yáñez-Cuna et al., 2012)
      ZELDA
      (Samee and Sinha, 2013)
      Zelda
      (Bellec et al., 2022, Bressan and Araujo, 2022, Bozek and Gompel, 2020, Eck et al., 2020, Huang and Saunders, 2020, Omura and Lott, 2020, Smits and Shvartsman, 2020, Lefebvre and Lécuyer, 2018, Mir et al., 2018, Mourad and Cuvier, 2018, Crocker et al., 2017, Kermi et al., 2017, Lefebvre et al., 2017, Liu and Grosshans, 2017, Rosa and Brivanlou, 2017, Elgart et al., 2016, Boija and Mannervik, 2015, Lee et al., 2014, Li et al., 2014, Rembold et al., 2014, Smith and Shilatifard, 2014, Xu et al., 2014, Chen et al., 2013, Darbo et al., 2013, Giannios and Tsitilou, 2013, Hombría and Sotillos, 2013, Saunders et al., 2013, Holmqvist et al., 2012, Kanodia et al., 2012, Kvon et al., 2012, Nien et al., 2011, Ozdemir et al., 2011, Tsurumi et al., 2011, Harrison et al., 2010, Lusk and Eisen, 2010)
      lethal (1) G0225
      lethal (1) G0353
      lethal (1) G0427
      vielfaltig
      (Wang et al., 2013, Staudt et al., 2006)
      vielfaltig/Zelda
      (Laver et al., 2013)
      zelda
      (Diaz-Cuadros et al., 2021, Blythe and Wieschaus, 2015, Dahlberg et al., 2015, Fu et al., 2014, Paris et al., 2013, Tsurumi et al., 2011, Liang et al., 2008, Liang et al., 2008)
      zinc-finger early Drosophila activator
      (Liang et al., 2008)
      Secondary FlyBase IDs
      • FBgn0028317
      • FBgn0031071
      • FBgn0040186
      • FBgn0027202
      • FBgn0043370
      Datasets (0)
      Study focus (0)
      Experimental Role
      Project
      Project Type
      Title
      Study result (0)
      Result
      Result Type
      Title
      External Crossreferences and Linkouts ( 52 )
      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/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
      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
      AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
      BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
      DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
      EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
      FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
      InterPro - A database of protein families, domains and functional sites
      KEGG Genes - Molecular building blocks of life in the genomic space.
      MARRVEL_MODEL - MARRVEL (model organism gene)
      Linkouts
      BioGRID - A database of protein and genetic interactions.
      Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
      DroID - A comprehensive database of gene and protein interactions.
      DRSC - Results frm RNAi screens
      Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
      FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
      FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
      Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
      Flygut - An atlas of the Drosophila adult midgut
      iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
      Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
      MIST (protein-protein) - An integrated Molecular Interaction Database
      Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
      References (204)