Affinage

Showing PDCD6IP — ALIX is a alias.

PDCD6IP

Programmed cell death 6-interacting protein · UniProt Q8WUM4

Length
868 aa
Mass
96.0 kDa
Annotated
2026-06-10
100 papers in source corpus 51 papers cited in narrative 50 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PDCD6IP/Alix is a conformationally regulated ESCRT-associated scaffold that links membrane cargo and accessory proteins to the ESCRT-III machinery to drive membrane fission across multiple cellular processes (PMID:17350572, PMID:32049272). Its architecture comprises an N-terminal Bro1 domain that binds CHMP4/ESCRT-III subunits through a hydrophobic code distinct from the MIT-domain partners of VPS4 (PMID:18511562), a central V domain that recognizes YPX(n)L late-domain motifs of retroviral Gag proteins and cellular GPCRs as well as K63-linked polyubiquitin chains (PMID:17350572, PMID:17277784, PMID:18066081, PMID:22547407, PMID:23201121), and a C-terminal proline-rich domain that engages ALG-2 in a Ca2+-dependent arginine-switch manner, TSG101, and endophilins (PMID:12034747, PMID:14999017, PMID:18940611, PMID:32917811). The proline-rich region folds back onto the Bro1/V domains to lock Alix in an autoinhibited closed conformation that is relieved by Ca2+/ALG-2, membrane recruitment, partner binding, or E3-ligase-mediated ubiquitination, thereby licensing CHMP4 recruitment and membrane association (PMID:21715492, PMID:20929444, PMID:27462417, PMID:22334701). Through these interactions Alix mediates enveloped virus budding (PMID:14505569, PMID:17350572), multivesicular body intraluminal vesicle formation and a tetraspanin-specific exosome pathway requiring lysobisphosphatidic acid (PMID:18768755, PMID:32049272), syndecan-syntenin-driven exosome biogenesis (PMID:24637612, PMID:25732677), ubiquitin-independent MVB/lysosomal sorting of GPCRs via YPX(n)L motifs (PMID:22547407, PMID:27301021), and cytokinetic abscission, where it is recruited to the midbody by CEP55 in mammals or centralspindlin in Drosophila and delivers CHMP4 to the abscission site (PMID:17853893, PMID:26929449, PMID:31607533). Alix additionally organizes the actin cytoskeleton and epithelial junctions; its loss in mice produces hydrocephalus from defects in epithelial polarity and tight-junction assembly (PMID:16966331, PMID:27336173), and it supports clathrin-independent endocytosis together with endophilin-A (PMID:27244115).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1999 Medium

    Established Alix's founding biochemical partnership, defining it as a Ca2+-regulated effector and hinting at a role in regulated death/signaling.

    Evidence Yeast two-hybrid and co-IP identifying ALG-2 as a calcium-dependent partner

    PMID:10200558

    Open questions at the time
    • Functional consequence of the ALG-2 interaction unresolved at this stage
    • No structural basis for Ca2+ dependence yet
    • No link to membrane trafficking established
  2. 2002 Medium

    Mapped the proline-rich domain as a protein-interaction hub by identifying endophilin binding, linking Alix to membrane-deforming machinery.

    Evidence Yeast two-hybrid, co-IP and overlay assays with PXRPPPP consensus mapping

    PMID:12034747

    Open questions at the time
    • Physiological process served by Alix-endophilin binding undefined
    • No structural detail of the interface
  3. 2003 High

    Placed Alix in the ESCRT pathway by showing it bridges retroviral Gag late domains to ESCRT-III and TSG101, defining its role in viral budding.

    Evidence Co-IP, binding and functional budding assays with HIV-1 p6, EIAV p9, and CHMP4b

    PMID:12588984 PMID:12860994 PMID:14505569

    Open questions at the time
    • Atomic basis of YPX(n)L and CHMP4 recognition not yet known
    • How budding is spatially coordinated unresolved
  4. 2004 Medium

    Defined the ALG-2 binding site on Alix and uncovered counter-roles in receptor downregulation, endosome organization, and non-apoptotic death, broadening Alix beyond budding.

    Evidence Mutagenesis/overlay/co-IP mapping ALG-2 to residues 794-827; siRNA and ubiquitination assays for EGFR/Cbl; Src kinase assays; paraptosis assays

    PMID:14999017 PMID:15086793 PMID:15195070 PMID:15456872 PMID:15557335

    Open questions at the time
    • Mechanistic link between EGFR regulation and ESCRT sorting unclear
    • Src phosphorylation site and structural effect not resolved at this stage
  5. 2005 Medium

    Connected Alix to endosome positioning and the lipid environment of MVBs, distinguishing its role from EGFR degradation and recycling.

    Evidence RNAi with immunofluorescence and immunoelectron microscopy measuring LBPA and endosome distribution

    PMID:15914539

    Open questions at the time
    • Molecular basis of LBPA dependence undefined
    • Mechanism linking Alix loss to aberrant actin/clathrin structures unclear
  6. 2006 Medium

    Demonstrated direct actin-cytoskeleton engagement, establishing a structural role for Alix beyond membrane scission.

    Evidence Cell-free direct binding to F-actin, cortactin, alpha-actinin; siRNA phenotypes on stress fibers

    PMID:16966331

    Open questions at the time
    • How cytoskeletal and ESCRT functions are coordinated unknown
    • In vivo relevance not addressed here
  7. 2007 High

    Solved the structural logic of Alix: defined Bro1 and V domain architecture, the YPX(n)L pocket, CHMP4 binding, and the CEP55-dependent midbody recruitment for abscission.

    Evidence Crystallography of Bro1/V domains and V-domain:late-domain complexes; proteomics, co-IP, siRNA and cytokinesis assays for CEP55/TSG101; NC-Bro1 packaging

    PMID:17277784 PMID:17350572 PMID:17853893 PMID:18032513 PMID:18066081

    Open questions at the time
    • Conformational regulation of these interactions not yet defined
    • Cargo-specific pathway selection unresolved
  8. 2008 High

    Defined the molecular grammar of partner selection and oligomeric scaffolding: the CHMP4 hydrophobic code, the ALG-2 arginine switch, V-domain dimerization, and a positive/negative regulatory split with TSG101 in ILV budding.

    Evidence Crystallography of Bro1:CHMP4 and Ca2+/ALG-2 states; SAXS dimer modeling and filament bridging; in vitro ILV reconstitution; abscission/HIV mutagenesis

    PMID:18511562 PMID:18641129 PMID:18768755 PMID:18940611 PMID:19523902

    Open questions at the time
    • How dimerization is triggered in cells unclear
    • Reconciliation of Alix as negative regulator in vitro vs positive elsewhere
  9. 2010 Medium

    Identified intramolecular autoinhibition as the master regulatory principle, with the PRD folding back to occlude V and Bro1 surfaces and ubiquitination/E3 ligases as activating inputs.

    Evidence Truncation/mutagenesis with co-IP, membrane fractionation, budding assays; Nedd4-1 and ALG-2 bridging studies

    PMID:19520058 PMID:20519395 PMID:20929444 PMID:21715492

    Open questions at the time
    • Quantitative dynamics of open/closed equilibrium not measured
    • Cell-type-specific activation inputs incompletely mapped
  10. 2012 Medium

    Extended Alix cargo recognition to ubiquitin-independent GPCR sorting via YPX(3)L motifs and to direct K63-polyubiquitin binding, and identified Ozz-E3 ubiquitination as a conformational/abundance regulator.

    Evidence siRNA/mutagenesis PAR1 sorting; in vitro polyUb binding and mutagenesis; endogenous co-IP, Ozz KO mouse and actin phenotypes

    PMID:22334701 PMID:22547407 PMID:23201121

    Open questions at the time
    • Relative contributions of ubiquitin-dependent vs -independent cargo capture unclear
    • Tissue-specific roles of Ozz regulation undefined
  11. 2013 High

    Established the ancestral V-domain ubiquitin-receptor function via the yeast Bro1 ortholog, paralleling ESCRT-0 in cargo sorting.

    Evidence Structural studies, mutagenesis, and synthetic genetic epistasis with ESCRT-0 in yeast

    PMID:23726974

    Open questions at the time
    • Conservation of this ubiquitin-receptor role in human Alix not formally proven here
    • Interplay with V-domain YPX(n)L binding unresolved
  12. 2014 Medium

    Defined the syndecan-syntenin-ALIX exosome pathway and its ARF6-PLD2 control, establishing a budding route distinct from viral budding.

    Evidence siRNA, overexpression, EM and exosome isolation

    PMID:24637612

    Open questions at the time
    • How ARF6-PLD2 lipids feed into Alix activation unclear
    • Cargo selectivity of this pathway incompletely defined
  13. 2015 Medium

    Integrated Alix into autophagy, basal endosomal exosome biogenesis, and demonstrated pathway-selective activation, while resolving exosome production regulators including heparanase and parallel abscission recruiters.

    Evidence ATG12-ATG3 co-IP/KO; ALG-2 activation specific to EGFR MVB sorting; heparanase exosome assays; Drosophila Shrub genetics; CHMP4C-specific abscission siRNA epistasis; ARRDC3/WWP2 ubiquitination

    PMID:25635693 PMID:25686249 PMID:25732677 PMID:26490116 PMID:26929449 PMID:27462417

    Open questions at the time
    • How a single scaffold achieves pathway-specific activation mechanistically unclear
    • Crosstalk between activating inputs (ALG-2, ubiquitination, lipids) unresolved
  14. 2016 Medium

    Broadened Alix's cellular roles to clathrin-independent endocytosis, epithelial junction/actin assembly with an in vivo hydrocephalus phenotype, additional GPCR sorting, and EV miRNA packaging.

    Evidence Alix KO endocytosis assays with endophilin-A; KO mouse epithelial polarity/tight-junction analysis; P2Y1 YPX3L sorting; Ago2 co-IP and EV miRNA quantification

    PMID:26935291 PMID:27244115 PMID:27301021 PMID:27336173

    Open questions at the time
    • Mechanistic link between Alix's actin-junction role and its ESCRT scaffold function unclear
    • Ago2/miRNA-packaging mechanism rests on a single low-confidence study
  15. 2018 Medium

    Linked Alix-dependent MVB sorting to immune regulation through PD-L1 trafficking and EV secretion, and identified S-palmitoylation as a lipid-modification input.

    Evidence siRNA, flow cytometry, exosome isolation and mouse tumor model for PD-L1; acyl-RAC palmitoylation and CD9 co-IP

    PMID:30021161 PMID:30251702

    Open questions at the time
    • Palmitoylation site and functional requirement rest on a single low-confidence study
    • How Alix selects PD-L1 cargo unresolved
  16. 2019 Medium

    Revealed an evolutionarily flexible recruitment logic — the V-domain LYPX(n)L pocket also captures the centralspindlin LxxLF motif for CEP55-independent midbody recruitment — and removal of phospho-MLKL via Alix-syntenin exocytosis.

    Evidence Co-IP, domain mutagenesis and Drosophila genetics for Pavarotti; cross-linking/AP and exocytosis assay for MLKL

    PMID:31138766 PMID:31607533

    Open questions at the time
    • Whether mammalian centralspindlin uses the same pocket unresolved
    • Regulation switching between viral and midbody ligands at the same pocket unclear
  17. 2020 High

    Reconstituted active Alix recruiting ESCRT-III on LBPA-containing bilayers for tetraspanin exosome sorting, defined the syndecan-4-syntenin-Alix module at the abscission site, and resolved a phosphorylation-controlled reversible amyloid in the PRD that regulates TSG101 competition.

    Evidence In vitro bilayer reconstitution and siRNA/EM for tetraspanins; Flemmingsome proteomics and abscission siRNA; NMR/relaxation/ThT and Src phosphorylation of the PRD

    PMID:32049272 PMID:32321914 PMID:32731849 PMID:32917811

    Open questions at the time
    • Physiological trigger of PRD amyloid formation in cells unclear
    • How the syndecan-4 module is distinguished from midbody recruitment incompletely defined
  18. 2021 Medium

    Showed Alix governs a polarity-specific exosome route, mediating apical exosome release distinct from ceramide-dependent basolateral release.

    Evidence siRNA and directional exosome assays in polarized epithelial cells

    PMID:33724661

    Open questions at the time
    • Molecular basis of apical vs basolateral sorting unresolved
    • Link to the epithelial junction role of Alix not connected
  19. 2023 Medium

    Identified a SUMO-dependent cargo-capture mode in which the Alix SIM recognizes SUMOylated hnRNPA2B1 to drive circRNA packaging into EVs with pro-metastatic consequences.

    Evidence Co-IP, SUMOylation assays, SIM mutation and in vivo metastasis model

    PMID:37925421

    Open questions at the time
    • Generality of the SIM-dependent cargo route beyond this circRNA unknown
    • How SUMO recognition integrates with autoinhibition release unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • How a single autoinhibited scaffold integrates Ca2+/ALG-2, ubiquitination, SUMO, lipid, and phosphorylation/amyloid inputs to select among virus budding, ILV/exosome biogenesis, abscission, endocytosis, and cytoskeletal roles in a given cell remains unresolved.
  • No unified quantitative model of conformational switching across pathways
  • Cargo- and pathway-specific activation inputs only partly mapped
  • Mammalian counterpart of several Drosophila/yeast findings not formally confirmed

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 5 GO:0008092 cytoskeletal protein binding 3 GO:0005198 structural molecule activity 2 GO:0098772 molecular function regulator activity 2
Localization
GO:0005768 endosome 4 GO:0005886 plasma membrane 3 GO:0005829 cytosol 2 GO:0005856 cytoskeleton 2
Pathway
R-HSA-5653656 Vesicle-mediated transport 4 R-HSA-162582 Signal Transduction 3 R-HSA-1640170 Cell Cycle 3 R-HSA-1643685 Disease 3 R-HSA-8953854 Metabolism of RNA 2 R-HSA-9612973 Autophagy 1
Partners
CEP55CHMP4BENDOPHILINNEDD4PAR1PDCD6 (ALG-2)SDCBP (SYNTENIN)TSG101
Complex memberships
ESCRT-III (CHMP4)actomyosin-tight junction complexsyndecan-syntenin-ALIX module

Evidence

Reading pass · 50 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1999 Alix (PDCD6IP) was identified as a novel binding partner of ALG-2 (apoptosis-linked gene 2) in a calcium-dependent manner, established by yeast two-hybrid screening and co-immunoprecipitation experiments. Yeast two-hybrid screen; co-immunoprecipitation Cell death and differentiation Medium 10200558
2002 Alix binds endophilins (SH3p4, SH3p8, SH3p13) through a proline-rich domain interaction (PXRPPPP consensus) at its C-terminal half, demonstrated by yeast two-hybrid, co-immunoprecipitation, and overlay experiments; overexpression of Alix-CT causes cytoplasmic vacuolization enhanced by endophilin co-expression. Yeast two-hybrid; co-immunoprecipitation; overlay assay; cell transfection/overexpression The Journal of biological chemistry Medium 12034747
2003 HIV-1 Gag p6 contains a second L-domain region that binds AIP1/Alix (PDCD6IP); Alix also interacts with Tsg101 and ESCRT-III homologs, and binds the EIAV p9 L domain, identifying Alix as a component of the viral budding machinery linking retroviral Gag proteins to ESCRT-III. Co-immunoprecipitation; binding assays; functional budding assays Cell High 14505569
2003 Alix interacts with CHMP4b (human Snf7 homolog) as demonstrated by GST pull-down and co-immunoprecipitation; CHMP4b and Alix co-localize at perinuclear puncta overlapping with endosomal markers, and cooperate in multivesicular body formation. Yeast two-hybrid; GST pull-down; co-immunoprecipitation; fluorescence microscopy The Journal of biological chemistry High 12860994
2003 Both fungal PalA and mammalian AIP1/Alix recognize the YPXL/I protein-protein binding motif, where Tyr, Pro, and Leu/Ile are crucial for binding, established by mutagenesis of the motif in PacC substrates and binding assays. Binding assays; site-directed mutagenesis of YPXL/I motif Molecular and cellular biology Medium 12588984
2004 Paraptosis (non-apoptotic cell death induced by IGFIR) is inhibited specifically by AIP-1/Alix overexpression; this inhibition does not affect apoptosis and is MAPK/JNK-1-mediated, placing Alix as a negative regulator of IGFIR-induced paraptosis. Cell transfection/overexpression; antisense oligonucleotides; MEK-2 inhibitors; cell death assays Cell death and differentiation Medium 15195070
2004 ALG-2 interacts with the C-terminal proline-rich region of Alix (residues 794–827) through four tandem PxY repeats in a Ca2+-dependent manner; specific proline and tyrosine residues within this region are critical for binding affinity, demonstrated by yeast two-hybrid, overlay assay, and co-immunoprecipitation with deletion/alanine-substitution mutants. Yeast two-hybrid; biotin-tagged overlay assay; co-immunoprecipitation; alanine mutagenesis Journal of biochemistry High 14999017
2004 Alix/AIP1 antagonizes EGFR downregulation by attenuating the Cbl-SETA/CIN85 interaction and inhibiting Cbl-mediated ubiquitination of EGFR; siRNA knockdown of Alix promotes EGFR internalization and downregulation. Co-immunoprecipitation; siRNA knockdown; ubiquitination assays; receptor internalization assay Molecular and cellular biology Medium 15456872
2004 Alix co-immunoprecipitates with exosomal transferrin receptor, and a peptide (P1, YTRFSLARQV) from the TfR cytosolic domain binds both hsc70 and Alix (via homology to its fungal homolog PalA), suggesting Alix contributes to exosomal sorting of TfR via the YTRF motif. Co-immunoprecipitation; peptide binding/affinity chromatography Traffic (Copenhagen, Denmark) Low 15086793
2004 Src phosphorylates Alix at its C-terminal tyrosine-rich region; this phosphorylation causes Alix to translocate from membrane/cytoskeleton to cytoplasm and reduces its interactions with SETA/CIN85, EGFR, and Pyk2, antagonizing Alix's negative regulation of receptor internalization and cell adhesion. In vitro kinase assay; co-immunoprecipitation; subcellular fractionation; siRNA/overexpression The Journal of biological chemistry Medium 15557335
2005 Alix depletion by RNAi causes redistribution of early endosomes from peripheral to perinuclear location, accumulation of unusual actin structures containing clathrin and cortactin, and reduction of lysobisphosphatidic acid levels in multivesicular endosomes, without affecting transferrin recycling or EGF receptor degradation. RNAi knockdown; immunofluorescence microscopy; quantitative immunoelectron microscopy Journal of cell science Medium 15914539
2006 Alix directly interacts with F-actin (via both Bro1 and proline-rich domains), cortactin (via the N-terminal half of the middle region), and alpha-actinin (via the C-terminal half of the middle region) in cell-free systems; Alix knockdown decreases F-actin content and inhibits stress fiber assembly and lamellipodial localization of cortactin. In vitro direct binding (cell-free); co-immunoprecipitation; siRNA knockdown; fluorescence microscopy The Journal of biological chemistry Medium 16966331
2007 Crystal structures of ALIX reveal an N-terminal Bro1 domain and a central V domain composed of two three-helix bundle arms; YPX(n)L late domains of retroviruses bind a conserved hydrophobic pocket on the second arm of the V domain; CHMP4/ESCRT-III proteins bind a conserved hydrophobic patch on the Bro1 domain; both interactions are required for virus budding. X-ray crystallography; mutagenesis; functional budding assays Cell High 17350572
2007 ALIX and TSG101/ESCRT-I are recruited to midbodies of dividing cells through direct interactions between CEP55 and GPP-based motifs within TSG101 and ALIX; ALIX point mutants blocking CEP55 and CHMP4/ESCRT-III binding inhibit cytokinetic abscission; depletion of ALIX inhibits the abscission step of HeLa cell cytokinesis. Proteomics; co-immunoprecipitation; siRNA depletion; fluorescence microscopy; cytokinesis assay The EMBO journal High 17853893
2007 Crystal structures of ALIX V domain in complex with YPX(n)L late domains from HIV-1 and EIAV reveal that both bind the same hydrophobic pocket but adopt different conformations; binding affinity and budding function require key interface residues identified by mutagenesis. X-ray crystallography; surface plasmon resonance; mutagenesis; functional budding assays Nature structural & molecular biology High 17277784 18066081
2007 ALIX also engages HIV-1 Gag through the nucleocapsid (NC) domain via the Bro1 domain (in addition to the V domain/p6 interaction); this NC-Bro1 interaction depends on NC zinc fingers and allows ALIX packaging into viral particles; zinc finger mutations affect particle production similarly to PTAP deletion. Co-immunoprecipitation; viral packaging assays; nuclease treatment; mutagenesis Journal of virology Medium 18032513
2008 Crystal structures of ALIX Bro1 domain in complex with C-terminal peptides of CHMP4A-C show that CHMP4 C-terminal peptides form amphipathic helices binding across the concave surface of Bro1; a hydrophobic code (M/L/IxxLxxW) distinguishes CHMP4 binding to Bro1-domain proteins from CHMP1-3 binding to MIT-domain proteins (VPS4); mutations in CHMP4 recognition residues block ALIX-dependent HIV-1 budding. X-ray crystallography; mutagenesis; HIV-1 budding assay Proceedings of the National Academy of Sciences of the United States of America High 18511562
2008 ALG-2 binds Ca2+ through EF3, triggering an 'arginine switch' (Arg125 movement) that opens the primary hydrophobic pocket on ALG-2 to accommodate the PPYP motif of Alix; crystal structures of Ca2+-free, Ca2+-bound, and Alix peptide-complexed ALG-2 forms established this mechanism. X-ray crystallography; in vitro binding assay with ALG-2 and Alix mutants Structure High 18940611
2008 Tsg101 acts as a positive regulator and Alix as a negative regulator of intraluminal vesicle budding into late endosomes, reconstituted in a novel quantitative biochemical in vitro budding assay; the process is ESCRT-dependent (inhibited by dominant-negative hVps4). In vitro reconstitution assay; dominant-negative VPS4; electron microscopy Molecular biology of the cell High 18768755
2008 ALIX Bro1-CHMP4 and ALIX V-domain interactions are both required for cytokinetic abscission; additionally the C-terminus of Alix encodes a multimerization activity essential for Alix-dependent HIV-1 release and interaction with Tsg101. Dominant-negative and deletion mutagenesis; cytokinesis assay; HIV budding assay Proceedings of the National Academy of Sciences of the United States of America Medium 18641129
2009 ALIX dimerizes via its middle V domain in solution; SAXS modeling shows dimeric ALIX adopts an elongated crescent shape; ALIX dimerization mutations prevent this and affect HIV-1 budding; dimeric ALIX bridges CHMP4B filaments formed in vitro, acting as a scaffold during membrane remodeling. Small-angle X-ray scattering (SAXS); dimerization mutagenesis; in vitro filament bridging assay; HIV budding assay Structure High 19523902
2009 ALG-2 bridges Alix and TSG101 as a Ca2+-dependent adaptor; the ALG-2-binding site on Alix (not the PSAP motif) is required for Ca2+-dependent pulldown of TSG101; dimeric ALG-2 is required for this bridging function. Strep-pulldown assays; ALG-2 knockdown; exogenous ALG-2 rescue; mutant analysis Biochemical and biophysical research communications Medium 19520058
2010 The ALIX C-terminal proline-rich region (PRR) folds back against upstream domains causing auto-inhibition of V domain binding to viral late domains; mutations destabilizing this closed conformation open the V domain, increase ALIX membrane association, and enhance virus budding. Mutagenesis; in vitro binding assays; virus budding assays; membrane association assay Journal of virology Medium 21715492
2010 ALIX contains an intramolecular interaction between Patch 2 of the Bro1 domain and the TSG101-docking site in the PRD that locks ALIX in a closed conformation, preventing CHMP4 and retroviral Gag protein binding; relieving this autoinhibition promotes ALIX interaction with partners and membrane association, and increases EIAV budding. ALIX truncation/mutation analysis; co-IP; membrane fractionation; EIAV budding assay The Biochemical journal Medium 20929444
2010 Nedd4-1 binds ALIX directly (co-IP and yeast two-hybrid); Nedd4-1 is required for ALIX ubiquitination; ALIX recruits Nedd4-1 through the LYPX(n)L motif to facilitate HIV-1 release; Nedd4-1 catalytic activity is required for this function. Co-immunoprecipitation; yeast two-hybrid; siRNA knockdown; HIV budding assay; catalytic mutant Journal of virology Medium 20519395
2012 ALIX binds a YPX(3)L motif in the GPCR PAR1 via its central V domain to mediate ubiquitin-independent, ESCRT-III-dependent MVB/lysosomal sorting of PAR1; depletion of ALIX impairs PAR1 sorting into ILVs and lysosomal degradation. siRNA depletion; site-directed mutagenesis of YPX(3)L; co-immunoprecipitation; MVB sorting assay The Journal of cell biology Medium 22547407
2012 The V domain of ALIX binds directly and selectively to K63-linked polyubiquitin chains (preference for chains >3 Ub); mutations in two putative Ub-binding sites on a helical surface of the V domain inhibit polyUb binding in vitro and impair lentiviral budding in cells. Affinity capture/mass spectrometry; in vitro binding assays; mutagenesis; lentiviral budding assay Developmental cell Medium 23201121
2013 Yeast Bro1 (Alix ortholog) functions as a ubiquitin receptor via the N-terminal trihelical arm of its V domain; Bro1 binds ubiquitin and sorts ubiquitinated cargo in parallel with ESCRT-0, demonstrated by structural studies, mutagenesis, and synthetic genetic interactions with ESCRT-0. Multiple structural approaches (X-ray/NMR); mutagenesis; genetic epistasis (synthetic lethality screen) Developmental cell High 23726974
2014 The syndecan-syntenin-ALIX pathway controls exosome biogenesis and ILV budding; ARF6 and its effector PLD2 regulate syntenin exosome production by controlling ILV budding into MVBs in an ALIX-dependent manner; this pathway does not affect HIV-1 budding. siRNA depletion; overexpression; electron microscopy; exosome isolation Nature communications Medium 24637612
2015 ATG12-ATG3 conjugate interacts with Alix (PDCD6IP) and controls multiple Alix-dependent processes including late endosome distribution, exosome biogenesis, and viral budding; Alix is functionally required for efficient basal (but not starvation-induced) autophagy. Co-immunoprecipitation; siRNA/knockout; autophagy flux assay; exosome isolation; electron microscopy Nature cell biology Medium 25686249
2015 Heparanase stimulates syndecan-syntenin-ALIX-dependent exosome production by trimming heparan sulfate chains on syndecans, promoting intraluminal budding of syndecan and syntenin-1 in an ALIX-dependent manner. siRNA depletion; heparanase treatment; exosome isolation and quantification; electron microscopy Cell research Medium 25732677
2015 ALG-2 calcium-dependent interaction with Alix relieves ALIX intramolecular autoinhibition, promoting CHMP4-dependent ALIX membrane association and MVB sorting of activated EGFR; this ALG-2 activation of ALIX is specific to MVB sorting and does not affect cytokinetic abscission or EIAV budding. Co-immunoprecipitation; membrane fractionation; MVB sorting assay; siRNA; mutagenesis Cell discovery Medium 27462417
2015 ARRDC3 (α-arrestin) regulates ALIX ubiquitination by WWP2 E3 ligase; ARRDC3 and WWP2 are required for ALIX interaction with activated PAR1 and CHMP4B, and for PAR1 lysosomal sorting; WWP2 interacts with ARRDC3 (not ALIX) to ubiquitinate ALIX. siRNA screen; co-immunoprecipitation; ubiquitination assay; GPCR sorting assay Molecular biology of the cell Medium 26490116
2015 ALIX is required for abscission completion in Drosophila female germline stem cells (fGSCs) in vivo; ALIX and ESCRT-III component Shrub interact directly, co-localize at midbody rings, and their direct interaction is required for normal abscission kinetics. Drosophila loss-of-function genetics; co-immunoprecipitation; fluorescence microscopy in vivo PLoS genetics Medium 25635693
2016 ALIX and ESCRT-I/II function as parallel ESCRT-III recruiters during cytokinetic abscission; ALIX specifically recruits CHMP4C (abscission checkpoint component) while ESCRT-II/CHMP6 cooperate with ESCRT-I to recruit CHMP4B; ALIX depletion causes furrow regression in cells with chromosome bridges, implicating ALIX in abscission checkpoint signaling. siRNA depletion; fluorescence microscopy; cytokinesis/abscission assay; epistasis analysis The Journal of cell biology Medium 26929449
2016 ALIX mediates ubiquitin-independent lysosomal sorting of the P2Y1 GPCR via a YPX3L motif in the second intracellular loop; site-directed mutation of this motif or ALIX depletion inhibits P2Y1 MVB/lysosomal sorting. siRNA depletion; site-directed mutagenesis; MVB sorting/lysosomal degradation assay PloS one Medium 27301021
2016 Alix binds Ago2 (Argonaute 2) by co-immunoprecipitation; ALIX knockdown decreases miRNA content in extracellular vesicles without reducing vesicle number, indicating a role for Alix in miRNA packaging into EVs. Co-immunoprecipitation; siRNA knockdown; RT-qPCR for miRNAs in EVs International journal of molecular medicine Low 26935291
2016 Alix is required for clathrin-independent endocytosis (CIE) but not clathrin-mediated endocytosis; in Alix knockout cells, fluid-phase endocytosis and CIE cargoes (cholera toxin, IL-2 receptor) are impaired; Alix acts with endophilin-A to promote CIE and regulate cell migration. Alix knockout cells; endocytosis assays; signaling assays; cell migration assay Scientific reports Medium 27244115
2016 Alix interacts with F-actin, the Par complex, and ZO-1 to assemble the apically restricted actomyosin-tight junction complex in epithelial cells; Alix knockout mice develop hydrocephalus due to defects in epithelial polarity, tight junction formation, and excessive cell extrusion in choroid plexus/ependyma. Knockout mouse model; co-immunoprecipitation; fluorescence/electron microscopy; phenotypic analysis Nature communications Medium 27336173
2018 Alix undergoes S-palmitoylation as a post-translational modification; this lipid modification influences Alix protein-protein interaction with CD9 (tetraspanin) and affects the structural organization of the lipid bilayer of extracellular vesicles. Palmitoylation assay (acyl-RAC); co-immunoprecipitation; biophysical lipid bilayer analysis Biochimica et biophysica acta. General subjects Low 30251702
2018 ALIX depletion results in prolonged EGFR activity after stimulation, defective PD-L1 trafficking through MVBs, reduced PD-L1 exosomal secretion, and redistribution of PD-L1 to the cell surface, conferring an EGFR-dependent immunosuppressive phenotype. siRNA depletion; EGFR signaling assays; flow cytometry for PD-L1; exosome isolation; mouse tumor model Cell reports Medium 30021161
2019 In Drosophila, ALIX is recruited to the midbody via its V domain through direct interaction with an LxxLF motif in centralspindlin component Pavarotti (MKLP1), using the same conserved hydrophobic V-domain pocket that binds LYPXnL virus proteins; this mechanism is CEP55-independent. Co-immunoprecipitation; domain mapping/mutagenesis; fluorescence microscopy; Drosophila genetics Current biology Medium 31607533
2019 Phosphorylated MLKL (necroptosis mediator) is removed from membranes via ALIX-syntenin-1-mediated exocytosis; ALIX and syntenin-1 are detected in membrane-localized MLKL immunoprecipitates by protein cross-linking/affinity purification. Protein cross-linking; affinity purification; functional exocytosis assay Science signaling Medium 31138766
2020 ALIX (active form) recruits ESCRT-III proteins to endosomes independently of other ESCRTs but requiring lysobisphosphatidic acid (LBPA) in vivo; this can be reconstituted on supported bilayers in vitro; this ALIX-ESCRT-III pathway promotes sorting of tetraspanins specifically to exosomes. In vitro reconstitution on supported bilayers; ALIX activation mutants; siRNA; electron microscopy The Journal of cell biology High 32049272
2020 The ESCRT-III machinery localizes to the abscission site via a plasma membrane-to-ESCRT module composed of syndecan-4 (transmembrane proteoglycan), ALIX, and syntenin; direct interactions between ALIX, syntenin, and syndecan-4 are essential for ESCRT-III enrichment at the abscission site (but not at the midbody); depletion of any of the three proteins delays abscission. Proteomics of isolated midbodies (Flemmingsome); siRNA depletion; co-immunoprecipitation; fluorescence microscopy Nature communications Medium 32321914
2020 ALIX and Syntenin-1 are required for proper subcellular localization and efficient EV secretion of APP; their knockdown sequesters APP in ER and endolysosomal compartments respectively; this pathway is ESCRT-independent. Inducible siRNA knockdown; subcellular localization; EV isolation and analysis BMC molecular and cell biology Low 32731849
2020 The ALIX proline-rich domain (PRD) contains three tandem proline-rich motifs that compete for a single binding site on TSG101-UEV; the C-terminal tyrosine-rich portion of ALIX-PRD forms amyloid fibrils that dissolve upon hyperphosphorylation by Src kinase, revealing a phosphorylation-controlled reversible amyloid mechanism regulating ALIX function. Heteronuclear NMR spectroscopy; relaxation dispersion; thioflavin T/Congo red assays; electron microscopy; Src kinase phosphorylation Proceedings of the National Academy of Sciences of the United States of America High 32917811
2021 ALIX (not the broader ESCRT machinery) is required specifically for apical exosome release in polarized epithelial cells via the ALIX-Syntenin1-Syndecan1 machinery, while basolateral exosome release depends on ceramide; these represent two independent polarized exosome release pathways. siRNA depletion; polarized epithelial cell model; exosome isolation; directional secretion assay EMBO reports Medium 33724661
2012 Alix is a substrate of the muscle-specific ubiquitin E3 ligase Ozz-E3; Ozz interaction with Alix changes Alix conformation and promotes its ubiquitination, regulating Alix levels at cortactin-containing compartments; Alix knockdown in muscle cells affects F-actin amount and distribution, impairs sarcolemmal protrusion formation and cell motility. Co-immunoprecipitation (endogenous proteins); ubiquitination assay; Ozz knockout mouse; siRNA knockdown; actin/morphology assays The Journal of biological chemistry Medium 22334701
2023 SUMOylated hnRNPA2B1 (SUMO2 at K108) is recognized by the SUMO interaction motif (SIM) of ALIX, activating ALIX to recruit ESCRT-III for sorting of circTLCD4-RWDD3 into extracellular vesicles; mutating the SIM in ALIX or K108 in hnRNPA2B1 blocks EV packaging and inhibits lymphatic metastasis in vivo. Co-immunoprecipitation; SUMOylation assay; SIM mutation in ALIX; in vivo tumor model Signal transduction and targeted therapy Medium 37925421

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2003 AIP1/ALIX is a binding partner for HIV-1 p6 and EIAV p9 functioning in virus budding. Cell 690 14505569
2007 Human ESCRT and ALIX proteins interact with proteins of the midbody and function in cytokinesis. The EMBO journal 595 17853893
2014 Syntenin-ALIX exosome biogenesis and budding into multivesicular bodies are controlled by ARF6 and PLD2. Nature communications 456 24637612
2020 ALIX- and ESCRT-III-dependent sorting of tetraspanins to exosomes. The Journal of cell biology 357 32049272
2004 Paraptosis: mediation by MAP kinases and inhibition by AIP-1/Alix. Cell death and differentiation 317 15195070
2007 Structural and biochemical studies of ALIX/AIP1 and its role in retrovirus budding. Cell 275 17350572
2015 Heparanase activates the syndecan-syntenin-ALIX exosome pathway. Cell research 272 25732677
2015 ATG12-ATG3 interacts with Alix to promote basal autophagic flux and late endosome function. Nature cell biology 259 25686249
2013 ALIX and the multivesicular endosome: ALIX in Wonderland. Trends in cell biology 257 24287454
1999 Alix, a novel mouse protein undergoing calcium-dependent interaction with the apoptosis-linked-gene 2 (ALG-2) protein. Cell death and differentiation 215 10200558
2008 ALIX-CHMP4 interactions in the human ESCRT pathway. Proceedings of the National Academy of Sciences of the United States of America 195 18511562
2003 The ALG-2-interacting protein Alix associates with CHMP4b, a human homologue of yeast Snf7 that is involved in multivesicular body sorting. The Journal of biological chemistry 184 12860994
2008 Differential requirements for Alix and ESCRT-III in cytokinesis and HIV-1 release. Proceedings of the National Academy of Sciences of the United States of America 182 18641129
2004 Degradation of AP2 during reticulocyte maturation enhances binding of hsc70 and Alix to a common site on TFR for sorting into exosomes. Traffic (Copenhagen, Denmark) 168 15086793
2012 Get on the exosome bus with ALIX. Nature cell biology 159 22743708
2006 The multiple personalities of Alix. Journal of cell science 157 16868030
2002 Alix (ALG-2-interacting protein X), a protein involved in apoptosis, binds to endophilins and induces cytoplasmic vacuolization. The Journal of biological chemistry 153 12034747
2007 Beyond Tsg101: the role of Alix in 'ESCRTing' HIV-1. Nature reviews. Microbiology 142 17982468
2012 ALIX binds a YPX(3)L motif of the GPCR PAR1 and mediates ubiquitin-independent ESCRT-III/MVB sorting. The Journal of cell biology 140 22547407
2016 Role of Alix in miRNA packaging during extracellular vesicle biogenesis. International journal of molecular medicine 139 26935291
2003 YPXL/I is a protein interaction motif recognized by aspergillus PalA and its human homologue, AIP1/Alix. Molecular and cellular biology 137 12588984
2007 Structural basis for viral late-domain binding to Alix. Nature structural & molecular biology 132 17277784
2007 Structural and functional studies of ALIX interactions with YPX(n)L late domains of HIV-1 and EIAV. Nature structural & molecular biology 129 18066081
2009 Herpes simplex virus type 1 production requires a functional ESCRT-III complex but is independent of TSG101 and ALIX expression. Journal of virology 127 19692479
2007 Potent rescue of human immunodeficiency virus type 1 late domain mutants by ALIX/AIP1 depends on its CHMP4 binding site. Journal of virology 127 17428861
2018 ALIX Regulates Tumor-Mediated Immunosuppression by Controlling EGFR Activity and PD-L1 Presentation. Cell reports 126 30021161
2016 ALIX and ESCRT-I/II function as parallel ESCRT-III recruiters in cytokinetic abscission. The Journal of cell biology 124 26929449
2009 A crescent-shaped ALIX dimer targets ESCRT-III CHMP4 filaments. Structure (London, England : 1993) 111 19523902
2000 The glioma-associated protein SETA interacts with AIP1/Alix and ALG-2 and modulates apoptosis in astrocytes. The Journal of biological chemistry 111 10858458
2001 Yeast PalA/AIP1/Alix homolog Rim20p associates with a PEST-like region and is required for its proteolytic cleavage. Journal of bacteriology 107 11698381
2004 Alix/AIP1 antagonizes epidermal growth factor receptor downregulation by the Cbl-SETA/CIN85 complex. Molecular and cellular biology 101 15456872
2005 Alix regulates cortical actin and the spatial distribution of endosomes. Journal of cell science 98 15914539
2012 Galectin-3 regulates intracellular trafficking of EGFR through Alix and promotes keratinocyte migration. The Journal of investigative dermatology 97 22785133
2019 Flotillin-mediated endocytosis and ALIX-syntenin-1-mediated exocytosis protect the cell membrane from damage caused by necroptosis. Science signaling 93 31138766
2007 Human immunodeficiency virus type 1 Gag engages the Bro1 domain of ALIX/AIP1 through the nucleocapsid. Journal of virology 93 18032513
2021 ALIX and ceramide differentially control polarized small extracellular vesicle release from epithelial cells. EMBO reports 90 33724661
2008 In vitro budding of intralumenal vesicles into late endosomes is regulated by Alix and Tsg101. Molecular biology of the cell 89 18768755
2006 An Alix fragment potently inhibits HIV-1 budding: characterization of binding to retroviral YPXL late domains. The Journal of biological chemistry 86 17158451
2003 Alix, a protein regulating endosomal trafficking, is involved in neuronal death. The Journal of biological chemistry 86 14585841
2010 The ESCRT-associated protein Alix recruits the ubiquitin ligase Nedd4-1 to facilitate HIV-1 release through the LYPXnL L domain motif. Journal of virology 82 20519395
2006 Alix, making a link between apoptosis-linked gene-2, the endosomal sorting complexes required for transport, and neuronal death in vivo. The Journal of neuroscience : the official journal of the Society for Neuroscience 78 16407552
2020 The Flemmingsome reveals an ESCRT-to-membrane coupling via ALIX/syntenin/syndecan-4 required for completion of cytokinesis. Nature communications 77 32321914
2009 Functional role of Alix in HIV-1 replication. Virology 77 19596386
2013 The yeast Alix homolog Bro1 functions as a ubiquitin receptor for protein sorting into multivesicular endosomes. Developmental cell 75 23726974
2005 AIP1/Alix is a binding partner of Sendai virus C protein and facilitates virus budding. Journal of virology 75 15994787
2015 The α-arrestin ARRDC3 mediates ALIX ubiquitination and G protein-coupled receptor lysosomal sorting. Molecular biology of the cell 71 26490116
2010 Alix regulates egress of hepatitis B virus naked capsid particles in an ESCRT-independent manner. Cellular microbiology 70 21129143
2014 Galectin-3 promotes HIV-1 budding via association with Alix and Gag p6. Glycobiology 68 24996823
2012 ALIX is a Lys63-specific polyubiquitin binding protein that functions in retrovirus budding. Developmental cell 67 23201121
2015 WASH and Tsg101/ALIX-dependent diversion of stress-internalized EGFR from the canonical endocytic pathway. Nature communications 66 26066081
2006 The YLDL sequence within Sendai virus M protein is critical for budding of virus-like particles and interacts with Alix/AIP1 independently of C protein. Journal of virology 64 17166905
2004 The penta-EF-hand protein ALG-2 interacts with a region containing PxY repeats in Alix/AIP1, which is required for the subcellular punctate distribution of the amino-terminal truncation form of Alix/AIP1. Journal of biochemistry 62 14999017
2008 Structural basis for Ca2+ -dependent formation of ALG-2/Alix peptide complex: Ca2+/EF3-driven arginine switch mechanism. Structure (London, England : 1993) 61 18940611
2007 A novel requirement for C. elegans Alix/ALX-1 in RME-1-mediated membrane transport. Current biology : CB 61 17997305
2004 Src phosphorylation of Alix/AIP1 modulates its interaction with binding partners and antagonizes its activities. The Journal of biological chemistry 60 15557335
2020 Hepatitis A virus structural protein pX interacts with ALIX and promotes the secretion of virions and foreign proteins through exosome-like vesicles. Journal of extracellular vesicles 58 32082513
2006 Involvement of the conserved adaptor protein Alix in actin cytoskeleton assembly. The Journal of biological chemistry 58 16966331
2008 Alix and ALG-2 are involved in tumor necrosis factor receptor 1-induced cell death. The Journal of biological chemistry 56 18936101
2006 Do Alix and ALG-2 really control endosomes for better or for worse? Biology of the cell 56 16354163
2006 HD-PTP and Alix share some membrane-traffic related proteins that interact with their Bro1 domains or proline-rich regions. Archives of biochemistry and biophysics 55 17174262
2004 CHMP4b is a major binding partner of the ALG-2-interacting protein Alix among the three CHMP4 isoforms. Archives of biochemistry and biophysics 55 14678797
2013 Structure-based in silico identification of ubiquitin-binding domains provides insights into the ALIX-V:ubiquitin complex and retrovirus budding. The EMBO journal 53 23361315
2015 ALIX Rescues Budding of a Double PTAP/PPEY L-Domain Deletion Mutant of Ebola VP40: A Role for ALIX in Ebola Virus Egress. The Journal of infectious diseases 52 25786915
2005 Functions of early (AP-2) and late (AIP1/ALIX) endocytic proteins in equine infectious anemia virus budding. The Journal of biological chemistry 52 16215227
2018 Palmitoylation is a post-translational modification of Alix regulating the membrane organization of exosome-like small extracellular vesicles. Biochimica et biophysica acta. General subjects 51 30251702
2016 ALIX Regulates the Ubiquitin-Independent Lysosomal Sorting of the P2Y1 Purinergic Receptor via a YPX3L Motif. PloS one 50 27301021
2012 AP-3 regulates PAR1 ubiquitin-independent MVB/lysosomal sorting via an ALIX-mediated pathway. Molecular biology of the cell 48 22833563
2009 Penta-EF-hand protein ALG-2 functions as a Ca2+-dependent adaptor that bridges Alix and TSG101. Biochemical and biophysical research communications 47 19520058
2023 SUMOylation-triggered ALIX activation modulates extracellular vesicles circTLCD4-RWDD3 to promote lymphatic metastasis of non-small cell lung cancer. Signal transduction and targeted therapy 46 37925421
2015 ALIX and ESCRT-III coordinately control cytokinetic abscission during germline stem cell division in vivo. PLoS genetics 46 25635693
2010 Interaction between the yellow fever virus nonstructural protein NS3 and the host protein Alix contributes to the release of infectious particles. Microbes and infection 44 21044891
2016 Alix-mediated assembly of the actomyosin-tight junction polarity complex preserves epithelial polarity and epithelial barrier. Nature communications 43 27336173
2021 Plasma-Derived Exosomal ALIX as a Novel Biomarker for Diagnosis and Classification of Pancreatic Cancer. Frontiers in oncology 42 34026608
2011 Activation of the retroviral budding factor ALIX. Journal of virology 42 21715492
2010 Identification and structural characterization of the ALIX-binding late domains of simian immunodeficiency virus SIVmac239 and SIVagmTan-1. Journal of virology 42 20962096
2007 Recruitment of Alix/AIP1 to the plasma membrane by Sendai virus C protein facilitates budding of virus-like particles. Virology 42 18028977
2006 Alix facilitates the interaction between c-Cbl and platelet-derived growth factor beta-receptor and thereby modulates receptor down-regulation. The Journal of biological chemistry 42 17082185
2008 Identification of Alix-type and Non-Alix-type ALG-2-binding sites in human phospholipid scramblase 3: differential binding to an alternatively spliced isoform and amino acid-substituted mutants. The Journal of biological chemistry 41 18256029
2002 Hp95 promotes anoikis and inhibits tumorigenicity of HeLa cells. Oncogene 41 12360406
2001 Overexpression of Hp95 induces G1 phase arrest in confluent HeLa cells. Differentiation; research in biological diversity 39 11683497
2011 ALIX/AIP1 is required for NP incorporation into Mopeia virus Z-induced virus-like particles. Journal of virology 38 21248028
2007 Sendai virus budding in the course of an infection does not require Alix and VPS4A host factors. Virology 38 17467023
2006 POSH, a scaffold protein for JNK signaling, binds to ALG-2 and ALIX in Drosophila. FEBS letters 38 16698022
2020 Detection of serum/salivary exosomal Alix in patients with oral squamous cell carcinoma. Oral diseases 37 32688445
2015 ALG-2 activates the MVB sorting function of ALIX through relieving its intramolecular interaction. Cell discovery 35 27462417
2010 Association of Alix with late endosomal lysobisphosphatidic acid is important for dengue virus infection in human endothelial cells. Journal of proteome research 35 20669987
2020 Alix and Syntenin-1 direct amyloid precursor protein trafficking into extracellular vesicles. BMC molecular and cell biology 34 32731849
2017 Alix is required during development for normal growth of the mouse brain. Scientific reports 34 28322231
2016 ALG-2 interacting protein-X (Alix) is essential for clathrin-independent endocytosis and signaling. Scientific reports 34 27244115
2010 Decoding the intrinsic mechanism that prohibits ALIX interaction with ESCRT and viral proteins. The Biochemical journal 34 20929444
2016 Exosomes in colorectal carcinoma formation: ALIX under the magnifying glass. Modern pathology : an official journal of the United States and Canadian Academy of Pathology, Inc 31 27150162
2015 Unravelling the pivotal role of Alix in MVB sorting and silencing of the activated EGFR. The Biochemical journal 31 25510652
2014 Interaction of HIV-1 Nef protein with the host protein Alix promotes lysosomal targeting of CD4 receptor. The Journal of biological chemistry 31 25118280
2019 ALIX increases protein content and protective function of iPSC-derived exosomes. Journal of molecular medicine (Berlin, Germany) 30 30944935
2004 Ca2+ binding to EF hands 1 and 3 is essential for the interaction of apoptosis-linked gene-2 with Alix/AIP1 in ocular melanoma. Biochemistry 29 15366927
2020 Proline-rich domain of human ALIX contains multiple TSG101-UEV interaction sites and forms phosphorylation-mediated reversible amyloids. Proceedings of the National Academy of Sciences of the United States of America 28 32917811
2012 Alix protein is substrate of Ozz-E3 ligase and modulates actin remodeling in skeletal muscle. The Journal of biological chemistry 28 22334701
2020 Occludin, caveolin-1, and Alix form a multi-protein complex and regulate HIV-1 infection of brain pericytes. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 27 33058236
2019 Centralspindlin Recruits ALIX to the Midbody during Cytokinetic Abscission in Drosophila via a Mechanism Analogous to Virus Budding. Current biology : CB 27 31607533
2022 Plant ESCRT protein ALIX coordinates with retromer complex in regulating receptor-mediated sorting of soluble vacuolar proteins. Proceedings of the National Academy of Sciences of the United States of America 26 35533279

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