Pathway Human Homo sapiens

Protein localization

R-HSA-9609507 in Reactome release 97: a top-level pathway, with 164 genes placed in it by the mapping files and 4 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-9609507 (mouse), R-RNO-9609507 (rat). Whether the event was inferred from this one is what the record says.

The record
The genes
Parents and children

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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-9609507.4
Class: TopLevelPathway
Species: Homo sapiens (human)
Inferred: no: a curated human pathway
Disease pathway: no
Released: 2018-09-12
Last updated: 2019-03-25
Events inside: 4, counted from the record

Protein localization encompasses the processes that establish and maintain proteins at specific locations. Mechanisms that target proteins to particular locations in the cell typically involve a motif in the targeted protein that interacts with proteins located at the destination (reviewed in Bauer et al. 2015).

Mitochondrial proteins encoded in the nucleus may be targeted to the outer membrane, intermembrane space, inner membrane, or the matrix (reviewed in Kutik et al. 2007, Milenkovic et al. 2007, Bolender et al. 2008, Ender and Yamano 2009, Wiedemann and Pfanner 2017, Kang et al. 2018). A presequence or an internal targeting sequence causes a protein in the cytosol to interact with the TOMM40:TOMM70 complex in the outer mitochondrial membrane. After passage across the outer membrane, sequence motifs cause proteins to be targeted to the outer membrane via the SAMM50 complex, to the inner membrane via the TIMM22 or TIMM23 complexes, to the matrix via the TIMM23 complex, or proteins may fold and remain in the intermembrane space.

All of the proteins contained in the peroxisomal matrix are imported from the cytosol by a unique mechanism that does not require the imported proteins to be unfolded as they cross the membrane (reviewed in Ma et al. 2011, Fujiki et al. 2014, Francisco et al. 2017). In the cytosol, receptor proteins, PEX5 and PEX7, bind to specific sequence motifs in cargo proteins and then interact with a protein complex containing PEX13, PEX14, PEX2, PEX10, and PEX12 in the peroxisome membrane. The cargo proteins then pass through a proteinaceous channel in the membrane and PEX5 is recycled by a mechanism involving ubiquitination and deubiquitination.

Most peroxisomal membrane proteins (PMPs) are inserted into the peroxisomal membrane by the receptor-chaperone PEX19 and the docking receptor PEX3 (reviewed in Ma et al. 2011, Fujiki et al. 2014). PEX19 binds the PMP as it is translated in the cytosol. The PEX19:PMP complex then interacts with PEX3 located in the peroxisomal membrane. Through a mechanism that is not yet clear, the PMP is inserted into the peroxisomal membrane and PEX19 dissociates from PEX3.

Literature Reactome attaches

Mechanisms Regulating Protein LocalizationTraffic, 2015, vol. 16, p. 1039-61PubMed 26172624
Cooperation of translocase complexes in mitochondrial protein importJ Cell Biol, 2007, vol. 179, p. 585-91PubMed 17998403
Diverse mechanisms and machineries for import of mitochondrial proteinsBiol Chem, 2007, vol. 388, p. 891-7PubMed 17696772
Multiple pathways for sorting mitochondrial precursor proteinsEMBO Rep, 2008, vol. 9, p. 42-9PubMed 18174896
Multiple pathways for mitochondrial protein trafficBiol Chem, 2009, vol. 390, p. 723-30PubMed 19453276
Protein transport into peroxisomes: Knowns and unknownsBioessays, 2017, vol. 39, p. 10.1002/bies.201700047PubMed 28787099
Peroxisome assembly: matrix and membrane protein biogenesisJ. Cell Biol., 2011, vol. 193, p. 7-16PubMed 21464226
Peroxisome biogenesis in mammalian cellsFront Physiol, 2014, vol. 5, p. 307PubMed 25177298
Mitochondrial protein transport in health and diseaseSemin. Cell Dev. Biol., 2018, vol. 76, p. 142-153PubMed 28765093
Mitochondrial Machineries for Protein Import and AssemblyAnnu. Rev. Biochem., 2017, vol. 86, p. 685-714PubMed 28301740

Reactome's record lists 14 orthologous events, R-RNO-9609507 (Rattus norvegicus), R-MMU-9609507 (Mus musculus) among the species this site serves, and 12 in species this site opens no page for.

Reactome ContentService, this pathway's record · Reactome v97 · read 2026-10-01 ET · Reactome pathway browser"Protein localization". Reactome, release 97, https://reactome.org/PathwayBrowser/#/R-HSA-9609507 with StableID: R-HSA-9609507.4 (accessed 2026-10-01 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 164 genes in this human pathway; showing 101 to 164, in pages of 100, sorted by symbol for reading. The order carries no ranking.

Genes Reactome places in this human pathway, page 2 of 2
Gene	Authority id	Mapping file id	Evidence codes
GenePEX2	AuthorityHGNC:9717	Mapping file id5828 NCBI file	EvidenceIEA, TAS
GenePEX26	AuthorityHGNC:22965	Mapping file id55670 NCBI file	EvidenceIEA, TAS
GenePEX3	AuthorityHGNC:8858	Mapping file id8504 NCBI file	EvidenceIEA, TAS
GenePEX5	AuthorityHGNC:9719	Mapping file id5830 NCBI file	EvidenceIEA, TAS
GenePEX6	AuthorityHGNC:8859	Mapping file id5190 NCBI file	EvidenceIEA, TAS
GenePEX7	AuthorityHGNC:8860	Mapping file id5191 NCBI file	EvidenceIEA, TAS
GenePHYH	AuthorityHGNC:8940	Mapping file id5264 NCBI file	EvidenceIEA, TAS
GenePIPOX	AuthorityHGNC:17804	Mapping file id51268 NCBI file	EvidenceIEA, TAS
GenePITRM1	AuthorityHGNC:17663	Mapping file id10531 NCBI file	EvidenceTAS
GenePMPCA	AuthorityHGNC:18667	Mapping file id23203 NCBI file	EvidenceIEA
GenePMPCB	AuthorityHGNC:9119	Mapping file id9512 NCBI file	EvidenceIEA
GenePRNP	AuthorityHGNC:9449	Mapping file id5621 NCBI file	EvidenceIEA, TAS
GenePXMP2	AuthorityHGNC:9716	Mapping file id5827 NCBI file	EvidenceTAS
GenePXMP4	AuthorityHGNC:15920	Mapping file id11264 NCBI file	EvidenceTAS
GeneRPS27A	AuthorityHGNC:10417	Mapping file id6233 NCBI file	EvidenceIEA, TAS
GeneSAMM50	AuthorityHGNC:24276	Mapping file id25813 NCBI file	EvidenceIEA
GeneSCP2	AuthorityHGNC:10606	Mapping file id6342 NCBI file	EvidenceIEA, TAS
GeneSEC61B	AuthorityHGNC:16993	Mapping file id10952 NCBI file	EvidenceIEA, TAS
GeneSEC61G	AuthorityHGNC:18277	Mapping file id23480 NCBI file	EvidenceIEA, TAS
GeneSERP1	AuthorityHGNC:10759	Mapping file id27230 NCBI file	EvidenceIEA, TAS
GeneSGTA	AuthorityHGNC:10819	Mapping file id6449 NCBI file	EvidenceIEA, TAS
GeneSLC25A12	AuthorityHGNC:10982	Mapping file id8604 NCBI file	EvidenceIEA
GeneSLC25A13	AuthorityHGNC:10983	Mapping file id10165 NCBI file	EvidenceIEA
GeneSLC25A17	AuthorityHGNC:10987	Mapping file id10478 NCBI file	EvidenceTAS
GeneSLC25A4	AuthorityHGNC:10990	Mapping file id291 NCBI file	EvidenceIEA, TAS
GeneSLC25A6	AuthorityHGNC:10992	Mapping file id293 NCBI file	EvidenceIEA, TAS
GeneSLC27A2	AuthorityHGNC:10996	Mapping file id11001 NCBI file	EvidenceIEA, TAS
GeneSTX1A	AuthorityHGNC:11433	Mapping file id6804 NCBI file	EvidenceIEA, TAS
GeneSTX5	AuthorityHGNC:11440	Mapping file id6811 NCBI file	EvidenceIEA, TAS
GeneTAFAZZIN	AuthorityHGNC:11577	Mapping file id6901 NCBI file	EvidenceIEA, TAS
GeneTIMM10	AuthorityHGNC:11814	Mapping file id26519 NCBI file	EvidenceIEA, TAS
GeneTIMM10B	AuthorityHGNC:4022	Mapping file id26515 NCBI file	EvidenceTAS
GeneTIMM13	AuthorityHGNC:11816	Mapping file id26517 NCBI file	EvidenceIEA
GeneTIMM17A	AuthorityHGNC:17315	Mapping file id10440 NCBI file	EvidenceIEA, TAS
GeneTIMM17B	AuthorityHGNC:17310	Mapping file id10245 NCBI file	EvidenceIEA, TAS
GeneTIMM21	AuthorityHGNC:25010	Mapping file id29090 NCBI file	EvidenceIEA
GeneTIMM22	AuthorityHGNC:17317	Mapping file id29928 NCBI file	EvidenceIEA, TAS
GeneTIMM23	AuthorityHGNC:17312	Mapping file id100287932 NCBI file	EvidenceIEA, TAS
GeneTIMM44	AuthorityHGNC:17316	Mapping file id10469 NCBI file	EvidenceIEA
GeneTIMM50	AuthorityHGNC:23656	Mapping file id92609 NCBI file	EvidenceIEA
GeneTIMM8A	AuthorityHGNC:11817	Mapping file id1678 NCBI file	EvidenceIEA
GeneTIMM8B	AuthorityHGNC:11818	Mapping file id26521 NCBI file	EvidenceIEA
GeneTIMM9	AuthorityHGNC:11819	Mapping file id26520 NCBI file	EvidenceIEA, TAS
GeneTOMM20	AuthorityHGNC:20947	Mapping file id9804 NCBI file	EvidenceIEA
GeneTOMM22	AuthorityHGNC:18002	Mapping file id56993 NCBI file	EvidenceIEA
GeneTOMM40	AuthorityHGNC:18001	Mapping file id10452 NCBI file	EvidenceIEA
GeneTOMM5	AuthorityHGNC:31369	Mapping file id401505 NCBI file	EvidenceIEA
GeneTOMM6	AuthorityHGNC:34528	Mapping file id100188893 NCBI file	EvidenceIEA
GeneTOMM7	AuthorityHGNC:21648	Mapping file id54543 NCBI file	EvidenceIEA
GeneTOMM70	AuthorityHGNC:11985	Mapping file id9868 NCBI file	EvidenceIEA
GeneTYSND1	AuthorityHGNC:28531	Mapping file id219743 NCBI file	EvidenceIEA, TAS
GeneUBA52	AuthorityHGNC:12458	Mapping file id7311 NCBI file	EvidenceIEA, TAS
GeneUBB	AuthorityHGNC:12463	Mapping file id7314 NCBI file	EvidenceIEA, TAS
GeneUBC	AuthorityHGNC:12468	Mapping file id7316 NCBI file	EvidenceIEA, TAS
GeneUBE2D1	AuthorityHGNC:12474	Mapping file id7321 NCBI file	EvidenceTAS
GeneUBE2D2	AuthorityHGNC:12475	Mapping file id7322 NCBI file	EvidenceTAS
GeneUBE2D3	AuthorityHGNC:12476	Mapping file id7323 NCBI file	EvidenceTAS
GeneUBE2J2	AuthorityHGNC:19268	Mapping file id118424 NCBI file	EvidenceIEA, TAS
GeneUBL4A	AuthorityHGNC:12505	Mapping file id8266 NCBI file	EvidenceIEA
GeneUSP9X	AuthorityHGNC:12632	Mapping file id8239 NCBI file	EvidenceIEA, TAS
GeneVAMP2	AuthorityHGNC:12643	Mapping file id6844 NCBI file	EvidenceIEA, TAS
GeneVAPA	AuthorityHGNC:12648	Mapping file id9218 NCBI file	EvidenceIEA, TAS
GeneVDAC1	AuthorityHGNC:12669	Mapping file id7416 NCBI file	EvidenceIEA
GeneZFAND6	AuthorityHGNC:30164	Mapping file id54469 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

None: this is a top-level pathway of the release.

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.