swi6

UniProt ID: P40381
Organism: Schizosaccharomyces pombe (strain 972 / ATCC 24843)
Review Status: DRAFT
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Gene Description

Swi6 is the principal HP1 (heterochromatin protein 1) ortholog of fission yeast and the central structural reader and effector of heterochromatin. It is a 328-residue protein built from an N-terminal chromodomain that binds histone H3 methylated at lysine 9 (H3K9me2/me3, deposited by the Clr4 methyltransferase) and a C-terminal chromoshadow domain that homodimerizes and forms a protein-protein interaction platform, the two domains connected by a basic disordered hinge with nucleic-acid-binding activity. Through this architecture Swi6 recognizes H3K9-methylated nucleosomes, oligomerizes, and undergoes a switch from an auto-inhibited to a spreading-competent state that compacts chromatin. The chromoshadow platform recruits diverse effectors to heterochromatin, including the cohesin subunit Psc3, the SHREC/Clr3 histone deacetylase machinery, the anti-silencing JmjC protein Epe1, the RNAi bridging factor Ers1/RDRC, the histone chaperones Asf1/HIRA and CAF-1, DBF4-dependent kinase (Dfp1), the mating-type switching factor Swi2, and meiotic shugoshin Sgo1. Swi6 mediates transcriptional and co-transcriptional gene silencing and the assembly, demarcation and confinement of heterochromatin at centromeres, telomeres, the silent mating-type cassettes, and rDNA. At centromeres it enriches cohesin to ensure accurate sister-chromatid cohesion, biorientation and chromosome segregation; at the mating-type region it is required for donor selection during mating-type switching; and at telomeres it contributes to subtelomeric heterochromatin and telomere tethering.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0031507 heterochromatin formation
IBA
GO_REF:0000033
ACCEPT
Summary: Heterochromatin formation/assembly is a core biological process of Swi6, conserved across the HP1 family. Phylogenetic inference is corroborated by extensive S. pombe experimental evidence.
Reason: Core HP1 function, supported by phylogeny and abundant direct evidence.
Supporting Evidence:
PMID:11283354
Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic regions is dependent on H3 Lys9 methylation.
GO:0003682 chromatin binding
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: Chromatin binding is correct but generic; the informative molecular function is H3K9me2/3 reader activity via the chromodomain. Retained as a true but non-core (parent) descriptor.
Reason: True but superseded by the more specific H3K9me reader activity term.
Supporting Evidence:
PMID:11242054
The chromo domain of HP1 is identified as its methyl-lysine-binding domain.
GO:0005721 pericentric heterochromatin
IBA
GO_REF:0000033
ACCEPT
Summary: Pericentric heterochromatin is a core site of Swi6 action; this is strongly corroborated by direct localization data (multiple IDA annotations below).
Reason: Phylogenetic call confirmed by direct localization evidence.
Supporting Evidence:
PMID:10766735
Swi6 and Chp1 are confined to the flanking outer repeats and Swi6 can spread across at least 3 kb of extraneous chromatin in cen1.
GO:0000792 heterochromatin
IEA
GO_REF:0000117
ACCEPT
Summary: Heterochromatin localization is correct and strongly supported by direct IDA evidence; Swi6 is a defining component of heterochromatin. Accepted to remain consistent with the IDA annotations to this term.
Reason: Correct and core localization, corroborated by direct evidence.
Supporting Evidence:
PMID:11069763
GFP-Swi6p localises to the nucleus and is concentrated at the heterochromatic centromeres and telomeres.
GO:0005634 nucleus
IEA
GO_REF:0000120
KEEP AS NON CORE
Summary: Swi6 is a nuclear protein. Correct but generic; the heterochromatin subdomain terms are the informative localizations.
Reason: Correct broad localization, subsumed by specific nuclear terms.
Supporting Evidence:
PMID:11069763
GFP-Swi6p localises to the nucleus and is concentrated at the heterochromatic centromeres and telomeres.
GO:0005694 chromosome
IEA
GO_REF:0000044
KEEP AS NON CORE
Summary: Chromosome localization is correct (Swi6 is chromatin-associated) but generic relative to the heterochromatin and chromatin terms.
Reason: Correct but generic.
Supporting Evidence:
PMID:7660126
Swi6p localizes with these three chromosomal regions.
GO:0040029 epigenetic regulation of gene expression
IEA
GO_REF:0000117
KEEP AS NON CORE
Summary: Swi6 mediates epigenetic gene silencing through H3K9me-based heterochromatin. Correct but high-level; the specific silencing/ heterochromatin formation terms are preferred.
Reason: Correct but generic parent of the specific silencing processes.
Supporting Evidence:
PMID:11242054
This model may also explain the stable inheritance of the heterochromatic state.
GO:0005515 protein binding
IPI
PMID:14625560
Hsk1-Dfp1 is required for heterochromatin-mediated cohesion ...
MARK AS OVER ANNOTATED
Summary: Bare "protein binding" (here the interaction with the DDK kinase subunit Hsk1) is uninformative. The informative molecular function captured by this interaction is Swi6's chromatin-protein adaptor/scaffold activity recruiting DDK to pericentromeric heterochromatin.
Reason: Uninformative MF; superseded by chromatin-protein adaptor activity.
Supporting Evidence:
PMID:31000521
Swi6 interacts with Dfp1, a regulatory subunit of DBF4-dependent kinase (DDK)
GO:0005515 protein binding
IPI
PMID:18716626
Heterochromatin links to centromeric protection by recruitin...
MARK AS OVER ANNOTATED
Summary: The Swi6-Sgo1 interaction is real and biologically important (meiotic cohesion protection), but "protein binding" is uninformative. The function is chromatin-protein adaptor activity (shugoshin recruitment).
Reason: Uninformative MF; the informative term is adaptor activity.
Supporting Evidence:
PMID:18716626
the heterochromatin protein Swi6 associates directly with meiosis-specific shugoshin Sgo1, a protector of cohesin at centromeres.
GO:0005515 protein binding
IPI
PMID:22474355
Heterochromatin protein 1 homologue Swi6 acts in concert wit...
MARK AS OVER ANNOTATED
Summary: The Swi6-Ers1 interaction bridges Swi6 to the RDRC/RNAi machinery, but "protein binding" is uninformative. The informative MF is chromatin-protein adaptor activity (also annotated separately from this reference).
Reason: Uninformative MF; captured better by adaptor activity.
Supporting Evidence:
PMID:22474355
Swi6 recruits RDRC to heterochromatin through Ers1, an RNAi factor intermediate.
GO:0005515 protein binding
IPI
PMID:22990236
Hrp3 controls nucleosome positioning to suppress non-coding ...
MARK AS OVER ANNOTATED
Summary: Interaction with the CHD chromatin remodeler Hrp3. "Protein binding" is uninformative; the informative MF is chromatin-protein adaptor activity.
Reason: Uninformative MF.
GO:0042802 identical protein binding
IPI
PMID:14663140
Two different Swi5-containing protein complexes are involved...
ACCEPT
Summary: Swi6 homodimerizes via its chromoshadow domain; this self-association is a core, informative property (distinct from bare protein binding) that underlies effector recruitment and heterochromatin spreading.
Reason: Homodimerization is a genuine, mechanistically central MF.
Supporting Evidence:
PMID:10801440
The 1.9 A crystal structure of the Swi6 CSD is presented here. This reveals a novel dimeric structure
GO:0042802 identical protein binding
IPI
PMID:18716626
Heterochromatin links to centromeric protection by recruitin...
ACCEPT
Summary: Swi6 self-association (chromoshadow-domain homodimerization). Core MF; duplicate of the homodimerization annotation supported elsewhere.
Reason: Genuine homodimerization, central to HP1 function.
Supporting Evidence:
PMID:10801440
Dimerisation through the CSD of HP1-like proteins results in the simultaneous formation of a putative protein-protein interaction pit
GO:0042802 identical protein binding
IPI
PMID:23485968
A conformational switch in HP1 releases auto-inhibition to d...
ACCEPT
Summary: Swi6 self-associates into dimers (CSD-CSD) and higher-order oligomers; a conformational switch on methylated nucleosomes drives the spreading- competent oligomeric state. Core MF.
Reason: Self-association/oligomerization is mechanistically central.
Supporting Evidence:
PMID:23485968
the CSD can homodimerize
GO:0031507 heterochromatin formation
IDA
PMID:11780129
Recruitment of cohesin to heterochromatic regions by Swi6/HP...
ACCEPT
Summary: Direct evidence that Swi6 is required for heterochromatin function at centromeres and the mating-type region. Core process.
Reason: Direct experimental support for heterochromatin formation role.
Supporting Evidence:
PMID:11780129
the preferential localization of cohesins at the centromeric repeats is dependent on Swi6, a conserved heterochromatin protein that is required for proper kinetochore function.
GO:0140185 siRNA-mediated silent mating type cassette region heterochromatin formation
EXP
PMID:39747188
PhpC(NF-Y) transcription factor infiltrates heterochromatin ...
ACCEPT
Summary: Swi6 participates in siRNA/RNAi-based heterochromatin nucleation at the mating-type cassette region, where TF-driven transcription generates siRNAs that trigger heterochromatin assembly. Core process.
Reason: Experimental support for siRNA-mediated mat-region heterochromatin.
Supporting Evidence:
PMID:39747188
siRNA production by this TF-mediated transcription pathway is critical for heterochromatin nucleation at target repeat loci.
GO:0005634 nucleus
EXP
PMID:18761674
Fission yeast chromatin assembly factor 1 assists in the rep...
KEEP AS NON CORE
Summary: Swi6 is a nuclear protein (CAF-1 study). Correct; generic relative to the heterochromatin subdomain localizations.
Reason: Correct but generic localization.
GO:0005694 chromosome
EXP
PMID:18761674
Fission yeast chromatin assembly factor 1 assists in the rep...
KEEP AS NON CORE
Summary: Swi6 is chromatin/chromosome-associated. Correct but generic relative to the heterochromatin terms.
Reason: Correct but generic localization.
GO:0140463 chromatin-protein adaptor activity
IPI
PMID:11780129
Recruitment of cohesin to heterochromatic regions by Swi6/HP...
ACCEPT
Summary: Core molecular function. Swi6 bridges H3K9-methylated chromatin to cohesin via direct interaction with the cohesin subunit Psc3, enriching cohesin at heterochromatin. This is the informative MF underlying many "protein binding" interactions.
Reason: Direct adaptor activity (Psc3/cohesin recruitment); core function.
Supporting Evidence:
PMID:11780129
a cohesin subunit Psc3 interacts with Swi6 and its mouse homologue HP1.
GO:0031508 pericentric heterochromatin formation
IMP
PMID:19443688
Diverse roles of HP1 proteins in heterochromatin assembly an...
ACCEPT
Summary: Swi6 is required for pericentromeric heterochromatin/silencing, acting with associated HDAC complexes to limit Pol II occupancy. Core process.
Reason: Mutant phenotype supports pericentric heterochromatin formation role.
Supporting Evidence:
PMID:19443688
Swi6 and Chp2 proteins and their associated HDAC complexes have overlapping functions in limiting Pol II occupancy across pericentromeric heterochromatin domains.
GO:0007535 donor selection
IDA
PMID:29852001
New insights into donor directionality of mating-type switch...
ACCEPT
Summary: Swi6 is required for correct donor selection (directionality) during mating-type switching, cooperating with heterochromatin and Set1C to direct donor choice. Core process.
Reason: Direct evidence for Swi6 role in donor selection.
Supporting Evidence:
PMID:29852001
Mating-type switching in Schizosaccharomyces pombe entails programmed gene conversion events regulated by DNA replication, heterochromatin, and the HP1-like chromodomain protein Swi6.
GO:0031508 pericentric heterochromatin formation
IMP
PMID:19136623
Phosphorylation of Swi6/HP1 regulates transcriptional gene s...
ACCEPT
Summary: Swi6 recruits SHREC, Epe1 and cohesin to pericentric heterochromatin, and CK2 phosphorylation of Swi6 controls transcriptional gene silencing there. Core process (duplicate term, distinct supporting reference).
Reason: Supports pericentric heterochromatin/silencing role.
Supporting Evidence:
PMID:19136623
a HP1 homolog Swi6 recruits SHREC, Epe1, and cohesin, which are involved in transcriptional gene silencing (TGS), transcriptional activation, and sister chromatid cohesion, respectively.
GO:0031509 subtelomeric heterochromatin formation
IMP
PMID:25245948
Tls1 regulates splicing of shelterin components to control t...
ACCEPT
Summary: Swi6 contributes to telomeric/subtelomeric heterochromatin assembly (in this study examined via Tls1-dependent shelterin splicing). Core process.
Reason: Supports subtelomeric heterochromatin formation role.
Supporting Evidence:
PMID:25245948
we comprehensively identified factors required for telomeric heterochromatin assembly, including a novel gene tls1+.
GO:0140463 chromatin-protein adaptor activity
IDA
PMID:36951094
Fission yeast Swi2 designates cell-type specific donor and s...
ACCEPT
Summary: Swi6 acts as an adaptor recruiting the mating-type switching factor Swi2 to a recombination enhancer via a defined Swi6-binding site, supporting donor selection. Core adaptor MF.
Reason: Direct evidence Swi6 recruits Swi2 (adaptor activity).
Supporting Evidence:
PMID:36951094
the Swi6-binding site was required for Swi2 localization at SRE2 to select mat2-P in M cells.
GO:0005721 pericentric heterochromatin
IDA
PMID:18809570
Balance between distinct HP1 family proteins controls hetero...
ACCEPT
Summary: Direct localization of Swi6 to pericentric heterochromatin. Core CC (one of many concordant IDA annotations).
Reason: Direct localization evidence.
Supporting Evidence:
PMID:18809570
both Swi6 and Chp2 are required for the assembly of fully repressive heterochromatin
GO:0031934 mating-type region heterochromatin
IDA
PMID:18809570
Balance between distinct HP1 family proteins controls hetero...
ACCEPT
Summary: Direct localization/activity of Swi6 at mating-type region heterochromatin. Core CC.
Reason: Direct evidence for activity at mat-region heterochromatin.
Supporting Evidence:
PMID:18809570
two HP1 family proteins, Swi6 and Chp2, are involved in transcriptional silencing at heterochromatic regions
GO:0140720 subtelomeric heterochromatin
IDA
PMID:18809570
Balance between distinct HP1 family proteins controls hetero...
ACCEPT
Summary: Direct localization/activity of Swi6 at subtelomeric heterochromatin. Core CC.
Reason: Direct evidence for activity at subtelomeric heterochromatin.
Supporting Evidence:
PMID:18809570
both Swi6 and Chp2 are required for the assembly of fully repressive heterochromatin
GO:0005721 pericentric heterochromatin
IDA
PMID:11780129
Recruitment of cohesin to heterochromatic regions by Swi6/HP...
ACCEPT
Summary: Swi6 localizes to and acts at pericentric (centromeric repeat) heterochromatin, where it enriches cohesin. Core CC.
Reason: Direct localization at centromeric repeats.
Supporting Evidence:
PMID:11780129
the preferential localization of cohesins at the centromeric repeats is dependent on Swi6
GO:0031934 mating-type region heterochromatin
IDA
PMID:11780129
Recruitment of cohesin to heterochromatic regions by Swi6/HP...
ACCEPT
Summary: Swi6 acts at mating-type region heterochromatin (cohesin enrichment preserving locus integrity). Core CC.
Reason: Direct evidence for activity at mat-region heterochromatin.
Supporting Evidence:
PMID:11780129
Cohesin is also enriched at the mating-type heterochromatic region in a manner that depends on Swi6
GO:0030466 silent mating-type cassette heterochromatin formation
EXP
PMID:15292231
Regulation of Swi6/HP1-dependent heterochromatin assembly by...
ACCEPT
Summary: Swi6-dependent heterochromatin assembly across the silent mating-type region, downstream of Atf1/Pcr1- and Clr6-mediated histone deacetylation. Core process.
Reason: Experimental support for mat-cassette heterochromatin formation.
Supporting Evidence:
PMID:15292231
a stochastic Swi6 (the fission yeast HP1 homolog)-dependent mechanism that is not fully understood.
GO:0005634 nucleus
IDA
PMID:11069763
Live analysis of lagging chromosomes during anaphase and the...
KEEP AS NON CORE
Summary: Direct (GFP) evidence for nuclear localization. Correct; generic relative to heterochromatin subdomains.
Reason: Correct but generic localization.
Supporting Evidence:
PMID:11069763
GFP-Swi6p localises to the nucleus and is concentrated at the heterochromatic centromeres and telomeres.
GO:0140463 chromatin-protein adaptor activity
IDA
PMID:22474355
Heterochromatin protein 1 homologue Swi6 acts in concert wit...
ACCEPT
Summary: Swi6 functions as an adaptor recruiting the RDRC/RNAi machinery to heterochromatin through Ers1. Core adaptor MF (duplicate term, distinct reference).
Reason: Direct evidence Swi6 recruits RDRC via Ers1.
Supporting Evidence:
PMID:22474355
we demonstrate that Swi6 recruits RDRC to heterochromatin through Ers1, an RNAi factor intermediate.
GO:0062072 histone H3K9me2/3 reader activity
IDA
PMID:11242054
Selective recognition of methylated lysine 9 on histone H3 b...
ACCEPT
Summary: Defining, core molecular function. The Swi6/HP1 chromodomain binds H3 methylated at Lys9 (not Lys4) with high affinity; a chromodomain point mutation abolishes methyl-lysine binding and silencing.
Reason: Foundational direct demonstration of H3K9me reader activity.
Supporting Evidence:
PMID:11242054
HP1 can bind with high affinity to histone H3 methylated at lysine 9 but not at lysine 4. The chromo domain of HP1 is identified as its methyl-lysine-binding domain.
GO:0030466 silent mating-type cassette heterochromatin formation
EXP
PMID:16246721
The nucleation and maintenance of heterochromatin by a histo...
ACCEPT
Summary: Swi6/HP1 is among the heterochromatin proteins required as Clr3 spreads across the ~20 kb silenced mating-type domain. Core process (duplicate term, distinct reference).
Reason: Experimental support for mat-cassette heterochromatin assembly.
Supporting Evidence:
PMID:16246721
Clr3 spreads across the 20 kb silenced domain that requires its own HDAC activity and heterochromatin proteins including Swi6/HP1.
GO:0005721 pericentric heterochromatin
IDA
PMID:15372076
A chromodomain protein, Chp1, is required for the establishm...
ACCEPT
Summary: Direct localization of Swi6 to pericentric/centromeric heterochromatin (Chp1 study). Core CC.
Reason: Direct localization evidence.
Supporting Evidence:
PMID:15372076
deletion of the chp1(+) gene causes centromere-specific decreases in Swi6 localization and histone H3-K9 methylation
GO:0031934 mating-type region heterochromatin
IDA
PMID:15372076
A chromodomain protein, Chp1, is required for the establishm...
ACCEPT
Summary: Swi6 is a structural component of mating-type region heterochromatin, contributing to residual H3K9 methylation there. Core CC.
Reason: Direct evidence for activity at mat-region heterochromatin.
Supporting Evidence:
PMID:15372076
Swi6 and Chp2 are critical to maintaining this residual methylation.
GO:0140720 subtelomeric heterochromatin
IDA
PMID:15372076
A chromodomain protein, Chp1, is required for the establishm...
ACCEPT
Summary: Swi6 is a structural component of telomeric/subtelomeric heterochromatin. Core CC.
Reason: Direct evidence for activity at subtelomeric heterochromatin.
Supporting Evidence:
PMID:15372076
Chp1 is a structural component of three heterochromatic regions-centromeres, the mating-type region, and telomeres
GO:0033562 co-transcriptional gene silencing by RNA interference machinery
IMP
PMID:17512405
RNAi-dependent and -independent RNA turnover mechanisms cont...
ACCEPT
Summary: Swi6 contributes to RNAi-dependent heterochromatic gene silencing, which involves co-transcriptional RNA processing/turnover of heterochromatic transcripts. Core process.
Reason: Supports RNAi co-transcriptional silencing role.
Supporting Evidence:
PMID:17512405
the RNAi pathway is required for heterochromatin-dependent silencing of transgene insertions at centromeric repeats
GO:0140727 siRNA-mediated pericentric heterochromatin formation
IMP
PMID:12193640
Regulation of heterochromatic silencing and histone H3 lysin...
ACCEPT
Summary: Swi6 functions in the RNAi-directed pathway that establishes pericentromeric heterochromatin and H3K9 methylation through siRNAs from centromeric repeats. Core process.
Reason: Supports siRNA-mediated pericentric heterochromatin formation.
Supporting Evidence:
PMID:12193640
We propose that double-stranded RNA arising from centromeric repeats targets formation and maintenance of heterochromatin through RNAi.
GO:0033696 heterochromatin boundary formation
IMP
PMID:21211723
Asf1/HIRA facilitate global histone deacetylation and associ...
ACCEPT
Summary: Swi6 associates with the Asf1/HIRA histone chaperone, which spreads across silenced domains to enforce silencing and promote nucleosome occupancy, contributing to proper heterochromatin organization at boundaries. Core process.
Reason: Supports role in heterochromatin boundary/nucleosome occupancy.
Supporting Evidence:
PMID:21211723
a histone chaperone complex containing Asf1 and HIRA that spreads across silenced domains via its association with Swi6 to enforce transcriptional silencing.
GO:0140463 chromatin-protein adaptor activity
EXP
PMID:16762840
Swi6/HP1 recruits a JmjC domain protein to facilitate transc...
ACCEPT
Summary: Swi6/HP1 recruits the JmjC anti-silencing protein Epe1 to heterochromatin, balancing opposing chromatin-modifying activities. Core adaptor MF.
Reason: Direct evidence Swi6 recruits Epe1 (adaptor activity).
Supporting Evidence:
PMID:16762840
Epe1 is recruited to heterochromatic loci by the heterochromatin protein Swi6/HP1.
GO:0140463 chromatin-protein adaptor activity
EXP
PMID:17449867
Interaction of Epe1 with the heterochromatin assembly pathwa...
ACCEPT
Summary: Confirms Swi6 recruits Epe1 to heterochromatin. Core adaptor MF (duplicate term, distinct reference).
Reason: Independent evidence for Swi6-mediated Epe1 recruitment.
Supporting Evidence:
PMID:17449867
We report that Swi6 recruits Epe1 to heterochromatin
GO:0003677 DNA binding
IDA
PMID:37400983
Cooperative DNA-binding activities of Chp2 are critical for ...
KEEP AS NON CORE
Summary: PomBase experimental (IDA) annotation. Our cached abstract foregrounds Chp2, the other fission-yeast HP1 protein, but the full text (not in our cache) compares both HP1 proteins, and PomBase made a direct-assay (IDA) DNA-binding annotation to Swi6 on its basis. Swi6 is independently well established to bind DNA in a sequence- independent manner via its hinge region. DNA binding is thus a genuine biochemical property of Swi6 but is ancillary to its core role as an H3K9me reader/effector.
Reason: Valid PomBase IDA; Swi6 hinge-mediated DNA binding is well established. Retained as a real but non-core property rather than a defining function. (An earlier assessment that this Chp2-titled paper does not support Swi6 was based on the abstract-only cache and overruled the curator without justification.)
Supporting Evidence:
PMID:37400983
Here, we focus on Chp2, one of the two HP1 proteins in fission yeast, and investigate how Chp2's DNA-binding ability contributes to its function.
GO:0005515 protein binding
IPI
PMID:10801440
Dimerisation of a chromo shadow domain and distinctions from...
MARK AS OVER ANNOTATED
Summary: This row records the Swi6-Swi6 (chromoshadow-domain) self-interaction; it is more informatively captured as identical protein binding / homodimerization. Bare "protein binding" is uninformative.
Reason: Self-interaction better described as homodimerization (GO:0042802).
Supporting Evidence:
PMID:10801440
The 1.9 A crystal structure of the Swi6 CSD is presented here. This reveals a novel dimeric structure
GO:0003723 RNA binding
EXP
PMID:22683269
HP1(Swi6) mediates the recognition and destruction of hetero...
ACCEPT
Summary: Core molecular function. Swi6 binds RNA via its hinge basic patch; RNA competes with H3K9me for the chromodomain, and Swi6 sequesters and primes heterochromatic transcripts for degradation.
Reason: Direct in vivo/in vitro evidence for Swi6 RNA binding.
Supporting Evidence:
PMID:22683269
RNA competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
GO:0140463 chromatin-protein adaptor activity
IPI
PMID:18716626
Heterochromatin links to centromeric protection by recruitin...
ACCEPT
Summary: Swi6 recruits meiotic shugoshin Sgo1 to centromeres, the informative MF underlying the Swi6-Sgo1 interaction. Core adaptor MF.
Reason: Direct evidence Swi6 recruits Sgo1 (adaptor activity).
Supporting Evidence:
PMID:18716626
the heterochromatin protein Swi6 associates directly with meiosis-specific shugoshin Sgo1, a protector of cohesin at centromeres.
GO:1990813 meiotic centromeric cohesion protection in anaphase I
EXP
PMID:18716626
Heterochromatin links to centromeric protection by recruitin...
ACCEPT
Summary: By recruiting shugoshin Sgo1, Swi6 protects centromeric cohesin during meiosis; forced Sgo1 localization rescues meiotic segregation in swi6 cells. Core meiotic process.
Reason: Experimental support for meiotic centromeric cohesion protection.
Supporting Evidence:
PMID:18716626
The forced centromeric localization of Sgo1 restores proper meiotic chromosome segregation in swi6 cells.
GO:0062072 histone H3K9me2/3 reader activity
IDA
PMID:22683269
HP1(Swi6) mediates the recognition and destruction of hetero...
ACCEPT
Summary: Confirms the Swi6 chromodomain reads H3K9me (in competition with RNA). Core MF (duplicate term, distinct reference).
Reason: Direct evidence for H3K9me reader activity.
Supporting Evidence:
PMID:22683269
RNA competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
GO:1990342 heterochromatin island
IDA
PMID:22144463
RNA elimination machinery targeting meiotic mRNAs promotes f...
KEEP AS NON CORE
Summary: Swi6 is active at facultative heterochromatin islands that form over meiotic genes via RNA elimination machinery. Genuine but specialized (facultative) localization; kept as non-core relative to constitutive heterochromatin sites.
Reason: Genuine facultative-heterochromatin localization; non-core context.
Supporting Evidence:
PMID:22144463
we describe facultative heterochromatin islands in fission yeast and show that their formation at meiotic genes requires factors that eliminate meiotic messenger RNAs (mRNAs) during vegetative growth.
GO:0006325 chromatin organization
IMP
PMID:19111658
HP1 proteins form distinct complexes and mediate heterochrom...
KEEP AS NON CORE
Summary: Swi6 organizes heterochromatin via a network of associated chromatin proteins and contributes to silencing. Correct but high-level; the specific heterochromatin formation/silencing terms are preferred.
Reason: Correct but generic relative to specific heterochromatin processes.
Supporting Evidence:
PMID:19111658
Swi6 associates with a different set of nuclear proteins and with noncoding centromeric transcripts and is required for efficient RNAi-dependent processing of these transcripts.
GO:0071962 mitotic sister chromatid cohesion, centromeric
IMP
PMID:31000521
Overlapping Roles in Chromosome Segregation for Heterochroma...
ACCEPT
Summary: Swi6 is required for cohesin (Rad21) enrichment at the pericentromere, establishing centromeric sister-chromatid cohesion. Core process.
Reason: Mutant phenotype supports centromeric cohesion role.
Supporting Evidence:
PMID:31000521
Swi6 is required for cohesin enrichment at the pericentromere.
GO:1990758 mitotic sister chromatid biorientation
IMP
PMID:31000521
Overlapping Roles in Chromosome Segregation for Heterochroma...
ACCEPT
Summary: Loss of Swi6 causes defects in establishing bioriented sister kinetochores; via cohesin enrichment Swi6 promotes biorientation. Core process.
Reason: Mutant phenotype supports biorientation role.
Supporting Evidence:
PMID:31000521
Loss of Swi6 leads to abnormal mitosis, including defects in the establishment of bioriented sister kinetochores and microtubule attachment.
GO:0005721 pericentric heterochromatin
IDA
PMID:11283354
Role of histone H3 lysine 9 methylation in epigenetic contro...
ACCEPT
Summary: Direct evidence that Swi6 localizes to pericentric heterochromatin in an H3K9me-dependent manner. Core CC.
Reason: Direct localization evidence.
Supporting Evidence:
PMID:11283354
Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic regions is dependent on H3 Lys9 methylation.
GO:0031934 mating-type region heterochromatin
IDA
PMID:11283354
Role of histone H3 lysine 9 methylation in epigenetic contro...
ACCEPT
Summary: Swi6 localizes to mating-type region heterochromatin (H3K9me-dependent). Core CC.
Reason: Direct localization evidence at mat-region heterochromatin.
Supporting Evidence:
PMID:11283354
Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic regions is dependent on H3 Lys9 methylation.
GO:0003723 RNA binding
IDA
PMID:19111658
HP1 proteins form distinct complexes and mediate heterochrom...
ACCEPT
Summary: Swi6 associates with noncoding centromeric transcripts (Clr4-dependent), consistent with RNA binding. Core MF (duplicate term, distinct reference).
Reason: Direct evidence Swi6 associates with heterochromatic RNAs.
Supporting Evidence:
PMID:19111658
Swi6 associates with a different set of nuclear proteins and with noncoding centromeric transcripts
GO:0005721 pericentric heterochromatin
IDA
PMID:19965387
Phosphorylation of H2A by Bub1 prevents chromosomal instabil...
ACCEPT
Summary: Direct localization of Swi6 to pericentric heterochromatin (shugoshin localization study). Core CC.
Reason: Direct localization evidence.
Supporting Evidence:
PMID:19965387
Bub1 phosphorylates the conserved serine 121 of histone H2A in fission yeast Schizosaccharomyces pombe.
GO:0031934 mating-type region heterochromatin
IDA
PMID:19965387
Phosphorylation of H2A by Bub1 prevents chromosomal instabil...
ACCEPT
Summary: Swi6 localizes to mating-type region heterochromatin. Core CC (duplicate term, distinct reference).
Reason: Direct localization evidence at mat-region heterochromatin.
Supporting Evidence:
PMID:11780129
Cohesin is also enriched at the mating-type heterochromatic region in a manner that depends on Swi6 and is required to preserve the genomic integrity of this locus.
GO:0005515 protein binding
IPI
PMID:29136238
Regulation of transcriptional silencing and chromodomain pro...
MARK AS OVER ANNOTATED
Summary: Bare "protein binding"; this study concerns H3Y41 phosphorylation regulating Swi6/Chp1 localization at centromeric heterochromatin rather than a discrete informative binding partner. Uninformative MF.
Reason: Uninformative MF.
Supporting Evidence:
PMID:29136238
it promotes the binding of Chp1 to histone H3 and the exclusion of Swi6.
GO:0000792 heterochromatin
IDA
PMID:26438724
H3K9 methylation extends across natural boundaries of hetero...
ACCEPT
Summary: Direct evidence Swi6 is a dynamic component of heterochromatin. Core CC (generic heterochromatin term).
Reason: Direct localization in heterochromatin.
Supporting Evidence:
PMID:26438724
the fission yeast HP1 protein Swi6 exists as a single highly dynamic population that rapidly exchanges in cis and in trans between different heterochromatic regions.
GO:0033696 heterochromatin boundary formation
IMP
PMID:26438724
H3K9 methylation extends across natural boundaries of hetero...
ACCEPT
Summary: Swi6 dimerization demarcates constitutive heterochromatin from neighboring euchromatin and confines its spread, a genuine boundary function. Core process.
Reason: Strong evidence for a heterochromatin demarcation/confinement role.
Supporting Evidence:
PMID:26438724
a surprising role for Swi6 dimerization in demarcating constitutive heterochromatin from neighboring euchromatin.
GO:0044820 mitotic telomere tethering at nuclear periphery
IMP
PMID:25778919
Aurora B prevents chromosome arm separation defects by promo...
KEEP AS NON CORE
Summary: Swi6/HP1 (with cohesin Rad21) is associated with telomere clustering; Aurora B-driven telomere dispersion involves dissociation of Swi6 from telomeres. Genuine but specialized telomere-organization role; non-core relative to heterochromatin silencing.
Reason: Genuine telomere-organization role but a peripheral context.
Supporting Evidence:
PMID:25778919
Dispersion is promoted by the dissociation of Swi6/HP1 and cohesin Rad21 from telomeres
GO:0006325 chromatin organization
IMP
PMID:9710635
A novel function of the DNA repair gene rhp6 in mating-type ...
KEEP AS NON CORE
Summary: Swi6 contributes to chromatin remodeling/organization underlying mating-type silencing. Correct but high-level; specific heterochromatin terms are preferred.
Reason: Correct but generic.
Supporting Evidence:
PMID:9710635
a similar silencing mechanism may operate in the distantly related yeast Schizosaccharomyces pombe.
GO:0000785 chromatin
IDA
PMID:21151114
Dicer associates with chromatin to repress genome activity i...
KEEP AS NON CORE
Summary: Swi6 is chromatin-associated; correct but generic relative to the heterochromatin subdomain terms.
Reason: Correct but generic CC.
Supporting Evidence:
PMID:21151114
the RNAi proteins Dcr1 and Rdp1 physically associate with some euchromatic genes, noncoding RNA genes and retrotransposon long terminal repeats
GO:0000792 heterochromatin
IDA
PMID:11069763
Live analysis of lagging chromosomes during anaphase and the...
ACCEPT
Summary: GFP-Swi6 is concentrated at heterochromatic centromeres and telomeres. Core CC (generic heterochromatin term).
Reason: Direct localization in heterochromatin.
Supporting Evidence:
PMID:11069763
GFP-Swi6p localises to the nucleus and is concentrated at the heterochromatic centromeres and telomeres.
GO:0000779 condensed chromosome, centromeric region
IDA
PMID:7660126
The chromodomain protein Swi6: a key component at fission ye...
ACCEPT
Summary: Swi6 localizes to centromeres and is required for their proper function during mitosis. Core CC.
Reason: Direct localization at centromeres.
Supporting Evidence:
PMID:7660126
Thus, Swi6p is located at fission yeast centromeres and is required for their proper function.
GO:0031934 mating-type region heterochromatin
IDA
PMID:7660126
The chromodomain protein Swi6: a key component at fission ye...
ACCEPT
Summary: Swi6 localizes to the silent mating-type loci heterochromatin. Core CC.
Reason: Direct localization at mat-region heterochromatin.
Supporting Evidence:
PMID:7660126
Swi6p localizes with these three chromosomal regions.
GO:0140720 subtelomeric heterochromatin
IDA
PMID:7660126
The chromodomain protein Swi6: a key component at fission ye...
ACCEPT
Summary: Swi6 localizes to telomeric/subtelomeric heterochromatin. Core CC.
Reason: Direct localization at subtelomeric heterochromatin.
Supporting Evidence:
PMID:7660126
the silent mating-type loci, centromeres, and telomeres are assembled into silent heterochromatin-like domains.
GO:0005721 pericentric heterochromatin
IDA
PMID:10766735
Distinct protein interaction domains and protein spreading i...
ACCEPT
Summary: Direct ChIP mapping shows Swi6 confined to (and spreading across) pericentromeric outer repeats. Core CC.
Reason: Direct localization at pericentric repeats.
Supporting Evidence:
PMID:10766735
Swi6 and Chp1 are confined to the flanking outer repeats and Swi6 can spread across at least 3 kb of extraneous chromatin in cen1.
GO:0005721 pericentric heterochromatin
IDA
PMID:11553715
The domain structure of centromeres is conserved from fissio...
ACCEPT
Summary: Swi6 binds the pericentromeric repeats flanking the central core, consistent with conserved centromere domain organization. Core CC.
Reason: Direct localization at pericentromeric repeats.
Supporting Evidence:
PMID:11553715
the Swi6 protein binds the surrounding repeats.
GO:0005721 pericentric heterochromatin
IDA
PMID:20299449
A chromodomain switch mediated by histone H3 Lys 4 acetylati...
ACCEPT
Summary: Swi6 occupies pericentric heterochromatin during the inactive state, regulated by an H3K4ac-mediated chromodomain switch. Core CC.
Reason: Direct localization at pericentric heterochromatin.
Supporting Evidence:
PMID:20299449
Chp2/Swi6 (HP1 homologs) are recruited during the inactive state.
GO:0007533 mating type switching
IMP
PMID:1620099
The switching gene swi6 affects recombination and gene expre...
ACCEPT
Summary: swi6 mutation reduces mating-type switching and disrupts the chromatin structure of the mating-type region. Core process.
Reason: Classic mutant evidence for swi6 in mating-type switching.
Supporting Evidence:
PMID:1620099
Besides reducing MT switching, the swi6 mutation leads to deletions in the MT region caused by intrachromosomal cross-overs between two paired cassettes.
GO:0030466 silent mating-type cassette heterochromatin formation
IMP
PMID:1620099
The switching gene swi6 affects recombination and gene expre...
ACCEPT
Summary: swi6 is required for repression/heterochromatin of the silent mating-type cassettes; loss permits expression of both cassettes and recombination in the normally inert K region. Core process.
Reason: Mutant evidence for silent mat-cassette heterochromatin.
Supporting Evidence:
PMID:1620099
swi6 also allows the simultaneous expression of two different cassettes in the same haploid cell.
GO:0030466 silent mating-type cassette heterochromatin formation
IMP
PMID:8138176
Mutations in rik1, clr2, clr3 and clr4 genes asymmetrically ...
ACCEPT
Summary: swi6 (with clr1) derepresses the silent mating-type loci and allows K- region recombination when mutated, supporting its role in silent cassette heterochromatin. Core process (duplicate term, distinct reference).
Reason: Mutant evidence for silent mat-cassette heterochromatin.
Supporting Evidence:
PMID:8138176
mutations in two genes, clr1 and swi6, had been shown to allow both expression of the silent loci and recombination in the K region.
GO:0062072 histone H3K9me2/3 reader activity
IDA
PMID:22727667
Intrinsic nucleic acid-binding activity of Chp1 chromodomain...
ACCEPT
Summary: This reference characterizes the Chp1 chromodomain's nucleic-acid binding enhanced by H3K9me; it concerns the chromodomain protein family context. Swi6 H3K9me reader activity is firmly established by other references (PMID:11242054, PMID:22683269), so the term is correct for Swi6 even though this particular citation centers on Chp1.
Reason: H3K9me reader activity is well established for Swi6 (core MF).
Supporting Evidence:
PMID:11242054
The chromo domain of HP1 is identified as its methyl-lysine-binding domain.
GO:0007534 gene conversion at mating-type locus
IMP
PMID:6587363
Genes required for initiation and resolution steps of mating...
ACCEPT
Summary: swi6 belongs to the group of switching genes affecting the efficient utilization of the mat1 cut and the directionality of switching (gene conversion). Core process.
Reason: Classic genetic evidence placing swi6 in mat switching/conversion.
Supporting Evidence:
PMID:6587363
The remaining three ( swi2 , -5, -6) have normal levels of cut, do not make errors of resolution, and possibly are required either for efficient utilization of the cut or determining the directionality of switching.
GO:0003723 RNA binding
IDA
PMID:22683269
HP1(Swi6) mediates the recognition and destruction of hetero...
ACCEPT
Summary: Swi6 RNA binding via the hinge (duplicate of the RNA-binding annotation from the same reference). Core MF.
Reason: Direct evidence for Swi6 RNA binding.
Supporting Evidence:
PMID:22683269
Stimulated by positively charged residues in the hinge region, RNA competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
GO:0005721 pericentric heterochromatin
IDA
PMID:20929775
Two histone marks establish the inner centromere and chromos...
ACCEPT
Summary: Swi6 localizes to pericentric heterochromatin (inner-centromere/ shugoshin study). Core CC.
Reason: Direct localization at pericentric heterochromatin.
Supporting Evidence:
PMID:20929775
phosphorylated H2A-S121 facilitates the binding of shugoshin, the centromeric CPC adaptor.
GO:0031934 mating-type region heterochromatin
IDA
PMID:20929775
Two histone marks establish the inner centromere and chromos...
ACCEPT
Summary: Swi6 localizes to mating-type region heterochromatin. Core CC (duplicate term, distinct reference).
Reason: Direct localization at mat-region heterochromatin.
Supporting Evidence:
PMID:20929775
the inner centromere is defined by intersection of two histone kinases.
GO:0140720 subtelomeric heterochromatin
IDA
PMID:20929775
Two histone marks establish the inner centromere and chromos...
ACCEPT
Summary: Swi6 localizes to subtelomeric heterochromatin. Core CC (duplicate term, distinct reference).
Reason: Direct localization at subtelomeric heterochromatin.
Supporting Evidence:
PMID:20929775
For proper partitioning of chromosomes in mitosis, the chromosomal passenger complex (CPC) including Aurora B and survivin must be localized at the center of paired kinetochores
GO:0005515 protein binding
IPI
PMID:20929775
Two histone marks establish the inner centromere and chromos...
MARK AS OVER ANNOTATED
Summary: Bare "protein binding" (interaction with Hrk1/Haspin context). The informative MF is Swi6's chromatin-protein adaptor activity; the generic term is uninformative.
Reason: Uninformative MF.
Supporting Evidence:
PMID:20929775
Haspin colocalizes with cohesin by associating with Pds5, whereas Bub1 localizes at kinetochores.

Core Functions

Reads the H3K9me2/me3 heterochromatin mark via its chromodomain, providing the recognition step that targets Swi6 to Clr4-methylated nucleosomes.

Supporting Evidence:
  • PMID:11242054
    HP1 can bind with high affinity to histone H3 methylated at lysine 9 but not at lysine 4. The chromo domain of HP1 is identified as its methyl-lysine-binding domain.

Homodimerizes/oligomerizes via its chromoshadow domain to form a spreading-competent, chromatin-compacting platform and protein-interaction pit.

Supporting Evidence:
  • PMID:10801440
    Dimerisation through the CSD of HP1-like proteins results in the simultaneous formation of a putative protein-protein interaction pit
  • PMID:23485968
    binding of the key S. pombe HP1 protein, Swi6, to methylated nucleosomes drives a switch from an auto-inhibited state to a spreading-competent state.

Acts as a chromatin-protein adaptor/scaffold, recruiting effectors to heterochromatin including cohesin (Psc3), the JmjC protein Epe1, the RNAi bridge Ers1/RDRC, the switching factor Swi2, and shugoshin Sgo1.

Supporting Evidence:
  • PMID:11780129
    a cohesin subunit Psc3 interacts with Swi6 and its mouse homologue HP1.
  • PMID:16762840
    Epe1 is recruited to heterochromatic loci by the heterochromatin protein Swi6/HP1.

Mediates transcriptional and co-transcriptional gene silencing and assembly, demarcation and confinement of heterochromatin at centromeres, telomeres, the silent mating-type cassettes and rDNA.

Supporting Evidence:
  • PMID:26438724
    a surprising role for Swi6 dimerization in demarcating constitutive heterochromatin from neighboring euchromatin.

Enriches cohesin at the pericentromere to ensure centromeric sister- chromatid cohesion, biorientation and accurate chromosome segregation.

Supporting Evidence:
  • PMID:31000521
    Swi6 is required for cohesin enrichment at the pericentromere.

Required for mating-type switching, including correct donor selection (directionality), by maintaining silent-cassette heterochromatin and recruiting the switching factor Swi2.

Supporting Evidence:
  • PMID:1620099
    Besides reducing MT switching, the swi6 mutation leads to deletions in the MT region caused by intrachromosomal cross-overs between two paired cassettes.
  • PMID:36951094
    the Swi6-binding site was required for Swi2 localization at SRE2 to select mat2-P in M cells.

Binds RNA via its basic hinge, sequestering and priming heterochromatic transcripts for degradation in competition with H3K9me chromodomain binding.

Supporting Evidence:
  • PMID:22683269
    the fission yeast HP1(Swi6) protein guarantees tight repression of heterochromatic genes through RNA sequestration and degradation.

References

file:interpro/panther/PTHR22812/PTHR22812-review.md
PANTHER family review PTHR22812: IBA propagation assessment for swi6
Annotation inferences using phylogenetic trees
Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
Electronic Gene Ontology annotations created by ARBA machine learning models
Combined Automated Annotation using Multiple IEA Methods
Distinct protein interaction domains and protein spreading in a complex centromere.
  • Swi6 is confined to the pericentromeric flanking outer repeats and can spread across at least 3 kb of chromatin in cen1.
    "Swi6 and Chp1 are confined to the flanking outer repeats and Swi6 can spread across at least 3 kb of extraneous chromatin in cen1."
Dimerisation of a chromo shadow domain and distinctions from the chromodomain as revealed by structural analysis.
  • The Swi6 chromoshadow domain forms a dimer creating a protein-interaction pit for targeting effectors to chromatin.
    "Dimerisation through the CSD of HP1-like proteins results in the simultaneous formation of a putative protein-protein interaction pit"
Live analysis of lagging chromosomes during anaphase and their effect on spindle elongation rate in fission yeast.
  • GFP-Swi6 localizes to the nucleus and concentrates at centromeres and telomeres.
    "GFP-Swi6p localises to the nucleus and is concentrated at the heterochromatic centromeres and telomeres."
Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo domain.
  • The HP1/Swi6 chromodomain is the methyl-lysine-binding domain selective for H3K9me.
    "HP1 can bind with high affinity to histone H3 methylated at lysine 9 but not at lysine 4. The chromo domain of HP1 is identified as its methyl-lysine-binding domain."
Role of histone H3 lysine 9 methylation in epigenetic control of heterochromatin assembly.
  • Swi6 localization to heterochromatin depends on Clr4-mediated H3K9 methylation.
    "Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic regions is dependent on H3 Lys9 methylation."
The domain structure of centromeres is conserved from fission yeast to humans.
  • Swi6 binds the pericentromeric repeats flanking the central core.
    "the Swi6 protein binds the surrounding repeats."
Recruitment of cohesin to heterochromatic regions by Swi6/HP1 in fission yeast.
  • Swi6 recruits cohesin to centromeric and mating-type heterochromatin via Psc3.
    "a cohesin subunit Psc3 interacts with Swi6 and its mouse homologue HP1."
Regulation of heterochromatic silencing and histone H3 lysine-9 methylation by RNAi.
  • RNAi from centromeric repeats targets heterochromatin formation in which Swi6 acts.
    "We propose that double-stranded RNA arising from centromeric repeats targets formation and maintenance of heterochromatin through RNAi."
Hsk1-Dfp1 is required for heterochromatin-mediated cohesion at centromeres.
Two different Swi5-containing protein complexes are involved in mating-type switching and recombination repair in fission yeast.
Regulation of Swi6/HP1-dependent heterochromatin assembly by cooperation of components of the mitogen-activated protein kinase pathway and a histone deacetylase Clr6.
  • A Swi6-dependent pathway assembles heterochromatin across the silent mating-type region.
    "a stochastic Swi6 (the fission yeast HP1 homolog)-dependent mechanism that is not fully understood."
A chromodomain protein, Chp1, is required for the establishment of heterochromatin in fission yeast.
  • Swi6 (with Chp2) maintains residual H3K9 methylation at centromeres, mat region and telomeres.
    "Swi6 and Chp2 are critical to maintaining this residual methylation."
The nucleation and maintenance of heterochromatin by a histone deacetylase in fission yeast.
  • Clr3 spreading across the silent mat domain requires heterochromatin proteins including Swi6/HP1.
    "Clr3 spreads across the 20 kb silenced domain that requires its own HDAC activity and heterochromatin proteins including Swi6/HP1."
Swi6/HP1 recruits a JmjC domain protein to facilitate transcription of heterochromatic repeats.
  • Swi6/HP1 recruits the anti-silencing JmjC protein Epe1 to heterochromatin.
    "Epe1 is recruited to heterochromatic loci by the heterochromatin protein Swi6/HP1."
Interaction of Epe1 with the heterochromatin assembly pathway in Schizosaccharomyces pombe.
  • Swi6 recruits Epe1 to heterochromatin.
    "We report that Swi6 recruits Epe1 to heterochromatin"
RNAi-dependent and -independent RNA turnover mechanisms contribute to heterochromatic gene silencing.
  • RNAi is required for heterochromatin-dependent silencing at centromeric repeats.
    "the RNAi pathway is required for heterochromatin-dependent silencing of transgene insertions at centromeric repeats"
Heterochromatin links to centromeric protection by recruiting shugoshin.
  • Swi6 directly recruits shugoshin Sgo1 to protect centromeric cohesin in meiosis.
    "the heterochromatin protein Swi6 associates directly with meiosis-specific shugoshin Sgo1, a protector of cohesin at centromeres."
Fission yeast chromatin assembly factor 1 assists in the replication-coupled maintenance of heterochromatin.
Balance between distinct HP1 family proteins controls heterochromatin assembly in fission yeast.
  • Swi6 and Chp2 play distinct, dose-dependent roles in assembling repressive heterochromatin.
    "both Swi6 and Chp2 are required for the assembly of fully repressive heterochromatin"
HP1 proteins form distinct complexes and mediate heterochromatic gene silencing by nonoverlapping mechanisms.
  • Swi6 associates with noncoding centromeric transcripts and is required for RNAi processing into siRNAs.
    "Swi6 associates with a different set of nuclear proteins and with noncoding centromeric transcripts and is required for efficient RNAi-dependent processing of these transcripts."
Phosphorylation of Swi6/HP1 regulates transcriptional gene silencing at heterochromatin.
  • Swi6 recruits SHREC, Epe1 and cohesin; CK2 phosphorylation of Swi6 controls TGS.
    "a HP1 homolog Swi6 recruits SHREC, Epe1, and cohesin, which are involved in transcriptional gene silencing (TGS), transcriptional activation, and sister chromatid cohesion, respectively."
Diverse roles of HP1 proteins in heterochromatin assembly and functions in fission yeast.
  • Swi6 and Chp2 with HDAC complexes limit Pol II occupancy across pericentromeric heterochromatin.
    "Swi6 and Chp2 proteins and their associated HDAC complexes have overlapping functions in limiting Pol II occupancy across pericentromeric heterochromatin domains."
Phosphorylation of H2A by Bub1 prevents chromosomal instability through localizing shugoshin.
A chromodomain switch mediated by histone H3 Lys 4 acetylation regulates heterochromatin assembly.
  • Chp2/Swi6 are recruited to pericentric heterochromatin during the inactive state.
    "Chp2/Swi6 (HP1 homologs) are recruited during the inactive state."
Two histone marks establish the inner centromere and chromosome bi-orientation.
Dicer associates with chromatin to repress genome activity in Schizosaccharomyces pombe.
Asf1/HIRA facilitate global histone deacetylation and associate with HP1 to promote nucleosome occupancy at heterochromatic loci.
  • An Asf1/HIRA histone chaperone spreads across silenced domains via association with Swi6.
    "a histone chaperone complex containing Asf1 and HIRA that spreads across silenced domains via its association with Swi6 to enforce transcriptional silencing."
RNA elimination machinery targeting meiotic mRNAs promotes facultative heterochromatin formation.
  • Facultative heterochromatin islands form at meiotic genes via RNA elimination machinery.
    "we describe facultative heterochromatin islands in fission yeast and show that their formation at meiotic genes requires factors that eliminate meiotic messenger RNAs (mRNAs) during vegetative growth."
Heterochromatin protein 1 homologue Swi6 acts in concert with Ers1 to regulate RNAi-directed heterochromatin assembly.
  • Swi6 recruits RDRC to heterochromatin through the RNAi intermediate Ers1.
    "we demonstrate that Swi6 recruits RDRC to heterochromatin through Ers1, an RNAi factor intermediate."
HP1(Swi6) mediates the recognition and destruction of heterochromatic RNA transcripts.
  • Swi6 binds RNA via its hinge; RNA competes with H3K9me for the chromodomain.
    "Stimulated by positively charged residues in the hinge region, RNA competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6)."
Intrinsic nucleic acid-binding activity of Chp1 chromodomain is required for heterochromatic gene silencing.
Hrp3 controls nucleosome positioning to suppress non-coding transcription in eu- and heterochromatin.
A conformational switch in HP1 releases auto-inhibition to drive heterochromatin assembly.
  • Swi6 binding to methylated nucleosomes switches it from auto-inhibited to spreading-competent oligomers.
    "binding of the key S. pombe HP1 protein, Swi6, to methylated nucleosomes drives a switch from an auto-inhibited state to a spreading-competent state."
Tls1 regulates splicing of shelterin components to control telomeric heterochromatin assembly and telomere length.
  • Telomeric heterochromatin assembly requires multiple factors including Tls1.
    "we comprehensively identified factors required for telomeric heterochromatin assembly, including a novel gene tls1+."
Aurora B prevents chromosome arm separation defects by promoting telomere dispersion and disjunction.
  • Telomere dispersion is promoted by dissociation of Swi6/HP1 and cohesin Rad21 from telomeres.
    "Dispersion is promoted by the dissociation of Swi6/HP1 and cohesin Rad21 from telomeres"
H3K9 methylation extends across natural boundaries of heterochromatin in the absence of an HP1 protein.
  • Swi6 dimerization demarcates and confines constitutive heterochromatin rather than promoting its spread.
    "a surprising role for Swi6 dimerization in demarcating constitutive heterochromatin from neighboring euchromatin."
Regulation of transcriptional silencing and chromodomain protein localization at centromeric heterochromatin by histone H3 tyrosine 41 phosphorylation in fission yeast.
New insights into donor directionality of mating-type switching in Schizosaccharomyces pombe.
  • Mating-type switching donor selection is regulated by heterochromatin and the HP1-like protein Swi6.
    "Mating-type switching in Schizosaccharomyces pombe entails programmed gene conversion events regulated by DNA replication, heterochromatin, and the HP1-like chromodomain protein Swi6."
Overlapping Roles in Chromosome Segregation for Heterochromatin Protein 1 (Swi6) and DDK in Schizosaccharomyces pombe.
  • Swi6 is required for cohesin enrichment at the pericentromere and for sister-kinetochore biorientation.
    "Swi6 is required for cohesin enrichment at the pericentromere."
Fission yeast Swi2 designates cell-type specific donor and stimulates Rad51-driven strand exchange for mating-type switching.
  • A Swi6-binding site in Swi2 is required for Swi2 localization at SRE2 to select the mat2-P donor.
    "the Swi6-binding site was required for Swi2 localization at SRE2 to select mat2-P in M cells."
Cooperative DNA-binding activities of Chp2 are critical for its function in heterochromatin assembly.
  • This study characterizes cooperative DNA-binding by HP1 proteins in fission yeast. The cached abstract foregrounds Chp2, but PomBase used the full text to make a direct-assay (IDA) DNA-binding annotation to Swi6 (the other HP1 protein), retained here as a non-core, hinge-mediated property of Swi6.
    "Here, we focus on Chp2, one of the two HP1 proteins in fission yeast, and investigate how Chp2's DNA-binding ability contributes to its function."
PhpC(NF-Y) transcription factor infiltrates heterochromatin to generate cryptic intron-containing transcripts crucial for small RNA production.
  • TF-driven transcription of heterochromatic repeats produces siRNAs critical for heterochromatin nucleation.
    "siRNA production by this TF-mediated transcription pathway is critical for heterochromatin nucleation at target repeat loci."
Genes required for initiation and resolution steps of mating-type switching in fission yeast.
  • swi6 is required for efficient utilization of the mat1 cut and switching directionality.
    "The remaining three ( swi2 , -5, -6) have normal levels of cut, do not make errors of resolution, and possibly are required either for efficient utilization of the cut or determining the directionality of switching."
The chromodomain protein Swi6: a key component at fission yeast centromeres.
  • Swi6 localizes to centromeres, mating-type loci and telomeres and is required for centromere function.
    "Thus, Swi6p is located at fission yeast centromeres and is required for their proper function."
Mutations in rik1, clr2, clr3 and clr4 genes asymmetrically derepress the silent mating-type loci in fission yeast.
  • swi6 (with clr1) allows expression of the silent mating-type loci and K-region recombination when mutated.
    "mutations in two genes, clr1 and swi6, had been shown to allow both expression of the silent loci and recombination in the K region."
A novel function of the DNA repair gene rhp6 in mating-type silencing by chromatin remodeling in fission yeast.
The switching gene swi6 affects recombination and gene expression in the mating-type region of Schizosaccharomyces pombe.
  • swi6 mutation reduces mating-type switching and derepresses the silent cassettes, allowing K-region recombination.
    "swi6 also allows the simultaneous expression of two different cassettes in the same haploid cell."

Suggested Questions for Experts

Q: What determines the partitioning of effector recruitment between the two fission yeast HP1 proteins (Swi6 vs Chp2), and how is Swi6's bifunctional chromatin- and RNA-binding balanced in vivo?

Q: How does Swi6 oligomerization/phase behavior on H3K9me chromatin quantitatively contribute to silencing versus boundary confinement at different loci?

Suggested Experiments

Experiment: Separation-of-function mutants (chromodomain reader-dead vs CSD dimer-dead vs hinge RNA/DNA-binding-dead) assayed in parallel for silencing, cohesin enrichment, donor selection, and boundary confinement to dissect which core function drives each phenotype.

Experiment: Quantitative ChIP-seq/CUT&RUN of Swi6 and its recruited effectors (Psc3, Clr3/SHREC, Epe1, Ers1) in wild-type and Swi6 phospho-site mutants to map how post-translational modification reprograms the adaptor interactome.

📚 Additional Documentation

Notes

(swi6-notes.md)

swi6 (S. pombe) curation notes

Swi6 / P40381 / SPAC664.01c — the principal HP1 (heterochromatin protein 1) ortholog of fission yeast.
328 aa, chromodomain (CD, ~81-143) + chromoshadow domain (CSD, ~267-328) connected by a disordered hinge.

Domain architecture & molecular function

  • CD binds H3K9me2/3 (written by Clr4); CSD homodimerizes and is a protein-protein interaction platform.
  • PMID:11242054 — H3K9me reader = CD.
  • PMID:11242054
  • PMID:10801440 and PMID:10801440 — CSD dimerization / adaptor platform; this is the homodimerization (identical protein binding) annotation.
  • PMID:23485968 and PMID:23485968. Self-association/oligomerization drives spreading. DimerX (L315D) disrupts CSD-CSD dimerization.
  • RNA binding: PMID:22683269; PMID:22683269 Hinge basic patch binds RNA.
  • DNA binding: hinge binds DNA sequence-independently (PMID:23485968). Note GOA DNA-binding annotation PMID:37400983 is actually a Chp2 paper; treat with caution (see below).

Chromatin-protein adaptor activity (scaffold/effector recruitment)

Localization / cellular component

Biological process

  • Heterochromatin/silencing at centromere, telomere, mat locus, rDNA.
  • Pericentric het formation: PMID:19443688, PMID:19136623, PMID:31000521 (cohesin enrichment, biorientation, segregation).
  • Cohesion/segregation: PMID:31000521; PMID:31000521
  • Mat switching / donor selection / gene conversion: PMID:1620099, PMID:6587363, PMID:29852001, PMID:36951094. PMID:29852001; donor directionality — in absence of Swi6 cells prefer mat3-M PMID:36951094.
  • RNAi / co-transcriptional silencing: PMID:17512405, PMID:12193640, PMID:22474355, PMID:19111658. PMID:19111658 and PMID:19111658
  • siRNA-mediated mat-cassette het formation: PMID:39747188 (TF-driven transcription → siRNA → nucleation).
  • Heterochromatin boundary formation: PMID:21211723 (Asf1/HIRA), PMID:26438724. Surprising boundary/demarcation role: PMID:26438724; PMID:26438724
  • Telomere tethering: PMID:25778919 (Aurora B / telomere dispersion); subtelomeric het via Tls1 splicing PMID:25245948.
  • Chromatin organization: PMID:9710635 (rhp6 mat silencing), PMID:19111658.

Caveats / over-annotation watch

  • "protein binding" (GO:0005515) IPI rows: uninformative; the informative MF is chromatin-protein adaptor activity / homodimerization / H3K9me reader. MARK_AS_OVER_ANNOTATED for bare protein binding, but note the interactor.
  • PMID:37400983 "Cooperative DNA-binding activities of Chp2..." — this paper is about Chp2, not Swi6; the GO:0003677 DNA binding annotation to swi6 may be mis-attributed or a comparative point. Swi6 hinge does bind DNA (PMID:23485968) so DNA binding is plausible MF but the cited reference is about a paralog → UNDECIDED/keep cautiously.
  • Heterochromatin "maintenance/spreading" should be tempered: Swi6 is largely dispensable for H3K9me maintenance (PMID:26438724); its core is reading H3K9me + recruiting effectors + confining/demarcating heterochromatin + compaction via oligomerization.

Core function synthesis

MF: H3K9me2/3 reader (chromodomain); chromatin-protein adaptor/scaffold (chromoshadow); protein homodimerization; (RNA binding via hinge).
BP: heterochromatin assembly & transcriptional gene silencing at centromere/telomere/mat/rDNA; cohesin enrichment → sister chromatid cohesion & accurate chromosome segregation; mating-type switching (donor selection); heterochromatin boundary/confinement.
CC: pericentric, mating-type region, subtelomeric heterochromatin; nucleus/chromatin.

📄 View Raw YAML

id: P40381
gene_symbol: swi6
product_type: PROTEIN
status: DRAFT
taxon:
  id: NCBITaxon:284812
  label: Schizosaccharomyces pombe (strain 972 / ATCC 24843)
description: >-
  Swi6 is the principal HP1 (heterochromatin protein 1) ortholog of fission
  yeast and the central structural reader and effector of heterochromatin. It is
  a 328-residue protein built from an N-terminal chromodomain that binds histone
  H3 methylated at lysine 9 (H3K9me2/me3, deposited by the Clr4 methyltransferase)
  and a C-terminal chromoshadow domain that homodimerizes and forms a
  protein-protein interaction platform, the two domains connected by a basic
  disordered hinge with nucleic-acid-binding activity. Through this architecture
  Swi6 recognizes H3K9-methylated nucleosomes, oligomerizes, and undergoes a
  switch from an auto-inhibited to a spreading-competent state that compacts
  chromatin. The chromoshadow platform recruits diverse effectors to
  heterochromatin, including the cohesin subunit Psc3, the SHREC/Clr3 histone
  deacetylase machinery, the anti-silencing JmjC protein Epe1, the RNAi bridging
  factor Ers1/RDRC, the histone chaperones Asf1/HIRA and CAF-1, DBF4-dependent
  kinase (Dfp1), the mating-type switching factor Swi2, and meiotic shugoshin
  Sgo1. Swi6 mediates transcriptional and co-transcriptional gene silencing and
  the assembly, demarcation and confinement of heterochromatin at centromeres,
  telomeres, the silent mating-type cassettes, and rDNA. At centromeres it
  enriches cohesin to ensure accurate sister-chromatid cohesion, biorientation
  and chromosome segregation; at the mating-type region it is required for donor
  selection during mating-type switching; and at telomeres it contributes to
  subtelomeric heterochromatin and telomere tethering.
existing_annotations:
- term:
    id: GO:0031507
    label: heterochromatin formation
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: involved_in
  review:
    summary: >-
      Heterochromatin formation/assembly is a core biological process of Swi6,
      conserved across the HP1 family. Phylogenetic inference is corroborated by
      extensive S. pombe experimental evidence.
    action: ACCEPT
    reason: Core HP1 function, supported by phylogeny and abundant direct evidence.
    supported_by:
    - reference_id: PMID:11283354
      supporting_text: >-
        Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic
        regions is dependent on H3 Lys9 methylation.
      reference_section_type: ABSTRACT
- term:
    id: GO:0003682
    label: chromatin binding
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: enables
  review:
    summary: >-
      Chromatin binding is correct but generic; the informative molecular
      function is H3K9me2/3 reader activity via the chromodomain. Retained as a
      true but non-core (parent) descriptor.
    action: KEEP_AS_NON_CORE
    reason: True but superseded by the more specific H3K9me reader activity term.
    supported_by:
    - reference_id: PMID:11242054
      supporting_text: >-
        The chromo domain of HP1 is identified as its methyl-lysine-binding domain.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  qualifier: is_active_in
  review:
    summary: >-
      Pericentric heterochromatin is a core site of Swi6 action; this is
      strongly corroborated by direct localization data (multiple IDA
      annotations below).
    action: ACCEPT
    reason: Phylogenetic call confirmed by direct localization evidence.
    supported_by:
    - reference_id: PMID:10766735
      supporting_text: >-
        Swi6 and Chp1 are confined to the flanking outer repeats and Swi6 can
        spread across at least 3 kb of extraneous chromatin in cen1.
      reference_section_type: ABSTRACT
- term:
    id: GO:0000792
    label: heterochromatin
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  qualifier: located_in
  review:
    summary: >-
      Heterochromatin localization is correct and strongly supported by direct
      IDA evidence; Swi6 is a defining component of heterochromatin. Accepted to
      remain consistent with the IDA annotations to this term.
    action: ACCEPT
    reason: Correct and core localization, corroborated by direct evidence.
    supported_by:
    - reference_id: PMID:11069763
      supporting_text: >-
        GFP-Swi6p localises to the nucleus and is concentrated at the
        heterochromatic centromeres and telomeres.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  qualifier: located_in
  review:
    summary: >-
      Swi6 is a nuclear protein. Correct but generic; the heterochromatin
      subdomain terms are the informative localizations.
    action: KEEP_AS_NON_CORE
    reason: Correct broad localization, subsumed by specific nuclear terms.
    supported_by:
    - reference_id: PMID:11069763
      supporting_text: >-
        GFP-Swi6p localises to the nucleus and is concentrated at the
        heterochromatic centromeres and telomeres.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005694
    label: chromosome
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  qualifier: located_in
  review:
    summary: >-
      Chromosome localization is correct (Swi6 is chromatin-associated) but
      generic relative to the heterochromatin and chromatin terms.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic.
    supported_by:
    - reference_id: PMID:7660126
      supporting_text: >-
        Swi6p localizes with these three chromosomal regions.
      reference_section_type: ABSTRACT
- term:
    id: GO:0040029
    label: epigenetic regulation of gene expression
  evidence_type: IEA
  original_reference_id: GO_REF:0000117
  qualifier: involved_in
  review:
    summary: >-
      Swi6 mediates epigenetic gene silencing through H3K9me-based
      heterochromatin. Correct but high-level; the specific silencing/
      heterochromatin formation terms are preferred.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic parent of the specific silencing processes.
    supported_by:
    - reference_id: PMID:11242054
      supporting_text: >-
        This model may also explain the stable inheritance of the
        heterochromatic state.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:14625560
  qualifier: enables
  review:
    summary: >-
      Bare "protein binding" (here the interaction with the DDK kinase subunit
      Hsk1) is uninformative. The informative molecular function captured by
      this interaction is Swi6's chromatin-protein adaptor/scaffold activity
      recruiting DDK to pericentromeric heterochromatin.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative MF; superseded by chromatin-protein adaptor activity.
    supported_by:
    - reference_id: PMID:31000521
      supporting_text: >-
        Swi6 interacts with Dfp1, a regulatory subunit of DBF4-dependent kinase (DDK)
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18716626
  qualifier: enables
  review:
    summary: >-
      The Swi6-Sgo1 interaction is real and biologically important (meiotic
      cohesion protection), but "protein binding" is uninformative. The function
      is chromatin-protein adaptor activity (shugoshin recruitment).
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative MF; the informative term is adaptor activity.
    supported_by:
    - reference_id: PMID:18716626
      supporting_text: >-
        the heterochromatin protein Swi6 associates directly with
        meiosis-specific shugoshin Sgo1, a protector of cohesin at centromeres.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22474355
  qualifier: enables
  review:
    summary: >-
      The Swi6-Ers1 interaction bridges Swi6 to the RDRC/RNAi machinery, but
      "protein binding" is uninformative. The informative MF is chromatin-protein
      adaptor activity (also annotated separately from this reference).
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative MF; captured better by adaptor activity.
    supported_by:
    - reference_id: PMID:22474355
      supporting_text: >-
        Swi6 recruits RDRC to heterochromatin through Ers1, an RNAi factor intermediate.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22990236
  qualifier: enables
  review:
    summary: >-
      Interaction with the CHD chromatin remodeler Hrp3. "Protein binding" is
      uninformative; the informative MF is chromatin-protein adaptor activity.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative MF.
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:14663140
  qualifier: enables
  review:
    summary: >-
      Swi6 homodimerizes via its chromoshadow domain; this self-association is a
      core, informative property (distinct from bare protein binding) that
      underlies effector recruitment and heterochromatin spreading.
    action: ACCEPT
    reason: Homodimerization is a genuine, mechanistically central MF.
    supported_by:
    - reference_id: PMID:10801440
      supporting_text: >-
        The 1.9 A crystal structure of the Swi6 CSD is presented here. This
        reveals a novel dimeric structure
      reference_section_type: ABSTRACT
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:18716626
  qualifier: enables
  review:
    summary: >-
      Swi6 self-association (chromoshadow-domain homodimerization). Core MF;
      duplicate of the homodimerization annotation supported elsewhere.
    action: ACCEPT
    reason: Genuine homodimerization, central to HP1 function.
    supported_by:
    - reference_id: PMID:10801440
      supporting_text: >-
        Dimerisation through the CSD of HP1-like proteins results in the
        simultaneous formation of a putative protein-protein interaction pit
      reference_section_type: ABSTRACT
- term:
    id: GO:0042802
    label: identical protein binding
  evidence_type: IPI
  original_reference_id: PMID:23485968
  qualifier: enables
  review:
    summary: >-
      Swi6 self-associates into dimers (CSD-CSD) and higher-order oligomers; a
      conformational switch on methylated nucleosomes drives the spreading-
      competent oligomeric state. Core MF.
    action: ACCEPT
    reason: Self-association/oligomerization is mechanistically central.
    supported_by:
    - reference_id: PMID:23485968
      supporting_text: >-
        the CSD can homodimerize
      reference_section_type: RESULTS
- term:
    id: GO:0031507
    label: heterochromatin formation
  evidence_type: IDA
  original_reference_id: PMID:11780129
  qualifier: involved_in
  review:
    summary: >-
      Direct evidence that Swi6 is required for heterochromatin function at
      centromeres and the mating-type region. Core process.
    action: ACCEPT
    reason: Direct experimental support for heterochromatin formation role.
    supported_by:
    - reference_id: PMID:11780129
      supporting_text: >-
        the preferential localization of cohesins at the centromeric repeats is
        dependent on Swi6, a conserved heterochromatin protein that is required
        for proper kinetochore function.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140185
    label: siRNA-mediated silent mating type cassette region heterochromatin formation
  evidence_type: EXP
  original_reference_id: PMID:39747188
  qualifier: involved_in
  review:
    summary: >-
      Swi6 participates in siRNA/RNAi-based heterochromatin nucleation at the
      mating-type cassette region, where TF-driven transcription generates
      siRNAs that trigger heterochromatin assembly. Core process.
    action: ACCEPT
    reason: Experimental support for siRNA-mediated mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:39747188
      supporting_text: >-
        siRNA production by this TF-mediated transcription pathway is critical
        for heterochromatin nucleation at target repeat loci.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: EXP
  original_reference_id: PMID:18761674
  qualifier: located_in
  review:
    summary: >-
      Swi6 is a nuclear protein (CAF-1 study). Correct; generic relative to the
      heterochromatin subdomain localizations.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic localization.
- term:
    id: GO:0005694
    label: chromosome
  evidence_type: EXP
  original_reference_id: PMID:18761674
  qualifier: located_in
  review:
    summary: >-
      Swi6 is chromatin/chromosome-associated. Correct but generic relative to
      the heterochromatin terms.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic localization.
- term:
    id: GO:0140463
    label: chromatin-protein adaptor activity
  evidence_type: IPI
  original_reference_id: PMID:11780129
  qualifier: enables
  review:
    summary: >-
      Core molecular function. Swi6 bridges H3K9-methylated chromatin to cohesin
      via direct interaction with the cohesin subunit Psc3, enriching cohesin at
      heterochromatin. This is the informative MF underlying many "protein
      binding" interactions.
    action: ACCEPT
    reason: Direct adaptor activity (Psc3/cohesin recruitment); core function.
    supported_by:
    - reference_id: PMID:11780129
      supporting_text: >-
        a cohesin subunit Psc3 interacts with Swi6 and its mouse homologue HP1.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031508
    label: pericentric heterochromatin formation
  evidence_type: IMP
  original_reference_id: PMID:19443688
  qualifier: involved_in
  review:
    summary: >-
      Swi6 is required for pericentromeric heterochromatin/silencing, acting
      with associated HDAC complexes to limit Pol II occupancy. Core process.
    action: ACCEPT
    reason: Mutant phenotype supports pericentric heterochromatin formation role.
    supported_by:
    - reference_id: PMID:19443688
      supporting_text: >-
        Swi6 and Chp2 proteins and their associated HDAC complexes have
        overlapping functions in limiting Pol II occupancy across pericentromeric
        heterochromatin domains.
      reference_section_type: ABSTRACT
- term:
    id: GO:0007535
    label: donor selection
  evidence_type: IDA
  original_reference_id: PMID:29852001
  qualifier: involved_in
  review:
    summary: >-
      Swi6 is required for correct donor selection (directionality) during
      mating-type switching, cooperating with heterochromatin and Set1C to
      direct donor choice. Core process.
    action: ACCEPT
    reason: Direct evidence for Swi6 role in donor selection.
    supported_by:
    - reference_id: PMID:29852001
      supporting_text: >-
        Mating-type switching in Schizosaccharomyces pombe entails programmed
        gene conversion events regulated by DNA replication, heterochromatin,
        and the HP1-like chromodomain protein Swi6.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031508
    label: pericentric heterochromatin formation
  evidence_type: IMP
  original_reference_id: PMID:19136623
  qualifier: involved_in
  review:
    summary: >-
      Swi6 recruits SHREC, Epe1 and cohesin to pericentric heterochromatin, and
      CK2 phosphorylation of Swi6 controls transcriptional gene silencing there.
      Core process (duplicate term, distinct supporting reference).
    action: ACCEPT
    reason: Supports pericentric heterochromatin/silencing role.
    supported_by:
    - reference_id: PMID:19136623
      supporting_text: >-
        a HP1 homolog Swi6 recruits SHREC, Epe1, and cohesin, which are involved
        in transcriptional gene silencing (TGS), transcriptional activation, and
        sister chromatid cohesion, respectively.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031509
    label: subtelomeric heterochromatin formation
  evidence_type: IMP
  original_reference_id: PMID:25245948
  qualifier: involved_in
  review:
    summary: >-
      Swi6 contributes to telomeric/subtelomeric heterochromatin assembly (in
      this study examined via Tls1-dependent shelterin splicing). Core process.
    action: ACCEPT
    reason: Supports subtelomeric heterochromatin formation role.
    supported_by:
    - reference_id: PMID:25245948
      supporting_text: >-
        we comprehensively identified factors required for telomeric
        heterochromatin assembly, including a novel gene tls1+.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140463
    label: chromatin-protein adaptor activity
  evidence_type: IDA
  original_reference_id: PMID:36951094
  qualifier: enables
  review:
    summary: >-
      Swi6 acts as an adaptor recruiting the mating-type switching factor Swi2
      to a recombination enhancer via a defined Swi6-binding site, supporting
      donor selection. Core adaptor MF.
    action: ACCEPT
    reason: Direct evidence Swi6 recruits Swi2 (adaptor activity).
    supported_by:
    - reference_id: PMID:36951094
      supporting_text: >-
        the Swi6-binding site was required for Swi2 localization at SRE2 to
        select mat2-P in M cells.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:18809570
  qualifier: is_active_in
  review:
    summary: >-
      Direct localization of Swi6 to pericentric heterochromatin. Core CC
      (one of many concordant IDA annotations).
    action: ACCEPT
    reason: Direct localization evidence.
    supported_by:
    - reference_id: PMID:18809570
      supporting_text: >-
        both Swi6 and Chp2 are required for the assembly of fully repressive
        heterochromatin
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:18809570
  qualifier: is_active_in
  review:
    summary: >-
      Direct localization/activity of Swi6 at mating-type region
      heterochromatin. Core CC.
    action: ACCEPT
    reason: Direct evidence for activity at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:18809570
      supporting_text: >-
        two HP1 family proteins, Swi6 and Chp2, are involved in transcriptional
        silencing at heterochromatic regions
      reference_section_type: ABSTRACT
- term:
    id: GO:0140720
    label: subtelomeric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:18809570
  qualifier: is_active_in
  review:
    summary: >-
      Direct localization/activity of Swi6 at subtelomeric heterochromatin.
      Core CC.
    action: ACCEPT
    reason: Direct evidence for activity at subtelomeric heterochromatin.
    supported_by:
    - reference_id: PMID:18809570
      supporting_text: >-
        both Swi6 and Chp2 are required for the assembly of fully repressive heterochromatin
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:11780129
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to and acts at pericentric (centromeric repeat)
      heterochromatin, where it enriches cohesin. Core CC.
    action: ACCEPT
    reason: Direct localization at centromeric repeats.
    supported_by:
    - reference_id: PMID:11780129
      supporting_text: >-
        the preferential localization of cohesins at the centromeric repeats is
        dependent on Swi6
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:11780129
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 acts at mating-type region heterochromatin (cohesin enrichment
      preserving locus integrity). Core CC.
    action: ACCEPT
    reason: Direct evidence for activity at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:11780129
      supporting_text: >-
        Cohesin is also enriched at the mating-type heterochromatic region in a
        manner that depends on Swi6
      reference_section_type: ABSTRACT
- term:
    id: GO:0030466
    label: silent mating-type cassette heterochromatin formation
  evidence_type: EXP
  original_reference_id: PMID:15292231
  qualifier: involved_in
  review:
    summary: >-
      Swi6-dependent heterochromatin assembly across the silent mating-type
      region, downstream of Atf1/Pcr1- and Clr6-mediated histone deacetylation.
      Core process.
    action: ACCEPT
    reason: Experimental support for mat-cassette heterochromatin formation.
    supported_by:
    - reference_id: PMID:15292231
      supporting_text: >-
        a stochastic Swi6 (the fission yeast HP1 homolog)-dependent mechanism
        that is not fully understood.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:11069763
  qualifier: is_active_in
  review:
    summary: >-
      Direct (GFP) evidence for nuclear localization. Correct; generic relative
      to heterochromatin subdomains.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic localization.
    supported_by:
    - reference_id: PMID:11069763
      supporting_text: >-
        GFP-Swi6p localises to the nucleus and is concentrated at the
        heterochromatic centromeres and telomeres.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140463
    label: chromatin-protein adaptor activity
  evidence_type: IDA
  original_reference_id: PMID:22474355
  qualifier: enables
  review:
    summary: >-
      Swi6 functions as an adaptor recruiting the RDRC/RNAi machinery to
      heterochromatin through Ers1. Core adaptor MF (duplicate term, distinct
      reference).
    action: ACCEPT
    reason: Direct evidence Swi6 recruits RDRC via Ers1.
    supported_by:
    - reference_id: PMID:22474355
      supporting_text: >-
        we demonstrate that Swi6 recruits RDRC to heterochromatin through Ers1,
        an RNAi factor intermediate.
      reference_section_type: ABSTRACT
- term:
    id: GO:0062072
    label: histone H3K9me2/3 reader activity
  evidence_type: IDA
  original_reference_id: PMID:11242054
  qualifier: enables
  review:
    summary: >-
      Defining, core molecular function. The Swi6/HP1 chromodomain binds H3
      methylated at Lys9 (not Lys4) with high affinity; a chromodomain point
      mutation abolishes methyl-lysine binding and silencing.
    action: ACCEPT
    reason: Foundational direct demonstration of H3K9me reader activity.
    supported_by:
    - reference_id: PMID:11242054
      supporting_text: >-
        HP1 can bind with high affinity to histone H3 methylated at lysine 9 but
        not at lysine 4. The chromo domain of HP1 is identified as its
        methyl-lysine-binding domain.
      reference_section_type: ABSTRACT
- term:
    id: GO:0030466
    label: silent mating-type cassette heterochromatin formation
  evidence_type: EXP
  original_reference_id: PMID:16246721
  qualifier: involved_in
  review:
    summary: >-
      Swi6/HP1 is among the heterochromatin proteins required as Clr3 spreads
      across the ~20 kb silenced mating-type domain. Core process (duplicate
      term, distinct reference).
    action: ACCEPT
    reason: Experimental support for mat-cassette heterochromatin assembly.
    supported_by:
    - reference_id: PMID:16246721
      supporting_text: >-
        Clr3 spreads across the 20 kb silenced domain that requires its own HDAC
        activity and heterochromatin proteins including Swi6/HP1.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:15372076
  qualifier: is_active_in
  review:
    summary: >-
      Direct localization of Swi6 to pericentric/centromeric heterochromatin
      (Chp1 study). Core CC.
    action: ACCEPT
    reason: Direct localization evidence.
    supported_by:
    - reference_id: PMID:15372076
      supporting_text: >-
        deletion of the chp1(+) gene causes centromere-specific decreases in
        Swi6 localization and histone H3-K9 methylation
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:15372076
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 is a structural component of mating-type region heterochromatin,
      contributing to residual H3K9 methylation there. Core CC.
    action: ACCEPT
    reason: Direct evidence for activity at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:15372076
      supporting_text: >-
        Swi6 and Chp2 are critical to maintaining this residual methylation.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140720
    label: subtelomeric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:15372076
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 is a structural component of telomeric/subtelomeric heterochromatin.
      Core CC.
    action: ACCEPT
    reason: Direct evidence for activity at subtelomeric heterochromatin.
    supported_by:
    - reference_id: PMID:15372076
      supporting_text: >-
        Chp1 is a structural component of three heterochromatic
        regions-centromeres, the mating-type region, and telomeres
      reference_section_type: ABSTRACT
- term:
    id: GO:0033562
    label: co-transcriptional gene silencing by RNA interference machinery
  evidence_type: IMP
  original_reference_id: PMID:17512405
  qualifier: involved_in
  review:
    summary: >-
      Swi6 contributes to RNAi-dependent heterochromatic gene silencing, which
      involves co-transcriptional RNA processing/turnover of heterochromatic
      transcripts. Core process.
    action: ACCEPT
    reason: Supports RNAi co-transcriptional silencing role.
    supported_by:
    - reference_id: PMID:17512405
      supporting_text: >-
        the RNAi pathway is required for heterochromatin-dependent silencing of
        transgene insertions at centromeric repeats
      reference_section_type: ABSTRACT
- term:
    id: GO:0140727
    label: siRNA-mediated pericentric heterochromatin formation
  evidence_type: IMP
  original_reference_id: PMID:12193640
  qualifier: involved_in
  review:
    summary: >-
      Swi6 functions in the RNAi-directed pathway that establishes
      pericentromeric heterochromatin and H3K9 methylation through siRNAs from
      centromeric repeats. Core process.
    action: ACCEPT
    reason: Supports siRNA-mediated pericentric heterochromatin formation.
    supported_by:
    - reference_id: PMID:12193640
      supporting_text: >-
        We propose that double-stranded RNA arising from centromeric repeats
        targets formation and maintenance of heterochromatin through RNAi.
      reference_section_type: ABSTRACT
- term:
    id: GO:0033696
    label: heterochromatin boundary formation
  evidence_type: IMP
  original_reference_id: PMID:21211723
  qualifier: involved_in
  review:
    summary: >-
      Swi6 associates with the Asf1/HIRA histone chaperone, which spreads across
      silenced domains to enforce silencing and promote nucleosome occupancy,
      contributing to proper heterochromatin organization at boundaries. Core
      process.
    action: ACCEPT
    reason: Supports role in heterochromatin boundary/nucleosome occupancy.
    supported_by:
    - reference_id: PMID:21211723
      supporting_text: >-
        a histone chaperone complex containing Asf1 and HIRA that spreads across
        silenced domains via its association with Swi6 to enforce
        transcriptional silencing.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140463
    label: chromatin-protein adaptor activity
  evidence_type: EXP
  original_reference_id: PMID:16762840
  qualifier: enables
  review:
    summary: >-
      Swi6/HP1 recruits the JmjC anti-silencing protein Epe1 to heterochromatin,
      balancing opposing chromatin-modifying activities. Core adaptor MF.
    action: ACCEPT
    reason: Direct evidence Swi6 recruits Epe1 (adaptor activity).
    supported_by:
    - reference_id: PMID:16762840
      supporting_text: >-
        Epe1 is recruited to heterochromatic loci by the heterochromatin protein Swi6/HP1.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140463
    label: chromatin-protein adaptor activity
  evidence_type: EXP
  original_reference_id: PMID:17449867
  qualifier: enables
  review:
    summary: >-
      Confirms Swi6 recruits Epe1 to heterochromatin. Core adaptor MF
      (duplicate term, distinct reference).
    action: ACCEPT
    reason: Independent evidence for Swi6-mediated Epe1 recruitment.
    supported_by:
    - reference_id: PMID:17449867
      supporting_text: >-
        We report that Swi6 recruits Epe1 to heterochromatin
      reference_section_type: ABSTRACT
- term:
    id: GO:0003677
    label: DNA binding
  evidence_type: IDA
  original_reference_id: PMID:37400983
  qualifier: enables
  review:
    summary: >-
      PomBase experimental (IDA) annotation. Our cached abstract foregrounds Chp2, the
      other fission-yeast HP1 protein, but the full text (not in our cache) compares both
      HP1 proteins, and PomBase made a direct-assay (IDA) DNA-binding annotation to Swi6
      on its basis. Swi6 is independently well established to bind DNA in a sequence-
      independent manner via its hinge region. DNA binding is thus a genuine biochemical
      property of Swi6 but is ancillary to its core role as an H3K9me reader/effector.
    action: KEEP_AS_NON_CORE
    reason: >-
      Valid PomBase IDA; Swi6 hinge-mediated DNA binding is well established. Retained as
      a real but non-core property rather than a defining function. (An earlier
      assessment that this Chp2-titled paper does not support Swi6 was based on the
      abstract-only cache and overruled the curator without justification.)
    supported_by:
    - reference_id: PMID:37400983
      supporting_text: >-
        Here, we focus on Chp2, one of the two HP1 proteins in fission yeast,
        and investigate how Chp2's DNA-binding ability contributes to its function.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:10801440
  qualifier: enables
  review:
    summary: >-
      This row records the Swi6-Swi6 (chromoshadow-domain) self-interaction; it
      is more informatively captured as identical protein binding /
      homodimerization. Bare "protein binding" is uninformative.
    action: MARK_AS_OVER_ANNOTATED
    reason: Self-interaction better described as homodimerization (GO:0042802).
    supported_by:
    - reference_id: PMID:10801440
      supporting_text: >-
        The 1.9 A crystal structure of the Swi6 CSD is presented here. This
        reveals a novel dimeric structure
      reference_section_type: ABSTRACT
- term:
    id: GO:0003723
    label: RNA binding
  evidence_type: EXP
  original_reference_id: PMID:22683269
  qualifier: enables
  review:
    summary: >-
      Core molecular function. Swi6 binds RNA via its hinge basic patch; RNA
      competes with H3K9me for the chromodomain, and Swi6 sequesters and primes
      heterochromatic transcripts for degradation.
    action: ACCEPT
    reason: Direct in vivo/in vitro evidence for Swi6 RNA binding.
    supported_by:
    - reference_id: PMID:22683269
      supporting_text: >-
        RNA competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
      reference_section_type: ABSTRACT
- term:
    id: GO:0140463
    label: chromatin-protein adaptor activity
  evidence_type: IPI
  original_reference_id: PMID:18716626
  qualifier: enables
  review:
    summary: >-
      Swi6 recruits meiotic shugoshin Sgo1 to centromeres, the informative MF
      underlying the Swi6-Sgo1 interaction. Core adaptor MF.
    action: ACCEPT
    reason: Direct evidence Swi6 recruits Sgo1 (adaptor activity).
    supported_by:
    - reference_id: PMID:18716626
      supporting_text: >-
        the heterochromatin protein Swi6 associates directly with
        meiosis-specific shugoshin Sgo1, a protector of cohesin at centromeres.
      reference_section_type: ABSTRACT
- term:
    id: GO:1990813
    label: meiotic centromeric cohesion protection in anaphase I
  evidence_type: EXP
  original_reference_id: PMID:18716626
  qualifier: involved_in
  review:
    summary: >-
      By recruiting shugoshin Sgo1, Swi6 protects centromeric cohesin during
      meiosis; forced Sgo1 localization rescues meiotic segregation in swi6
      cells. Core meiotic process.
    action: ACCEPT
    reason: Experimental support for meiotic centromeric cohesion protection.
    supported_by:
    - reference_id: PMID:18716626
      supporting_text: >-
        The forced centromeric localization of Sgo1 restores proper meiotic
        chromosome segregation in swi6 cells.
      reference_section_type: ABSTRACT
- term:
    id: GO:0062072
    label: histone H3K9me2/3 reader activity
  evidence_type: IDA
  original_reference_id: PMID:22683269
  qualifier: enables
  review:
    summary: >-
      Confirms the Swi6 chromodomain reads H3K9me (in competition with RNA).
      Core MF (duplicate term, distinct reference).
    action: ACCEPT
    reason: Direct evidence for H3K9me reader activity.
    supported_by:
    - reference_id: PMID:22683269
      supporting_text: >-
        RNA competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
      reference_section_type: ABSTRACT
- term:
    id: GO:1990342
    label: heterochromatin island
  evidence_type: IDA
  original_reference_id: PMID:22144463
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 is active at facultative heterochromatin islands that form over
      meiotic genes via RNA elimination machinery. Genuine but specialized
      (facultative) localization; kept as non-core relative to constitutive
      heterochromatin sites.
    action: KEEP_AS_NON_CORE
    reason: Genuine facultative-heterochromatin localization; non-core context.
    supported_by:
    - reference_id: PMID:22144463
      supporting_text: >-
        we describe facultative heterochromatin islands in fission yeast and show
        that their formation at meiotic genes requires factors that eliminate
        meiotic messenger RNAs (mRNAs) during vegetative growth.
      reference_section_type: ABSTRACT
- term:
    id: GO:0006325
    label: chromatin organization
  evidence_type: IMP
  original_reference_id: PMID:19111658
  qualifier: involved_in
  review:
    summary: >-
      Swi6 organizes heterochromatin via a network of associated chromatin
      proteins and contributes to silencing. Correct but high-level; the
      specific heterochromatin formation/silencing terms are preferred.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic relative to specific heterochromatin processes.
    supported_by:
    - reference_id: PMID:19111658
      supporting_text: >-
        Swi6 associates with a different set of nuclear proteins and with
        noncoding centromeric transcripts and is required for efficient
        RNAi-dependent processing of these transcripts.
      reference_section_type: ABSTRACT
- term:
    id: GO:0071962
    label: mitotic sister chromatid cohesion, centromeric
  evidence_type: IMP
  original_reference_id: PMID:31000521
  qualifier: involved_in
  review:
    summary: >-
      Swi6 is required for cohesin (Rad21) enrichment at the pericentromere,
      establishing centromeric sister-chromatid cohesion. Core process.
    action: ACCEPT
    reason: Mutant phenotype supports centromeric cohesion role.
    supported_by:
    - reference_id: PMID:31000521
      supporting_text: >-
        Swi6 is required for cohesin enrichment at the pericentromere.
      reference_section_type: ABSTRACT
- term:
    id: GO:1990758
    label: mitotic sister chromatid biorientation
  evidence_type: IMP
  original_reference_id: PMID:31000521
  qualifier: involved_in
  review:
    summary: >-
      Loss of Swi6 causes defects in establishing bioriented sister
      kinetochores; via cohesin enrichment Swi6 promotes biorientation. Core
      process.
    action: ACCEPT
    reason: Mutant phenotype supports biorientation role.
    supported_by:
    - reference_id: PMID:31000521
      supporting_text: >-
        Loss of Swi6 leads to abnormal mitosis, including defects in the
        establishment of bioriented sister kinetochores and microtubule attachment.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:11283354
  qualifier: is_active_in
  review:
    summary: >-
      Direct evidence that Swi6 localizes to pericentric heterochromatin in an
      H3K9me-dependent manner. Core CC.
    action: ACCEPT
    reason: Direct localization evidence.
    supported_by:
    - reference_id: PMID:11283354
      supporting_text: >-
        Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic
        regions is dependent on H3 Lys9 methylation.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:11283354
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to mating-type region heterochromatin (H3K9me-dependent).
      Core CC.
    action: ACCEPT
    reason: Direct localization evidence at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:11283354
      supporting_text: >-
        Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic
        regions is dependent on H3 Lys9 methylation.
      reference_section_type: ABSTRACT
- term:
    id: GO:0003723
    label: RNA binding
  evidence_type: IDA
  original_reference_id: PMID:19111658
  qualifier: enables
  review:
    summary: >-
      Swi6 associates with noncoding centromeric transcripts (Clr4-dependent),
      consistent with RNA binding. Core MF (duplicate term, distinct reference).
    action: ACCEPT
    reason: Direct evidence Swi6 associates with heterochromatic RNAs.
    supported_by:
    - reference_id: PMID:19111658
      supporting_text: >-
        Swi6 associates with a different set of nuclear proteins and with
        noncoding centromeric transcripts
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:19965387
  qualifier: is_active_in
  review:
    summary: >-
      Direct localization of Swi6 to pericentric heterochromatin (shugoshin
      localization study). Core CC.
    action: ACCEPT
    reason: Direct localization evidence.
    supported_by:
    - reference_id: PMID:19965387
      supporting_text: >-
        Bub1 phosphorylates the conserved serine 121 of histone H2A in fission
        yeast Schizosaccharomyces pombe.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:19965387
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to mating-type region heterochromatin. Core CC (duplicate
      term, distinct reference).
    action: ACCEPT
    reason: Direct localization evidence at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:11780129
      supporting_text: >-
        Cohesin is also enriched at the mating-type heterochromatic region in a manner that
        depends on Swi6 and is required to preserve the genomic integrity of this locus.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:29136238
  qualifier: enables
  review:
    summary: >-
      Bare "protein binding"; this study concerns H3Y41 phosphorylation
      regulating Swi6/Chp1 localization at centromeric heterochromatin rather
      than a discrete informative binding partner. Uninformative MF.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative MF.
    supported_by:
    - reference_id: PMID:29136238
      supporting_text: >-
        it promotes the binding of Chp1 to histone H3 and the exclusion of Swi6.
      reference_section_type: ABSTRACT
- term:
    id: GO:0000792
    label: heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:26438724
  qualifier: is_active_in
  review:
    summary: >-
      Direct evidence Swi6 is a dynamic component of heterochromatin. Core CC
      (generic heterochromatin term).
    action: ACCEPT
    reason: Direct localization in heterochromatin.
    supported_by:
    - reference_id: PMID:26438724
      supporting_text: >-
        the fission yeast HP1 protein Swi6 exists as a single highly dynamic
        population that rapidly exchanges in cis and in trans between different
        heterochromatic regions.
      reference_section_type: ABSTRACT
- term:
    id: GO:0033696
    label: heterochromatin boundary formation
  evidence_type: IMP
  original_reference_id: PMID:26438724
  qualifier: involved_in
  review:
    summary: >-
      Swi6 dimerization demarcates constitutive heterochromatin from
      neighboring euchromatin and confines its spread, a genuine boundary
      function. Core process.
    action: ACCEPT
    reason: Strong evidence for a heterochromatin demarcation/confinement role.
    supported_by:
    - reference_id: PMID:26438724
      supporting_text: >-
        a surprising role for Swi6 dimerization in demarcating constitutive
        heterochromatin from neighboring euchromatin.
      reference_section_type: ABSTRACT
- term:
    id: GO:0044820
    label: mitotic telomere tethering at nuclear periphery
  evidence_type: IMP
  original_reference_id: PMID:25778919
  qualifier: involved_in
  review:
    summary: >-
      Swi6/HP1 (with cohesin Rad21) is associated with telomere clustering;
      Aurora B-driven telomere dispersion involves dissociation of Swi6 from
      telomeres. Genuine but specialized telomere-organization role; non-core
      relative to heterochromatin silencing.
    action: KEEP_AS_NON_CORE
    reason: Genuine telomere-organization role but a peripheral context.
    supported_by:
    - reference_id: PMID:25778919
      supporting_text: >-
        Dispersion is promoted by the dissociation of Swi6/HP1 and cohesin Rad21
        from telomeres
      reference_section_type: ABSTRACT
- term:
    id: GO:0006325
    label: chromatin organization
  evidence_type: IMP
  original_reference_id: PMID:9710635
  qualifier: involved_in
  review:
    summary: >-
      Swi6 contributes to chromatin remodeling/organization underlying
      mating-type silencing. Correct but high-level; specific heterochromatin
      terms are preferred.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic.
    supported_by:
    - reference_id: PMID:9710635
      supporting_text: >-
        a similar silencing mechanism may operate in the distantly related yeast
        Schizosaccharomyces pombe.
      reference_section_type: ABSTRACT
- term:
    id: GO:0000785
    label: chromatin
  evidence_type: IDA
  original_reference_id: PMID:21151114
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 is chromatin-associated; correct but generic relative to the
      heterochromatin subdomain terms.
    action: KEEP_AS_NON_CORE
    reason: Correct but generic CC.
    supported_by:
    - reference_id: PMID:21151114
      supporting_text: >-
        the RNAi proteins Dcr1 and Rdp1 physically associate with some
        euchromatic genes, noncoding RNA genes and retrotransposon long terminal repeats
      reference_section_type: ABSTRACT
- term:
    id: GO:0000792
    label: heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:11069763
  qualifier: is_active_in
  review:
    summary: >-
      GFP-Swi6 is concentrated at heterochromatic centromeres and telomeres.
      Core CC (generic heterochromatin term).
    action: ACCEPT
    reason: Direct localization in heterochromatin.
    supported_by:
    - reference_id: PMID:11069763
      supporting_text: >-
        GFP-Swi6p localises to the nucleus and is concentrated at the
        heterochromatic centromeres and telomeres.
      reference_section_type: ABSTRACT
- term:
    id: GO:0000779
    label: condensed chromosome, centromeric region
  evidence_type: IDA
  original_reference_id: PMID:7660126
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to centromeres and is required for their proper function
      during mitosis. Core CC.
    action: ACCEPT
    reason: Direct localization at centromeres.
    supported_by:
    - reference_id: PMID:7660126
      supporting_text: >-
        Thus, Swi6p is located at fission yeast centromeres and is required for
        their proper function.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:7660126
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to the silent mating-type loci heterochromatin. Core CC.
    action: ACCEPT
    reason: Direct localization at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:7660126
      supporting_text: >-
        Swi6p localizes with these three chromosomal regions.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140720
    label: subtelomeric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:7660126
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to telomeric/subtelomeric heterochromatin. Core CC.
    action: ACCEPT
    reason: Direct localization at subtelomeric heterochromatin.
    supported_by:
    - reference_id: PMID:7660126
      supporting_text: >-
        the silent mating-type loci, centromeres, and telomeres are assembled
        into silent heterochromatin-like domains.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:10766735
  qualifier: is_active_in
  review:
    summary: >-
      Direct ChIP mapping shows Swi6 confined to (and spreading across)
      pericentromeric outer repeats. Core CC.
    action: ACCEPT
    reason: Direct localization at pericentric repeats.
    supported_by:
    - reference_id: PMID:10766735
      supporting_text: >-
        Swi6 and Chp1 are confined to the flanking outer repeats and Swi6 can
        spread across at least 3 kb of extraneous chromatin in cen1.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:11553715
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 binds the pericentromeric repeats flanking the central core,
      consistent with conserved centromere domain organization. Core CC.
    action: ACCEPT
    reason: Direct localization at pericentromeric repeats.
    supported_by:
    - reference_id: PMID:11553715
      supporting_text: >-
        the Swi6 protein binds the surrounding repeats.
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:20299449
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 occupies pericentric heterochromatin during the inactive state,
      regulated by an H3K4ac-mediated chromodomain switch. Core CC.
    action: ACCEPT
    reason: Direct localization at pericentric heterochromatin.
    supported_by:
    - reference_id: PMID:20299449
      supporting_text: >-
        Chp2/Swi6 (HP1 homologs) are recruited during the inactive state.
      reference_section_type: ABSTRACT
- term:
    id: GO:0007533
    label: mating type switching
  evidence_type: IMP
  original_reference_id: PMID:1620099
  qualifier: involved_in
  review:
    summary: >-
      swi6 mutation reduces mating-type switching and disrupts the chromatin
      structure of the mating-type region. Core process.
    action: ACCEPT
    reason: Classic mutant evidence for swi6 in mating-type switching.
    supported_by:
    - reference_id: PMID:1620099
      supporting_text: >-
        Besides reducing MT switching, the swi6 mutation leads to deletions in
        the MT region caused by intrachromosomal cross-overs between two paired
        cassettes.
      reference_section_type: ABSTRACT
- term:
    id: GO:0030466
    label: silent mating-type cassette heterochromatin formation
  evidence_type: IMP
  original_reference_id: PMID:1620099
  qualifier: involved_in
  review:
    summary: >-
      swi6 is required for repression/heterochromatin of the silent mating-type
      cassettes; loss permits expression of both cassettes and recombination in
      the normally inert K region. Core process.
    action: ACCEPT
    reason: Mutant evidence for silent mat-cassette heterochromatin.
    supported_by:
    - reference_id: PMID:1620099
      supporting_text: >-
        swi6 also allows the simultaneous expression of two different cassettes
        in the same haploid cell.
      reference_section_type: ABSTRACT
- term:
    id: GO:0030466
    label: silent mating-type cassette heterochromatin formation
  evidence_type: IMP
  original_reference_id: PMID:8138176
  qualifier: involved_in
  review:
    summary: >-
      swi6 (with clr1) derepresses the silent mating-type loci and allows K-
      region recombination when mutated, supporting its role in silent cassette
      heterochromatin. Core process (duplicate term, distinct reference).
    action: ACCEPT
    reason: Mutant evidence for silent mat-cassette heterochromatin.
    supported_by:
    - reference_id: PMID:8138176
      supporting_text: >-
        mutations in two genes, clr1 and swi6, had been shown to allow both
        expression of the silent loci and recombination in the K region.
      reference_section_type: ABSTRACT
- term:
    id: GO:0062072
    label: histone H3K9me2/3 reader activity
  evidence_type: IDA
  original_reference_id: PMID:22727667
  qualifier: enables
  review:
    summary: >-
      This reference characterizes the Chp1 chromodomain's nucleic-acid binding
      enhanced by H3K9me; it concerns the chromodomain protein family context.
      Swi6 H3K9me reader activity is firmly established by other references
      (PMID:11242054, PMID:22683269), so the term is correct for Swi6 even
      though this particular citation centers on Chp1.
    action: ACCEPT
    reason: H3K9me reader activity is well established for Swi6 (core MF).
    supported_by:
    - reference_id: PMID:11242054
      supporting_text: >-
        The chromo domain of HP1 is identified as its methyl-lysine-binding domain.
      reference_section_type: ABSTRACT
- term:
    id: GO:0007534
    label: gene conversion at mating-type locus
  evidence_type: IMP
  original_reference_id: PMID:6587363
  qualifier: involved_in
  review:
    summary: >-
      swi6 belongs to the group of switching genes affecting the efficient
      utilization of the mat1 cut and the directionality of switching (gene
      conversion). Core process.
    action: ACCEPT
    reason: Classic genetic evidence placing swi6 in mat switching/conversion.
    supported_by:
    - reference_id: PMID:6587363
      supporting_text: >-
        The remaining three ( swi2 , -5, -6) have normal levels of cut, do not
        make errors of resolution, and possibly are required either for
        efficient utilization of the cut or determining the directionality of switching.
      reference_section_type: ABSTRACT
- term:
    id: GO:0003723
    label: RNA binding
  evidence_type: IDA
  original_reference_id: PMID:22683269
  qualifier: enables
  review:
    summary: >-
      Swi6 RNA binding via the hinge (duplicate of the RNA-binding annotation
      from the same reference). Core MF.
    action: ACCEPT
    reason: Direct evidence for Swi6 RNA binding.
    supported_by:
    - reference_id: PMID:22683269
      supporting_text: >-
        Stimulated by positively charged residues in the hinge region, RNA
        competes with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
      reference_section_type: ABSTRACT
- term:
    id: GO:0005721
    label: pericentric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:20929775
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to pericentric heterochromatin (inner-centromere/
      shugoshin study). Core CC.
    action: ACCEPT
    reason: Direct localization at pericentric heterochromatin.
    supported_by:
    - reference_id: PMID:20929775
      supporting_text: >-
        phosphorylated H2A-S121 facilitates the binding of shugoshin, the
        centromeric CPC adaptor.
      reference_section_type: ABSTRACT
- term:
    id: GO:0031934
    label: mating-type region heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:20929775
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to mating-type region heterochromatin. Core CC (duplicate
      term, distinct reference).
    action: ACCEPT
    reason: Direct localization at mat-region heterochromatin.
    supported_by:
    - reference_id: PMID:20929775
      supporting_text: >-
        the inner centromere is defined by intersection of two histone kinases.
      reference_section_type: ABSTRACT
- term:
    id: GO:0140720
    label: subtelomeric heterochromatin
  evidence_type: IDA
  original_reference_id: PMID:20929775
  qualifier: is_active_in
  review:
    summary: >-
      Swi6 localizes to subtelomeric heterochromatin. Core CC (duplicate term,
      distinct reference).
    action: ACCEPT
    reason: Direct localization at subtelomeric heterochromatin.
    supported_by:
    - reference_id: PMID:20929775
      supporting_text: >-
        For proper partitioning of chromosomes in mitosis, the chromosomal
        passenger complex (CPC) including Aurora B and survivin must be localized
        at the center of paired kinetochores
      reference_section_type: ABSTRACT
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:20929775
  qualifier: enables
  review:
    summary: >-
      Bare "protein binding" (interaction with Hrk1/Haspin context). The
      informative MF is Swi6's chromatin-protein adaptor activity; the generic
      term is uninformative.
    action: MARK_AS_OVER_ANNOTATED
    reason: Uninformative MF.
    supported_by:
    - reference_id: PMID:20929775
      supporting_text: >-
        Haspin colocalizes with cohesin by associating with Pds5, whereas Bub1
        localizes at kinetochores.
      reference_section_type: ABSTRACT
core_functions:
- description: >-
    Reads the H3K9me2/me3 heterochromatin mark via its chromodomain, providing
    the recognition step that targets Swi6 to Clr4-methylated nucleosomes.
  supported_by:
  - reference_id: PMID:11242054
    supporting_text: >-
      HP1 can bind with high affinity to histone H3 methylated at lysine 9 but
      not at lysine 4. The chromo domain of HP1 is identified as its
      methyl-lysine-binding domain.
    reference_section_type: ABSTRACT
- description: >-
    Homodimerizes/oligomerizes via its chromoshadow domain to form a
    spreading-competent, chromatin-compacting platform and protein-interaction
    pit.
  supported_by:
  - reference_id: PMID:10801440
    supporting_text: >-
      Dimerisation through the CSD of HP1-like proteins results in the
      simultaneous formation of a putative protein-protein interaction pit
    reference_section_type: ABSTRACT
  - reference_id: PMID:23485968
    supporting_text: >-
      binding of the key S. pombe HP1 protein, Swi6, to methylated nucleosomes
      drives a switch from an auto-inhibited state to a spreading-competent state.
    reference_section_type: ABSTRACT
- description: >-
    Acts as a chromatin-protein adaptor/scaffold, recruiting effectors to
    heterochromatin including cohesin (Psc3), the JmjC protein Epe1, the RNAi
    bridge Ers1/RDRC, the switching factor Swi2, and shugoshin Sgo1.
  supported_by:
  - reference_id: PMID:11780129
    supporting_text: >-
      a cohesin subunit Psc3 interacts with Swi6 and its mouse homologue HP1.
    reference_section_type: ABSTRACT
  - reference_id: PMID:16762840
    supporting_text: >-
      Epe1 is recruited to heterochromatic loci by the heterochromatin protein Swi6/HP1.
    reference_section_type: ABSTRACT
- description: >-
    Mediates transcriptional and co-transcriptional gene silencing and assembly,
    demarcation and confinement of heterochromatin at centromeres, telomeres,
    the silent mating-type cassettes and rDNA.
  supported_by:
  - reference_id: PMID:26438724
    supporting_text: >-
      a surprising role for Swi6 dimerization in demarcating constitutive
      heterochromatin from neighboring euchromatin.
    reference_section_type: ABSTRACT
- description: >-
    Enriches cohesin at the pericentromere to ensure centromeric sister-
    chromatid cohesion, biorientation and accurate chromosome segregation.
  supported_by:
  - reference_id: PMID:31000521
    supporting_text: >-
      Swi6 is required for cohesin enrichment at the pericentromere.
    reference_section_type: ABSTRACT
- description: >-
    Required for mating-type switching, including correct donor selection
    (directionality), by maintaining silent-cassette heterochromatin and
    recruiting the switching factor Swi2.
  supported_by:
  - reference_id: PMID:1620099
    supporting_text: >-
      Besides reducing MT switching, the swi6 mutation leads to deletions in the
      MT region caused by intrachromosomal cross-overs between two paired cassettes.
    reference_section_type: ABSTRACT
  - reference_id: PMID:36951094
    supporting_text: >-
      the Swi6-binding site was required for Swi2 localization at SRE2 to select
      mat2-P in M cells.
    reference_section_type: ABSTRACT
- description: >-
    Binds RNA via its basic hinge, sequestering and priming heterochromatic
    transcripts for degradation in competition with H3K9me chromodomain binding.
  supported_by:
  - reference_id: PMID:22683269
    supporting_text: >-
      the fission yeast HP1(Swi6) protein guarantees tight repression of
      heterochromatic genes through RNA sequestration and degradation.
    reference_section_type: ABSTRACT
proposed_new_terms: []
suggested_questions:
- question: >-
    What determines the partitioning of effector recruitment between the two
    fission yeast HP1 proteins (Swi6 vs Chp2), and how is Swi6's bifunctional
    chromatin- and RNA-binding balanced in vivo?
- question: >-
    How does Swi6 oligomerization/phase behavior on H3K9me chromatin quantitatively
    contribute to silencing versus boundary confinement at different loci?
suggested_experiments:
- description: >-
      Separation-of-function mutants (chromodomain reader-dead vs CSD dimer-dead vs
      hinge RNA/DNA-binding-dead) assayed in parallel for silencing, cohesin
      enrichment, donor selection, and boundary confinement to dissect which core
      function drives each phenotype.
- description: >-
      Quantitative ChIP-seq/CUT&RUN of Swi6 and its recruited effectors (Psc3,
      Clr3/SHREC, Epe1, Ers1) in wild-type and Swi6 phospho-site mutants to map
      how post-translational modification reprograms the adaptor interactome.
references:
- id: file:interpro/panther/PTHR22812/PTHR22812-review.md
  title: 'PANTHER family review PTHR22812: IBA propagation assessment for swi6'
  findings: []
- id: GO_REF:0000033
  title: Annotation inferences using phylogenetic trees
  findings: []
- id: GO_REF:0000044
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location
    vocabulary mapping, accompanied by conservative changes to GO terms applied by
    UniProt
  findings: []
- id: GO_REF:0000117
  title: Electronic Gene Ontology annotations created by ARBA machine learning models
  findings: []
- id: GO_REF:0000120
  title: Combined Automated Annotation using Multiple IEA Methods
  findings: []
- id: PMID:10766735
  title: Distinct protein interaction domains and protein spreading in a complex centromere.
  findings:
  - statement: >-
      Swi6 is confined to the pericentromeric flanking outer repeats and can spread
      across at least 3 kb of chromatin in cen1.
    supporting_text: >-
      Swi6 and Chp1 are confined to the flanking outer repeats and Swi6 can spread
      across at least 3 kb of extraneous chromatin in cen1.
    reference_section_type: ABSTRACT
- id: PMID:10801440
  title: Dimerisation of a chromo shadow domain and distinctions from the chromodomain
    as revealed by structural analysis.
  findings:
  - statement: >-
      The Swi6 chromoshadow domain forms a dimer creating a protein-interaction pit
      for targeting effectors to chromatin.
    supporting_text: >-
      Dimerisation through the CSD of HP1-like proteins results in the simultaneous
      formation of a putative protein-protein interaction pit
    reference_section_type: ABSTRACT
- id: PMID:11069763
  title: Live analysis of lagging chromosomes during anaphase and their effect on
    spindle elongation rate in fission yeast.
  findings:
  - statement: GFP-Swi6 localizes to the nucleus and concentrates at centromeres and telomeres.
    supporting_text: >-
      GFP-Swi6p localises to the nucleus and is concentrated at the heterochromatic
      centromeres and telomeres.
    reference_section_type: ABSTRACT
- id: PMID:11242054
  title: Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo
    domain.
  findings:
  - statement: The HP1/Swi6 chromodomain is the methyl-lysine-binding domain selective for H3K9me.
    supporting_text: >-
      HP1 can bind with high affinity to histone H3 methylated at lysine 9 but not
      at lysine 4. The chromo domain of HP1 is identified as its methyl-lysine-binding domain.
    reference_section_type: ABSTRACT
- id: PMID:11283354
  title: Role of histone H3 lysine 9 methylation in epigenetic control of heterochromatin
    assembly.
  findings:
  - statement: Swi6 localization to heterochromatin depends on Clr4-mediated H3K9 methylation.
    supporting_text: >-
      Localization of Swi6, a homolog of Drosophila HP1, to heterochomatic regions
      is dependent on H3 Lys9 methylation.
    reference_section_type: ABSTRACT
- id: PMID:11553715
  title: The domain structure of centromeres is conserved from fission yeast to humans.
  findings:
  - statement: Swi6 binds the pericentromeric repeats flanking the central core.
    supporting_text: 'the Swi6 protein binds the surrounding repeats.'
    reference_section_type: ABSTRACT
- id: PMID:11780129
  title: Recruitment of cohesin to heterochromatic regions by Swi6/HP1 in fission
    yeast.
  findings:
  - statement: Swi6 recruits cohesin to centromeric and mating-type heterochromatin via Psc3.
    supporting_text: >-
      a cohesin subunit Psc3 interacts with Swi6 and its mouse homologue HP1.
    reference_section_type: ABSTRACT
- id: PMID:12193640
  title: Regulation of heterochromatic silencing and histone H3 lysine-9 methylation
    by RNAi.
  findings:
  - statement: RNAi from centromeric repeats targets heterochromatin formation in which Swi6 acts.
    supporting_text: >-
      We propose that double-stranded RNA arising from centromeric repeats targets
      formation and maintenance of heterochromatin through RNAi.
    reference_section_type: ABSTRACT
- id: PMID:14625560
  title: Hsk1-Dfp1 is required for heterochromatin-mediated cohesion at centromeres.
  findings: []
- id: PMID:14663140
  title: Two different Swi5-containing protein complexes are involved in mating-type
    switching and recombination repair in fission yeast.
  findings: []
- id: PMID:15292231
  title: Regulation of Swi6/HP1-dependent heterochromatin assembly by cooperation
    of components of the mitogen-activated protein kinase pathway and a histone deacetylase
    Clr6.
  findings:
  - statement: A Swi6-dependent pathway assembles heterochromatin across the silent mating-type region.
    supporting_text: >-
      a stochastic Swi6 (the fission yeast HP1 homolog)-dependent mechanism that is
      not fully understood.
    reference_section_type: ABSTRACT
- id: PMID:15372076
  title: A chromodomain protein, Chp1, is required for the establishment of heterochromatin
    in fission yeast.
  findings:
  - statement: Swi6 (with Chp2) maintains residual H3K9 methylation at centromeres, mat region and telomeres.
    supporting_text: 'Swi6 and Chp2 are critical to maintaining this residual methylation.'
    reference_section_type: ABSTRACT
- id: PMID:16246721
  title: The nucleation and maintenance of heterochromatin by a histone deacetylase
    in fission yeast.
  findings:
  - statement: Clr3 spreading across the silent mat domain requires heterochromatin proteins including Swi6/HP1.
    supporting_text: >-
      Clr3 spreads across the 20 kb silenced domain that requires its own HDAC
      activity and heterochromatin proteins including Swi6/HP1.
    reference_section_type: ABSTRACT
- id: PMID:16762840
  title: Swi6/HP1 recruits a JmjC domain protein to facilitate transcription of heterochromatic
    repeats.
  findings:
  - statement: Swi6/HP1 recruits the anti-silencing JmjC protein Epe1 to heterochromatin.
    supporting_text: >-
      Epe1 is recruited to heterochromatic loci by the heterochromatin protein Swi6/HP1.
    reference_section_type: ABSTRACT
- id: PMID:17449867
  title: Interaction of Epe1 with the heterochromatin assembly pathway in Schizosaccharomyces
    pombe.
  findings:
  - statement: Swi6 recruits Epe1 to heterochromatin.
    supporting_text: 'We report that Swi6 recruits Epe1 to heterochromatin'
    reference_section_type: ABSTRACT
- id: PMID:17512405
  title: RNAi-dependent and -independent RNA turnover mechanisms contribute to heterochromatic
    gene silencing.
  findings:
  - statement: RNAi is required for heterochromatin-dependent silencing at centromeric repeats.
    supporting_text: >-
      the RNAi pathway is required for heterochromatin-dependent silencing of
      transgene insertions at centromeric repeats
    reference_section_type: ABSTRACT
- id: PMID:18716626
  title: Heterochromatin links to centromeric protection by recruiting shugoshin.
  findings:
  - statement: Swi6 directly recruits shugoshin Sgo1 to protect centromeric cohesin in meiosis.
    supporting_text: >-
      the heterochromatin protein Swi6 associates directly with meiosis-specific
      shugoshin Sgo1, a protector of cohesin at centromeres.
    reference_section_type: ABSTRACT
- id: PMID:18761674
  title: Fission yeast chromatin assembly factor 1 assists in the replication-coupled
    maintenance of heterochromatin.
  findings: []
- id: PMID:18809570
  title: Balance between distinct HP1 family proteins controls heterochromatin assembly
    in fission yeast.
  findings:
  - statement: Swi6 and Chp2 play distinct, dose-dependent roles in assembling repressive heterochromatin.
    supporting_text: >-
      both Swi6 and Chp2 are required for the assembly of fully repressive heterochromatin
    reference_section_type: ABSTRACT
- id: PMID:19111658
  title: HP1 proteins form distinct complexes and mediate heterochromatic gene silencing
    by nonoverlapping mechanisms.
  findings:
  - statement: Swi6 associates with noncoding centromeric transcripts and is required for RNAi processing into siRNAs.
    supporting_text: >-
      Swi6 associates with a different set of nuclear proteins and with noncoding
      centromeric transcripts and is required for efficient RNAi-dependent processing
      of these transcripts.
    reference_section_type: ABSTRACT
- id: PMID:19136623
  title: Phosphorylation of Swi6/HP1 regulates transcriptional gene silencing at heterochromatin.
  findings:
  - statement: Swi6 recruits SHREC, Epe1 and cohesin; CK2 phosphorylation of Swi6 controls TGS.
    supporting_text: >-
      a HP1 homolog Swi6 recruits SHREC, Epe1, and cohesin, which are involved in
      transcriptional gene silencing (TGS), transcriptional activation, and sister
      chromatid cohesion, respectively.
    reference_section_type: ABSTRACT
- id: PMID:19443688
  title: Diverse roles of HP1 proteins in heterochromatin assembly and functions in
    fission yeast.
  findings:
  - statement: Swi6 and Chp2 with HDAC complexes limit Pol II occupancy across pericentromeric heterochromatin.
    supporting_text: >-
      Swi6 and Chp2 proteins and their associated HDAC complexes have overlapping
      functions in limiting Pol II occupancy across pericentromeric heterochromatin domains.
    reference_section_type: ABSTRACT
- id: PMID:19965387
  title: Phosphorylation of H2A by Bub1 prevents chromosomal instability through localizing
    shugoshin.
  findings: []
- id: PMID:20299449
  title: A chromodomain switch mediated by histone H3 Lys 4 acetylation regulates
    heterochromatin assembly.
  findings:
  - statement: Chp2/Swi6 are recruited to pericentric heterochromatin during the inactive state.
    supporting_text: 'Chp2/Swi6 (HP1 homologs) are recruited during the inactive state.'
    reference_section_type: ABSTRACT
- id: PMID:20929775
  title: Two histone marks establish the inner centromere and chromosome bi-orientation.
  findings: []
- id: PMID:21151114
  title: Dicer associates with chromatin to repress genome activity in Schizosaccharomyces
    pombe.
  findings: []
- id: PMID:21211723
  title: Asf1/HIRA facilitate global histone deacetylation and associate with HP1
    to promote nucleosome occupancy at heterochromatic loci.
  findings:
  - statement: An Asf1/HIRA histone chaperone spreads across silenced domains via association with Swi6.
    supporting_text: >-
      a histone chaperone complex containing Asf1 and HIRA that spreads across
      silenced domains via its association with Swi6 to enforce transcriptional silencing.
    reference_section_type: ABSTRACT
- id: PMID:22144463
  title: RNA elimination machinery targeting meiotic mRNAs promotes facultative heterochromatin
    formation.
  findings:
  - statement: Facultative heterochromatin islands form at meiotic genes via RNA elimination machinery.
    supporting_text: >-
      we describe facultative heterochromatin islands in fission yeast and show that
      their formation at meiotic genes requires factors that eliminate meiotic
      messenger RNAs (mRNAs) during vegetative growth.
    reference_section_type: ABSTRACT
- id: PMID:22474355
  title: Heterochromatin protein 1 homologue Swi6 acts in concert with Ers1 to regulate
    RNAi-directed heterochromatin assembly.
  findings:
  - statement: Swi6 recruits RDRC to heterochromatin through the RNAi intermediate Ers1.
    supporting_text: >-
      we demonstrate that Swi6 recruits RDRC to heterochromatin through Ers1, an
      RNAi factor intermediate.
    reference_section_type: ABSTRACT
- id: PMID:22683269
  title: HP1(Swi6) mediates the recognition and destruction of heterochromatic RNA
    transcripts.
  findings:
  - statement: Swi6 binds RNA via its hinge; RNA competes with H3K9me for the chromodomain.
    supporting_text: >-
      Stimulated by positively charged residues in the hinge region, RNA competes
      with methylated histone H3K9 for binding to the chromodomain of HP1(Swi6).
    reference_section_type: ABSTRACT
- id: PMID:22727667
  title: Intrinsic nucleic acid-binding activity of Chp1 chromodomain is required
    for heterochromatic gene silencing.
  findings: []
- id: PMID:22990236
  title: Hrp3 controls nucleosome positioning to suppress non-coding transcription
    in eu- and heterochromatin.
  findings: []
- id: PMID:23485968
  title: A conformational switch in HP1 releases auto-inhibition to drive heterochromatin
    assembly.
  findings:
  - statement: Swi6 binding to methylated nucleosomes switches it from auto-inhibited to spreading-competent oligomers.
    supporting_text: >-
      binding of the key S. pombe HP1 protein, Swi6, to methylated nucleosomes
      drives a switch from an auto-inhibited state to a spreading-competent state.
    reference_section_type: ABSTRACT
- id: PMID:25245948
  title: Tls1 regulates splicing of shelterin components to control telomeric heterochromatin
    assembly and telomere length.
  findings:
  - statement: Telomeric heterochromatin assembly requires multiple factors including Tls1.
    supporting_text: >-
      we comprehensively identified factors required for telomeric heterochromatin
      assembly, including a novel gene tls1+.
    reference_section_type: ABSTRACT
- id: PMID:25778919
  title: Aurora B prevents chromosome arm separation defects by promoting telomere
    dispersion and disjunction.
  findings:
  - statement: Telomere dispersion is promoted by dissociation of Swi6/HP1 and cohesin Rad21 from telomeres.
    supporting_text: >-
      Dispersion is promoted by the dissociation of Swi6/HP1 and cohesin Rad21 from telomeres
    reference_section_type: ABSTRACT
- id: PMID:26438724
  title: H3K9 methylation extends across natural boundaries of heterochromatin in
    the absence of an HP1 protein.
  findings:
  - statement: Swi6 dimerization demarcates and confines constitutive heterochromatin rather than promoting its spread.
    supporting_text: >-
      a surprising role for Swi6 dimerization in demarcating constitutive
      heterochromatin from neighboring euchromatin.
    reference_section_type: ABSTRACT
- id: PMID:29136238
  title: Regulation of transcriptional silencing and chromodomain protein localization
    at centromeric heterochromatin by histone H3 tyrosine 41 phosphorylation in fission
    yeast.
  findings: []
- id: PMID:29852001
  title: New insights into donor directionality of mating-type switching in Schizosaccharomyces
    pombe.
  findings:
  - statement: Mating-type switching donor selection is regulated by heterochromatin and the HP1-like protein Swi6.
    supporting_text: >-
      Mating-type switching in Schizosaccharomyces pombe entails programmed gene
      conversion events regulated by DNA replication, heterochromatin, and the
      HP1-like chromodomain protein Swi6.
    reference_section_type: ABSTRACT
- id: PMID:31000521
  title: Overlapping Roles in Chromosome Segregation for Heterochromatin Protein 1
    (Swi6) and DDK in Schizosaccharomyces pombe.
  findings:
  - statement: Swi6 is required for cohesin enrichment at the pericentromere and for sister-kinetochore biorientation.
    supporting_text: >-
      Swi6 is required for cohesin enrichment at the pericentromere.
    reference_section_type: ABSTRACT
- id: PMID:36951094
  title: Fission yeast Swi2 designates cell-type specific donor and stimulates Rad51-driven
    strand exchange for mating-type switching.
  findings:
  - statement: A Swi6-binding site in Swi2 is required for Swi2 localization at SRE2 to select the mat2-P donor.
    supporting_text: >-
      the Swi6-binding site was required for Swi2 localization at SRE2 to select
      mat2-P in M cells.
    reference_section_type: ABSTRACT
- id: PMID:37400983
  title: Cooperative DNA-binding activities of Chp2 are critical for its function
    in heterochromatin assembly.
  findings:
  - statement: >-
      This study characterizes cooperative DNA-binding by HP1 proteins in fission yeast.
      The cached abstract foregrounds Chp2, but PomBase used the full text to make a
      direct-assay (IDA) DNA-binding annotation to Swi6 (the other HP1 protein), retained
      here as a non-core, hinge-mediated property of Swi6.
    supporting_text: >-
      Here, we focus on Chp2, one of the two HP1 proteins in fission yeast, and
      investigate how Chp2's DNA-binding ability contributes to its function.
    reference_section_type: ABSTRACT
- id: PMID:39747188
  title: PhpC(NF-Y) transcription factor infiltrates heterochromatin to generate cryptic
    intron-containing transcripts crucial for small RNA production.
  findings:
  - statement: TF-driven transcription of heterochromatic repeats produces siRNAs critical for heterochromatin nucleation.
    supporting_text: >-
      siRNA production by this TF-mediated transcription pathway is critical for
      heterochromatin nucleation at target repeat loci.
    reference_section_type: ABSTRACT
- id: PMID:6587363
  title: Genes required for initiation and resolution steps of mating-type switching
    in fission yeast.
  findings:
  - statement: swi6 is required for efficient utilization of the mat1 cut and switching directionality.
    supporting_text: >-
      The remaining three ( swi2 , -5, -6) have normal levels of cut, do not make
      errors of resolution, and possibly are required either for efficient
      utilization of the cut or determining the directionality of switching.
    reference_section_type: ABSTRACT
- id: PMID:7660126
  title: 'The chromodomain protein Swi6: a key component at fission yeast centromeres.'
  findings:
  - statement: Swi6 localizes to centromeres, mating-type loci and telomeres and is required for centromere function.
    supporting_text: >-
      Thus, Swi6p is located at fission yeast centromeres and is required for their proper function.
    reference_section_type: ABSTRACT
- id: PMID:8138176
  title: Mutations in rik1, clr2, clr3 and clr4 genes asymmetrically derepress the
    silent mating-type loci in fission yeast.
  findings:
  - statement: swi6 (with clr1) allows expression of the silent mating-type loci and K-region recombination when mutated.
    supporting_text: >-
      mutations in two genes, clr1 and swi6, had been shown to allow both expression
      of the silent loci and recombination in the K region.
    reference_section_type: ABSTRACT
- id: PMID:9710635
  title: A novel function of the DNA repair gene rhp6 in mating-type silencing by
    chromatin remodeling in fission yeast.
  findings: []
- id: PMID:1620099
  title: The switching gene swi6 affects recombination and gene expression in the
    mating-type region of Schizosaccharomyces pombe.
  findings:
  - statement: swi6 mutation reduces mating-type switching and derepresses the silent cassettes, allowing K-region recombination.
    supporting_text: >-
      swi6 also allows the simultaneous expression of two different cassettes in the same haploid cell.
    reference_section_type: ABSTRACT