FB2021_05, released October 15, 2021

Gene: Dmel\Pc

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General Information
Symbol
Dmel\Pc
Species
D. melanogaster
Name
Polycomb
Annotation Symbol
CG32443
Feature Type
protein_coding_gene
FlyBase ID
FBgn0003042
Gene Model Status
Current
Stock Availability
36 publicly available
Gene Summary
Polycomb (Pc) encodes a chromatin binding protein that interacts with histone H3K27me3. It is involved in epigenetic silencing and its target genes are predominantly transcription factors or receptors that control cell fate and most developmental decisions. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
All Summaries
Also Known As

PcG, Pc-G

Key Links
Genomic Location
Cytogenetic map
78C6-78C7
Sequence location
Recombination map
3-47
RefSeq locus
NT_037436 REGION:21306138..21318020
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
GO Summary Ribbons
Gene Ontology (GO) Annotations (24 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
enables chromatin binding
inferred from direct assay
(Enderle et al., 2011, Negre et al., 2006)
enables DNA binding
inferred from direct assay
(Ogiyama et al., 2018)
enables methylated histone binding
inferred from direct assay
(Czermin et al., 2002)
enables protein binding
inferred from physical interaction with UniProtKB:P42124,UniProtKB:Q9XZ06
(assigned by UniProt )
(Gozalo et al., 2020)
enables sequence-specific DNA binding
inferred from direct assay
(Ogiyama et al., 2018)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (13 terms)
Terms Based on Experimental Evidence (13 terms)
CV Term
Evidence
References
involved_in anterior/posterior axis specification
inferred from mutant phenotype
(assigned by UniProt )
(Simon et al., 1992)
involved_in gene silencing
inferred from mutant phenotype
(Roseman et al., 2001)
inferred from genetic interaction with FLYBASE:ph-p; FB:FBgn0004861
(Roseman et al., 2001)
involved_in negative regulation of gene expression
inferred from mutant phenotype
(Nishioka et al., 2018, Zhang et al., 2017)
involved_in negative regulation of response to gamma radiation
inferred from mutant phenotype
(Zhang et al., 2008)
involved_in negative regulation of transcription, DNA-templated
inferred from mutant phenotype
(Enderle et al., 2011)
involved_in neurogenesis
inferred from mutant phenotype
(Bello et al., 2007)
involved_in neuron remodeling
inferred from mutant phenotype
(Wang et al., 2006)
involved_in regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(assigned by UniProt )
(Simon et al., 1992)
involved_in regulation of transcription, DNA-templated
inferred from mutant phenotype
(Goodliffe et al., 2007)
involved_in specification of segmental identity, abdomen
inferred from mutant phenotype
(Chen et al., 2005)
involved_in syncytial blastoderm mitotic cell cycle
inferred from mutant phenotype
(O'Dor et al., 2006)
involved_in ventral cord development
inferred from high throughput mutant phenotype
(Bivik et al., 2015)
involved_in wound healing
inferred from high throughput mutant phenotype
(Campos et al., 2010)
Terms Based on Predictions or Assertions (0 terms)
Cellular Component (6 terms)
Terms Based on Experimental Evidence (6 terms)
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
(Chen et al., 2005)
located_in nucleolus
inferred from direct assay
(Chen et al., 2011, Chen et al., 2005)
located_in nucleus
inferred from direct assay
(Grimaud et al., 2006, Mulholland et al., 2003)
located_in polytene chromosome band
inferred from direct assay
(Marchetti et al., 2003)
located_in polytene chromosome
inferred from direct assay
(Franke et al., 1992)
part_of PRC1 complex
inferred from direct assay
(Mulholland et al., 2003)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
Polycomb (Pc) encodes a chromatin binding protein that interacts with histone H3K27me3. It is involved in epigenetic silencing and its target genes are predominantly transcription factors or receptors that control cell fate and most developmental decisions. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
POLYCOMB REPRESSIVE COMPLEX 1 (CORE SUBUNITS) -
The Polycomb repressive complex 1 (PRC1) is a Polycomb group complex. It binds to histone H3K27me3 (histone H3 K27 methylation is catalyzed by PRC2) and monoubiquitinates H2A on K119. This leads to chromatin compaction, RNA polymerase II stalling and gene silencing. (Adapted from FBrf0179742 and FBrf0228921).
Protein Function (UniProtKB)
Polycomb group (PcG) protein. PcG proteins act by forming multiprotein complexes, which are required to maintain the transcriptionally repressive state of homeotic genes throughout development. PcG proteins are not required to initiate repression, but to maintain it during later stages of development. Component of the PcG multiprotein PRC1 complex, a complex that acts via chromatin remodeling and modification of histones; it mediates monoubiquitination of histone H2A 'Lys-118', rendering chromatin heritably changed in its expressibility. Promotes locus-specific chromatin compaction.
(UniProt, P26017)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
Pc: Polycomb
Pc+ may be considered a negative regulator of the bithorax complex (BXC) and the Antennapedia complex (ANTC), with a decreasing gradient of activity from anterior to posterior. When homozygous or hemizygous, Pc mutants are late embryonic lethals. Embryos with at least one dose of the BXC show incomplete head development and caudad transformations, the thoracic and first seven abdominal segments being partially transformed into the eighth abdominal segment (Lewis, 1978; Denell, 1982; Haynie, 1983, Dev. Biol. 100: 399-411; Denell and Frederick, 1983, Dev. Biol. 97: 34-47). This homeotic effect in homozygotes is enhanced by increasing the dosage of the BXC. Transformations involve brain and ventral nerve cord as well as epidermis (Jimenez and Campos-Ortega, 1981). Pc+ alleles in the mother weaken the homeotic effect (Denell, 1982; Lawrence, Johnson, and Struhl, 1983, Cell 35: 27-34). Pc2/Pc2 or Pc3/Pc3 clones induced in leg and eye-antennal tissue during larval development also show similar posteriorly-directed transformations (Struhl, 1981; Duncan and Lewis, 1982). Pc/+ flies carrying at least one dose of the BXC show caudad transformations, i.e. partial conversion of wings into halteres and of anterior abdominal segments into more posterior ones. Some Pc heterozygotes show phenotypes characteristic of ANTC mutants, i.e. partial conversion of antennae into legs and of second and third legs into first legs (with sex combs in males) (Hannah-Alava, 1958; Duncan, 1982). The frequency of wing transformations varies directly with the BXC dosage, but does not seem to be changed by variation in ANTC dosage (Duncan and Lewis, 1982; Botas et al., 1982). The number of abdominal transformations, however, varies inversely with the doses of the BXC while it increases as the doses of the ANTC are increased (Duncan, 1982; Duncan and Lewis, 1982). Other changes observed in Pc/+ flies include a transformation of ventral to dorsal wing (Tiong; Sato et al., 1983), elevated, divergent, or crinkled wings, terminal gaps in the L4 wing vein, bent humeral or notopleural bristles, and defective sternopleural bristles, all abnormalities being less extreme in males than in females (sometimes absent in males). When doubly heterozygous with AntpYu and AntpB, Pc enhances Antp. The expression of all Pc mutant heterozygotes (including deficiencies for the locus) is enhanced by the second chromosome dominant, E(Pc) (Sato et al., 1983, 1984). Pc3/Pc3/Dp(1;3;4)7 flies (carrying a Pc+ duplication) show stronger leg and wing transformations than E(Pc)/+;Pc3/+ flies (Duncan and Lewis, 1982; Sato et al., 1983).
Summary (Interactive Fly)

transcription factor - chromo domain - Polycomb group - chromatin associated protein - involved in gene silencing chromatin binding protein that interacts with histone H3K27me3 - Pc target genes are predominantly transcription factors or receptors that control cell fate and most developmental decisions

Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Pc 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 5.45

Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Pc-RA
FBtr0078405
NM_079475
2462
390
Additional Transcript Data and Comments
Reported size (kB)

2.5, 2.0 (northern blot)

(Paro and Hogness, 1991)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
Pc-PA
FBpp0078059
44.0
390
7.64
NP_524199
AAF51707
Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments
Reported size (kDa)

390 (aa); 44 (kD predicted)

(Paro and Hogness, 1991)
Comments

A variation on the in vivo formaldehyde cross-linking technique was used to map Pc-interacting sites along the chromatin in the BX-C region. Pc was found to quantitatively cover large regulatory regions of repressed BX-C genes. Pc did not bind the transcriptionally active Abd-B region.

(Orlando and Paro, 1993)

Pc protein and polyhomeotic protein are coprecipitated by antibodies to either showing that they are constituents of a multimeric protein complex. Pc and polyhomeotic complexes can be coprecipitated over the entire period of embryogenesis. The molecular weight of the complex is ~2-5x103kD. Double-immunofluorescence labeling was used to show that Pc protein and polyhomeotic protein have exactly the same binding patterns on polytene chromosomes.

(Franke et al., 1992)

Pc protein is associated with ~60 sites on the

polytene chromosomes. It is associated with loci shown to interact with

Pc by genetic methods. It is abundant at the ANT-C and the BX-C and is

found at several locations of Pc-group genes. The Pc protein expressed

in salivary glands is encoded by a 1.0kb Pc transcript and is smaller

than the 390aa Pc protein.

(Zink and Paro, 1989)
External Data
Subunit Structure (UniProtKB)

Component of PRC1 complex, which contains many PcG proteins like Pc, ph, Scm, Psc, Sce and also chromatin-remodeling proteins such as histone deacetylases. This complex is distinct from the Esc/E(z) complex, at least composed of esc, E(z), Su(z)12, HDAC1/Rpd3 and Caf1-55. The 2 complexes however cooperate and interact together during the first 3 hours of development to establish PcG silencing (PubMed:11493925, PubMed:11583617). Interacts with stx; the interaction targets Pc for ubiquitin-independent proteasomal degradation (PubMed:27326929). Interacts with Nup93-1 (PubMed:31784359).

(UniProt, P26017)
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\Pc 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
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
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
larval stage
embryonic/larval salivary gland
(Paro and Hogness, 1991)
unfertilized egg stage
unfertilized egg
(Paro and Hogness, 1991)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
organism | ubiquitous
(Paro and Hogness, 1991)
embryonic stage 1 -- 3
organism | ubiquitous

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4
organism | ubiquitous
(Paro and Hogness, 1991)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 9 -- 10
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 11 -- 12
anterior midgut primordium
(Fisher et al., 2012)
central brain primordium
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic brain
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
embryonic/larval nervous system
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
embryonic stage 16 -- 17
embryonic/larval central nervous system
(Paro and Hogness, 1991)
unfertilized egg stage
unfertilized egg
(Paro and Hogness, 1991)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 1 -- 12

Comment: reference states 0-8 hr AEL

(Paro and Zink, 1992)
unfertilized egg stage
unfertilized egg
(Paro and Zink, 1992)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
embryonic/larval salivary gland
(Paro and Zink, 1992)
embryonic/larval pharynx
(Paro and Zink, 1992)
presumptive embryonic/larval central nervous system
(Franke et al., 1992, Paro and Zink, 1992)
adult ventral nerve cord
(Paro and Zink, 1992)
gnathal ganglion
(Paro and Zink, 1992)
supraesophageal ganglion
(Paro and Zink, 1992)
embryonic stage 4 -- 9
organism | ubiquitous
(Paro and Zink, 1992)
embryonic stage 7 -- 9
mitotic domain
(Franke et al., 1992)
embryonic stage 16
embryonic/larval central nervous system
(Franke et al., 1992)
larval stage
embryonic/larval fat body | anterior
(Marchetti et al., 2003)
third instar larval stage
embryonic/larval salivary gland
(Franke et al., 1992)
adult stage & oogenesis
germarium
(Paro and Zink, 1992)
adult stageoogenesis stage S1 -- S2
egg chamber
(Paro and Zink, 1992)
adult stageoogenesis stage S9 -- S14
nurse cell
(Paro and Zink, 1992)
adult stageoogenesis stage S10 -- S14
follicle cell
(Paro and Zink, 1992)
Additional Descriptive Data

The Pc protein accumulates in the region of the bithorax complex genes on polytene chromosomes from anterior regions of larval fat bodies where the homeotic genes are repressed.

(Marchetti et al., 2003)

The distribution of polyhomeotic protein and Pc protein is nearly indistinguishable in embryos.

(Franke et al., 1992)

Pc protein is expressed in ovaries and during all stages of embryogenesis. Protein levels increase with developmental time in embryos. Two phases of Pc expression are observed in ovaries. First, substantial Pc protein is found in the germarium and in stage S1 and S2 follicles. Protein levels decline during stages S3-S8 and rise again in stages S9 and S10. In the germline, most of the protein is found in the polyploid nurse cell nuclei. Some Pc protein is deposited in the oocyte but none is observed in the nucleus. A very dynamic pattern of expression involving overlapping gradients is observed in follicle cells during stages S10-S14. In embryos, Pc transcripts are homogenously distributed over the length of the embryo at the cellular blastoderm stage and in germ band extended embryos. In late stage embryos, transcripts are found predominantly in the CNS. Particularly strong expression is observed in the supra-, suboesophageal and anterior thoracic ganglia. Strong staining is also observed in the pharynx, salivary glands, and other internal tissues.

(Paro and Zink, 1992)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in chromatin
inferred from direct assay
(Chen et al., 2005)
located_in nucleolus
inferred from direct assay
(Chen et al., 2011, Chen et al., 2005)
located_in nucleus
inferred from direct assay
(Grimaud et al., 2006, Mulholland et al., 2003)
located_in polytene chromosome band
inferred from direct assay
(Marchetti et al., 2003)
located_in polytene chromosome
inferred from direct assay
(Franke et al., 1992)
part_of PRC1 complex
inferred from direct assay
(Mulholland et al., 2003)
Expression Deduced from Reporters
Reporter: P{Pc-EGFP}
Stage
Tissue/Position (including subcellular localization)
Reference
spermatogenesis
primary gonial cell
(Chen et al., 2005)
spermatocyte
(Chen et al., 2005)
spermatid

Comment: round spermatid

(Chen et al., 2005)
elongation stage spermatid
(Chen et al., 2005)
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Pc in GBrowse 2
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 ( 52 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 39 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Pc
Transgenic constructs containing regulatory region of Pc
Aberrations (Deficiencies and Duplications) ( 20 )
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal | heat sensitive
lethal | male, with Scer\GAL4Act5C.PI
lethal | recessive
lethal - all die before end of pupal stage | heat sensitive, with tsrUAS.int/tsrΔ96
lethal - all die before end of pupal stage | heat sensitive, with tsrΔ96, tsrUAS.int
lethal - all die during embryonic stage | heat sensitive
lethal - all die during embryonic stage | homeotic | recessive
lethal - all die during embryonic stage | recessive
Other Phenotypes
Allele
abnormal mitotic cell cycle | embryonic stage
abnormal mitotic cell cycle | somatic clone
abnormal neuroanatomy
abnormal neuroanatomy | somatic clone
decreased cell number | somatic clone
decreased cell number | spermatogenesis, with Scer\GAL4C587
decreased occurrence of cell division | embryonic stage
enhancer of variegation, with P{418}E(Pc)17BT2.2.31
enhancer of variegation, with P{418}I7.1
enhancer of variegation, with P{418}I33.1
enhancer of variegation, with P{418}T3
enhancer of variegation, with P{418}T13.4
enhancer of variegation, with P{418}T13.5
enhancer of variegation, with P{418}T26
enhancer of variegation, with P{418}T30
enhancer of variegation, with P{418}T33.1.2
enhancer of variegation, with P{418}T33.1.24j
enhancer of variegation, with P{418}T40
enhancer of variegation, with P{418}T42.1
enhancer of variegation, with P{418}T43
enhancer of variegation, with P{418}T72.1
enhancer of variegation, with P{419}6.1
homeotic
homeotic | adult stage
homeotic | adult stage, with Scer\GAL4Dll.PU
homeotic | dominant
homeotic | dominant | male
homeotic | embryonic stage
increased cell number | embryonic stage
increased cell number | somatic clone | spermatogenesis
increased cell number | spermatogenesis, with Scer\GAL4C587
increased occurrence of cell division | embryonic stage
increased occurrence of cell division | spermatogenesis, with Scer\GAL4C587
melanotic mass phenotype, with Scer\GAL4SPARC-MI00329-GAL4
neoplasia | somatic clone
non-enhancer of variegation, with P{418}e(Pc)84DET66.1
non-enhancer of variegation, with P{418}I2.1.2
non-enhancer of variegation, with P{418}T15.2
non-enhancer of variegation, with P{418}T33.1.5
non-enhancer of variegation, with P{418}T44
non-enhancer of variegation, with P{418}T53.3
non-enhancer of variegation, with P{418}T63
non-enhancer of variegation, with P{418}T69.3
non-enhancer of variegation, with P{418}T72.2
non-enhancer of variegation, with P{hsp26-pt-T}39C-5
non-suppressor of variegation, with P{hsp26-pt-T}39C-5
non-suppressor of variegation | dominant, with In(1)wm4
non-suppressor of variegation | dominant, with P{hsp26-pt-T}39C-5
some die during embryonic stage | heat sensitive
some die during embryonic stage | recessive
some die during pupal stage | heat sensitive, with tsrUAS.int/tsrΔ96
some die during pupal stage | heat sensitive, with tsrΔ96, tsrUAS.int
suppressor of variegation, with P{UAS-lacZ.Abd-B.24F6}GCD-6/+:
suppressor of variegation | dominant, with P{hsp26-pt-T}39C-5
suppressor of variegation | dominant, with P{hsp26-pt-T}39C-27
suppressor of variegation | dominant, with P{hsp26-pt-T}39C-31
suppressor of variegation | dominant, with P{SUPor-P.PRE#6}69M3
suppressor of variegation | dominant, with P{UAS-lacZ.Abd-B.5F24}25,2a
suppressor of variegation | dominant, with P{wA}4-4
visible
visible, with Scer\GAL4Tub.PU
visible | adult stage, with Scer\GAL4Dll.PU
visible | adult stage | dominant
visible | dominant
visible | dominant | heat sensitive
visible | dominant | homeotic
visible | dominant | homeotic | heat sensitive
visible | embryonic stage
visible | homeotic
visible | recessive
visible | somatic clone
wild-type
Phenotype manifest in
Allele
abdominal segment
abdominal segment 4 | male
abdominal segment 5 | ectopic | male
abdominal tergite 4 | male
adult abdominal segment 1
adult abdominal segment 4
adult abdominal segment 4 | male
adult abdominal segment 5
adult abdominal segment 5 | ectopic | male
adult abdominal segment 6
adult external mesothorax
adult external prothorax | ectopic
adult optic lobe, with tsrUAS.int/tsrΔ96
adult optic lobe, with tsrΔ96, tsrUAS.int
anaphase | embryonic stage
antenna
antenna, with Scer\GAL4Dll.PU
apterous-expressing neuron of the lateral cluster of the thorax
apterous-expressing neuron of the lateral cluster of the thorax Ap4
cyst cell of testis, with Scer\GAL4C587
cyst progenitor cell | somatic clone | spermatogenesis
cyst progenitor cell | spermatogenesis, with Scer\GAL4C587
dendrite & dorsal multidendritic neuron ddaC | somatic clone
dendrite | somatic clone
denticle belt | embryonic stage
embryonic/larval brain | embryonic stage
embryonic abdominal segment
embryonic abdominal segment 1
embryonic abdominal segment 2
embryonic abdominal segment 3
embryonic abdominal segment 4
embryonic abdominal segment 5
embryonic abdominal segment 6
embryonic abdominal segment 7
embryonic abdominal segment 8
embryonic abdominal segment 9
embryonic abdominal segment 10
embryonic abdominal segment 11
embryonic head
embryonic midgut constriction
embryonic thoracic segment
eye, with tsrUAS.int/tsrΔ96
eye, with tsrΔ96, tsrUAS.int
eye | somatic clone
eye disc | somatic clone
fat body, with Scer\GAL4SPARC-MI00329-GAL4
genital disc primordium | ectopic
germinal proliferation center | somatic clone | spermatogenesis
germinal proliferation center | spermatogenesis, with Scer\GAL4C587
gonadal sheath proper primordium (with Pc3)
gonadal sheath proper primordium (with Pc15)
haltere | ectopic
haltere | ectopic | female
lamina, with tsrUAS.int/tsrΔ96
lamina, with tsrΔ96, tsrUAS.int
larval abdominal segment 4
larval abdominal segment 5 | ectopic
larval head
larval segment
leg
leg | ectopic
male germline stem cell | somatic clone | spermatogenesis
male germline stem cell | spermatogenesis, with Scer\GAL4C587
melanotic mass, with Scer\GAL4SPARC-MI00329-GAL4
mesothoracic leg
mesothoracic leg & sex comb | ectopic
mesothoracic leg & sex comb tooth | ectopic
mesothoracic leg | male
mesothoracic segment
metathoracic leg
metathoracic leg & sex comb | ectopic
metathoracic leg & sex comb tooth | ectopic
metathoracic segment
morphogenetic furrow | somatic clone
neuroblast | somatic clone
ommatidium, with tsrUAS.int/tsrΔ96
ommatidium, with tsrΔ96, tsrUAS.int
photoreceptor cell | somatic clone
presumptive embryonic/larval central nervous system
primary gonial cell | somatic clone | spermatogenesis
primary gonial cell | spermatogenesis, with Scer\GAL4C587
prothoracic leg | ectopic
prothoracic leg | ectopic | male
retina, with tsrUAS.int/tsrΔ96
retina, with tsrΔ96, tsrUAS.int
rhabdomere, with tsrUAS.int/tsrΔ96
rhabdomere, with tsrΔ96, tsrUAS.int
rhabdomere membrane, with tsrUAS.int/tsrΔ96
rhabdomere membrane, with tsrΔ96, tsrUAS.int
sex comb
sex comb (with Df(3R)ry27)
sex comb (with Df(3R)ry619)
sex comb | ectopic
sex comb | ectopic | heat sensitive
sex comb | ectopic | male
sex comb | increased number
sex comb tooth
sex comb tooth | ectopic
sternopleural bristle
telophase | embryonic stage
tergite, with Scer\GAL4Tub.PU
ventral mesothoracic disc primordium
wing
wing | female
wing | somatic clone
Orthologs
Downloads
Download All DIOPT Orthologs
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (9)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\CBX8
9 of 15
Yes
Yes
Hsap\CBX6
8 of 15
No
Yes
Hsap\CBX4
6 of 15
No
Yes
Hsap\CBX7
6 of 15
No
Yes
Hsap\CBX2
5 of 15
No
Yes
Hsap\CBX1
1 of 15
No
No
2  
Hsap\CBX3
1 of 15
No
No
3  
Hsap\CBX5
1 of 15
No
No
4  
Hsap\MPHOSPH8
1 of 15
No
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (9)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Cbx8
9 of 15
Yes
Yes
Mmus\Cbx6
7 of 15
No
Yes
Mmus\Cbx4
6 of 15
No
Yes
Mmus\Cbx7
6 of 15
No
Yes
Mmus\Cbx2
4 of 15
No
Yes
 
1  
Mmus\Npcd
4 of 15
No
Yes
Mmus\Cbx1
1 of 15
No
No
1  
Mmus\Cbx5
1 of 15
No
No
Mmus\Mphosph8
1 of 15
No
No
Rattus norvegicus (Norway rat) (9)
Rnor\Cbx8
6 of 13
Yes
Yes
Rnor\Cbx7
5 of 13
No
Yes
Rnor\Cbx2
4 of 13
No
Yes
Rnor\Cbx6
3 of 13
No
Yes
Rnor\Cbx4
2 of 13
No
Yes
Rnor\Cbx1
1 of 13
No
No
Rnor\Cbx3
1 of 13
No
No
Rnor\Cbx5
1 of 13
No
No
Rnor\Mphosph8
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (10)
Xtro\cbx7
6 of 12
Yes
Yes
Xtro\cbx8
6 of 12
Yes
Yes
Xtro\cbx6
4 of 12
No
Yes
Xtro\cbx2
3 of 12
No
Yes
Xtro\cbx4
3 of 12
No
Yes
Xtro\cbx1
1 of 12
No
No
Xtro\cbx3
1 of 12
No
No
Xtro\cbx5
1 of 12
No
No
Xtro\LOC108646164
1 of 12
No
No
Xtro\mphosph8
1 of 12
No
Yes
Danio rerio (Zebrafish) (16)
Drer\cbx8a
8 of 15
Yes
Yes
Drer\cbx6a
7 of 15
No
Yes
Drer\cbx7b
5 of 15
No
Yes
Drer\cbx8b
5 of 15
No
Yes
Drer\cbx4
4 of 15
No
Yes
Drer\cbx6b
4 of 15
No
Yes
Drer\cbx2
3 of 15
No
Yes
Drer\cbx7a
3 of 15
No
Yes
Drer\cbx1a
1 of 15
No
No
Drer\cbx1b
1 of 15
No
No
Drer\cbx3a
1 of 15
No
No
Drer\cbx3b
1 of 15
No
No
Drer\cbx5
1 of 15
No
No
Drer\mphosph8
1 of 15
No
No
Drer\nptxra
1 of 15
No
Yes
Drer\nptxrb
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (9)
Cele\cec-1
4 of 15
Yes
Yes
Cele\C38D4.10
1 of 15
No
Yes
Cele\cec-4
1 of 15
No
No
Cele\cec-5
1 of 15
No
Yes
Cele\cec-8
1 of 15
No
No
Cele\H04D03.2
1 of 15
No
Yes
Cele\heri-1
1 of 15
No
No
Cele\hpl-1
1 of 15
No
No
Cele\hpl-2
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (1)
Atha\TFL2
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (0)
No records found.
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG09190DG5 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00004250
Drosophila simulans
Dsim\GD12124
Drosophila sechellia
Dsec\GM22147
Drosophila erecta
Dere\GG13243
Drosophila yakuba
Dyak\GE22343
Drosophila yakuba
Dyak\GE22965
Drosophila ananassae
Dana\GF24200
Drosophila pseudoobscura pseudoobscura
Dpse\GA16908
Drosophila persimilis
Dper\GL24588
Drosophila willistoni
Dwil\GK20423
Drosophila virilis
Dvir\GJ12080
Drosophila mojavensis
Dmoj\GI13310
Drosophila grimshawi
Dgri\GH16158
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150CWN )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17020037
Glossina morsitans
Tsetse fly
Gmor\GMOY009614
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL014098
Anopheles darlingi
American malaria mosquito
Adar\ADAC000737
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ008739
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0ESN )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bmor\BGIBMGA006904
Danaus plexippus
Monarch butterfly
Dple\DPOGS212988-PA
Heliconius melpomene
Postman butterfly
Hmel\HMEL007695
Apis florea
Little honeybee
Aflo\2209622596
Apis mellifera
Western honey bee
Amel\Pc
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP17250
Bombus terrestris
Buff-tailed bumblebee
Bter\15210334
Linepithema humile
Argentine ant
Lhum\LH25013
Megachile rotundata
Alfalfa leafcutting bee
Mrot\MROTU:002a47
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG002385
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_07949
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_06936
Tribolium castaneum
Red flour beetle
Tcas\Pc
Pediculus humanus
Human body louse
Phum\PHUM001810
Rhodnius prolixus
Kissing bug
Rpro\RPRC011889
Cimex lectularius
Bed bug
Clec\CLEC010099
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI002201
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_09211
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0ES0 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR012867
Strigamia maritima
European centipede
Smar\SMAR002003
Ixodes scapularis
Black-legged tick
Isca\ISCW012843
Stegodyphus mimosarum
African social velvet spider
Smim\KFM78333.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM67605.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur05g04900
Tetranychus urticae
Two-spotted spider mite
Turt\tetur14g03640
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_191862
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0NOT )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Cbxh4
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (8)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
CG8289
1 of 10
HP1b
1 of 10
HP1c
1 of 10
HP1e
1 of 10
HP6
1 of 10
Oxp
1 of 10
rhi
1 of 10
Su(var)205
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
Disease Model Summary Ribbon
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Allele
Disease
Evidence
References
Pc15
model of  cancer
CEA
(Classen et al., 2009)
Potential Models Based on Orthology ( 1 )
Human Ortholog
Disease
Evidence
References
CBX2; chromobox 2
model of  46,XY sex reversal 5
IEA
(FlyBase, 2019-)
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?
CBX8; chromobox 8
9 of 15
617354
      CBX6; chromobox 6
      8 of 15
      617438
          CBX4; chromobox 4
          6 of 15
          603079
              CBX7; chromobox 7
              6 of 15
              608457
                  CBX2; chromobox 2
                  5 of 15
                  602770
                  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.
                  Dmel gene
                  Ortholog showing functional complementation
                  Supporting References
                  Pc
                  Mmus\Cbx2
                  (Muller et al., 1995)
                  Interactions
                  Summary of Physical Interactions
                  esyN Network Diagram
                  Show neighbor-neighbor interactions:
                  Select Layout:
                  Legend:
                  Protein
                  RNA
                  Selected Interactor(s)
                  Interactions Browser
                  View in Interactions Browser
                  Please see the Physical Interaction reports below for full details
                  protein-protein
                  Physical Interaction
                  Assay
                  References
                  Pc - Adf1
                  anti bait coimmunoprecipitation, western blot
                  (Orsi et al., 2014)
                  Pc - Br140
                  tandem affinity purification, Identification by mass spectrometry, pull down, peptide massfingerprinting
                  (Kang et al., 2015, Strübbe et al., 2011)
                  Pc - E(z)
                  anti bait coimmunoprecipitation, western blot
                  (Poux et al., 2001)
                  Pc - Eaf6
                  pull down, peptide massfingerprinting
                  (Strübbe et al., 2011)
                  Pc - HDAC1
                  anti bait coimmunoprecipitation, autoradiography, western blot
                  (Tie et al., 2016, Poux et al., 2001)
                  Pc - His2A
                  affinity technology, peptide array, western blot
                  (Stein et al., 2013)
                  Pc - His2Av
                  affinity technology, peptide array, western blot
                  (Stein et al., 2013)
                  Pc - His2B
                  affinity technology, peptide array, western blot
                  (Stein et al., 2013)
                  Pc - His3
                  pull down, autoradiography, western blot, affinity technology, peptide array
                  (Stein et al., 2013, Tie et al., 2007, Cao et al., 2002, Czermin et al., 2002)
                  Pc - His4
                  affinity technology, peptide array, western blot
                  (Stein et al., 2013)
                  Pc - Nup93-1
                  anti bait coimmunoprecipitation, western blot
                  (Gozalo et al., 2020)
                  Pc - Nup93-2
                  anti bait coimmunoprecipitation, western blot
                  (Gozalo et al., 2020)
                  Pc - PR-Set7
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Lv et al., 2016)
                  Pc - Polr2A
                  anti bait coimmunoprecipitation, western blot
                  (Tie et al., 2016)
                  Pc - Pp1-87B
                  pull down, peptide massfingerprinting
                  (Strübbe et al., 2011)
                  Pc - Psc
                  anti tag coimmunoprecipitation, western blot, anti bait coimmunoprecipitation, two hybrid, Identification by mass spectrometry, tandem affinity purification, pull down, autoradiography, molecular weight estimation by staining, affinity chromatography technology, peptide massfingerprinting, molecular sieving
                  (Kang et al., 2020, Kang et al., 2017, Tie et al., 2016, Kang et al., 2015, Fereres et al., 2014, Mohd-Sarip et al., 2012, Strübbe et al., 2011, Lagarou et al., 2008, Peterson et al., 2004, Breiling et al., 2001, Poux et al., 2001, Shao et al., 1999, Kyba and Brock, 1998, Strutt and Paro, 1997)
                  Pc - SMC1
                  pull down, peptide massfingerprinting, anti bait coimmunoprecipitation, western blot
                  (Strübbe et al., 2011)
                  Pc - SMC3
                  pull down, peptide massfingerprinting, anti bait coimmunoprecipitation, western blot
                  (Strübbe et al., 2011)
                  Pc - Sce
                  anti bait coimmunoprecipitation, Identification by mass spectrometry, western blot, pull down, experimental knowledge based, peptide massfingerprinting, autoradiography, tandem affinity purification
                  (Kang et al., 2015, Fereres et al., 2014, Gambetta and Müller, 2014, Mohd-Sarip et al., 2012, Guruharsha et al., 2011, Strübbe et al., 2011, Lo et al., 2009, Lagarou et al., 2008, Gorfinkiel et al., 2004)
                  Pc - Scm
                  tandem affinity purification, Identification by mass spectrometry, anti tag coimmunoprecipitation, western blot, molecular sieving
                  (Kang et al., 2015, Shao et al., 1999)
                  Pc - Sfmbt
                  pull down, western blot, tandem affinity purification, Identification by mass spectrometry, peptide massfingerprinting
                  (Kang et al., 2015, Strübbe et al., 2011)
                  Pc - Su(var)205
                  anti bait coimmunoprecipitation, western blot
                  (Cabrera et al., 2015)
                  Pc - Su(z)2
                  anti bait coimmunoprecipitation, western blot, affinity chromatography technology, molecular weight estimation by staining, tandem affinity purification, Identification by mass spectrometry, pull down, peptide massfingerprinting
                  (Kang et al., 2015, Strübbe et al., 2011, Lo et al., 2009)
                  Pc - Taf4
                  anti bait coimmunoprecipitation, western blot
                  (Saurin et al., 2001)
                  Pc - Taf5
                  anti bait coimmunoprecipitation, western blot
                  (Saurin et al., 2001)
                  Pc - Taf9
                  anti bait coimmunoprecipitation, western blot
                  (Saurin et al., 2001)
                  Pc - Tbp
                  anti bait coimmunoprecipitation, western blot
                  (Breiling et al., 2001, Saurin et al., 2001)
                  Pc - TfIIB
                  anti bait coimmunoprecipitation, western blot
                  (Breiling et al., 2001)
                  Pc - Trl
                  anti bait coimmunoprecipitation, autoradiography
                  (Poux et al., 2001)
                  Pc - br
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Lv et al., 2016)
                  Pc - cg
                  anti bait coimmunoprecipitation, western blot
                  (Ray et al., 2016)
                  Pc - corto
                  anti tag coimmunoprecipitation, western blot
                  (Salvaing et al., 2003)
                  Pc - ebi
                  pull down, peptide massfingerprinting
                  (Strübbe et al., 2011)
                  Pc - enok
                  tandem affinity purification, Identification by mass spectrometry, pull down, peptide massfingerprinting
                  (Kang et al., 2015, Strübbe et al., 2011)
                  Pc - esc
                  anti bait coimmunoprecipitation, western blot, autoradiography
                  (Poux et al., 2001)
                  Pc - fs(1)h
                  pull down, peptide massfingerprinting, tandem affinity purification, Identification by mass spectrometry, western blot
                  (Kang et al., 2015, Strübbe et al., 2011)
                  Pc - grh
                  pull down, peptide massfingerprinting
                  (Strübbe et al., 2011)
                  Pc - nej
                  anti bait coimmunoprecipitation, western blot, pull down, enzymatic study
                  (Tie et al., 2016)
                  Pc - ph-d
                  anti bait coimmunoprecipitation, western blot, pull down, peptide massfingerprinting, tandem affinity purification, Identification by mass spectrometry, anti tag coimmunoprecipitation
                  (Frey et al., 2016, Kang et al., 2015, Fereres et al., 2014, Gambetta and Müller, 2014, Mohd-Sarip et al., 2012, Strübbe et al., 2011, Lagarou et al., 2008, Lupo et al., 2001, Shao et al., 1999, Kyba and Brock, 1998, Franke et al., 1992)
                  Pc - ph-p
                  tandem affinity purification, Identification by mass spectrometry, anti bait coimmunoprecipitation, anti tag coimmunoprecipitation, ion exchange chromatography, molecular sieving, western blot, autoradiography, pull down, peptide massfingerprinting, molecular weight estimation by staining, affinity chromatography technology
                  (Frey et al., 2016, Kang et al., 2015, Fereres et al., 2014, Gambetta and Müller, 2014, Mohd-Sarip et al., 2012, Strübbe et al., 2011, Lo et al., 2009, Lagarou et al., 2008, Peterson et al., 2004, Breiling et al., 2001, Francis et al., 2001, Lupo et al., 2001, Poux et al., 2001, Saurin et al., 2001, Shao et al., 1999, Kyba and Brock, 1998, Strutt and Paro, 1997, Franke et al., 1992)
                  Pc - pho
                  pull down, western blot, anti bait coimmunoprecipitation, peptide massfingerprinting
                  (Strübbe et al., 2011, Poux et al., 2001)
                  Pc - psq
                  anti bait coimmunoprecipitation, western blot, pull down
                  (Huang et al., 2002)
                  Pc - rdx
                  pull down, peptide massfingerprinting
                  (Strübbe et al., 2011)
                  Pc - stx
                  anti tag coimmunoprecipitation, western blot, pull down, anti tag western blot
                  (Du et al., 2016)
                  Pc - trx
                  anti bait coimmunoprecipitation, western blot
                  (Tie et al., 2016)
                  Pc - vtd
                  pull down, peptide massfingerprinting, anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation
                  (Strübbe et al., 2011)
                  Pc - wts
                  anti tag coimmunoprecipitation, anti tag western blot
                  (Parrish et al., 2007)
                  RNA-protein
                  Physical Interaction
                  Assay
                  References
                  Pc - brat
                  anti tag coimmunoprecipitation, primer specific pcr
                  (Loedige et al., 2015)
                  Pc - stau
                  anti bait coimmunoprecipitation, partial DNA sequence identification by hybridization, anti tag coimmunoprecipitation
                  (Laver et al., 2013)
                  Summary of Genetic Interactions
                  esyN Network Diagram
                  esyN Network Key:
                  Suppression
                  Enhancement
                  View in Interactions Browser
                  Please look at the allele data for full details of the genetic interactions
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  Pc
                  suppressible
                  Abd-B
                  (Zhang et al., 2017, Chen et al., 2005)
                  Pc
                  enhanceable
                  Abd-B|chm
                  (Grienenberger et al., 2002)
                  Pc
                  enhanceable
                  Adf1
                  (Orsi et al., 2014)
                  Pc
                  enhanceable
                  Asx
                  (Milne et al., 1999, Breen and Duncan, 1986, Jurgens, 1985)
                  Pc
                  suppressible
                  Bap170|polybromo
                  (Chalkley et al., 2008)
                  Pc
                  suppressible
                  Bap170|e(y)3
                  (Chalkley et al., 2008)
                  Antp|Pc
                  suppressible
                  brm
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  brm
                  (Tamkun et al., 1992, Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  btl
                  (Lindsley and Zimm, 1992, Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  bur
                  (van der Knaap et al., 2005)
                  Pc
                  suppressible
                  bur
                  (van der Knaap et al., 2005)
                  Pc
                  suppressible
                  Caf1-55
                  (Anderson et al., 2011)
                  Pc
                  enhanceable
                  Caf1-180
                  (Song et al., 2007)
                  Pc
                  enhanceable
                  chm
                  (Grienenberger et al., 2002)
                  Pc
                  enhanceable
                  corto
                  (Kodjabachian et al., 1998)
                  Pc
                  suppressible
                  Dll
                  (Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  dom
                  (Ruhf et al., 2001)
                  Pc
                  suppressible
                  dome
                  (Classen et al., 2009)
                  Pc
                  enhanceable
                  Dsp1
                  (Decoville et al., 2001)
                  Pc
                  suppressible
                  Dsp1
                  (Decoville et al., 2001)
                  Pc|ph-p
                  enhanceable
                  E(Pc)17B
                  (Fauvarque and Dura, 1993)
                  Pc
                  enhanceable
                  E(Pc)
                  (Sinclair et al., 1998, Stankunas et al., 1998, Lindsley and Zimm, 1992)
                  Pc
                  enhanceable
                  E(z)
                  (Campbell et al., 1995)
                  Pc
                  suppressible
                  ebi
                  (Strübbe et al., 2011)
                  Pc
                  enhanceable
                  eff
                  (Fauvarque et al., 2001, Fauvarque and Dura, 1993)
                  ph-p|Pc
                  enhanceable
                  eff
                  (Fauvarque and Dura, 1993)
                  Pc
                  suppressible
                  enok
                  (Umer et al., 2019)
                  Pc
                  enhanceable
                  esc
                  (Parrish et al., 2007)
                  tsr|Pc
                  enhanceable
                  flr
                  (Pham et al., 2008)
                  Pc
                  enhanceable
                  gpp
                  (Shanower et al., 2005)
                  Pc
                  suppressible
                  gpp
                  (Shanower et al., 2005)
                  Pc
                  enhanceable
                  Haspin
                  (Fresán et al., 2020)
                  Pc
                  enhanceable
                  Hcf
                  (Rodriguez-Jato et al., 2011)
                  Pc
                  enhanceable
                  HDAC1
                  (Chang et al., 2001)
                  Pc
                  suppressible
                  HDAC1
                  (Chang et al., 2001)
                  Pc
                  enhanceable
                  His3
                  (McKay et al., 2015)
                  Pc
                  enhanceable
                  hpo
                  (Parrish et al., 2007)
                  Pc
                  suppressible
                  Hsc70-4
                  (Mollaaghababa et al., 2001)
                  Pc
                  enhanceable
                  Hsc70-4
                  (Wang and Brock, 2003, Mollaaghababa et al., 2001)
                  Pc
                  suppressible
                  Hsp83
                  (Tariq et al., 2009)
                  Pc
                  enhanceable
                  jing
                  (Culi et al., 2006)
                  Pc
                  enhanceable
                  Kdm2
                  (Lagarou et al., 2008)
                  Pc
                  suppressible
                  kis
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  kto
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  l(3)80Fj
                  (Schulze et al., 2001)
                  Pc
                  suppressible
                  lawc
                  (Zorin et al., 1999)
                  Pc
                  suppressible
                  lolal
                  (Faucheux et al., 2003)
                  Pc
                  enhanceable
                  lolal
                  (Faucheux et al., 2003)
                  Pc
                  suppressible
                  M(2)56F
                  (Sinclair et al., 1984)
                  Pc
                  suppressible
                  M(3)80
                  (Sinclair et al., 1984)
                  Pc|ph-p
                  suppressible
                  Marcal1
                  (Morimoto et al., 2016)
                  Pc
                  suppressible
                  Marcal1
                  (Morimoto et al., 2016)
                  Pc
                  enhanceable
                  mbf1
                  (Nishioka et al., 2018)
                  Pc
                  enhanceable
                  mbf1|pcm
                  (Nishioka et al., 2018)
                  Pc
                  enhanceable
                  Mi-2
                  (Kehle et al., 1998)
                  Pc
                  suppressible
                  mor
                  (Brizuela and Kennison, 1997, Kennison and Tamkun, 1988)
                  Antp|Pc
                  suppressible
                  mor
                  (Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  mxc
                  (Docquier et al., 1996)
                  Pc
                  enhanceable
                  nej
                  (Tie et al., 2009)
                  Pc
                  suppressible
                  Nipped-B
                  (Schaaf et al., 2013, Hallson et al., 2008)
                  Pc
                  suppressible
                  Nsf2
                  (Kennison and Tamkun, 1988)
                  Antp|Pc
                  suppressible
                  Nsf2
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  osa
                  (Kennison and Tamkun, 1988)
                  Antp|Pc
                  suppressible
                  osa
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  Pcl
                  (Campbell et al., 1995)
                  Pc
                  enhanceable
                  Pcl
                  (Kehle et al., 1998, Kennison and Tamkun, 1988, Duncan, 1982)
                  Pc|Antp
                  enhanceable
                  Pcl
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  pcm
                  (Nishioka et al., 2018)
                  Pc
                  enhanceable
                  pds5
                  (Schaaf et al., 2013)
                  Pc|Haspin
                  enhanceable
                  pds5
                  (Fresán et al., 2020)
                  Pc
                  enhanceable
                  ph-d
                  (Dura et al., 1985)
                  Pc|su(Hw)|w
                  suppressible
                  ph-p
                  (Roseman et al., 2001)
                  Pc
                  enhanceable
                  ph-p
                  (Fauvarque and Dura, 1993, Dura et al., 1985)
                  Pc
                  enhanceable
                  psq
                  (Huang et al., 2002, Schwendemann and Lehmann, 2002)
                  Pc
                  suppressible
                  rdx
                  (Strübbe et al., 2011)
                  Pc
                  enhanceable
                  Rm62
                  (Lamiable et al., 2010)
                  Pc
                  enhanceable
                  salm
                  (Landecker et al., 1994)
                  Pc
                  enhanceable
                  Sam-S
                  (Larsson et al., 1996)
                  Pc
                  suppressible
                  scf
                  (Ogasawara et al., 2007)
                  Pc
                  enhanceable
                  Scm
                  (Bornemann et al., 1998, Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  Scr
                  (Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  Sirt1
                  (Furuyama et al., 2004)
                  Pc
                  suppressible
                  skd
                  (Lindsley and Zimm, 1992, Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  sls
                  (Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  SMC1
                  (Schaaf et al., 2013)
                  tsr|Pc
                  enhanceable
                  ssh
                  (Pham et al., 2008)
                  Pc|ph-p
                  suppressible
                  Su(Pc)35CD
                  (Fauvarque and Dura, 1993)
                  Pc
                  suppressible
                  Su(Pc)35CD
                  (Fauvarque and Dura, 1993)
                  Pc
                  suppressible
                  Su(Pc)37D
                  (Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  sxc
                  (Mariappa et al., 2018)
                  Pc
                  suppressible
                  tara
                  (Calgaro et al., 2002, Fauvarque et al., 2001)
                  Pc
                  enhanceable
                  tara
                  (Dutta and Li, 2017)
                  Pc
                  suppressible
                  tna
                  (Gutierrez et al., 2003)
                  Pc
                  suppressible
                  tou
                  (Fauvarque et al., 2001)
                  Pc
                  enhanceable
                  Trl
                  (Busturia et al., 2001, Strutt et al., 1997)
                  Antp|Pc
                  suppressible
                  trx
                  (Kennison and Tamkun, 1988)
                  Pc|ss
                  suppressible
                  trx
                  (Kuzin et al., 1997)
                  Pc
                  suppressible
                  trx
                  (Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  upSET
                  (Rincon-Arano et al., 2012)
                  Pc
                  suppressible
                  urd
                  (Kennison and Tamkun, 1988)
                  Pc
                  enhanceable
                  Usp7
                  (van der Knaap et al., 2005)
                  Pc
                  suppressible
                  Vha55
                  (Davies et al., 1996, Kennison and Tamkun, 1988)
                  Pc
                  suppressible
                  vtd
                  (Schaaf et al., 2013, Strübbe et al., 2011, Schulze et al., 2001, Sinclair et al., 2000, Kennison and Tamkun, 1988)
                  Antp|Pc
                  suppressible
                  vtd
                  (Kennison and Tamkun, 1988)
                  Pc|Haspin
                  enhanceable
                  vtd
                  (Fresán et al., 2020)
                  Pc
                  suppressible
                  wg
                  (Zhang et al., 2017)
                  Pc
                  enhanceable
                  wts
                  (Parrish et al., 2007)
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  Abd-B
                  enhanceable
                  Pc|chm
                  (Grienenberger et al., 2002)
                  Abd-B
                  enhanceable
                  Pc
                  (Mihaly et al., 1997)
                  Antp
                  enhanceable
                  Pc
                  (Decoville et al., 2001, Hannah-Alava, 1964)
                  ash1|trx
                  suppressible
                  Pc
                  (Gildea et al., 2000)
                  brm|trx
                  suppressible
                  Pc
                  (Gildea et al., 2000)
                  chm
                  enhanceable
                  Abd-B|Pc
                  (Grienenberger et al., 2002)
                  CycE
                  suppressible
                  Pc
                  (Brumby et al., 2002)
                  Dsp1
                  suppressible
                  Pc
                  (Decoville et al., 2001)
                  E2f1
                  suppressible
                  Pc
                  (Ji et al., 2012)
                  E2f1|Dp
                  enhanceable
                  Pc
                  (Ji et al., 2012)
                  E(z)
                  suppressible
                  Pc
                  (Bajusz et al., 2001)
                  eya
                  enhanceable
                  Pc
                  (Tian et al., 2019)
                  eya
                  suppressible
                  Pc
                  (Tian et al., 2019)
                  gcm
                  enhanceable
                  Pc
                  (Popkova et al., 2012)
                  Hcf
                  suppressible
                  Pc
                  (Rodriguez-Jato et al., 2011)
                  His3
                  enhanceable
                  Pc
                  (McKay et al., 2015)
                  Marcal1
                  enhanceable
                  Pc
                  (Morimoto et al., 2016)
                  mxc
                  enhanceable
                  Pc
                  (Saget et al., 1998, Docquier et al., 1996)
                  ph-p
                  enhanceable
                  Pc
                  (Fauvarque and Dura, 1993)
                  rho
                  enhanceable
                  Pc
                  (Marenda et al., 2004)
                  RYBP
                  enhanceable
                  Pc
                  (Bejarano et al., 2005)
                  RYBP|Ubx
                  suppressible
                  Pc
                  (Bejarano et al., 2005)
                  sd
                  suppressible
                  Pc
                  (Bantignies et al., 2003)
                  sens
                  suppressible
                  Pc
                  (Anderson et al., 2011)
                  stx
                  suppressible
                  Pc
                  (Du et al., 2016)
                  su(Hw)|w
                  enhanceable
                  Pc
                  (Roseman et al., 2001)
                  su(Hw)|y
                  enhanceable
                  Pc
                  (Roseman et al., 2001)
                  tara
                  enhanceable
                  Pc
                  (Dutta and Li, 2017)
                  Ubx
                  enhanceable
                  Pc
                  (Dutta and Li, 2017)
                  Utx
                  suppressible
                  Pc
                  (Herz et al., 2010)
                  vg
                  enhanceable
                  Pc
                  (Ahmad and Spens, 2019)
                  w
                  suppressible
                  Pc
                  (Classen et al., 2009, Martinez et al., 2006, Kelley and Kuroda, 2003, Poux et al., 2002, Netter et al., 1998)
                  wg
                  enhanceable
                  Pc
                  (Lee et al., 2005)
                  External Data
                  Subunit Structure (UniProtKB)
                  Component of PRC1 complex, which contains many PcG proteins like Pc, ph, Scm, Psc, Sce and also chromatin-remodeling proteins such as histone deacetylases. This complex is distinct from the Esc/E(z) complex, at least composed of esc, E(z), Su(z)12, HDAC1/Rpd3 and Caf1-55. The 2 complexes however cooperate and interact together during the first 3 hours of development to establish PcG silencing (PubMed:11493925, PubMed:11583617). Interacts with stx; the interaction targets Pc for ubiquitin-independent proteasomal degradation (PubMed:27326929). Interacts with Nup93-1 (PubMed:31784359).
                  (UniProt, P26017 )
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  DroID - A comprehensive database of gene and protein interactions.
                  MIST (genetic) - An integrated Molecular Interaction Database
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Pathways
                  Signaling Pathways (FlyBase)
                  Metabolic Pathways
                  FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                  External Data
                  Genomic Location and Detailed Mapping Data
                  Chromosome (arm)
                  3L
                  Recombination map
                  3-47
                  Cytogenetic map
                  78C6-78C7
                  Sequence location
                  FlyBase Computed Cytological Location
                  Cytogenetic map
                  Evidence for location
                  78C6-78C7
                  Limits computationally determined from genome sequence between P{lacW}l(3)j1B10j1B10 and P{lacW}l(3)j2C4j2C4
                  Experimentally Determined Cytological Location
                  Cytogenetic map
                  Notes
                  References
                  78D7-78D8
                  (determined by in situ hybridisation)
                  (Harding et al., 1995)
                  78E-78E
                  (Kennison and Tamkun, 1988)
                  Location from complementation analysis with deficiency and duplication chromosomes (details unspecified).
                  (Kennison and Tamkun, 1988)
                  Experimentally Determined Recombination Data
                  Location

                  3-47

                  (FlyBase, 2016)

                  3-43.8

                  (Comeron et al., 2012)

                  3-47.1

                  (Kennison and Tamkun, 1988, Puro and Nygren, 1975)

                  3-48

                  (Tearle and Nusslein-Volhard, 1987)

                  3-47.7

                  (Duncan and Kaufman, 1975)
                  Left of (cM)
                  (Puro and Nygren, 1975)
                  Right of (cM)
                  (Puro and Nygren, 1975)
                  Notes
                  Stocks and Reagents
                  Stocks (36)
                  Genomic Clones (13)
                   

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

                  cDNA Clones (163)
                   

                  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)
                  BDGP DGC clones
                  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
                   

                  polyclonal

                  (Nishioka et al., 2018, Lv et al., 2016, Tie et al., 2016, Mason-Suares et al., 2013, Zink and Paro, 1989)
                  Commercially Available Antibodies
                   
                  Other Information
                  Relationship to Other Genes
                  Source for database identify of
                  Source for database merge of
                  Additional comments
                  Other Comments

                  Haploinsufficient locus (not associated with strong haplolethality or haplosterility).

                  (Cook et al., 2012)

                  DNA-protein interactions: genome-wide binding profile assayed for Pc protein in Kc167 cells; see Chromatin_types_NKI collection report. Individual protein-binding experiments listed under "Samples" at GEO_GSE22069 (http://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE22069).

                  (Filion et al., 2010)

                  Pc is required for normal neuroblast survival and proliferation in postembryonic central nervous system development.

                  (Bello et al., 2007)

                  Pc is required for for the maintenance of dendritic fields in class IV dendrite arborisation (da) neurons.

                  (Parrish et al., 2007)

                  dsRNA made from templates generated with primers directed against Pc used to treat S2 cells results in a decreased proportion of cells in G1 and S phase, with an accompanying increase in G2/M cells. This modification of the cell cycle profile occurs in the absence of any apparent growth defect.

                  (Martinez et al., 2006)

                  Mutant allele fails to complement a QTL affecting male mating behaviour.

                  (Moehring and Mackay, 2004)

                  Pc-G proteins can silence gene expression at a large number of chromosomal locations, affecting both enhancer-activated and basal w transcription. Repression is observed even with separation distances of up to 3.0kb between target promoters and binding sites for tethered Pc-G proteins.

                  (Roseman et al., 2001)

                  Mutant analysis demonstrates functional interactions between Pc, sxc and Asx in vivo.

                  (Milne et al., 1999)

                  Pc protein is associated with genes silenced as part of the cosuppression phenomenon.

                  (Pal-Bhadra et al., 1999)

                  Pc, Scm, Psc, ph-p and ph-d contribute to the PRC1 (Polycomb repressive complex 1). PRC1 directly antagonizes ATP-dependent remodeling of nucleosomal arrays in a purified system and may directly modulate (and be modified by) SWI/SNF (brm/mor) activity.

                  (Shao et al., 1999)

                  Evidence of physical interaction between esc and E(z) in vitro and in vivo and coimmunoprecipitation in vivo suggests the proteins are direct partners in Pc-G mediated repression and this relationship has been evolutionarily conserved.

                  (Jones et al., 1998)

                  ph-p, Psc and Pc proteins coimmunoprecipitate from nuclear extracts. Interacting domains are identified and delimited using the two-hybrid system, the interactions are shown to be direct by using an in vitro binding assay.

                  (Kyba and Brock, 1998)

                  The Pc and trx gene products bind to PRE target sequences by cellular blastoderm, when the bithorax complex transcription begins. At the same stage trx but not Pc gene product is strongly associated with core promoters. At germ band extension, the time of derepression in Pc mutants, Pc gene product is found outside core PREs.

                  (Orlando et al., 1998)

                  Abd-B MCP725 element is a silencer that functions throughout proliferation of the imaginal discs. MCP725-mediated silencing requires the Pc and Pcl proteins (it is likely that other members of the PcG also interact with the MCP725 silencer).

                  (Busturia et al., 1997)

                  Mutant phenotype suppressed by alleles of kis and (weakly) EcR.

                  (Gellon et al., 1997)

                  Silencing activity of the iab-7PRE in the bithorax complex is dependent upon proteins from the Polycomb group.

                  (Hagstrom et al., 1997)

                  The chromodomain of Pc is necessary for protein-protein interactions within a Pc-polyhomeotic complex. In addition, Psc protein coimmunoprecipitates Pc and polyhomeotic indicating they are members of a common multimeric protein complex. These three proteins are associated with identical regulatory elements of en in tissue culture cells and differentially distributed on regulatory sequences of inv.

                  (Strutt and Paro, 1997)

                  The distribution of Pc protein on the bithorax complex of Drosophila tissue culture cells has been mapped. The protein is not distributed homogeneously on the regulatory regions of the repressed Ubx and abd-A genes, but is highly enriched at discrete sequence elements, many of which coincide with previously mapped Pc response elements (PREs). Trl protein is also bound at those PREs which contain Trl consensus-binding sites.

                  (Strutt et al., 1997)

                  Using reporter gene constructs the Pc group response elements (PRE) behaves as an orientation-dependent silencer capable of inducing mosaic gene expression on neighbouring genes.

                  (Breiling et al., 1995)

                  In an effort to subdivide the Pc-group genes functionally, the phenotypes of adult flies heterozygous for every pairwise combination of Pc-group mutation were examined. Asx, Pc, Pcl, Psc, Scm and Sce have similar functions in some imaginal tissues. Genetic interactions have been demonstrated between esc, Asx, E(Pc), Pcl, E(z) and sxc. Most duplications of Pc-group genes neither exhibit anterior transformations nor suppress the extra sex comb phenotype of Pc-group mutations, suggesting that not all Pc-group genes behave as predicted by the mass action model.

                  (Campbell et al., 1995)

                  Proper regulation of both trx and Pc can be established by transient assays with a haploid cell line expressing Ubx promoters fused to a Ecol\lacZ reporter gene. This system has been used to map the regulatory sequences in the Ubx promoter to a 440bp region. The Ubx proximal promoter is essential for trx-dependent activation.

                  (Chang et al., 1995)

                  Pc associates with multiple sites in the bithorax complex and these sites all contain maintenance elements, as defined by a Ecol\lacZ expression assay.

                  (Chiang et al., 1995)

                  The negative autoregulation of ph-p starts at the blastoderm stage and is partly mediated by a transvection effect. As the number of functional copies of ph-p increases a concomitant reduction of the transcription in each copy is observed. This regulation is ensured positively by the trx group and negatively by the Pc group gene products.

                  (Fauvarque et al., 1995)

                  Sections of the Scr regulatory region may be important for regulation of Scr by Polycomb- and trithorax-group genes.

                  (Gindhart and Kaufman, 1995)

                  Mutations of Pc interact with Dfd to reduce the viability of the Dfd3/Dfd13 combination.

                  (Harding et al., 1995)

                  A Pc-response element has been identified in an intron of abd-A.

                  (Lunde et al., 1995)

                  Phenotypic rescue of Pc- larvae cuticle phenotype and Pc- clones of adult cells demonstrates the Mmus\Cbx2 protein can emulate Pc both early and late in development.

                  (Muller et al., 1995)

                  Pc acts as a transcriptional silencer in embryos if tethered to reporter genes by the DNA binding domain of Scer\GAL4 (as a Scer\GAL4-Pc fusion gene). Silencing by Scer\GAL4-Pc requires the C-terminal portion, but not the chromodomain.

                  (Muller, 1995)

                  A method based on in-vivo formaldehyde crosslinking and chromatin immunoprecipitation has allowed the determination of the in vivo distribution of Pc, brm and Abd-B products at their target sites.

                  (Orlando et al., 1995)

                  A chimeric Su(var)205-Pc protein (the chromo domain of Pc in the context of Su(var)205) causes mislocalisation of Su(var)205 to Pc binding sites and expression in transgenic flies promotes heterochromatin mediated gene silencing. Results support the view that the chromo domain homology reflects a common mechanistic basis for homeotic and heterochromatic silencing.

                  (Platero et al., 1995)

                  The ventral to dorsal effect of Pc mutants is mediated through Dlw.

                  (Tiong et al., 1995)

                  Pc group response elements found in the Fab and Mcp regions of Abd-B induce mosaic expression of w+mC, this is dependent on Pc.

                  (Zink and Paro, 1995)

                  Members of the Pc group function as potent repressors in mammalian cells.

                  (Bunker and Kingston, 1994)

                  Duplication of ph-d suppress homeotic transformations of Pc and Pcl, this data supports the conclusion that ph-d is at equilibrium with a multimeric complex containing Pc group genes.

                  (Cheng et al., 1994)

                  Pc product is found on larval salivary chromosomes at about 100 specific loci, in the same positions as the products of Pcl, ph-d and ph-p genes.

                  (Lonie et al., 1994)

                  Mutation analysis and en expression patterns demonstrate a role for Pc gene in regulation of some segmentation genes.

                  (McKeon et al., 1994)

                  P elements bearing ph-p regulatory sequences preferentially insert into regions containing known Pc and ph-p protein binding sites, perhaps because shared regulatory proteins bring transposons and insertion sites into close proximity.

                  (Fauvarque and Dura, 1993)

                  Pc binds polytene chromosomes at 45 locations where other Pc-group proteins, encoded by the Psc and polyhomeotic genes, are present.

                  (Martin and Adler, 1993)

                  Distribution of the Pc protein in the bithorax complex was determined, and Pc protein quantitatively covers large regulatory regions of repressed genes.

                  (Orlando and Paro, 1993)

                  There is considerable overlap of chromosomal binding sites for Psc, Su(z)2, z, Pc and the polyhomeotic proteins.

                  (Rastelli et al., 1993)

                  The bithorax complex genes are regulated by the Pc group of genes, acting via 'Pc group response elements' (PREs), that can work even when removed from the normal bithorax complex context. The Pc group products apparently provide stable memory or imprinting of boundaries which are specified by gap and pair-rule regulators.

                  (Simon et al., 1993)

                  Extensive overlap exists between the binding sites of ph-p and Pc on polytene chromosomes.

                  (DeCamillis et al., 1992)

                  Pc and polyhomeotic proteins are constituents of a soluble multimeric nuclear protein complex.

                  (Franke et al., 1992)

                  When homozygous or hemizygous, Pc mutants are late embryonic lethals. Embryos with at least one dose of the BXC show incomplete head development and caudal transformations, the thoracic and first seven abdominal segments being partially transformed into the eighth abdominal segment (FBrf0032262; FBrf0063402; FBrf0039217; FBrf0039211). This homeotic effect in homozygotes is enhanced by increasing the dosage of the BXC. Transformations involve brain and ventral nerve cord as well as epidermis (FBrf0037316). Pc+ alleles in the mother weaken the homeotic effect (FBrf0063402; FBrf0038997). Pc2/Pc2 or Pc3/Pc3 clones induced in leg and eye-antennal tissue during larval development also show similar posteriorly-directed transformations (FBrf0037148; FBrf0063405). Pc/+ flies carrying at least one dose of the BXC show caudal transformations, i.e. partial conversion of wings into halteres and of anterior abdominal segments into more posterior ones. Some Pc heterozygotes show phenotypes characteristic of ANTC mutants, i.e. partial conversion of antennae into legs and of second and third legs into first legs (with sex combs in males) (FBrf0012037; FBrf0038059). The frequency of wing transformations varies directly with the BXC dosage, but does not seem to be changed by variation in ANTC dosage (FBrf0063405; FBrf0037909). The number of abdominal transformations, however, varies inversely with the doses of the BXC while it increases as the doses of the ANTC are increased (FBrf0038059; FBrf0063405). Other changes observed in Pc/+ flies include a transformation of ventral to dorsal wing (FBrf0051513; FBrf0039451), elevated, divergent, or crinkled wings, terminal gaps in the L4 wing vein, bent humeral or notopleural bristles, and defective sternopleural bristles, all abnormalities being less extreme in males than in females (sometimes absent in males). When doubly heterozygous with AntpYu and AntpB, Pc enhances Antp. The expression of all Pc mutant heterozygotes (including deficiencies for the locus) is enhanced by the second chromosome dominant, E(Pc) (FBrf0039451; FBrf0040741). Pc3/Pc3/Dp(1;3;4)7 flies (carrying a Pc+ duplication) show stronger leg and wing transformations than E(Pc)/+; Pc3/+ flies (FBrf0063405; FBrf0039451).

                  (Lindsley and Zimm, 1992)

                  Pc+ may be considered a negative regulator of the bithorax complex (BXC) and the Antennapedia complex (ANTC), with a decreasing gradient of activity from anterior to posterior.

                  (Lindsley and Zimm, 1992)

                  The chromo domain is a homologous protein motif between Pc and Su(var)205. Transgenic lines and transient assays in culture cells have been used to determine the functional role of the Pc chromo domain, it is important for the function of Pc and is absolutely required for binding of Pc protein to chromatin. The chromo domain could be involved in a packaging mechanism essential for compacting chromosomal proteins within heterochromatin or heterochromatin-like complexes.

                  (Messmer et al., 1992)

                  Embryos mutant for two or more Pc group genes (Pc, Scm, Pcl, Psc, Asx, E(Pc), E(z), ph-d, pho and esc) show strong ectopic en expression, but only weak derepression occurs if embryo is mutant at only one of the Pc group genes. This effect is independent of the function of en itself, and wg.

                  (Moazed and O'Farrell, 1992)

                  Developmental profile of Pc gene products demonstrates that Pc is down-regulated by one of its own targets, Ubx.

                  (Paro and Zink, 1992)

                  Mutations of genes in the polycomb group (esc, E(z), Pc, ph-p, ph-d, Scm, Pcl, Sce, Asx, Psc, pho and Antp) cause abnormal segmental development due to the ectopic expression of abd-A and Abd-B. Embryos lacking both maternal and zygotic Pc product were generated to determine abd-A and Abd-B expression patterns.

                  (Simon et al., 1992)

                  Pc dominant loss-of-function mutants cause widespread alterations in cell fates by derepressing genes in the BXC and ANTC.

                  (Tamkun et al., 1992)

                  The Pc group genes are negative regulators of homeotic genes.

                  (McKeon and Brock, 1991)

                  Mutations in maternal class gene Pc do not interact with RpII140wimp.

                  (Parkhurst and Ish-Horowicz, 1991)

                  Pc has been cloned and sequenced.

                  (Paro and Hogness, 1991)

                  Homeotic gene activity programs primordia as either discs or histoblast nests by the early extended germ band stage. In Pc- embryos homeotic gene expression is no longer restricted to the normal expression domains.

                  (Simcox et al., 1991)

                  The Pc gene product regulates Antp transcription directly at the chromosomal level. Regulation of Antp P2 differs from Antp P1 in that in that it can be repressed by transacting factors other than Pc.

                  (Zink et al., 1991)

                  The role of Pc in maintaining stable patterns of homeotic gene transcription has been studied.

                  (Gould et al., 1990)

                  Interactions with a mutation of the Pc gene were used to confirm that ash1 and ash2 are members of a functionally related class of genes whose alleles have similar transformation properties to mutations of trx. The dominant extra sex combs phenotype of adult males heterozygous for Pc mutations is sensitive to the gene dosage of trx, and is suppressed by heterozygosity for a trx deletion and enhanced by heterozygosity for a trx duplication.

                  (Shearn, 1989)

                  Cell clones deficient for Pc and the BXC genes have abnormal wings and legs, Scr and en are derepressed in the absence of Pc and BXC function. By using the Pc- mutation and various BXC mutant combinations imaginal cell clones possessing different combinations of active homeotic genes have been generated. In the absence of BXC genes Pc- clones develop prothoracic patterns: Scr activity overrules Antp. Adding contributions of Ubx, abd-A and Abd-B results in thoracic or abdominal patterns.

                  (Busturia and Morata, 1988)

                  Mutations in 18 complementation groups identify modifiers of Pc and/or Antp phenotypes. All 18 complementation groups are required for viability. Alleles of Pc, Pcl, Scm, Dll, brm, kto, Scr and trx show clear dominant enhancement or suppression of AntpScx, whereas alleles of vtd, Vha55, Su(Pc)37D, urd, mor, skd and osa do not.

                  (Kennison and Tamkun, 1988)

                  Mutants of Pc exhibit a reduction in sex comb teeth on the second and third legs.

                  (Kennison and Russell, 1987)

                  Genetic analysis of the Pc locus suggests it is a complex gene. Insufficiency of Pc products can be corrected by insufficiency of hb products or be exaggerated by the excess of the same products. Pc+ products regulate the expression Of the Ubx protein coding region.

                  (Capdevila et al., 1986)

                  The effect of various mutations in the bithorax and antennapedia complexes on the Pc mutant phenotype has been studied.

                  (Sato and Denell, 1985)

                  Pc mutants display homeotic transformation of head and thoracic segments towards A8.

                  (Jurgens et al., 1984)

                  Mutant males display sex combs in the second and third legs.

                  (Duncan and Kaufman, 1975)
                  Origin and Etymology
                  Discoverer

                  P.H. Lewis.

                  (Lindsley and Zimm, 1992)
                  Etymology
                  Identification
                  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/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
                  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
                  Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                  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
                  FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.
                  GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
                  iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                  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)
                  modMine - A data warehouse for the modENCODE project
                  SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  DroID - A comprehensive database of gene and protein interactions.
                  DRSC - Results frm RNAi screens
                  FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                  FlyMine - An integrated database for Drosophila genomics
                  Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                  MIST (genetic) - An integrated Molecular Interaction Database
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                  Synonyms and Secondary IDs (12)
                  Reported As
                  Symbol Synonym
                  CG32443
                  (Meadows, 2018.4.4, Christis and Munro, 2012, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Ni et al., 2008, Klebes et al., 2005)
                  CG7618
                  DmPc
                  (Bertram and Pereira-Smith, 2001)
                  PC
                  (Srinivasan and Mishra, 2020, Decourcelle et al., 2019, Ogiyama et al., 2018, L Black et al., 2016, Pindyurin et al., 2016, Tie et al., 2016, Cesarini et al., 2015, Bowman et al., 2014, Grossniklaus and Paro, 2014, Ho et al., 2014, Lavington et al., 2014, Schuettengruber et al., 2014, Lo Sardo et al., 2013, Schuettengruber and Cavalli, 2013, Steffen et al., 2013, Follmer et al., 2012, Lanzuolo and Orlando, 2012, Mohd-Sarip et al., 2012, Riddle et al., 2012, Tie et al., 2012, Demakov et al., 2011, Dufourt et al., 2011, Grau et al., 2011, Islam et al., 2011, Kharchenko et al., 2011, Moon et al., 2011, Tolhuis et al., 2011, Vatolina et al., 2011, Vatolina et al., 2011, Filion et al., 2010, Johnson et al., 2010, Mills, 2010, Schwartz et al., 2010, Suganuma and Workman, 2010, van Steensel et al., 2010, Wang et al., 2010, Schuettengruber et al., 2009, Fanti et al., 2008, Pindyurin et al., 2008, Srinivasan et al., 2008, Lanzuolo et al., 2007, Ringrose and Paro, 2007, Tie et al., 2007, Cavalli, 2006, Comet et al., 2006, Grimaud et al., 2006, Hill, 2006, Kahn et al., 2006, Mohd-Sarip et al., 2006, Negre et al., 2006, Salvaing et al., 2006, Schwartz et al., 2006, Andreyeva et al., 2005, Chopra and Mishra, 2005, Schwartz et al., 2005, Ringrose et al., 2004, Czermin et al., 2002, Mohd-Sarip et al., 2002, Hodgson et al., 2001)
                  Pc
                  (Chaouch and Lasko, 2021, Khan et al., 2021, Lam Wong and Verheyen, 2021, Ponrathnam et al., 2021, Ulianov et al., 2021, Vidaurre and Chen, 2021, Barbour et al., 2020, Chetverina et al., 2020, DeLuca et al., 2020, Fresán et al., 2020, Geigges et al., 2020, Gozalo et al., 2020, Harris et al., 2020, Kang et al., 2020, Liu and Hyeon, 2020, Liu et al., 2020, Loubiere et al., 2020, Meyer-Nava et al., 2020, Morgan and Shilatifard, 2020, Seif et al., 2020, Ahmad and Spens, 2019, Bonnet et al., 2019, Chetverina et al., 2019, Cheutin and Cavalli, 2019, Delandre and Marshall, 2019, Gervais et al., 2019, Gutierrez-Perez et al., 2019, Hall et al., 2019, Leatham-Jensen et al., 2019, Luhur et al., 2019, Parey and Crombach, 2019, Pherson et al., 2019, Umer et al., 2019, Dasari et al., 2018, Davis and Rebay, 2018, La Fortezza et al., 2018, Lv et al., 2018, Nishioka et al., 2018, Rosales-Vega et al., 2018, Rybina et al., 2018, Sadasivam and Huang, 2018, Yao et al., 2018, Akishina et al., 2017, Dutta and Li, 2017, Eagen et al., 2017, Maini et al., 2017, Marshall and Brand, 2017, Monribot-Villanueva et al., 2017, Schwartz and Cavalli, 2017, Sharma et al., 2017, Transgenic RNAi Project members, 2017-, Aggarwal et al., 2016, Cruz-Becerra et al., 2016, Du et al., 2016, Frey et al., 2016, Harris et al., 2016, Kahn et al., 2016, Lv et al., 2016, Morata and Herrera, 2016, Morimoto et al., 2016, Peng et al., 2016, Piunti and Shilatifard, 2016, Ray et al., 2016, Shih et al., 2016, Shlyueva et al., 2016, Wani et al., 2016, Bivik et al., 2015, Dietz et al., 2015, Dupont et al., 2015, Erokhin et al., 2015, Finley et al., 2015, Grifoni et al., 2015, Hsu et al., 2015, Kang et al., 2015, Kim et al., 2015, Lee et al., 2015, Li et al., 2015, McKay et al., 2015, Ríos-Barrera et al., 2015, Saberi et al., 2015, Yung et al., 2015, Zang et al., 2015, Comoglio and Paro, 2014, Fereres et al., 2014, Gambetta and Müller, 2014, Gonzalez et al., 2014, Herrera and Morata, 2014, Liu et al., 2014, Maksimenko et al., 2014, McElroy et al., 2014, Ost et al., 2014, Pascual-Garcia et al., 2014, Stern et al., 2014, Tie et al., 2014, Aloia et al., 2013, Debruhl et al., 2013, Di Croce and Helin, 2013, Iovino et al., 2013, Lim et al., 2013, Mason-Suares et al., 2013, Mulero et al., 2013, Schaaf et al., 2013, Vaque et al., 2013, Vasanthi et al., 2013, Antao et al., 2012, Cheutin and Cavalli, 2012, Cook et al., 2012, Cuddapah et al., 2012, Gutiérrez et al., 2012, Hainaut et al., 2012, Herz et al., 2012, Ji et al., 2012, Langlais et al., 2012, Lim et al., 2012, Meier et al., 2012, Park et al., 2012, Petruk et al., 2012, Popkova et al., 2012, Pushpavalli et al., 2012, Rincon-Arano et al., 2012, Sakoparnig et al., 2012, Stern et al., 2012, Wen et al., 2012, Anderson et al., 2011, Bantignies et al., 2011, Cavalli, 2011, Chen et al., 2011, Choo et al., 2011, Davis et al., 2011, Enderle et al., 2011, Grau et al., 2011, Katsuyama and Paro, 2011, Mikhaylova and Nurminsky, 2011, Okulski et al., 2011, Pruteanu-Malinici et al., 2011, Rodriguez-Jato et al., 2011, Strübbe et al., 2011, Sultana et al., 2011, Ahn et al., 2010, Baig et al., 2010, Basu and Atchison, 2010, Bhatia et al., 2010, Brown and Kassis, 2010, Campos et al., 2010, Gan et al., 2010, Grau et al., 2010, Guertin and Lis, 2010, Herz et al., 2010, Huen and Russell, 2010, Lamiable et al., 2010, Lee et al., 2010, Li et al., 2010, modENCODE Consortium et al., 2010, Saj et al., 2010, Smulders-Srinivasan et al., 2010, Wang et al., 2010, Wilkinson et al., 2010, Classen et al., 2009, Fang et al., 2009, Francis et al., 2009, Gambetta et al., 2009, Gambetta et al., 2009, González and Busturia, 2009, Khan et al., 2009, Lo et al., 2009, Schaaf et al., 2009, Shevelyov et al., 2009, Sinclair et al., 2009, Sinclair et al., 2009, Sing et al., 2009, Tariq et al., 2009, Tie et al., 2009, Zhai et al., 2009, Chalkley et al., 2008, Chen et al., 2008, de Wit et al., 2008, Diop et al., 2008, Dixit et al., 2008, Doheny et al., 2008, Estella et al., 2008, Fedorova et al., 2008, Fujioka et al., 2008, Garaulet et al., 2008, González et al., 2008, Hallson et al., 2008, Hauenschild et al., 2008, Jung et al., 2008, Kwong et al., 2008, Lagarou et al., 2008, Lin et al., 2008, McDermott and Kliman, 2008, Pauli et al., 2008, Pérez-Lluch et al., 2008, Petruk et al., 2008, Pham et al., 2008, Zhang et al., 2008, Bai et al., 2007, Bello et al., 2007, de Ayala Alonso et al., 2007, de Wit et al., 2007, Engström et al., 2007, Goodliffe et al., 2007, Goodliffe et al., 2007, Mishra et al., 2007, Ogasawara et al., 2007, Parrish et al., 2007, Pindyurin et al., 2007, Schuettengruber et al., 2007, Schwartz and Pirrotta, 2007, Song et al., 2007, Chopra and Mishra, 2006, Culi et al., 2006, de Navas et al., 2006, Kang et al., 2006, Kavi et al., 2006, Klymenko et al., 2006, Le Bras and Van Doren, 2006, Liu et al., 2006, Maeda and Karch, 2006, Muller and Kassis, 2006, Papp and Muller, 2006, Qi et al., 2006, Ringrose, 2006, Tolhuis et al., 2006, Wang et al., 2006, Wang et al., 2006, Bejarano et al., 2005, Brown et al., 2005, Canudas et al., 2005, Chanas and Maschat, 2005, Chen et al., 2005, Chen et al., 2005, King et al., 2005, Ali and Bender, 2004, Breiling et al., 2004, Dellino et al., 2004, Gutierrez et al., 2004, Fischle et al., 2003, Kwon et al., 2003, Balasov, 2002, Huang et al., 2002, Joanis and Lloyd, 2002, King et al., 2002, Fitzgerald and Bender, 2001, Hirose et al., 2001, Choi et al., 2000, Franke et al., 1995)
                  Pc-G
                  (Akbari et al., 2006, Chopra and Mishra, 2003)
                  PcG
                  (Sneppen and Ringrose, 2019, Dasari et al., 2018, Kingston and Tamkun, 2014, Follmer et al., 2012, Worley et al., 2012, Jung et al., 2007)
                  dPC
                  (Simms et al., 2003)
                  pc
                  (Loedige et al., 2015)
                  Name Synonyms
                  POLYCOMB
                  (Bowman et al., 2014)
                  Polycomb
                  (Bender, 2020, El-Sharnouby et al., 2017, Schwartz and Cavalli, 2017, Piunti and Shilatifard, 2016, Tie et al., 2016, Wani et al., 2016, Wieschaus and Nüsslein-Volhard, 2016, Bajusz et al., 2015, Kim et al., 2015, Comoglio and Paro, 2014, Gambetta and Müller, 2014, Liu et al., 2014, McElroy et al., 2014, Orsi et al., 2014, Stern et al., 2014, Laver et al., 2013, Lo Sardo et al., 2013, Mulero et al., 2013, Schuettengruber and Cavalli, 2013, Follmer et al., 2012, Herz et al., 2012, Leser et al., 2012, Lim et al., 2012, Mohd-Sarip et al., 2012, Nowak et al., 2012, Petruk et al., 2012, Riddle et al., 2012, Stern et al., 2012, Wen et al., 2012, Anderson et al., 2011, Bantignies et al., 2011, Chen et al., 2011, Crozatier and Vincent, 2011, Davis et al., 2011, Katsuyama and Paro, 2011, Okulski et al., 2011, Strübbe et al., 2011, Tolhuis et al., 2011, Ahn et al., 2010, Baig et al., 2010, Basu and Atchison, 2010, Bhatia et al., 2010, Grau et al., 2010, Herz et al., 2010, Huen and Russell, 2010, Johnson et al., 2010, Kim et al., 2010, Schwartz et al., 2010, Suganuma and Workman, 2010, van Steensel et al., 2010, Wang et al., 2010, Wang et al., 2010, Classen et al., 2009, Fang et al., 2009, Lau et al., 2009, Schaaf et al., 2009, Shevelyov et al., 2009, Sinclair et al., 2009, Sing et al., 2009, Tariq et al., 2009, Zhai et al., 2009, Zhao et al., 2009, Chalkley et al., 2008, Chen et al., 2008, Chen et al., 2008, Chopra and Levine, 2008, Diop et al., 2008, Estella et al., 2008, Fanti et al., 2008, Garaulet et al., 2008, Jung et al., 2008, Kwong et al., 2008, Lu et al., 2008, Pauli et al., 2008, Pindyurin et al., 2008, Zhang et al., 2008, Bello et al., 2007, Bird, 2007, de Wit et al., 2007, Engström et al., 2007, Lanzuolo et al., 2007, Mishra et al., 2007, Pindyurin et al., 2007, Redhouse and White, 2007, Riaz et al., 2007, Ringrose and Paro, 2007, Schuettengruber et al., 2007, Song et al., 2007, Cavalli, 2006, Chopra and Mishra, 2006, Eissenberg, 2006, Kavi et al., 2006, Maeda and Karch, 2006, Maurange et al., 2006, Muller and Kassis, 2006, Negre et al., 2006, Nystul and Spradling, 2006, O'Dor et al., 2006, Ringrose, 2006, Schwartz et al., 2006, Bejarano et al., 2005, Brown et al., 2005, Chen et al., 2005, King et al., 2005, Schwartz et al., 2005, Ali and Bender, 2004, Fischle et al., 2003, Kwon et al., 2003, Levine and Tjian, 2003, Min et al., 2003, Wang and Brock, 2003, Balasov, 2002, Czermin et al., 2002, Huang et al., 2002, Mohd-Sarip et al., 2002, Saller and Bienz, 2001, Choi et al., 2000)
                  polycomb
                  (Wang et al., 2018, Aggarwal et al., 2016, Schuettengruber et al., 2014, Stein et al., 2013)
                  Secondary FlyBase IDs
                  • FBgn0015223
                  • FBgn0052443
                  Datasets (3)
                  Study focus (3)
                  Experimental Role
                  Project
                  Project Type
                  Title
                  • transgene_used
                  FBGSE22069
                  Protein profiling reveals five principal chromatin types in Drosophila cells.
                  • bait_protein
                  modENCODE_chromatin_proteins_cells
                  Genome-wide localization of chromosomal proteins in cell lines by ChIP-chip and ChIP-Seq.
                  • bait_protein
                  modENCODE_chromatin_proteins_flies
                  Genome-wide localization of chromosomal proteins in fly tissues by ChIP-chip and ChIP-Seq.
                  References (940)