Pathway Human Homo sapiens

ATP-dependent chromatin remodelers

R-HSA-9932444 in Reactome release 97: under Chromatin organization, with 195 genes placed in it by the mapping files and 2 child pathways in the hierarchy.

The same number in the other species

Reactome writes the mouse and rat events it infers from a human pathway under the human number with the species token. A door opens only where that species' own list in this release holds the id: R-MMU-9932444 (mouse), R-RNO-9932444 (rat). Whether the event was inferred from this one is what the record says.

The record
The genes
Parents and children

Scripts are off

This page reads the pathway's record from its source while you read the rest; with scripts off the card stays at its loading line. Reload with scripts on to see it, or open the source's own page from the link in the card.

01The record

Reactome's own record of this pathway

What this tells you

The pathway list, the hierarchy and the genes on this page are read from the files Reactome publishes for Reactome release 97, built into this site on 2026-09-09: the pathway list, the hierarchy relationship file and the two gene mapping files. The record card is the one live read, Reactome's own record of this pathway.

Reactome's download page describes the mapping files: Mapping files link the source database identifier to the lowest level pathway diagram or subset of the pathway, all levels of the pathway hierarchy or database identifier to all reactions. [R13] The gene list here is the all-levels mapping of NCBI Gene ids, filled from the all-levels mapping of Ensembl ids where the NCBI file has no row for a gene; each row names which file it came from by the shape of its source id. Each row carries the evidence codes the file writes for it, as written and unranked, and both where the file writes both; no page of Reactome's documentation the survey read defines the codes, so this page does not expand them. A gene id the identity files do not name is kept as the source's own id with no door rather than turned into a symbol.

On citing what a search finds, Reactome says: It should be remembered that while we strive to contain the most current and accurate data, Reactome should not be used in citations where other primary sources of information are available. [R12] The record card prints the literature Reactome attaches to a curated pathway for that reason. The data are public domain: All data in the Reactome database and files derived from that data are licensed under the Creative Commons Public Domain Dedication (CC0). User may copy, modify, and distribute these data, even for commercial purposes, without asking for permission. Attribution is encouraged but not required. [R10] The pathway illustrations are licensed apart, under CC BY 4.0, and none is shown here.

A gene on this list is one Reactome places in this pathway in this release, and that is all the row says: it does not say the gene is expressed in a tissue or associated with a disease, which are the other two explorers' questions, read from other sources. Reactome's papers of record are [R01] [R02].

[R01] Ragueneau E, Gong C, Sinquin P, Sevilla C, Beavers D, Grentner A, et al. (2026). The Reactome Knowledgebase 2026. Nucleic Acids Research 54:D673-D681. PMID 41251150, doi 10.1093/nar/gkaf1223.
[R02] Milacic M, Beavers D, Conley P, Gong C, Gillespie M, Griss J, et al. (2024). The Reactome Pathway Knowledgebase 2024. Nucleic Acids Research 52:D672-D678. PMID 37941124, doi 10.1093/nar/gkad1025.
[R10] Reactome, reactome.org. License Agreement. https://reactome.org/license, read 2026-09-09.
[R12] Reactome, reactome.org. Citing us. https://reactome.org/cite, read 2026-09-09.
[R13] Reactome, reactome.org. Download. https://reactome.org/download-data, read 2026-09-09.

Stable id: R-HSA-9932444.3
Class: Pathway
Species: Homo sapiens (human)
Inferred: no: a curated human pathway
Disease pathway: no
Released: 2025-03-26
Events inside: 2, counted from the record

ATP-dependent chromatin remodelers use the energy from ATP hydrolysis to organize or rearrange nucleosomes by altering histone-DNA contacts (reviewed in de la Serna et al, 2006; Hargreaves and Crabtree, 2011; Narlikar et al, 2013; Clapier et al, 2017; Eustermann et al, 2024). There are four main classes of mammalian ATP-dependent chromatin remodellers: SWI/SNF (for switch/sucrose non-fermenting, initially characterized in S. cerevisiae); ISWI (imitation switch), chromodomain helicase DNA-binding (CHD) and INO80 (inositol requiring mutant 80). Each of these complexes is characterized by a SWI2/SNF2-family ATPase domain that contributes the catalytic function and translocates the DNA along the histone core of the nucleosome (reviewed in Narlikar et al, 2013; Clapier et al, 2017; Eustermann et al, 2024). The remainder of the 1 - 1.5MDa complexes are made up of accessory proteins that modulate ATPase activity and contribute to target recognition and specificity by interacting with nucleosomes and histone tails (reviewed in Hargreaves and Crabtree, 2011; Narlikar et al, 2013; Clapier et al, 2017; Eustermann et al, 2024).

Human SWI/SNF ATPases bind acetylated nucleosomes through the bromodomains of their two alternative catalytic subunits, SMARCA2 (also known as BRM) or SMARCA4 (BRG1) (Hassan et al, 2002; Singh et al, 2006; Singh et al, 2007; reviewed in Euskirchen et al 2012). Histone and DNA contacts are also mediated by other protein components of the SWI/SNF complex (reviewed in Euskirchen et al, 2012; Cenik and Shilatifard, 2021). SWI/SNF chromatin remodellers are generally involved in gene activation or repression and provide access to chromatin by repositioning or ejecting nucleosomes (reviewed in Eustermann et al, 2024).

Human CHD family ATPases contain one of nine catalytic CHD protein paralogs. DNA-binding is mediated by contacts between methylated histones and the chromodomain of CHD1-9, as well as through other protein constituents of the ATPase complex (Bannister et al, 2001; Lachner et al, 2001; Flanagan et al, 2005; Sims et al, 2005; reviewed in Eustermann et al, 2024). CHD family ATPases are involved in diverse chromatin remodelling activities, including nucleosome assembly and spacing, access to regulatory elements and changes to histone composition (reviewed in Clapier et al, 2017).

Human ISWI family ATPases contain one of two potential catalytic subunits, SMARCA5 (also known as SNF2H) or SMARCA1 (SNF2L), that interact with unmodified histone tails through their SANT domains (Grune et al, 2003; Boyer et al, 2004). ISWI complexes are generally involved in nucleosome maturation and regulation of spacing and assembly, often resulting in repressive gene states, but some are also involved in transcriptional activation (Xiao et al, 2001; reviewed in Clapier et al, 2017; Eustermann et al, 2024).

Human INO80 family ATPases contain one of three catalytic subunits, INO80, SRCAP or EP400. INO80 complexes generally contribute more to histone exchange than to nucleosome spacing, assembly or access (reviewed in Clapier et al, 2017; Eustermann et al, 2024).

Literature Reactome attaches

Chromatin remodelling in mammalian differentiation: lessons from ATP-dependent remodellersNat Rev Genet, 2006, vol. 7, p. 461-73PubMed 16708073
ATP-dependent chromatin remodeling: genetics, genomics and mechanismsCell Res, 2011, vol. 21, p. 396-420PubMed 21358755
Mechanisms and functions of ATP-dependent chromatin-remodeling enzymesCell, 2013, vol. 154, p. 490-503PubMed 23911317
Mechanisms of action and regulation of ATP-dependent chromatin-remodelling complexesNat Rev Mol Cell Biol, 2017, vol. 18, p. 407-422PubMed 28512350
Energy-driven genome regulation by ATP-dependent chromatin remodellersNat Rev Mol Cell Biol, 2024, vol. 25, p. 309-332PubMed 38081975
Function and selectivity of bromodomains in anchoring chromatin-modifying complexes to promoter nucleosomesCell, 2002, vol. 111, p. 369-79PubMed 12419247
Structural ramification for acetyl-lysine recognition by the bromodomain of human BRG1 protein, a central ATPase of the SWI/SNF remodeling complexChembiochem, 2007, vol. 8, p. 1308-16

Reactome's record lists 13 orthologous events, R-RNO-9932444 (Rattus norvegicus), R-MMU-9932444 (Mus musculus) among the species this site serves, and 11 in species this site opens no page for.

Reactome ContentService, this pathway's record · Reactome v97 · read 2026-09-30 ET · Reactome pathway browser"ATP-dependent chromatin remodelers". Reactome, release 97, https://reactome.org/PathwayBrowser/#/R-HSA-9932444 with StableID: R-HSA-9932444.3 (accessed 2026-09-30 ET). Reactome version 97 (CC0).

Reading this pathway's record from Reactome (the pathway record).Still reading. A first read of a pathway can take a while; this page waits up to 35 seconds for it, and its scripts then bring in the panel, or a line saying what did not arrive.

02The genes

Genes Reactome places in this human pathway

The mapping files place 195 genes in this human pathway; showing 1 to 100, in pages of 100, sorted by symbol for reading. The order carries no ranking.

Genes Reactome places in this human pathway, page 1 of 2
Gene	Authority id	Mapping file id	Evidence codes
GeneACTB	AuthorityHGNC:132	Mapping file id60 NCBI file	EvidenceIEA, TAS
GeneACTL6A	AuthorityHGNC:24124	Mapping file id86 NCBI file	EvidenceIEA, TAS
GeneACTL6B	AuthorityHGNC:160	Mapping file id51412 NCBI file	EvidenceIEA
GeneADNP	AuthorityHGNC:15766	Mapping file id23394 NCBI file	EvidenceIEA, TAS
GeneADNP2	AuthorityHGNC:23803	Mapping file id22850 NCBI file	EvidenceIEA
GeneARID1A	AuthorityHGNC:11110	Mapping file id8289 NCBI file	EvidenceIEA, TAS
GeneARID1B	AuthorityHGNC:18040	Mapping file id57492 NCBI file	EvidenceIEA, TAS
GeneARID2	AuthorityHGNC:18037	Mapping file id196528 NCBI file	EvidenceTAS
GeneAXIN2	AuthorityHGNC:904	Mapping file id8313 NCBI file	EvidenceTAS
GeneBCL11A	AuthorityHGNC:13221	Mapping file id53335 NCBI file	EvidenceIEA, TAS
GeneBCL11B	AuthorityHGNC:13222	Mapping file id64919 NCBI file	EvidenceIEA, TAS
GeneBCL7A	AuthorityHGNC:1004	Mapping file id605 NCBI file	EvidenceIEA, TAS
GeneBCL7B	AuthorityHGNC:1005	Mapping file id9275 NCBI file	EvidenceIEA, TAS
GeneBCL7C	AuthorityHGNC:1006	Mapping file id9274 NCBI file	EvidenceIEA, TAS
GeneBICRA	AuthorityHGNC:4332	Mapping file id29998 NCBI file	EvidenceTAS
GeneBICRAL	AuthorityHGNC:21111	Mapping file id23506 NCBI file	EvidenceTAS
GeneBRD7	AuthorityHGNC:14310	Mapping file id29117 NCBI file	EvidenceTAS
GeneBRD9	AuthorityHGNC:25818	Mapping file id65980 NCBI file	EvidenceTAS
GeneCBX1	AuthorityHGNC:1551	Mapping file id10951 NCBI file	EvidenceIEA, TAS
GeneCBX3	AuthorityHGNC:1553	Mapping file id11335 NCBI file	EvidenceIEA, TAS
GeneCDC73	AuthorityHGNC:16783	Mapping file id79577 NCBI file	EvidenceTAS
GeneCDK2AP1	AuthorityHGNC:14002	Mapping file id8099 NCBI file	EvidenceTAS
GeneCDK2AP2	AuthorityHGNC:30833	Mapping file id10263 NCBI file	EvidenceTAS
GeneCHD1	AuthorityHGNC:1915	Mapping file id1105 NCBI file	EvidenceTAS
GeneCHD2	AuthorityHGNC:1917	Mapping file id1106 NCBI file	EvidenceIEA, TAS
GeneCHD3	AuthorityHGNC:1918	Mapping file id1107 NCBI file	EvidenceTAS
GeneCHD4	AuthorityHGNC:1919	Mapping file id1108 NCBI file	EvidenceIEA, TAS
GeneCHD5	AuthorityHGNC:16816	Mapping file id26038 NCBI file	EvidenceTAS
GeneCHD6	AuthorityHGNC:19057	Mapping file id84181 NCBI file	EvidenceIEA, TAS
GeneCHD7	AuthorityHGNC:20626	Mapping file id55636 NCBI file	EvidenceTAS
GeneCHD8	AuthorityHGNC:20153	Mapping file id57680 NCBI file	EvidenceTAS
GeneCHD9	AuthorityHGNC:25701	Mapping file id80205 NCBI file	EvidenceTAS
GeneCHERP	AuthorityHGNC:16930	Mapping file id10523 NCBI file	EvidenceTAS
GeneCREBBP	AuthorityHGNC:2348	Mapping file id1387 NCBI file	EvidenceIEA, TAS
GeneCTCF	AuthorityHGNC:13723	Mapping file id10664 NCBI file	EvidenceTAS
GeneCTNNB1	AuthorityHGNC:2514	Mapping file id1499 NCBI file	EvidenceTAS
GeneCTR9	AuthorityHGNC:16850	Mapping file id9646 NCBI file	EvidenceTAS
GeneDDX42	AuthorityHGNC:18676	Mapping file id11325 NCBI file	EvidenceTAS
GeneDDX46	AuthorityHGNC:18681	Mapping file id9879 NCBI file	EvidenceTAS
GeneDHX15	AuthorityHGNC:2738	Mapping file id1665 NCBI file	EvidenceTAS
GeneDKK2	AuthorityHGNC:2892	Mapping file id27123 NCBI file	EvidenceTAS
GeneDPF1	AuthorityHGNC:20225	Mapping file id8193 NCBI file	EvidenceIEA, TAS
GeneDPF2	AuthorityHGNC:9964	Mapping file id5977 NCBI file	EvidenceIEA, TAS
GeneDPF3	AuthorityHGNC:17427	Mapping file id8110 NCBI file	EvidenceIEA, TAS
GeneEP300	AuthorityHGNC:3373	Mapping file id2033 NCBI file	EvidenceIEA, TAS
GeneFAM124B	AuthorityHGNC:26224	Mapping file id79843 NCBI file	EvidenceTAS
GeneFBP1	AuthorityHGNC:3606	Mapping file id2203 NCBI file	EvidenceTAS
GeneG6PC1	AuthorityHGNC:4056	Mapping file id2538 NCBI file	EvidenceTAS
GeneGATAD2A	AuthorityHGNC:29989	Mapping file id54815 NCBI file	EvidenceTAS
GeneGATAD2B	AuthorityHGNC:30778	Mapping file id57459 NCBI file	EvidenceTAS
GeneH2AB1	AuthorityHGNC:22516	Mapping file id474382 NCBI file	EvidenceIEA, TAS
GeneH2AC14	AuthorityHGNC:4727	Mapping file id8331 NCBI file	EvidenceIEA, TAS
GeneH2AC18	AuthorityHGNC:4736	Mapping file id8337 NCBI file	EvidenceIEA, TAS
GeneH2AC19	AuthorityHGNC:29668	Mapping file id723790 NCBI file	EvidenceIEA, TAS
GeneH2AC20	AuthorityHGNC:4738	Mapping file id8338 NCBI file	EvidenceIEA, TAS
GeneH2AC4	AuthorityHGNC:4734	Mapping file id8335 NCBI file	EvidenceIEA, TAS
GeneH2AC6	AuthorityHGNC:4733	Mapping file id8334 NCBI file	EvidenceIEA, TAS
GeneH2AC7	AuthorityHGNC:4729	Mapping file id3013 NCBI file	EvidenceIEA, TAS
GeneH2AC8	AuthorityHGNC:4724	Mapping file id3012 NCBI file	EvidenceIEA, TAS
GeneH2AJ	AuthorityHGNC:14456	Mapping file id55766 NCBI file	EvidenceIEA, TAS
GeneH2AX	AuthorityHGNC:4739	Mapping file id3014 NCBI file	EvidenceIEA, TAS
GeneH2AZ2	AuthorityHGNC:20664	Mapping file id94239 NCBI file	EvidenceIEA, TAS
GeneH2BC1	AuthorityHGNC:18730	Mapping file id255626 NCBI file	EvidenceIEA, TAS
GeneH2BC10	AuthorityHGNC:4756	Mapping file id8346 NCBI file	EvidenceIEA, TAS
GeneH2BC11	AuthorityHGNC:4761	Mapping file id8970 NCBI file	EvidenceIEA, TAS
GeneH2BC12	AuthorityHGNC:13954	Mapping file id85236 NCBI file	EvidenceIEA, TAS
GeneH2BC12L	AuthorityHGNC:4762	Mapping file id54145 NCBI file	EvidenceIEA, TAS
GeneH2BC13	AuthorityHGNC:4748	Mapping file id8340 NCBI file	EvidenceIEA, TAS
GeneH2BC14	AuthorityHGNC:4750	Mapping file id8342 NCBI file	EvidenceIEA, TAS
GeneH2BC15	AuthorityHGNC:4749	Mapping file id8341 NCBI file	EvidenceIEA, TAS
GeneH2BC17	AuthorityHGNC:4758	Mapping file id8348 NCBI file	EvidenceIEA, TAS
GeneH2BC21	AuthorityHGNC:4760	Mapping file id8349 NCBI file	EvidenceIEA, TAS
GeneH2BC26	AuthorityHGNC:20514	Mapping file id128312 NCBI file	EvidenceIEA, TAS
GeneH2BC3	AuthorityHGNC:4751	Mapping file id3018 NCBI file	EvidenceIEA, TAS
GeneH2BC4	AuthorityHGNC:4757	Mapping file id8347 NCBI file	EvidenceIEA, TAS
GeneH2BC5	AuthorityHGNC:4747	Mapping file id3017 NCBI file	EvidenceIEA, TAS
GeneH2BC6	AuthorityHGNC:4753	Mapping file id8344 NCBI file	EvidenceIEA, TAS
GeneH2BC7	AuthorityHGNC:4752	Mapping file id8343 NCBI file	EvidenceIEA, TAS
GeneH2BC8	AuthorityHGNC:4746	Mapping file id8339 NCBI file	EvidenceIEA, TAS
GeneH2BC9	AuthorityHGNC:4755	Mapping file id8345 NCBI file	EvidenceIEA, TAS
GeneH3-3A	AuthorityHGNC:4764	Mapping file id3020 NCBI file	EvidenceIEA, TAS
GeneH3-3B	AuthorityHGNC:4765	Mapping file id3021 NCBI file	EvidenceIEA, TAS
GeneH3C1	AuthorityHGNC:4766	Mapping file id8350 NCBI file	EvidenceIEA, TAS
GeneH3C10	AuthorityHGNC:4775	Mapping file id8357 NCBI file	EvidenceIEA, TAS
GeneH3C11	AuthorityHGNC:4771	Mapping file id8354 NCBI file	EvidenceIEA, TAS
GeneH3C12	AuthorityHGNC:4774	Mapping file id8356 NCBI file	EvidenceIEA, TAS
GeneH3C13	AuthorityHGNC:25311	Mapping file id653604 NCBI file	EvidenceIEA, TAS
GeneH3C14	AuthorityHGNC:20503	Mapping file id126961 NCBI file	EvidenceIEA, TAS
GeneH3C15	AuthorityHGNC:20505	Mapping file id333932 NCBI file	EvidenceIEA, TAS
GeneH3C2	AuthorityHGNC:4776	Mapping file id8358 NCBI file	EvidenceIEA, TAS
GeneH3C3	AuthorityHGNC:4768	Mapping file id8352 NCBI file	EvidenceIEA, TAS
GeneH3C4	AuthorityHGNC:4767	Mapping file id8351 NCBI file	EvidenceIEA, TAS
GeneH3C6	AuthorityHGNC:4769	Mapping file id8353 NCBI file	EvidenceIEA, TAS
GeneH3C7	AuthorityHGNC:4773	Mapping file id8968 NCBI file	EvidenceIEA, TAS
GeneH3C8	AuthorityHGNC:4772	Mapping file id8355 NCBI file	EvidenceIEA, TAS
GeneH4C1	AuthorityHGNC:4781	Mapping file id8359 NCBI file	EvidenceIEA, TAS
GeneH4C11	AuthorityHGNC:4785	Mapping file id8363 NCBI file	EvidenceIEA, TAS
GeneH4C12	AuthorityHGNC:4784	Mapping file id8362 NCBI file	EvidenceIEA, TAS
GeneH4C13	AuthorityHGNC:4791	Mapping file id8368 NCBI file	EvidenceIEA, TAS
GeneH4C14	AuthorityHGNC:4794	Mapping file id8370 NCBI file	EvidenceIEA, TAS

Evidence codes on this page: IEA, TAS, as the mapping file writes them; a row carrying two was written twice by the file, once under each. The mapping file id says which file placed the gene: an NCBI Gene id from NCBI2Reactome_All_Levels.txt, an Ensembl gene id from Ensembl2Reactome_All_Levels.txt. The two files can disagree about a code, so a row's codes are the one file's view. Measured by this site's build over this release on 2026-09-09: of the 138,908 human pathway-gene pairs both files place, 207 (0.149 per cent, over 47 genes) carry TAS in the Ensembl file where the NCBI rows carry IEA alone.

Reactome mapping files, the human rows for this pathway · Reactome release 97 · read 2026-09-09 · Reactome downloads"NCBI2Reactome_All_Levels.txt" and "Ensembl2Reactome_All_Levels.txt", Reactome, release 97, https://reactome.org/download-data/ (date of access 2026-09-09). Data CC0; attribution encouraged.

03The hierarchy

Parents and children in this release's hierarchy

Parents

Chromatin organizationR-HSA-4839726380 genes

Children

Reactome's hierarchy is a graph rather than a tree: a pathway can sit under more than one parent, and the gene counts are each pathway's own placements at every level under it, so a parent's count is not the sum of its children's.

Reactome, the human pathway list and hierarchy relationship file · Reactome release 97 · read 2026-09-09 · Reactome downloads"ReactomePathways.txt" and "ReactomePathwaysRelation.txt", Reactome, release 97, https://reactome.org/download-data/ (date of access 2026-09-09). Data CC0; attribution encouraged.