Affinage

SNX4

Sorting nexin-4 · UniProt O95219

Length
450 aa
Mass
51.9 kDa
Annotated
2026-06-10
11 papers in source corpus 11 papers cited in narrative 11 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SNX4 is a PI(3)P-associated sorting nexin that coordinates endosomal cargo sorting, tubulation-driven membrane recycling, and autophagosome biogenesis (PMID:17994011, PMID:32513819). Through its PX and BAR domains it associates with tubular and vesicular elements of early endosomes and the juxtanuclear endocytic recycling compartment, where it sorts cargo such as the transferrin receptor away from lysosomal degradation (PMID:17994011). SNX4 couples this tubulation-based sorting to long-range microtubule-based transport by interacting with KIBRA to engage minus-end-directed dynein and by binding clathrin through a clathrin-box variant motif, with both interactions dependent on PI3-kinase activity (PMID:17994011, PMID:19529763). In yeast the ortholog forms two functionally distinct heterodimers, Snx4-Atg20 and Snx4-Snx41, that coat separate endosome-derived tubules to retrieve distinct cargoes (Snc1 and Atg27), with Vps1 promoting tubule fission (PMID:12554655, PMID:28026081). In mammalian cells SNX4 assembles analogous heterodimers with SNX7 or SNX30; the autophagy-specific SNX4-SNX7 complex supports LC3 lipidation and recycles the lipid scramblase ATG9A from endolysosomes to early endosomes, sustaining the ATG9A pool required for starvation-induced autophagosome assembly (PMID:32513819, PMID:33468622). In yeast SNX4 additionally drives selective autophagy of the 26S proteasome and of transcription factors (Ssn2/Med13, Rim15, Msn2) by delivering cargo to Atg17-initiated phagophores (PMID:29109144, PMID:33678121, PMID:37733395). In neurons SNX4 acts as a negative regulator of synaptic vesicle docking, limiting docked vesicle density and neurotransmitter release at the active zone (PMID:39699951).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2003 High

    Established that the SNX4 ortholog defines a retromer-independent retrieval route, answering whether SNX4-family proteins sort specific cargo back to the Golgi from early endosomes.

    Evidence Chemical cross-linking of Snc1p to Snx4p with yeast deletion mutants and fractionation

    PMID:12554655

    Open questions at the time
    • Did not define the structural basis of cargo selection
    • Mammalian relevance not established at this point
  2. 2007 High

    Defined SNX4 as an endosomal sorting/recycling factor that protects the transferrin receptor from degradation and linked its membrane tubulation to dynein-based transport, answering how sorted cargo reaches the recycling compartment.

    Evidence siRNA knockdown with TfnR degradation phenotype plus reciprocal Co-IP of SNX4-KIBRA-dynein in mammalian cells

    PMID:17994011

    Open questions at the time
    • Mechanism of KIBRA-mediated motor recruitment not resolved at molecular level
    • Did not address autophagy roles
  3. 2009 Medium

    Identified the clathrin-binding determinant on SNX4 and showed PI3-kinase dependence of its dynein/clathrin associations, clarifying how SNX4 carrier transport is regulated.

    Evidence Peptide pulldown defining a clathrin-box variant motif, Co-IP, and wortmannin inhibition with toxin trafficking assays

    PMID:19529763

    Open questions at the time
    • Single lab
    • Functional consequence of clathrin regulation on physiological cargo not fully defined
  4. 2013 Medium

    Placed reggie-1/flotillin-2 upstream of the Rab11a/SNX4 recycling pathway, addressing what regulates SNX4-dependent recycling of TfR and E-cadherin.

    Evidence Direct Co-IP plus shRNA knockdown with overexpression rescue by Rab11a or SNX4 in mammalian cells

    PMID:23825023

    Open questions at the time
    • Single lab
    • Direct biochemical hierarchy among reggie-1, Rab11a, and SNX4 not fully resolved
  5. 2017 Medium

    Resolved SNX4 into two cargo-specific heterodimers and linked their tubules to Vps1-mediated fission, answering how a single nexin sorts distinct cargoes.

    Evidence Live-cell imaging of distinct tubules with cargo-specific trafficking assays and deletions in yeast

    PMID:28026081

    Open questions at the time
    • Single lab
    • Did not establish whether mammalian heterodimers segregate cargo identically
  6. 2017 Medium

    Extended SNX4 function to selective autophagy of large complexes by showing it is required for cytoplasmic proteasome agglomeration prior to autophagic delivery.

    Evidence Targeted autophagy gene screen and fluorescence microscopy with proteasome puncta readout in yeast

    PMID:29109144

    Open questions at the time
    • Mechanism of proteasome capture by Snx4 not defined
    • Single organism
  7. 2020 High

    Established the mammalian SNX4-SNX7 heterodimer as an autophagy-specific complex required for LC3 lipidation and proper ATG9A trafficking, answering whether SNX4 contributes to autophagosome assembly in human cells.

    Evidence CRISPR knockout and siRNA with autophagy phenotypes, Co-IP heterodimer identification, and ATG9A imaging

    PMID:32513819

    Open questions at the time
    • Did not define the precise membrane source supported by SNX4-SNX7
    • Structural basis of heterodimer specificity unresolved
  8. 2021 Medium

    Defined the directionality of SNX4-dependent ATG9A recycling (endolysosome to early endosome, then retromer to TGN), explaining how SNX4 sustains the ATG9A pool for autophagy.

    Evidence siRNA knockdown with compartment-specific ATG9A accumulation and autophagic flux assays

    PMID:33468622

    Open questions at the time
    • Single lab
    • Did not resolve how SNX4 and retromer hand off ATG9A
  9. 2021 Medium

    Defined a selective autophagy pathway (SAA-TF) in which Snx4-Atg20 binds Atg17 and delivers transcription-factor cargo to vacuole-anchored phagophores.

    Evidence Yeast deletion analysis, fluorescence microscopy, and Co-IP for Snx4-Atg17 interaction

    PMID:33678121

    Open questions at the time
    • Single lab
    • Cargo recognition mechanism for individual transcription factors not defined
  10. 2024 Medium

    Ordered the temporal steps of the SAA-TF pathway, showing receptor (Ksp1) recruitment precedes Snx4-mediated cargo delivery to phagophores.

    Evidence Yeast two-hybrid, genetic epistasis, and AIM/LIR motif mutagenesis in yeast

    PMID:37733395

    Open questions at the time
    • Single lab
    • Conservation of this temporal logic in mammals unknown
  11. 2024 High

    Revealed a neuron-specific role, establishing SNX4 as a negative regulator of synaptic vesicle docking and neurotransmitter release independent of its autophagy and recycling roles.

    Evidence Conditional knockout mouse with EM ultrastructure and electrophysiology of evoked release

    PMID:39699951

    Open questions at the time
    • Molecular mechanism by which SNX4 restrains docking not defined
    • Cargo or partner mediating active-zone effect unidentified

Open questions

Synthesis pass · forward-looking unresolved questions
  • How SNX4 heterodimer composition, lipid binding, and partner engagement are switched between its recycling, autophagy, and synaptic-regulatory functions remains unresolved.
  • No structural model connecting heterodimer choice to cargo selection
  • Mechanism linking SNX4 to synaptic vesicle docking unknown
  • Integration of dynein/clathrin transport with autophagic membrane supply not established

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 3 GO:0008289 lipid binding 2 GO:0008092 cytoskeletal protein binding 1
Localization
GO:0005768 endosome 5 GO:0005794 Golgi apparatus 2 GO:0031410 cytoplasmic vesicle 2 GO:0005764 lysosome 1
Pathway
R-HSA-5653656 Vesicle-mediated transport 4 R-HSA-9612973 Autophagy 4 R-HSA-9609507 Protein localization 3 R-HSA-112316 Neuronal System 1
Complex memberships
SNX4-SNX30 heterodimerSNX4-SNX7 heterodimerSnx4-Atg20 heterodimerSnx4-Snx41 heterodimer

Evidence

Reading pass · 11 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2007 SNX4, through its PX and BAR domains, associates with tubular and vesicular elements of early endosomes and the juxtanuclear endocytic recycling compartment (ERC). SNX4 suppression causes lysosomal degradation of the transferrin receptor (TfnR), establishing its role in endosomal sorting and recycling. SNX4 interacts with KIBRA (which binds dynein light chain 1) to associate with the minus-end-directed microtubule motor dynein, linking membrane tubulation-driven cargo sorting to long-range carrier transport from early endosomes to the ERC. siRNA knockdown, co-immunoprecipitation, fluorescence microscopy, endosomal fractionation Nature cell biology High 17994011
2003 In yeast, Snx4p (ortholog of SNX4) mediates retrieval of the exocytic v-SNARE Snc1p from post-Golgi endosomes back to the Golgi. Snc1p can be chemically cross-linked to Snx4p. Snx4p physically binds Snx41p and Snx42p, and all three sorting nexins are required for efficient Snc1p sorting, defining a distinct retrieval pathway from early endosomes that is separable from the retromer pathway operating at pre-vacuolar endosomes. Chemical cross-linking, yeast genetics (deletion mutants), subcellular fractionation, fluorescence microscopy The EMBO journal High 12554655
2009 SNX4 forms complexes with clathrin and dynein on endosomes; these interactions depend on PI3-kinase activity (inhibited by wortmannin), indicating they occur when SNX4 is PI(3)P-associated. A clathrin-box variant motif on SNX4 was identified as the clathrin-interacting site; a short peptide containing this motif is sufficient to pull down both clathrin and dynein. Clathrin knockdown is not required for the SNX4/dynein interaction but leads to increased retrograde Golgi transport of ricin and redistribution of endosomes, suggesting clathrin regulates SNX4-dependent transport. Co-immunoprecipitation, peptide pulldown, siRNA knockdown, pharmacological inhibition (wortmannin), fluorescence microscopy, toxin trafficking assay PloS one Medium 19529763
2013 Reggie-1 (flotillin-2) directly interacts with SNX4 and Rab11a at the tubulovesicular recycling compartment. Reggie-1 depletion impairs TfR recycling to the plasma membrane and E-cadherin recycling after internalization; both defects are rescued by overexpression of constitutively active Rab11a or SNX4, placing reggie-1 as a regulator upstream of the Rab11a/SNX4-controlled recycling pathway. Co-immunoprecipitation (direct interaction), shRNA knockdown, overexpression rescue experiments, fluorescence microscopy Molecular biology of the cell Medium 23825023
2017 Yeast Snx4 forms two functionally distinct heterodimers: Snx4-Atg20 and Snx4-Snx41. Each heterodimer coats a distinct endosome-derived tubule mediating retrograde sorting of different cargo: Snc1 is sorted by Snx4-Atg20, and Atg27 is sorted by Snx4-Snx41. The dynamin-family GTPase Vps1, which promotes fission of retromer-coated tubules, also promotes fission of Snx4-Atg20-coated tubules, linking tubule scission to Snx4-dependent retrograde trafficking. Live-cell fluorescence microscopy, genetic deletion, cargo-specific trafficking assays, co-localization Traffic (Copenhagen, Denmark) Medium 28026081
2017 Yeast Snx4 cooperates with Snx41 and Snx42 to mediate autophagic turnover of the 26S proteasome and other large multisubunit complexes during nitrogen starvation. Snx4 is required for the formation of cytoplasmic proteasome puncta that accumulate when autophagosome formation is blocked, placing Snx4 at the step of cytoplasmic agglomeration of proteasomes prior to autophagic delivery. Yeast deletion mutants, fluorescence microscopy, pharmacological tethering, targeted autophagy gene screen The Journal of biological chemistry Medium 29109144
2020 In mammalian cells, SNX4 is required for efficient LC3 lipidation and autophagosome assembly. SNX4 forms functional heterodimers with either SNX7 or SNX30 on tubulovesicular endocytic membranes. SNX4-SNX7 is an autophagy-specific heterodimer required for recruitment/retention of core autophagy regulators at the nascent isolation membrane. SNX4 partially co-localizes with juxtanuclear ATG9A-positive membranes, and SNX4 disruption causes mis-trafficking/retention of ATG9A in the Golgi region, linking the SNX4-SNX7 complex to ATG9A trafficking during autophagosome assembly. siRNA knockdown, CRISPR-Cas9 knockout, quantitative fluorescence microscopy, co-immunoprecipitation (heterodimer identification) Journal of cell science High 32513819
2021 SNX4 mediates recycling of the lipid scramblase ATG9A from endolysosomes to early endosomes, from where ATG9A is further recycled to the trans-Golgi network in a retromer (VPS35)-dependent manner. SNX4 depletion causes accumulation of ATG9A on endolysosomes; VPS35 depletion causes accumulation on early endosomes. SNX4-mediated ATG9A recycling is required for starvation-induced autophagosome biogenesis and autophagic flux, likely by preventing exhaustion of the ATG9A pool. siRNA knockdown, fluorescence microscopy, autophagy flux assays Journal of cell science Medium 33468622
2021 In yeast, the sorting nexin heterodimer Snx4/Atg24-Atg20 is required for a selective autophagy pathway (Snx4-assisted autophagy of transcription factors, SAA-TF) that targets transcriptional regulators Ssn2/Med13, Rim15, and Msn2 for vacuolar proteolysis upon nitrogen starvation. Snx4-Atg20 binds Atg17, relocates to the perinucleus upon starvation, and is required for efficient transfer of cargo (Ssn2/Med13) to Atg17-initiated phagophores anchored to the vacuole. Yeast genetics (deletion analysis), fluorescence microscopy, co-immunoprecipitation Autophagy Medium 33678121
2024 In yeast, the Snx4-Atg20 heterodimer delivers the cargo Ssn2/Med13 to phagophores in the SAA-TF pathway. Ksp1, an autophagic receptor, is recruited early to phagophores via Atg29 independently of both Atg8 and Snx4; Snx4 delivers Ssn2/Med13 to phagophores thereafter, defining the temporal order of receptor and cargo delivery in this pathway. Yeast two-hybrid, genetic deletion analysis, fluorescence microscopy, mutagenesis of AIM/LIR motifs Autophagy Medium 37733395
2024 In neurons, SNX4 conditional knockout leads to an increase in docked synaptic vesicles at the active zone and decreased active zone length, resulting in increased docked vesicle density per release site. This causes enhanced neurotransmission during train stimulation without affecting vesicle recycling, autophagic flux, or VAMP2/synaptobrevin-2 levels or localization, establishing SNX4 as a negative regulator of synaptic vesicle docking and release. Conditional knockout mouse model, electron microscopy (ultrastructure), electrophysiology (evoked EPSCs), Western blot, fluorescence microscopy eLife High 39699951

Source papers

Stage 0 corpus · 11 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 SNX4 coordinates endosomal sorting of TfnR with dynein-mediated transport into the endocytic recycling compartment. Nature cell biology 222 17994011
2003 Retromer and the sorting nexins Snx4/41/42 mediate distinct retrieval pathways from yeast endosomes. The EMBO journal 175 12554655
2017 Autophagic clearance of proteasomes in yeast requires the conserved sorting nexin Snx4. The Journal of biological chemistry 65 29109144
2013 Reggies/flotillins interact with Rab11a and SNX4 at the tubulovesicular recycling compartment and function in transferrin receptor and E-cadherin trafficking. Molecular biology of the cell 65 23825023
2017 Distinct complexes of yeast Snx4 family SNX-BARs mediate retrograde trafficking of Snc1 and Atg27. Traffic (Copenhagen, Denmark) 52 28026081
2020 A heterodimeric SNX4--SNX7 SNX-BAR autophagy complex coordinates ATG9A trafficking for efficient autophagosome assembly. Journal of cell science 37 32513819
2009 SNX4 in complex with clathrin and dynein: implications for endosome movement. PloS one 35 19529763
2021 The phosphatidylinositol 3-phosphate-binding protein SNX4 controls ATG9A recycling and autophagy. Journal of cell science 33 33468622
2021 Snx4-assisted vacuolar targeting of transcription factors defines a new autophagy pathway for controlling ATG expression. Autophagy 15 33678121
2024 Ksp1 is an autophagic receptor protein for the Snx4-assisted autophagy of Ssn2/Med13. Autophagy 5 37733395
2024 Endosomal sorting protein SNX4 limits synaptic vesicle docking and release. eLife 1 39699951

Missed literature

Know a paper Affinage missed for SNX4? Flag it for the maintainers and the community.

No submissions yet.