FB2025_05 , released December 11, 2025
Gene: Dmel\rhi
Open Close
General Information
Symbol
Dmel\rhi
Species
D. melanogaster
Name
rhino
Annotation Symbol
CG10683
Feature Type
protein_coding_gene
FlyBase ID
FBgn0004400
Gene Model Status
Current
Stock Availability
10 publicly available
Gene Summary
rhino (rhi) encodes a germline-restricted heterochromatin protein 1 required for fertility. It silences germline transposable elements by promoting transcription of the small RNAs encoded in Piwi-RNA clusters. rhi (as well as its interacting partners del and cuff) shows ovary-biased expression, but it is also expressed and functional in the testis. [Date last reviewed: 2021-09-16] (FlyBase Gene Snapshot)
All Summaries
Also Known As

HP1D, HP1

Key Links
Genomic Location
Cytogenetic map
54D2-54D2
Sequence location
Recombination map
2-83
RefSeq locus
NT_033778 REGION:17628491..17633516
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (12 terms)
Molecular Function (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
enables chromatin binding
inferred from direct assay
(Baumgartner et al., 2022, Klattenhoff et al., 2009)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
enables chromatin binding
inferred from biological aspect of ancestor with PANTHER:PTN000517042
(GO Reference Genome Project, 2011-)
inferred from sequence or structural similarity with FLYBASE:Su(var)205; FB:FBgn0003607
(Levine et al., 2012)
Biological Process (7 terms)
Terms Based on Experimental Evidence (5 terms)
CV Term
Evidence
References
involved_in chorion-containing eggshell pattern formation
inferred from mutant phenotype
(Volpe et al., 2001)
involved_in piRNA transcription
inferred from mutant phenotype
(Chang et al., 2019)
involved_in positive regulation of piRNA transcription
inferred from mutant phenotype
(Baumgartner et al., 2022, Akkouche et al., 2017, Andersen et al., 2017)
inferred from direct assay
(Klattenhoff et al., 2009)
involved_in positive regulation of snRNA transcription by RNA polymerase II
inferred from mutant phenotype
(Pritykin et al., 2017)
involved_in positive regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Andersen et al., 2017)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in chromosome organization
inferred from sequence or structural similarity with FLYBASE:Su(var)205; FB:FBgn0003607
(Levine et al., 2012)
involved_in heterochromatin formation
inferred from biological aspect of ancestor with PANTHER:PTN000517042
(GO Reference Genome Project, 2011-)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
colocalizes_with heterochromatin
inferred from direct assay
(Klattenhoff et al., 2009)
located_in heterochromatin
inferred from direct assay
(Andersen et al., 2017)
is_active_in nucleus
inferred from direct assay
(Baumgartner et al., 2022)
located_in nucleus
inferred from direct assay
(Klattenhoff et al., 2009, Vermaak et al., 2005)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
located_in chromatin
inferred from sequence or structural similarity with FLYBASE:Su(var)205; FB:FBgn0003607
(Levine et al., 2012)
is_active_in pericentric heterochromatin
inferred from biological aspect of ancestor with PANTHER:PTN000517042
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
rhino (rhi) encodes a germline-restricted heterochromatin protein 1 required for fertility. It silences germline transposable elements by promoting transcription of the small RNAs encoded in Piwi-RNA clusters. rhi (as well as its interacting partners del and cuff) shows ovary-biased expression, but it is also expressed and functional in the testis. [Date last reviewed: 2021-09-16]
Gene Group (FlyBase)
HETEROCHROMATIN PROTEIN 1 FAMILY -
Heterochromatin Protein 1 (HP1, Su(var)205) was first discovered in Drosophila as a dominant suppressor of position-effect variegation and a major component of heterochromatin. Members of the HP1 family are characterised by the presence of an N-terminal chromodomain and a C-terminal chromoshadow domain. In addition to heterochromatin packaging and silencing, HP1 proteins have diverse roles in the nucleus, including the regulation of euchromatic genes. (Adapted from FBrf0193304).
RHINO-DEADLOCK-CUTOFF COMPLEX -
The Rhino-Deadlock-Cutoff (RDC) complex is a trimer that binds H3K9me3 and anchors itself onto dual-strand clusters. RDC complex regulates the non-canonical transcription and co-transcriptional processing of PIWI interacting RNA (piRNA) precursors and transposon silencing. (Adapted from FBrf0225218 and FBrf0250952).
Protein Function (UniProtKB)
Involved in piRNA (piwi-interacting RNA)-mediated transposon repression (PubMed:19732946, PubMed:36193674). May be involved in formation of the perinuclear nuage, a subcellular structure implicated in RNA processing that may be involved in transposon RNA surveillance and silencing (PubMed:19732946, PubMed:36193674). Required for ping-pong amplification during piRNA biogenesis, probably by promoting transcription of piRNA precursors (PubMed:19732946, PubMed:25085419, PubMed:36193674). As part of the Rhino-Deadlock-Cutoff (RDC) Complex associates with, and drives non-canonical transcription of germline specific dual-strand piRNA clusters 80F, 38C and 42AB, but not single-stranded piRNA cluster 20A (PubMed:19732946, PubMed:24906153, PubMed:25085419, PubMed:36193674). Induction of piRNA expression is potentially achieved through a mechanism that prevents transcriptional termination and leads to readthrough from flanking transcription units (PubMed:24906153). Recruited to specific chromatin regions by a combination of H3K9me2/3 histone methylation and differentially expressed sequence-specific recruitment factors (PubMed:24906153, PubMed:25085419, PubMed:25613572, PubMed:36193674). This association may involve direct interaction with DNA (PubMed:25613572). Associates with chromatin upon exposure to homologous piRNA and facilitates transcriptional read-through (PubMed:27292797). As part of the RDC complex, involved in suppression of splicing (PubMed:27292797). In ovaries, recruitment to specific heterochromatin clusters is nucleated and stabilized by kipf/kipferl (PubMed:36193674). During oogenesis, involved in axis specification and may regulate chromosome condensation at the onset of a mitotic-like phase that occurs during nurse cell chromosome duplication (PubMed:11729157). Involved in the distribution of mRNAs for proteins that play a role in anterior-posterior and dorsal-ventral axes specification during development of the oocyte, including grk/gurken, osk/oskar and vas/vasa (PubMed:11729157, PubMed:19732946). Mitigates meiotic double strand breaks and interacts with DNA damage signaling to mediate axis specification (PubMed:19732946).
(UniProt, Q7JXA8)
Summary (Interactive Fly)

HP1 homolog - required for nuage organization, transposon silencing, and ping-pong amplification of Piwi-interacting RNAs

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

Please see the JBrowse view of Dmel\rhi for information on other features

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

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry Q7JXA8)

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

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

Experimentally Determined Structures
Crossreferences
PDB - An information portal to biological macromolecular structures
Comments on Gene Model

Gene model reviewed during 5.50

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
rhi-RA
FBtr0086897
NM_080533
1570
418
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
rhi-PA
FBpp0086056
46.3
418
9.90
Q7JXA8
NP_536794
AAF57822
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)
Comments
External Data
Subunit Structure (UniProtKB)

Homodimer in solution (PubMed:25085419, PubMed:25613572). Dimerization is essential for chromatin binding (PubMed:25613572). Component of the Rhino-Deadlock-Cutoff (RDC) complex, composed of rhi/rhino, del/deadlock and cuff/cutoff (PubMed:24906153). Interacts (via C-terminus) with del/deadlock (via N-terminus); this interaction is direct (PubMed:24906153, PubMed:29858487). Two copies of del/deadlock associate with each rhi/rhino dimer (PubMed:29858487). Interacts with cuff/cutoff; this interaction is indirect and is mediated by del/deadlock (PubMed:24906153, PubMed:25085419). Interacts (via Chromo domain) with kipf/kipferl (via C2H2 type zinc finger 4) (PubMed:36193674). Interacts (via Chromo domain) with His3/histone H3 (via N-terminus di- or tri-methylated on 'Lys-10' (H3K9me2/3)); this interaction is direct (PubMed:24906153, PubMed:25085419, PubMed:25613572). Two His3 N-terminal tails oriented anti-parallel to each other are required for dimer binding to His3 (PubMed:25613572).

(UniProt, Q7JXA8)
Domain

The Chromo domain mediates binding to chromatin through interaction with the histone H3 N-terminal tail. Binding requires di- or tri- methylation of histone H3 'Lys-10' (H3K9me2/3).

(UniProt, Q7JXA8)
Crossreferences
InterPro - A database of protein families, domains and functional sites
PDB - An information portal to biological macromolecular structures
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\rhi using the Feature Mapper tool.

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

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

-0.15

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

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 7 -- 8
organism | ubiquitous
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic dorsal epidermis
(Fisher et al., 2012)
embryonic head epidermis
(Fisher et al., 2012)
embryonic ventral epidermis
(Fisher et al., 2012)
embryonic/larval posterior spiracle
(Fisher et al., 2012)
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage | female
ovary
(Levine et al., 2012)
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
Additional Descriptive Data
Marker for
 
Subcellular Localization
CV Term
Evidence
References
colocalizes_with heterochromatin
inferred from direct assay
(Klattenhoff et al., 2009)
located_in heterochromatin
inferred from direct assay
(Andersen et al., 2017)
is_active_in nucleus
inferred from direct assay
(Baumgartner et al., 2022)
located_in nucleus
inferred from direct assay
(Klattenhoff et al., 2009, Vermaak et al., 2005)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\rhi in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 1-3
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 24 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 19 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of rhi
Transgenic constructs containing regulatory region of rhi
Aberrations (Deficiencies and Duplications) ( 3 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
viable
Sterility
Allele
female fertile, with Scer\GAL4nanos.PG
female sterile
female sterile, with Scer\GAL4VP16.nanos.UTR, Scer\GAL4VP16.otu, Scer\GAL4nanos.PG
female sterile | recessive
fertile
Other Phenotypes
Allele
abnormal cell cycle | oogenesis, with Scer\GAL4nanos.PG
abnormal stress response | oogenesis, with Scer\GAL4nanos.PG
decreased cell number | progressive | spermatogenesis (with rhi02086)
decreased cell number | progressive | spermatogenesis (with rhiKG00910)
Phenotype manifest in
Allele
dorsal appendage
dorsal appendage (with rhi2-S18)
dorsal appendage (with rhi02086)
dorsal appendage | maternal effect (with rhi02086)
dorsal appendage | maternal effect (with rhiKG00910)
egg
egg chamber (with rhi1)
egg chamber (with rhi1-S6)
egg chamber (with rhi2-L12)
egg chamber (with rhi2-sL15)
egg chamber (with rhi02086)
female germline stem cell, with Scer\GAL4nanos.PG
germarium, with Scer\GAL4nanos.PU
male germline stem cell | decreased number (with rhi02086)
male germline stem cell | decreased number (with rhiKG00910)
nurse cell (with rhi1)
nurse cell (with rhi1-S6)
nurse cell (with rhi2-L12)
nurse cell (with rhi2-sL15)
nurse cell (with rhi02086)
nurse cell & chromatin (with rhi1)
nurse cell & chromatin (with rhi1-S6)
nurse cell & chromatin (with rhi2-L12)
nurse cell & chromatin (with rhi2-sL15)
nurse cell & chromatin (with rhi02086)
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (18)
Hsap\CBX1
5 of 14
Yes
No
2  
Hsap\CBX5
5 of 14
Yes
No
4  
Hsap\CBX3
4 of 14
No
No
3  
Hsap\CBX7
3 of 14
No
No
1  
Hsap\CBX2
2 of 14
No
No
Hsap\CBX4
2 of 14
No
No
Hsap\CBX6
2 of 14
No
No
Hsap\CBX8
2 of 14
No
No
Hsap\CDYL2
2 of 14
No
No
1  
Hsap\CDYL
2 of 14
No
No
Hsap\MPHOSPH8
2 of 14
No
No
Hsap\CDY1B
1 of 14
No
No
Hsap\CDY1
1 of 14
No
No
Hsap\CDY2A
1 of 14
No
No
Hsap\CDY2B
1 of 14
No
No
Hsap\RTL1
1 of 14
No
Yes
Hsap\SUV39H1
1 of 14
No
No
4  
Hsap\SUV39H2
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (15)
Rnor\Cbx1
5 of 14
Yes
No
Rnor\Cbx5
5 of 14
Yes
No
Rnor\Cbx3
4 of 14
No
No
Rnor\Cbx7
3 of 14
No
No
Rnor\Mphosph8
3 of 14
No
Yes
Rnor\Cbx6
2 of 14
No
No
Rnor\Cdyl2
2 of 14
No
No
Rnor\Cdyl
2 of 14
No
No
Rnor\LOC688980
2 of 14
No
No
Rnor\Cbx2
1 of 14
No
No
Rnor\Cbx4
1 of 14
No
No
Rnor\Cbx8
1 of 14
No
No
Rnor\LOC100363193
1 of 14
No
Yes
Rnor\Suv39h1l1
1 of 14
No
No
Rnor\Suv39h2
1 of 14
No
No
Mus musculus (laboratory mouse) (14)
Mmus\Cbx1
5 of 14
Yes
No
1  
Mmus\Cbx5
5 of 14
Yes
No
Mmus\Cbx3
3 of 14
No
No
Mmus\Cbx7
3 of 14
No
No
Mmus\Mphosph8
3 of 14
No
Yes
Mmus\Cbx6
2 of 14
No
No
Mmus\Cbx8
2 of 14
No
No
Mmus\Cdyl2
2 of 14
No
No
Mmus\Cdyl
2 of 14
No
No
Mmus\Cbx2
1 of 14
No
No
1  
Mmus\Cbx4
1 of 14
No
No
Mmus\Rtl1
1 of 14
No
Yes
Mmus\Suv39h1
1 of 14
No
No
Mmus\Suv39h2
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (16)
Xtro\cbx1
4 of 13
Yes
No
Xtro\cbx3
3 of 13
No
No
Xtro\cbx7
3 of 13
No
No
Xtro\cbx2
2 of 13
No
No
Xtro\cbx4
2 of 13
No
No
Xtro\cbx5
2 of 13
No
No
Xtro\zfp36l2.1
2 of 13
No
No
Xtro\cbx6
1 of 13
No
No
Xtro\cbx8
1 of 13
No
No
Xtro\cdyl2
1 of 13
No
No
Xtro\cdyl
1 of 13
No
No
Xtro\hoxd4
1 of 13
No
No
Xtro\mphosph8
1 of 13
No
Yes
Xtro\suv39h2
1 of 13
No
No
Xtro\syngr2
1 of 13
No
No
Xtro\ucn3
1 of 13
No
Yes
Danio rerio (Zebrafish) (17)
Drer\cbx1a
5 of 14
Yes
No
Drer\cbx1b
5 of 14
Yes
No
Drer\cbx3b
4 of 14
No
No
Drer\cbx5
4 of 14
No
No
Drer\cbx3a
3 of 14
No
No
Drer\cbx7b
3 of 14
No
No
Drer\mphosph8
3 of 14
No
Yes
Drer\cbx7a
2 of 14
No
No
Drer\cdyl
2 of 14
No
No
Drer\cbx2
1 of 14
No
No
Drer\cbx4
1 of 14
No
No
Drer\cbx6a
1 of 14
No
No
Drer\cbx6b
1 of 14
No
No
Drer\cbx8a
1 of 14
No
No
Drer\cbx8b
1 of 14
No
No
Drer\suv39h1a
1 of 14
No
No
Drer\suv39h1b
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (14)
Cele\cec-1
5 of 14
Yes
Yes
Cele\hpl-1
4 of 14
No
No
Cele\cec-3
3 of 14
No
No
Cele\cec-4
3 of 14
No
No
Cele\cec-5
2 of 14
No
No
Cele\cec-6
2 of 14
No
Yes
Cele\cec-8
2 of 14
No
No
Cele\hpl-2
2 of 14
No
No
Cele\C35E7.9
1 of 14
No
Yes
Cele\C38D4.10
1 of 14
No
No
Cele\F36H12.3
1 of 14
No
Yes
Cele\H04D03.6
1 of 14
No
No
Cele\heri-1
1 of 14
No
Yes
Cele\W05F2.8
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (4)
Agam\AgaP_AGAP004723
4 of 12
Yes
No
Agam\AgaP_AGAP009444
4 of 12
Yes
No
Agam\AgaP_AGAP003597
1 of 12
No
No
Agam\AgaP_AGAP012013
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (2)
Atha\TFL2
3 of 13
Yes
No
Atha\CMT1
1 of 13
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (3)
Scer\YGR109W-A
1 of 13
Yes
Yes
Scer\YGR109W-B
1 of 13
Yes
Yes
Scer\YIL082W-A
1 of 13
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (16)
Spom\chp1
3 of 12
Yes
No
Spom\chp2
3 of 12
Yes
No
Spom\swi6
2 of 12
No
No
Spom\Tf2-1
1 of 12
No
Yes
Spom\Tf2-2
1 of 12
No
Yes
Spom\Tf2-3
1 of 12
No
Yes
Spom\Tf2-4
1 of 12
No
Yes
Spom\Tf2-5
1 of 12
No
Yes
Spom\Tf2-6
1 of 12
No
Yes
Spom\Tf2-7
1 of 12
No
Yes
Spom\Tf2-8
1 of 12
No
Yes
Spom\Tf2-9
1 of 12
No
Yes
Spom\Tf2-10-pseudo
1 of 12
No
Yes
Spom\Tf2-11
1 of 12
No
Yes
Spom\Tf2-12
1 of 12
No
Yes
Spom\Tf2-13-pseudo
1 of 12
No
Yes
Escherichia coli (enterobacterium) (1)
Ecol\yihI
1 of 11
Yes
Yes
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:rhi. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (12)
HP1b
5 of 13
HP1c
5 of 13
HP1e
5 of 13
HP6
5 of 13
Oxp
5 of 13
Su(var)205
5 of 13
CG8289
3 of 13
Pc
2 of 13
Skadu
2 of 13
HP1D3csd
1 of 13
HP1Lcsd
1 of 13
Su(var)3-9
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 2 )
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    CBX1; chromobox 1
    5 of 14
    604511
         
        CBX5; chromobox 5
        5 of 14
        604478
             
            CBX3; chromobox 3
            4 of 14
            604477
                 
                CBX7; chromobox 7
                3 of 14
                608457
                     
                    Functional Complementation Data
                    Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
                    Interactions
                    Summary of Physical Interactions
                    Summary of Genetic Interactions
                    Interaction Browsers
                    View in FlyBase Interactions Browser   View in MIST tool (external link)
                    Please look at the allele data for full details of the genetic interactions
                    Starting gene(s)
                    Interaction type
                    Interacting gene(s)
                    Reference
                    rhi
                    suppressible
                    lok
                    (Klattenhoff et al., 2009)
                    rhi
                    suppressible
                    mei-41
                    (Klattenhoff et al., 2009)
                    Starting gene(s)
                    Interaction type
                    Interacting gene(s)
                    Reference
                    vas
                    suppressible
                    rhi
                    (Adashev et al., 2024)
                    External Data
                    Subunit Structure (UniProtKB)
                    Homodimer in solution (PubMed:25085419, PubMed:25613572). Dimerization is essential for chromatin binding (PubMed:25613572). Component of the Rhino-Deadlock-Cutoff (RDC) complex, composed of rhi/rhino, del/deadlock and cuff/cutoff (PubMed:24906153). Interacts (via C-terminus) with del/deadlock (via N-terminus); this interaction is direct (PubMed:24906153, PubMed:29858487). Two copies of del/deadlock associate with each rhi/rhino dimer (PubMed:29858487). Interacts with cuff/cutoff; this interaction is indirect and is mediated by del/deadlock (PubMed:24906153, PubMed:25085419). Interacts (via Chromo domain) with kipf/kipferl (via C2H2 type zinc finger 4) (PubMed:36193674). Interacts (via Chromo domain) with His3/histone H3 (via N-terminus di- or tri-methylated on 'Lys-10' (H3K9me2/3)); this interaction is direct (PubMed:24906153, PubMed:25085419, PubMed:25613572). Two His3 N-terminal tails oriented anti-parallel to each other are required for dimer binding to His3 (PubMed:25613572).
                    (UniProt, Q7JXA8 )
                    Linkouts
                    BioGRID - A database of protein and genetic interactions.
                    DroID - A comprehensive database of gene and protein interactions.
                    MIST (protein-protein) - An integrated Molecular Interaction Database
                    Pathways
                    Signaling Pathways (FlyBase)
                    Metabolic Pathways
                    FlyBase
                    External Links
                    External Data
                    Linkouts
                    Class of Gene
                    Genomic Location and Detailed Mapping Data
                    Chromosome (arm)
                    2R
                    Recombination map
                    2-83
                    Cytogenetic map
                    54D2-54D2
                    Sequence location
                    FlyBase Computed Cytological Location
                    Cytogenetic map
                    Evidence for location
                    54D2-54D2
                    Limits computationally determined from genome sequence between P{PZ}l(2)1050510505 and P{lacW}l(2)k07509k07509&P{EP}CG10934EP538
                    Experimentally Determined Cytological Location
                    Cytogenetic map
                    Notes
                    References
                    54C-54D
                    (determined by in situ hybridisation)
                    (Volpe et al., 2001)
                    54C1-54C12
                    (determined by in situ hybridisation)
                    (Spradling et al., 1999)
                    54C-54C
                    (determined by in situ hybridisation)
                    (Berg, 1994)
                    The insertions in rhi1 and rhi02086 both map to the 54C--54D border.
                    (Volpe et al., 2001)
                    Experimentally Determined Recombination Data
                    Location

                    2-83

                    (FlyBase, 2016)

                    2-86.5

                    (Comeron et al., 2012)
                    Left of (cM)
                    Right of (cM)
                    Notes
                    Stocks and Reagents
                    Stocks (10)
                    Genomic Clones (33)
                    cDNA Clones (21)
                     

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

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

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

                    cDNA Clones, End Sequenced (ESTs)
                    BDGP DGC clones
                    Other clones
                    RNAi and Array Information
                    Linkouts
                    DRSC - Results frm RNAi screens
                    Antibody Information
                    Laboratory Generated Antibodies
                     

                    monoclonal

                    (Baumgartner et al., 2022)

                    polyclonal

                    (Akulenko et al., 2018, Radion et al., 2017, Mohn et al., 2014, Zhang et al., 2014, Klattenhoff et al., 2009)
                    Commercially Available Antibodies
                     
                    Cell Line Information
                    Publicly Available Cell Lines
                     
                      Other Stable Cell Lines
                       
                        Other Comments

                        rhi is required for transposon silencing and piRNA production by dual-strand clusters.

                        (Klattenhoff et al., 2009)

                        rhi is required for proper chromosome conformation in the nurse cells during oogenesis and for the correct patterning of the eggshell and embryo.

                        (Volpe et al., 2001)

                        rhi is required for the formation of the two dorsolateral respiratory appendages of the eggshell from follicle cells.

                        (Volpe and Berg, 1995)

                        Mutant rhi embryos display altered dorsal/ventral polarity and altered dorsal appendage structure.

                        (Gillespie and Berg, 1991)

                        rhi mutants produce eggs with fused or thin filaments.

                        (Berg and Spradling, 1989)
                        Relationship to Other Genes
                        Source for database merge of

                        Source for merge of: rhi CG10683

                        (FlyBase, 1996-)
                        Additional comments

                        Source for merge of rhi CG10683 was sequence comparison ( date:010922 ).

                        (FlyBase, 1996-)
                        Nomenclature History
                        Source for database identify of
                        Nomenclature comments
                        Etymology

                        The gene is named "rhino" after the mutant phenotype; many eggs laid by mutant females have a single dorsal appendage.

                        (Volpe et al., 2001)
                        Synonyms and Secondary IDs (14)
                        Reported As
                        Symbol Synonym
                        CG10683
                        (Baumgartner et al., 2022, Meadows, 2017.1.11, Chen et al., 2012, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Zhang et al., 2010, Keleman et al., 2009.8.5)
                        HP1
                        (Le Thomas et al., 2014, Yamamoto et al., 2014)
                        HP1D
                        (Levine et al., 2016, Beck and Llopart, 2015, Levine and Malik, 2013, Levine et al., 2012, Levine et al., 2012, Vermaak et al., 2005)
                        HP1d
                        (Comoglio and Paro, 2014)
                        RHI
                        (Gamez et al., 2020, Kolaczkowski et al., 2011)
                        Rhi
                        (Suyama and Kai, 2025, Patel et al., 2024, Jin et al., 2023, Lu et al., 2023, Huang and Wong, 2021, Onishi et al., 2021, Cacchione et al., 2020, Parhad et al., 2020, Sato and Siomi, 2020, Kneuss et al., 2019, DeLuca and Spradling, 2018, Rashpa et al., 2017, Ryazansky et al., 2017, Ryazansky et al., 2017, Yang and Xi, 2017, Hirakata and Siomi, 2016, Malone et al., 2015, Guzzardo et al., 2013, Levine et al., 2012)
                        Rhino
                        (Pritam and Signor, 2025, Richards et al., 2022, Gibert and Peronnet, 2021, Liu et al., 2020, Soleimani et al., 2020, Casier et al., 2019, Zhang et al., 2018, Chen et al., 2016, Eissenberg and Elgin, 2014)
                        hp1d
                        (Lavrenov et al., 2014)
                        rhi
                        (Stanek et al., 2025, Baumgartner et al., 2024, Kotov et al., 2024, Iki et al., 2023, Nieken et al., 2023, Saha et al., 2023, McCarthy et al., 2022, Chen et al., 2021, Chen et al., 2021, Gebert et al., 2021, Kelleher, 2021, Kim et al., 2021, Ellison et al., 2020, Saint-Leandre et al., 2020, Chang et al., 2019, Courret et al., 2019, ElMaghraby et al., 2019, Kneuss et al., 2019, Story et al., 2019, Wakisaka et al., 2019, Gene Disruption Project members, 2018-, Théron et al., 2018, Akkouche et al., 2017, Cheng et al., 2017, Parhad et al., 2017, Pritykin et al., 2017, Hermant et al., 2015, Mohn et al., 2014, Zhang et al., 2014, Czech et al., 2013, Sen et al., 2013, Kelleher et al., 2012, Langley et al., 2012, Baxley et al., 2011, Pane et al., 2011, Khurana et al., 2010, Klattenhoff et al., 2009, Neuman-Silberberg, 2007, Tao et al., 2007, Xing et al., 2007)
                        rhino
                        (Chen and Aravin, 2015, Han et al., 2015, Le Thomas et al., 2014, Mani and Juliano, 2013, Kelleher et al., 2012, Zhang et al., 2012)
                        rno
                        Name Synonyms
                        Heterochromatin protein 1
                        (Yamamoto et al., 2014)
                        RHINO
                        (Ho et al., 2014)
                        Rhino
                        (Ho et al., 2024, Baumgartner et al., 2022, Kotov et al., 2022, Casale et al., 2021, Wang et al., 2021, Kordyukova et al., 2020, Parhad et al., 2020, Hirakata and Siomi, 2019, Radion et al., 2019, Saha and Mishra, 2019, Cheng et al., 2017, Huang et al., 2017, Pritykin et al., 2017, Ryazansky et al., 2017, Hirakata and Siomi, 2016, Molla-Herman et al., 2015, Yu et al., 2015, Dufourt et al., 2014, Kumar and Chen, 2012, Levine et al., 2012, Brideau and Barbash, 2011, Pek and Kai, 2011, Klattenhoff et al., 2009)
                        heterochromatin protein 1
                        (Le Thomas et al., 2014)
                        rhino
                        (Shapiro-Kulnane et al., 2022, Zhang et al., 2021, Andersen et al., 2017, Marie et al., 2017, Specchia et al., 2017, Blumenstiel et al., 2016, Chen and Aravin, 2015, Comoglio and Paro, 2014, Dufourt et al., 2014, Handler et al., 2013, Levine and Malik, 2013, Simkin et al., 2013, Baxley et al., 2011, Olivieri et al., 2010, McGraw et al., 2008, Neuman-Silberberg, 2007, Tao et al., 2007, Xing et al., 2007, Vermaak et al., 2005)
                        Secondary FlyBase IDs
                        • FBgn0034254
                        Datasets (1)
                        Study focus (1)
                        Experimental Role
                        Project
                        Project Type
                        Title
                        • bait_protein
                        modENCODE_chromatin_proteins_cells
                        Genome-wide localization of chromosomal proteins in cell lines by ChIP-chip and ChIP-Seq.
                        Study result (0)
                        Result
                        Result Type
                        Title
                        External Crossreferences and Linkouts ( 87 )
                        Sequence Crossreferences
                        NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                        GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                        GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                        RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                        UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                        UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                        UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                        Other crossreferences
                        AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                        BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                        DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                        EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                        FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                        FlyMine - An integrated database for Drosophila genomics
                        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)
                        PDB - An information portal to biological macromolecular structures
                        Linkouts
                        BioGRID - A database of protein and genetic interactions.
                        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                        DroID - A comprehensive database of gene and protein interactions.
                        DRSC - Results frm RNAi screens
                        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                        Flygut - An atlas of the Drosophila adult midgut
                        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                        MIST (protein-protein) - An integrated Molecular Interaction Database
                        References (192)