Affinage

TOMM5

Mitochondrial import receptor subunit TOM5 homolog · UniProt Q8N4H5

Round 2 corrected
Length
51 aa
Mass
6.0 kDa
Annotated
2026-04-28
73 papers in source corpus 30 papers cited in narrative 30 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

TOMM5 is a small, C-tail-anchored subunit of the translocase of the outer mitochondrial membrane (TOM) complex that contributes to the structural integrity of the general import pore (GIP) and facilitates preprotein transfer from surface receptors to the Tom40 translocation channel (PMID:9774667, PMID:10397776). Its transmembrane segment surrounds the Tom40 β-barrel in the dimeric TOM core complex, as demonstrated by cryo-EM structures across fungi, Drosophila, and humans (PMID:28802041, PMID:33083003, PMID:39575538). TOMM5 acts as an assembly factor that promotes Tom40 biogenesis at the SAM complex, interacts with EMC proteins to form an ER–mitochondria tether important for phosphatidylserine transfer, and stabilizes PINK1 at the outer membrane during PINK1-Parkin-mediated mitophagy (PMID:20668160, PMID:25313861, PMID:40080546). Tomm5-knockout mice develop cryptogenic organizing pneumonia, linked to dysregulated mitochondrial membrane potential in alveolar epithelial cells (PMID:22688586, PMID:38794801).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 1998 High

    Establishing Tom5 as a core subunit of the TOM GIP complex resolved the identity of the small proteins surrounding Tom40 and showed Tom5 participates in preprotein transfer from receptors to the channel.

    Evidence Biochemical characterization, blue native PAGE, co-immunoprecipitation, and yeast mutant analysis of TOM complex components

    PMID:9603986 PMID:9774667

    Open questions at the time
    • Precise binding site on Tom40 unknown
    • Contribution of Tom5 versus other small Toms not individually resolved
  2. 1999 High

    Demonstration that Tom5 is essential for import of small Tim IMS proteins—but dispensable for cytochrome c import—defined substrate-selective roles within the GIP complex.

    Evidence In organello import assays in yeast mutants lacking individual Tom proteins

    PMID:10397776

    Open questions at the time
    • Mechanism of selectivity at Tom5 not structurally resolved
    • Whether selectivity is conserved in mammals unknown
  3. 2001 High

    Identification of a Tom5-containing ~250 kDa assembly intermediate showed that Tom5 participates early in Tom40 biogenesis, prior to formation of the mature GIP.

    Evidence Pulse-chase import assays with native PAGE in yeast

    PMID:11259583 PMID:11276259

    Open questions at the time
    • Order of small Tom association during assembly not fully kinetically resolved
    • Role of lipid environment in assembly intermediate formation unclear
  4. 2002 High

    Systematic mutagenesis of the Tom5 C-tail anchor defined the targeting determinants (TMS length, internal proline, charged C-terminal residues) for outer membrane insertion of C-tail-anchored mitochondrial proteins.

    Evidence GFP fusions with systematic deletions/mutations expressed in yeast and mammalian COS-7 cells; confocal microscopy and fractionation

    PMID:12006657 PMID:12896971

    Open questions at the time
    • Receptor or insertase for Tom5 biogenesis not identified
    • Whether MIM/Mim1 pathway inserts Tom5 itself not tested
  5. 2008 High

    Identification of human TOMM5 and TOMM6 as bona fide TOM complex subunits, and demonstration that combinatorial knockdown of small Tom proteins impairs matrix import, extended the functional framework from yeast to humans.

    Evidence FLAG-immunoisolation and mass spectrometry from HeLa cells; siRNA double knockdowns with import assays

    PMID:18331822

    Open questions at the time
    • Individual contribution of human TOMM5 versus TOMM6/TOMM7 not separately quantified
    • Human assembly intermediates not characterized
  6. 2010 High

    Genetic epistasis showed Tom5 acts downstream of Mim1 to promote the second SAM-stage interaction during Tom40 biogenesis, and that Tom5/Tom6 stimulate while Tom7 antagonizes this step, revealing opposing modulatory roles of the small Toms.

    Evidence Blue native PAGE, import assays, epistasis analysis in yeast single and double mutants

    PMID:20668160 PMID:21059357

    Open questions at the time
    • Direct structural contacts between Tom5 and SAM complex not resolved
    • Mechanism by which Tom7 opposes Tom5 function unknown
  7. 2012 Medium

    Tomm5-knockout mice developed lung-specific cryptogenic organizing pneumonia, establishing the first in vivo mammalian phenotype and suggesting a tissue-selective role for TOMM5 in alveolar homeostasis.

    Evidence Knockout mouse model with histopathology

    PMID:22688586

    Open questions at the time
    • Molecular mechanism linking TOMM5 loss to pneumonia not established
    • Import defects in alveolar cells not directly measured
    • Single mouse model without independent replication
  8. 2014 High

    Discovery that Tom5 interacts with all EMC subunits and that this interaction mediates phosphatidylserine transfer from ER to mitochondria revealed a non-canonical tethering function for a TOM subunit at ER–mitochondria contact sites.

    Evidence Genetic screen, reciprocal co-immunoprecipitation, lipid transfer assays, and growth assays in yeast

    PMID:25313861

    Open questions at the time
    • Whether the EMC-Tom5 tether is conserved in mammals not tested
    • Structural basis of Tom5–EMC interaction unknown
    • Relative contribution to total ER–mitochondria PS flux unclear
  9. 2020 High

    Atomic-resolution cryo-EM structures of the human and Neurospora TOM core complexes placed TOMM5 at the periphery of the Tom40 β-barrel and showed the Tom40 N-terminal segment contacts Tom5 at the IMS face, providing a structural rationale for preprotein hand-off.

    Evidence Single-particle cryo-EM at near-atomic resolution

    PMID:28802041 PMID:33083003

    Open questions at the time
    • Dynamics of Tom5 during active translocation not captured
    • Lipid interactions at the Tom5–Tom40 interface not fully modeled
  10. 2024 Medium

    Functional studies in alveolar epithelial cells showed that TOMM5 regulates mitochondrial membrane potential, suppresses early apoptosis, and promotes proliferation, mechanistically linking TOMM5 to the organizing pneumonia phenotype observed in knockout mice.

    Evidence In vitro knockdown/overexpression with membrane potential assays, flow cytometry for apoptosis, and bleomycin mouse model

    PMID:38794801

    Open questions at the time
    • Whether membrane potential effect is direct or secondary to import defects not resolved
    • Specific import substrates affected in alveolar cells unknown
  11. 2025 High

    High-resolution cryo-EM of PINK1 at a TOM-VDAC array showed TOMM5 directly contacts the PINK1 kinase C-lobe, and genetic ablation confirmed the TOM complex is required for PINK1 retention during mitophagy, establishing a direct structural role for TOMM5 in the PINK1-Parkin pathway.

    Evidence Cryo-EM at 3.1 Å of endogenous TOM-VDAC complex (Science); CRISPR screen and genetic ablation with PINK1 retention assays (EMBO J); Drosophila cryo-EM confirming conserved Tom5 architecture (IUCrJ)

    PMID:39575538 PMID:40080546 PMID:41266657

    Open questions at the time
    • Whether TOMM5 loss alone (versus full TOM ablation) is sufficient to prevent PINK1 retention not individually tested
    • How TOMM5–PINK1 interaction is regulated by membrane potential not known

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key open questions include: whether the Tom5–EMC tethering function is conserved in mammals; the structural basis for substrate selectivity at the Tom5 stage of import; and how TOMM5-dependent PINK1 stabilization is coordinated with chaperone release and membrane potential sensing.
  • No mammalian EMC–TOMM5 interaction data
  • No time-resolved structure of translocation through Tom5-containing pore
  • Individual TOMM5 knockout in human cells for PINK1 pathway not reported

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005198 structural molecule activity 5 GO:0005215 transporter activity 3 GO:0060090 molecular adaptor activity 3
Localization
GO:0005739 mitochondrion 8
Pathway
R-HSA-1852241 Organelle biogenesis and maintenance 6 R-HSA-9609507 Protein localization 5 R-HSA-392499 Metabolism of proteins 3 R-HSA-9612973 Autophagy 2 R-HSA-382551 Transport of small molecules 1
Partners
EMC1PINK1TOMM22TOMM40TOMM6TOMM7VDAC2
Complex memberships
SAM-Tom5-Tom40 assembly intermediateTOM complex (GIP/TOM core)

Evidence

Reading pass · 30 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1998 Tom5 in the yeast TOM complex recognizes the mitochondria-targeting sequence (MTS) of preproteins, functioning as a component of the outer membrane protein translocation machinery that mediates transfer of preproteins from receptors to the Tom40 channel. Biochemical characterization of TOM complex components; reconstitution of preprotein import pathway in yeast Journal of biochemistry Medium 9603986
1998 Tom5, Tom6, and Tom7 are small subunits of the ~400 kDa general import pore (GIP) complex of yeast mitochondria, which also contains Tom40 and Tom22. Tom6 promotes stable association of Tom22 with Tom40, and its absence causes dissociation of Tom22 and formation of a ~100 kDa subcomplex containing Tom40, Tom7, and Tom5. Blue native PAGE, co-immunoprecipitation, yeast mutant analysis Molecular and cellular biology High 9774667
2001 Tom5 participates in the assembly of the yeast Tom40 import channel: the Tom40 precursor first assembles with Tom5 to form a ~250 kDa intermediate exposed to the intermembrane space, before progression to the mature ~400 kDa GIP complex. Pulse-chase import assays, native PAGE, sequential assembly intermediate analysis Nature structural biology High 11276259
2001 Tom5 is part of the stable TOM GIP core complex together with Tom40 and Tom22. Under stringent detergent conditions, Tom5 (along with Tom20 and other small Toms) is released while preprotein remains in the GIP, indicating Tom5 is not essential for preprotein holding but contributes to complex architecture. Urea/alkaline resistance assays, detergent fractionation, preprotein arrest experiments, electrophysiology Molecular and cellular biology High 11259583
2001 Biogenesis of yeast porin (VDAC) depends on Tom5 of the GIP complex, in addition to Tom20, Tom22, and Tom40, as shown by import competition and mutant analysis. In organello import assays, competition assays, yeast mutants lacking individual Tom proteins The Journal of cell biology High 11266446
1999 Import of small Tim proteins of the mitochondrial IMS uses a novel pathway where surface receptors Tom20 and Tom70 are dispensable, but Tom5 of the GIP complex is crucial, defining a third import route. In organello import assays in yeast mutants lacking individual Tom proteins Molecular biology of the cell High 10397776
2002 Insertion of bacterial porin PorB into the mitochondrial outer membrane in vitro depends on Tom5, Tom20, and Tom40, but is independent of Tom70, demonstrating a shared import mechanism with VDAC. In vitro import assays into isolated mitochondria; antibody inhibition of specific TOM subunits The EMBO journal Medium 11953311
2002 Bcl-2alpha insertion into the yeast mitochondrial outer membrane does not require Tom5 or Tom40, demonstrating that Bcl-2alpha bypasses the GIP and follows a pathway distinct from that requiring Tom5. In organello import assays in yeast tom5 and tom40 mutants Journal of molecular biology Medium 12419260
2002 Yeast Tom5 is a C-tail-anchored protein; the signal directing it to the mitochondrial outer membrane requires an appropriate TMS length, a proline at a correct position within the TMS, and specific surrounding residues, but (unlike dispersed outer membrane proteins) does not require a positive C-terminal segment. GFP reporter fusions with systematic deletions/mutations, confocal microscopy, cell fractionation, blue native PAGE complementation in tom5-ts yeast The Journal of biological chemistry High 12896971
2002 The mitochondrial targeting signal for C-tail-anchored proteins in mammals, using yeast Tom5 as a model in COS-7 cells, requires three basic amino acid residues in the C-terminal five-residue segment and an appropriate TMS length; elongation of TMS or separation of TMS and C-segment impairs targeting. GFP reporter fusions expressed in COS-7 cells, confocal microscopy, cell fractionation Molecular biology of the cell High 12006657
2003 Import of cytochrome c into yeast mitochondria does not require Tom5, Tom6, or Tom7, establishing that these small Tom proteins are dispensable for this particular import pathway. In organello import assays in yeast mutants lacking individual Tom proteins Journal of molecular biology Medium 12628251
2005 Identification of Neurospora crassa Tom5 as a TOM complex subunit with its C-terminus facing the IMS. In yeast, Tom5 is required for structural stability of the TOM complex and efficient protein import, but Neurospora Tom5 knockout shows no growth or import defect, indicating organism-specific roles. Yeast TOM5 deletion can be rescued by overexpression of Neurospora Tom5. Identification by sequence analysis and biochemistry; tom5 deletion in both yeast and Neurospora; import assays; blue native PAGE; complementation experiments The Journal of biological chemistry High 15701639
2005 Import of yeast Taz1 (tafazzin) into mitochondria depends on the receptor Tom5 of the TOM complex and the small Tim proteins of the IMS, but is independent of the SAM complex. In organello import assays in yeast mutants; submitochondrial fractionation Molecular biology of the cell Medium 16135531
2000 The cytosolic domain of yeast TOM5 forms a stable helical structure: CD spectroscopy shows a pH-invariant helical conformation, and NMR NOESY data reveal a stable helical core between residues E11 and R15 with a less rigid helix extending to the C-terminus. Circular dichroism (CD) spectroscopy, NMR (NOESY) FEBS letters Medium 10683449
2009 A subcomplex of Tom5 and Tom40 associates with the SAM core complex to form a large SAM complex involved in biogenesis of the alpha-helical Tom6 protein after Mim1-dependent membrane insertion. Blue native PAGE, co-immunoprecipitation, import assays in yeast mutants Journal of molecular biology High 20026336
2010 Tom5 promotes the second stage of Tom40 assembly at the SAM complex. Mim1-deficient mitochondria accumulate Tom40 at the first SAM stage like Tom5-deficient mitochondria; overexpression of Tom5 suppresses the Tom40 assembly defect of mim1Δ mitochondria, placing Tom5 downstream of Mim1 in the Tom40 biogenesis pathway. Import assays, blue native PAGE, epistasis analysis in yeast mutants Molecular biology of the cell High 20668160
2010 Tom5 and Tom6 play a stimulatory role in biogenesis of Tom40 at the SAM complex, antagonized by Tom7; Tom5 and Tom6 associate with the Tom40 precursor at an early assembly stage, while Tom7 inhibits this step and additionally promotes dissociation of the SAM-Mdm10 complex. Import assays, blue native PAGE, genetic epistasis in yeast mutants Journal of molecular biology High 21059357
2008 Tom5 (along with Tom22, Tom7, and Tom6) functions as a modulator of the pore dynamics of Tom40, significantly reducing the energy barrier between different conformational states of the TOM channel. Planar lipid bilayer electrophysiology with purified TOM core complex and Tom40 from Neurospora crassa Biophysical journal Medium 18456827
2008 Human Tom5 and Tom6 were identified as components of the human TOM complex by immunoisolation from HeLa cells. They associate with Tom40 in the TOM complex. Knockdown of hTom40 decreases levels of all small Tom proteins. Double knockdown of any combination of small Tom proteins (Tom5, Tom6, Tom7) affects matrix import of preproteins. FLAG-immunoisolation of TOM complex from HeLa cells, mass spectrometry identification, siRNA knockdown, import assays Biochemical and biophysical research communications High 18331822
2014 Yeast Tom5 of the TOM complex interacts with all Emc proteins of the ER membrane protein complex (EMC), and this interaction is important for phosphatidylserine (PS) transfer from the ER to mitochondria and for cell growth, suggesting that the EMC forms an ER-mitochondria tether by associating with the TOM complex through Tom5. Genetic screen, co-immunoprecipitation, lipid transfer assays, growth assays in yeast PLoS biology High 25313861
2017 Cryo-EM structure of the Neurospora crassa TOM core complex at ~10 Å resolution shows Tom5, Tom6, and Tom7 transmembrane segments surrounding each Tom40 β-barrel pore in the dimeric complex, with Tom22 connecting the two Tom40 pores at the dimer interface. Cryo-electron microscopy, single particle analysis Cell High 28802041
2020 Atomic resolution cryo-EM structure of the dimeric human TOM core complex shows TOMM5, TOMM6, and TOMM7 surrounding the Tom40 channels at the periphery of the dimer; the N-terminal segment of Tom40 spans from cytosol to IMS to interact with Tom5 at the dimer periphery, providing insight into preprotein translocation paths. Single-particle cryo-EM at near-atomic resolution Cell discovery High 33083003
2012 Tomm5 knockout mice (Tomm5−/−) develop a lung-specific phenotype of cryptogenic organizing pneumonia (COP/BOOP), characterized by intra-alveolar fibrosis with fibroblasts/myofibroblasts in alveolar lumina and eosinophilic inflammation, while other organ systems appear normal. Knockout mouse model; histopathology Veterinary pathology Medium 22688586
2024 TOMM5 regulates mitochondrial membrane potential in alveolar epithelial cells. In a bleomycin-induced murine model of organizing pneumonia, TOMM5 levels increase with lung fibrosis. In vitro, TOMM5 reduces the proportion of early apoptotic cells and promotes cell proliferation. In vitro knockdown/overexpression in alveolar epithelial cells; mitochondrial membrane potential assays; flow cytometry for apoptosis; bleomycin mouse model Redox report Medium 38794801
2025 Cryo-EM structure of human PINK1 at a TOM-VDAC array shows that TOM5 participates in symmetric arrangement of two TOM core complexes around a central VDAC2 dimer, and that TOM5 binds the PINK1 kinase C-lobe, stabilizing PINK1 at the TOM complex during mitophagy. Single-particle cryo-EM at 3.1 Å resolution of endogenous TOM-VDAC complex; structural analysis of PINK1 interaction Science High 40080546
2025 TOMM5 (as a TOM complex subunit) is required for PINK1 retention on the mitochondrial surface during mitophagy. Ablation of TOM (including TOMM5) prevents PINK1 accumulation at the outer membrane when membrane potential is lost, establishing TOM as the platform for PINK1 stabilization. Genome-wide CRISPR screen with novel Parkin reporter; genetic ablation of TOM subunits; PINK1 import/retention assays The EMBO journal Medium 41266657
2025 The HSP90-CDC37 chaperone complex holds PINK1 in a partially unfolded state; the C-terminal extension (CTE) of PINK1 is covered by HSP90 in a region that overlaps with the TOM5 and TOM20 interaction sites, suggesting that chaperone release is coupled to TOM engagement for PINK1 import/stabilization. Cryo-EM structure of human PINK1-HSP90-CDC37 complex bioRxivpreprint Low bio_10.1101_2025.10.17.682828
2025 The Drosophila TOM complex cryo-EM structure at 3.3 Å shows Tom5 as one of four endogenous TOM components co-assembled with Tom40, confirming evolutionary conservation of Tom5's position surrounding the Tom40 β-barrel. Small conformational differences at subunit interfaces relative to human TOM are attributable to lipid-binding residue variation. Single-particle cryo-EM of ex vivo Drosophila TOM complex IUCrJ High 39575538
2011 A CTCF-mediated insulator loop encompassing the TOMM5 gene resides between synthetically interacting genetic elements of the breast cancer susceptibility locus MCS5A/Mcs5a, suggesting TOMM5 is located within a higher-order chromatin structure relevant to locus regulation. CTCF ChIP, chromatin conformation capture (3C), transgenic rat models Nucleic acids research Low 21914726
2011 Proapoptotic fusion protein p53-Tom5, in which wild-type p53 is fused to the mitochondrial transmembrane domain of Tom5, exclusively localizes to mitochondria in ARF-null A549 lung cancer cells, induces mitochondrial dysfunction and cytochrome c release, and suppresses cell proliferation — effects not seen with wild-type p53 alone. Plasmid transfection; confocal microscopy localization; cell proliferation assays; cytochrome c release assay Biological & pharmaceutical bulletin Low 21467644

Source papers

Stage 0 corpus · 73 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2002 Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences. Proceedings of the National Academy of Sciences of the United States of America 1479 12477932
2015 A human interactome in three quantitative dimensions organized by stoichiometries and abundances. Cell 1015 26496610
2021 Dual proteome-scale networks reveal cell-specific remodeling of the human interactome. Cell 705 33961781
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2004 The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC). Genome research 438 15489334
2022 OpenCell: Endogenous tagging for the cartography of human cellular organization. Science (New York, N.Y.) 432 35271311
1996 Normalization and subtraction: two approaches to facilitate gene discovery. Genome research 401 8889548
2021 A proximity-dependent biotinylation map of a human cell. Nature 339 34079125
2014 A conserved endoplasmic reticulum membrane protein complex (EMC) facilitates phospholipid transfer from the ER to mitochondria. PLoS biology 262 25313861
2021 Quantitative high-confidence human mitochondrial proteome and its dynamics in cellular context. Cell metabolism 239 34800366
2016 Mitochondrial Protein Interaction Mapping Identifies Regulators of Respiratory Chain Function. Molecular cell 220 27499296
1993 Identification and genetic analysis of normal and mutant phytoene synthase genes of tomato by sequencing, complementation and co-suppression. Plant molecular biology 213 8343597
2016 Ubiquilins Chaperone and Triage Mitochondrial Membrane Proteins for Degradation. Molecular cell 210 27345149
1998 Preprotein translocase of the outer mitochondrial membrane: molecular dissection and assembly of the general import pore complex. Molecular and cellular biology 210 9774667
2005 Taz1, an outer mitochondrial membrane protein, affects stability and assembly of inner membrane protein complexes: implications for Barth Syndrome. Molecular biology of the cell 174 16135531
2001 Multistep assembly of the protein import channel of the mitochondrial outer membrane. Nature structural biology 165 11276259
2020 A High-Density Human Mitochondrial Proximity Interaction Network. Cell metabolism 148 32877691
2001 Protein import channel of the outer mitochondrial membrane: a highly stable Tom40-Tom22 core structure differentially interacts with preproteins, small tom proteins, and import receptors. Molecular and cellular biology 145 11259583
2017 Cryo-EM Structure of the TOM Core Complex from Neurospora crassa. Cell 141 28802041
2001 Biogenesis of porin of the outer mitochondrial membrane involves an import pathway via receptors and the general import pore of the TOM complex. The Journal of cell biology 134 11266446
2011 Interactions of pathological hallmark proteins: tubulin polymerization promoting protein/p25, beta-amyloid, and alpha-synuclein. The Journal of biological chemistry 131 21832049
1998 Mitochondria-targeting sequence, a multi-role sorting sequence recognized at all steps of protein import into mitochondria. Journal of biochemistry 128 9603986
2002 Characterization of signal that directs C-tail-anchored proteins to mammalian mitochondrial outer membrane. Molecular biology of the cell 127 12006657
2007 Toward a confocal subcellular atlas of the human proteome. Molecular & cellular proteomics : MCP 114 18029348
2021 Protein interaction landscapes revealed by advanced in vivo cross-linking-mass spectrometry. Proceedings of the National Academy of Sciences of the United States of America 113 34349018
2020 Atomic structure of human TOM core complex. Cell discovery 104 33083003
2018 Histone Interaction Landscapes Visualized by Crosslinking Mass Spectrometry in Intact Cell Nuclei. Molecular & cellular proteomics : MCP 101 30021884
1999 Biogenesis of Tim proteins of the mitochondrial carrier import pathway: differential targeting mechanisms and crossing over with the main import pathway. Molecular biology of the cell 101 10397776
2021 SARS-CoV-2-host proteome interactions for antiviral drug discovery. Molecular systems biology 86 34709727
2004 DNA sequence and analysis of human chromosome 9. Nature 86 15164053
2009 Two modular forms of the mitochondrial sorting and assembly machinery are involved in biogenesis of alpha-helical outer membrane proteins. Journal of molecular biology 85 20026336
2010 Biogenesis of mitochondria: dual role of Tom7 in modulating assembly of the preprotein translocase of the outer membrane. Journal of molecular biology 74 21059357
2002 VDAC and the bacterial porin PorB of Neisseria gonorrhoeae share mitochondrial import pathways. The EMBO journal 73 11953311
2002 Bcl-2 and porin follow different pathways of TOM-dependent insertion into the mitochondrial outer membrane. Journal of molecular biology 65 12419260
2023 Intratumoral microbial heterogeneity affected tumor immune microenvironment and determined clinical outcome of HBV-related HCC. Hepatology (Baltimore, Md.) 58 37114494
2010 Assembly of the mitochondrial protein import channel: role of Tom5 in two-stage interaction of Tom40 with the SAM complex. Molecular biology of the cell 56 20668160
2022 Therapeutic targeting of the USP2-E2F4 axis inhibits autophagic machinery essential for zinc homeostasis in cancer progression. Autophagy 53 35253629
2005 Role of Tom5 in maintaining the structural stability of the TOM complex of mitochondria. The Journal of biological chemistry 50 15701639
2008 Identification of Tom5 and Tom6 in the preprotein translocase complex of human mitochondrial outer membrane. Biochemical and biophysical research communications 47 18331822
2003 Targeting and assembly of mitochondrial tail-anchored protein Tom5 to the TOM complex depend on a signal distinct from that of tail-anchored proteins dispersed in the membrane. The Journal of biological chemistry 47 12896971
2019 Rewiring of the Human Mitochondrial Interactome during Neuronal Reprogramming Reveals Regulators of the Respirasome and Neurogenesis. iScience 45 31536960
2021 An antibody-based proximity labeling map reveals mechanisms of SARS-CoV-2 inhibition of antiviral immunity. Cell chemical biology 35 34672954
2025 Structure of human PINK1 at a mitochondrial TOM-VDAC array. Science (New York, N.Y.) 33 40080546
2003 Biogenesis of yeast mitochondrial cytochrome c: a unique relationship to the TOM machinery. Journal of molecular biology 32 12628251
2000 The transport machinery for the import of preproteins across the outer mitochondrial membrane. The international journal of biochemistry & cell biology 32 10661891
2021 The receptor subunit Tom20 is dynamically associated with the TOM complex in mitochondria of human cells. Molecular biology of the cell 30 34347503
1995 Isolation and characterisation of a melon cDNA clone encoding phytoene synthase. Plant molecular biology 25 7766896
2016 Nucleo-mitochondrial interaction of yeast in response to cadmium sulfide quantum dot exposure. Journal of hazardous materials 23 27890358
2008 Dynamics of the preprotein translocation channel of the outer membrane of mitochondria. Biophysical journal 22 18456827
2020 Structural snapshot of the mitochondrial protein import gate. The FEBS journal 18 33305524
2017 Pharmacogenetic meta-analysis of baseline risk factors, pharmacodynamic, efficacy and tolerability endpoints from two large global cardiovascular outcomes trials for darapladib. PloS one 17 28753643
2024 Ribosome Profiling and Mass Spectrometry Reveal Widespread Mitochondrial Translation Defects in a Striatal Cell Model of Huntington Disease. Molecular & cellular proteomics : MCP 15 38447791
2011 An insulator loop resides between the synthetically interacting elements of the human/rat conserved breast cancer susceptibility locus MCS5A/Mcs5a. Nucleic acids research 15 21914726
2022 CLEC16A interacts with retromer and TRIM27, and its loss impairs endosomal trafficking and neurodevelopment. Human genetics 14 36538041
2010 LILBID-mass spectrometry of the mitochondrial preprotein translocase TOM. Journal of physics. Condensed matter : an Institute of Physics journal 14 21339618
2011 Genetic and epigenetic variations contributed by Alu retrotransposition. BMC genomics 13 22185517
2012 Alterations in metabolism-related genes induced in SHSY5Y cells by okadaic acid exposure. Journal of toxicology and environmental health. Part A 12 22788371
2022 Structural overview of the translocase of the mitochondrial outer membrane complex. Biophysics and physicobiology 10 35859989
2022 Interaction of LATS1 with SMAC links the MST2/Hippo pathway with apoptosis in an IAP-dependent manner. Cell death & disease 10 35941108
2012 Cryptogenic organizing pneumonia in Tomm5(-/-) mice. Veterinary pathology 9 22688586
2024 Identification of MAP1LC3A as a promising mitophagy-related gene in polycystic ovary syndrome. Scientific reports 7 39043888
2000 Structure of the cytosolic domain of TOM5, a mitochondrial import protein. FEBS letters 7 10683449
2023 Diagnostic model based on key autophagy-related genes in intervertebral disc degeneration. BMC musculoskeletal disorders 6 38041088
2011 Proapoptotic action of p53-Tom5 in p53-resistant A549 human non-small cell lung cancer cells through direct mitochondrial dysfunction. Biological & pharmaceutical bulletin 5 21467644
2025 Structure of an ex vivoDrosophila TOM complex determined by single-particle cryoEM. IUCrJ 2 39575538
2025 Human protein interaction networks of ancestral and variant SARS-CoV-2 in organ-specific cells and bodily fluids. Nature communications 2 40593736
2025 A unified mechanism for mitochondrial damage sensing in PINK1-Parkin-mediated mitophagy. The EMBO journal 1 41266657
2024 TOM5 regulates the mitochondrial membrane potential of alveolar epithelial cells in organizing pneumonia. Redox report : communications in free radical research 1 38794801
2026 Differentiation-dependent proximity proteomics identifies novel host factors linked to HPV16 E2 function. mBio 0 41524403
2025 Mapping the intracellular HMGB1 interactome and alterations induced by Toll-like receptor 4 activation. The Journal of biological chemistry 0 41161382
2024 Identification of new therapeutic targets related to endoplasmic reticulum stress and mitochondrial dysfunction to reduce the risk of rupture in degenerative ascending aortic aneurysm. Clinica e investigacion en arteriosclerosis : publicacion oficial de la Sociedad Espanola de Arteriosclerosis 0 39424523
2022 Isolation of Plant Mitochondria Using Affinity Purification. Methods in molecular biology (Clifton, N.J.) 0 34545483
2022 In Vivo Epitope Tagging of Plant Mitochondria. Methods in molecular biology (Clifton, N.J.) 0 35188666