Affinage

DYNLL2

Dynein light chain 2, cytoplasmic · UniProt Q96FJ2

Length
89 aa
Mass
10.3 kDa
Annotated
2026-06-09
27 papers in source corpus 15 papers cited in narrative 15 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

DYNLL2 is a conserved homodimeric hub protein that functions not as a simple motor cargo adaptor but as a dimerization engine, recognizing short linear motifs (KXTQT and XG(I/V)QVD types) in intrinsically disordered segments of structurally diverse partners and templating their dimerization (PMID:21777386, PMID:20889982). Structural analysis shows that a disordered partner motif with nascent helical character folds into a β-strand occupying each of the two parallel grooves of the DYNLL2 homodimer, with flanking residues contacting the dimer interface and the symmetry-related partner peptide; recognition follows a conformational-selection mechanism, binding affinity is tuned by phosphorylation (the S88E phosphomimetic raises affinity for dimeric myosin Va), and bivalent ligands gain nanomolar avidity (PMID:20889982, PMID:25312846). Through this activity DYNLL2 is incorporated as a shared subunit of the cytoplasmic dynein-2 complex via the WDR34 intermediate chain, an association required for retrograde intraflagellar transport and ciliogenesis (PMID:25205765, PMID:30649997). The same dimerization activity organizes other systems: it drives oligomerization of the postsynaptic scaffold GKAP to stabilize the postsynaptic density and enhance NMDA receptor activity (PMID:24938595), induces homo- and hetero-dimerization of the BH3-only proteins Bim and Bmf and stabilizes Bmf against 20S proteasomal degradation to modulate apoptosis (PMID:31189926), and mediates MBNL1-dependent retrograde axonal transport of the BDNF–TrkB signaling complex (PMID:36922901). DYNLL2 also interacts with PAK1 to promote endocytosis of bacterial outer membrane vesicles, enabling cytosolic LPS sensing and Caspase-11-dependent pyroptosis (PMID:41360226).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2010 High

    Established the biophysical logic of DYNLL2 ligand recognition—whether it binds defined motifs and how affinity is regulated—resolving that it engages two consensus motifs by conformational selection and is tuned by phosphorylation and ligand valency.

    Evidence In vitro ITC/SPR binding kinetics with S88E mutagenesis on recombinant DYNLL1/DYNLL2

    PMID:20889982

    Open questions at the time
    • Does S88 phosphorylation occur and regulate binding in cells?
    • Kinase responsible for S88 not identified
  2. 2011 High

    Reframed DYNLL1/2 from cargo adaptors to dimerization hubs that act on disordered partner motifs, redefining the conceptual role of the protein family.

    Evidence Review synthesizing structural, peptide-binding, Y2H and co-IP data across labs

    PMID:21777386

    Open questions at the time
    • Review-level synthesis; DYNLL2-specific partner repertoire not exhaustively distinguished from DYNLL1
  3. 2012 Medium

    Identified GKAP as a postsynaptic DYNLL2 partner, linking the hub to synaptic scaffolding and NMDA receptor function.

    Evidence BRET imaging, electrophysiology and interaction-deficient mutants in neurons

    PMID:22328512

    Open questions at the time
    • Stoichiometry and dimerization requirement not yet resolved at this stage
    • Direct vs indirect interaction not structurally defined
  4. 2014 High

    Determined the atomic basis of motif recognition—how a disordered partner peptide is captured—showing it folds into a β-strand across both grooves of the homodimer.

    Evidence NMR, X-ray crystallography and MD on the myosin 5a tail motif bound to DYNLL2

    PMID:25312846

    Open questions at the time
    • Structures of other partner complexes not solved
    • How flanking-region contacts vary across partners unknown
  5. 2014 High

    Placed DYNLL2 as a shared subunit of cytoplasmic dynein-2 via WDR34, connecting the hub to retrograde IFT and ciliogenesis.

    Evidence Co-IP/MS complex definition, cilia localization, WDR34 co-IP and rescue of ATMIN-depletion ciliogenesis defects

    PMID:25205765 PMID:25294941

    Open questions at the time
    • Whether DYNLL2 dimerizes WDR34 specifically not directly shown
    • Functional redundancy with DYNLL1 in cilia not separated
  6. 2014 High

    Demonstrated that DYNLL2 acts as a dimerization engine in a physiological scaffold, quantifying GKAP–DLC2 oligomer composition and showing disruption inhibits NMDA receptor activity.

    Evidence Two-photon sN&B fluorescence fluctuation microscopy with interaction-deficient mutants in living neurons

    PMID:24938595

    Open questions at the time
    • What limits oligomer size in spines vs shafts is unknown
    • Upstream signals controlling activity-dependent assembly not defined
  7. 2019 High

    Extended the dimerization-hub role to apoptosis, showing DYNLL2 drives Bim/Bmf oligomerization, stabilizes Bmf against the 20S proteasome, and mediates Bmf-induced Bim degradation.

    Evidence Cell-free and cellular co-IP, proteasome degradation assays, DYNLL-binding-deficient mutants, cell death assays

    PMID:31189926

    Open questions at the time
    • Relative contribution of DYNLL1 vs DYNLL2 to apoptotic regulation not separated
    • Physiological context where this axis dominates unclear
  8. 2018 Medium

    Mapped DYNLL2 to a defined dynein-2 subcomplex architecture, clarifying which intermediate/light chains it co-assembles with.

    Evidence Visible immunoprecipitation (VIP) interaction mapping of dynein-2 subunits

    PMID:29742051

    Open questions at the time
    • Quantitative stoichiometry within the subcomplex not established
  9. 2019 High

    Showed the WDR34 N-terminal DYNLL-binding region is functionally required, since its deletion acts dominant-negatively to block retrograde ciliary trafficking.

    Evidence KO cell lines with WDR34 deletion/mutant constructs, dominant-negative analysis, ciliary trafficking assays

    PMID:30649997

    Open questions at the time
    • Whether DYNLL binding promotes WDR34 dimerization in this context not directly tested
  10. 2023 Medium

    Linked DYNLL2 to retrograde neurotrophin signaling, showing an RNA-independent MBNL1 interaction is needed for retrograde axonal transport of the BDNF–TrkB complex, with disease relevance in DM1.

    Evidence RNA-independent co-IP, live imaging of co-movement, shRNA knockdown, fractionation, DM1 mouse and human tissue

    PMID:36922901

    Open questions at the time
    • Direct binding motif on MBNL1 not mapped
    • Mechanistic coupling to dynein motor not shown
  11. 2025 Medium

    Implicated DYNLL2 in innate immune signaling, showing a PAK1 interaction drives OMV endocytosis enabling cytosolic LPS sensing and Caspase-11-dependent pyroptosis, with a small-molecule disruptor protective in endotoxemia.

    Evidence Co-IP, siRNA/shRNA knockdown, OMV internalization and Caspase-11/GSDMD cleavage assays, virtual screening, murine endotoxemia model

    PMID:41360226

    Open questions at the time
    • Direct binding interface with PAK1 not structurally defined
    • Whether DYNLL2 dimerizes PAK1 as for other partners untested

Open questions

Synthesis pass · forward-looking unresolved questions
  • How DYNLL2's single dimerization activity is selectively deployed across ciliary, synaptic, apoptotic, transport and immune contexts—and which roles are unique to DYNLL2 versus shared with DYNLL1—remains unresolved.
  • Isoform-specific (DYNLL2 vs DYNLL1) partner selectivity not systematically mapped
  • Cellular regulation of S88 phosphorylation in each context unknown
  • No structures of most partner complexes

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 5
Localization
GO:0005929 cilium 2 GO:0005856 cytoskeleton 1
Pathway
R-HSA-9609507 Protein localization 3 R-HSA-112316 Neuronal System 2 R-HSA-5357801 Programmed Cell Death 2
Complex memberships
cytoplasmic dynein-2

Evidence

Reading pass · 15 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2011 DYNLL1 and DYNLL2 (LC8 family) function as regulatory hub proteins that interact with short linear motifs (KXTQT and XG(I/V)QVD) in intrinsically disordered segments of binding partners, promoting dimerization of those partners rather than acting as simple cargo adaptors for dynein/myosin motors. Binding partners include the dynein intermediate chain, myosin 5a, and diverse scaffold proteins. Review synthesizing structural and functional studies; yeast two-hybrid, peptide binding, co-IP, structural data from multiple labs The FEBS journal High 21777386
2010 DYNLL2 binds two putative consensus motifs (KXTQTX and XG(I/V)QVD) with micromolar affinity, with the KXTQTX motif showing slower off-rate constants. Binding kinetics best fit a conformational selection model (slow isomerization + rapid binding). A phosphomimetic S88E mutant of DYNLL2 shows significantly lower apparent Kd for dimeric myosin Va, suggesting phosphorylation modulates binding affinity. Bivalent (dimeric) ligands show strong avidity effects (nanomolar Kd). In vitro binding assays (isothermal titration calorimetry, surface plasmon resonance), site-directed mutagenesis of DYNLL2 (S88E), comparison of both DYNLL1 and DYNLL2 isoforms The Journal of biological chemistry High 20889982
2014 DYNLL2 binds to an extended linear motif in the intrinsically disordered tail of myosin 5a. In the free form the motif has nascent helical character; upon DYNLL2 binding it folds into a β-strand occupying each of the two parallel binding grooves of the DYNLL2 homodimer. Flanking residues make additional contacts: the N-terminal extension folds back to partially block the free β-sheet edge, and the C-terminal extension contacts the dimer interface and interacts with the symmetry-related second myosin 5a peptide. NMR spectroscopy, X-ray crystallography, molecular dynamics simulations Biochemistry High 25312846
2014 DYNLL1 and DYNLL2 are subunits of the human cytoplasmic dynein-2 complex, which mediates retrograde intraflagellar transport (IFT). They associate with the dynein-2 intermediate chain WDR34. The dynein-2 complex composition was defined to include DYNLL1/DYNLL2 as shared subunits. Co-immunoprecipitation/mass spectrometry to define complex composition; siRNA knockdown functional assays for cilia Journal of cell science High 25205765
2014 DYNLL1 and DYNLL2 localize to cilia in puncta consistent with IFT particles and physically interact with WDR34, a dynein-2 intermediate chain. Expression of DYNLL2 (as well as DYNLL1) rescued ciliogenesis defects caused by ATMIN/DYNLL1 depletion, placing DYNLL2 downstream of ATMIN in ciliogenesis. Immunofluorescence localization to cilia, co-immunoprecipitation with WDR34, rescue experiments by exogenous DYNLL2 expression in ATMIN-depleted cells Development (Cambridge, England) Medium 25294941
2018 DYNLL1 and DYNLL2 associate with the WDR34 intermediate chain subcomplex of dynein-2 (WDR34-DYNLL1/DYNLL2-DYNLRB1/DYNLRB2). This subcomplex is one of three defined subcomplexes in dynein-2. Visible immunoprecipitation (VIP) assay mapping pairwise and higher-order interactions among dynein-2 subunits Molecular biology of the cell Medium 29742051
2019 The WDR34 intermediate chain interacts with DYNLL1/DYNLL2 and DYNLRB1/DYNLRB2 via distinct sites. These interactions are required for retrograde ciliary protein trafficking; WDR34 constructs lacking the DYNLL-binding N-terminal region act in a dominant-negative manner, inhibiting ciliary biogenesis and retrograde trafficking. Co-immunoprecipitation, knockout cell lines expressing WDR34 deletion/mutant constructs, dominant-negative analysis, ciliary trafficking assays Molecular biology of the cell High 30649997
2012 DYNLL2 (DLC2) interacts with the postsynaptic scaffold protein GKAP in dendritic spines. This interaction stabilizes scaffolding protein expression at the postsynaptic density and enhances synaptic NMDA receptor activity. The interaction is favored by sustained synaptic activity. BRET imaging, immunostaining, electrophysiological recording of NMDA receptor-mediated currents, co-expression of interaction-deficient mutants Journal of cell science Medium 22328512
2014 DYNLL2 (DLC2) dimerization is required for its interaction with GKAP. DLC2-GKAP hetero-oligomeric complexes in dendritic shafts comprise ~2 DLC2 and ~2 GKAP monomers, while in dendritic spines the complexes are much larger (~16 DLC2 and ~13 GKAP monomers). Disruption of the GKAP-DLC2 interaction destabilizes these oligomers, decreases spine-preferential localization of GKAP, and inhibits NMDA receptor activity. DLC2 thus acts as a dimerization engine for GKAP to organize the postsynaptic scaffold. Two-photon scanning number and brightness (sN&B) fluorescence fluctuation microscopy in living neurons, GKAP-DLC2 interaction-deficient mutants Journal of cell science High 24938595
2019 DYNLL1 and DYNLL2 both interact with the BH3-only proteins Bim and Bmf (not exclusively isoform-specific). DYNLL1/2 induces homodimerization of Bim and Bmf and promotes formation of ternary Bim-DYNLL-Bmf complexes in cell-free and cellular systems. DYNLL-induced oligomerization stabilizes Bmf by inhibiting its degradation by the ubiquitin-independent 20S proteasome. A DYNLL-binding-deficient Bmf mutant fails to induce Bim degradation upon overexpression, demonstrating that Bmf-induced Bim degradation is mediated through modulation of Bim-DYNLL association rather than displacement from anti-apoptotic Bcl-2 proteins. Co-immunoprecipitation in cell-free and cellular systems, proteasome degradation assays, overexpression of DYNLL-binding-deficient mutants, cell death assays Cell death and differentiation High 31189926
2011 DYNLL1 (but not DYNLL2) is expressed in retinal ganglion cells (RGCs), while DYNLL2 is expressed in glia. DYNLL1 is persistently downregulated in glaucomatous RGCs at all measured timepoints. DYNLL2 is upregulated in glia at 2 weeks after glaucoma induction. These findings indicate non-redundant, cell-type-specific expression patterns for the two isoforms in the retina. Laser capture microdissection of RGCs and GCL, quantitative PCR, immunostaining, rat chronic glaucoma model Experimental eye research Medium 21145319
2023 DYNLL2 interacts with MBNL1 in an RNA-independent manner. In neurons, overexpressed MBNL1 and DYNLL2 co-traffic along axonal processes. MBNL1 knockdown impairs retrograde movement of DYNLL2, and in DM1 model brains with reduced cytoplasmic MBNL1, DYNLL2 and activated pTrkB accumulate in the synaptosomal fraction, indicating impaired DYNLL2-dependent retrograde transport of the BDNF-TrkB signaling complex. Co-immunoprecipitation (RNA-independent), time-lapse live imaging of MBNL1 and DYNLL2 co-movement, shRNA knockdown, subcellular fractionation, DM1 mouse model and postmortem human tissue analysis Acta neuropathologica communications Medium 36922901
2025 DYNLL2 interacts with PAK1 to regulate endocytosis of Gram-negative bacterial outer membrane vesicles (OMVs), facilitating cytosolic LPS release and subsequent Caspase-11 inflammasome activation, thereby triggering pyroptosis (GSDMD cleavage). Depletion of DYNLL2 suppresses OMV internalization, Caspase-11/GSDMD cleavage, and proinflammatory cytokine release. Oroxylin A was identified as an inhibitor of the DYNLL2-PAK1 interaction, blocking Caspase-11-dependent pyroptosis in vitro and improving survival in murine endotoxemia models. Co-immunoprecipitation (DYNLL2-PAK1 interaction), siRNA/shRNA knockdown of DYNLL2 and PAK1, OMV internalization assays, caspase-11/GSDMD cleavage assays, virtual screening, in vitro pharmacological inhibition, murine endotoxemia model Biochemical pharmacology Medium 41360226
2015 DYNLL2 levels are regulated by fed/fasted states in oxytocin neurons of the hypothalamic paraventricular nucleus, with higher DYNLL2 expression under fed conditions correlating with increased NMDA receptor-mediated excitatory postsynaptic currents, suggesting DYNLL2 participates in NMDA receptor trafficking to the postsynaptic site under metabolic regulation. Electrophysiological recording (mEPSC and NMDA-EPSCs), western blot/immunostaining for DYNLL2 in PVN oxytocin neurons, fed vs. fasted paradigm Neuropeptides Low 26344333
2011 DYNLL1 binds to multiple SQ/TQ motifs in the C-terminal domain of ATMIN via KXTQT-type linear motifs. Co-expression of mCherry-DYNLL1 and GFP-ATMIN mutually affects intracellular localization: DYNLL1 presence partly impedes DNA-damage-induced nuclear foci formation by ATMIN. (Note: DYNLL2 is implied to share similar binding capacity but DYNLL1 was the primary focus.) Yeast two-hybrid screen, pepscan, recombinant expression/gel filtration, NMR-modeled complexes, co-expression fluorescence localization, DNA damage assay Biochemical and biophysical research communications Medium 21971545

Source papers

Stage 0 corpus · 27 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2011 DYNLL/LC8: a light chain subunit of the dynein motor complex and beyond. The FEBS journal 108 21777386
2021 Role of Calcium Signaling Pathway-Related Gene Regulatory Networks in Ischemic Stroke Based on Multiple WGCNA and Single-Cell Analysis. Oxidative medicine and cellular longevity 89 34987704
2014 Subunit composition of the human cytoplasmic dynein-2 complex. Journal of cell science 83 25205765
2011 Convergent functional genomics of anxiety disorders: translational identification of genes, biomarkers, pathways and mechanisms. Translational psychiatry 72 22832404
2018 Interaction of WDR60 intermediate chain with TCTEX1D2 light chain of the dynein-2 complex is crucial for ciliary protein trafficking. Molecular biology of the cell 54 29742051
2019 Interactions of the dynein-2 intermediate chain WDR34 with the light chains are required for ciliary retrograde protein trafficking. Molecular biology of the cell 40 30649997
2014 ATMIN is a transcriptional regulator of both lung morphogenesis and ciliogenesis. Development (Cambridge, England) 39 25294941
2008 An association analysis of murine anxiety genes in humans implicates novel candidate genes for anxiety disorders. Biological psychiatry 39 18639233
2011 LC8 dynein light chain (DYNLL1) binds to the C-terminal domain of ATM-interacting protein (ATMIN/ASCIZ) and regulates its subcellular localization. Biochemical and biophysical research communications 31 21971545
2010 Affinity, avidity, and kinetics of target sequence binding to LC8 dynein light chain isoforms. The Journal of biological chemistry 29 20889982
2019 Dynein light chain binding determines complex formation and posttranslational stability of the Bcl-2 family members Bmf and Bim. Cell death and differentiation 25 31189926
2013 The novel zinc finger protein dASCIZ regulates mitosis in Drosophila via an essential role in dynein light-chain expression. Genetics 25 24336747
2012 GKAP-DLC2 interaction organizes the postsynaptic scaffold complex to enhance synaptic NMDA receptor activity. Journal of cell science 19 22328512
2015 Fasted/fed states regulate postsynaptic hub protein DYNLL2 and glutamatergic transmission in oxytocin neurons in the hypothalamic paraventricular nucleus. Neuropeptides 18 26344333
2014 DYNLL2 dynein light chain binds to an extended linear motif of myosin 5a tail that has structural plasticity. Biochemistry 17 25312846
2021 LncRNAs and their regulatory networks in breast muscle tissue of Chinese Gushi chickens during late postnatal development. BMC genomics 16 33422015
2021 Integrative bioinformatics approaches for identifying potential biomarkers and pathways involved in non-obstructive azoospermia. Translational andrology and urology 14 33532314
2022 Weighted gene co-expression network indicates that the DYNLL2 is an important regulator of chicken breast muscle development and is regulated by miR-148a-3p. BMC genomics 13 35379193
2014 The stoichiometry of scaffold complexes in living neurons - DLC2 functions as a dimerization engine for GKAP. Journal of cell science 9 24938595
2017 The Drosophila LC8 homolog cut up specifies the axonal transport of proteasomes. Journal of cell science 8 28808087
2024 Aggrephagy-related gene signature correlates with survival and tumor-associated macrophages in glioma: Insights from single-cell and bulk RNA sequencing. Mathematical biosciences and engineering : MBE 7 38454689
2010 The expression of dynein light chain DYNLL1 (LC8-1) is persistently downregulated in glaucomatous rat retinal ganglion cells. Experimental eye research 7 21145319
2023 Machine learning and integrative analysis identify the common pathogenesis of azoospermia complicated with COVID-19. Frontiers in immunology 6 37283758
2023 Loss of MBNL1-mediated retrograde BDNF signaling in the myotonic dystrophy brain. Acta neuropathologica communications 3 36922901
2018 Neospora caninum cytoplasmic dynein LC8 light chain 2 (NcDYNLL2) is differentially produced by pathogenically distinct isolates and regulates the host immune response. Parasitology 3 30561290
2025 Targeting the DYNLL2-PAK1 axis inhibits caspase-11-dependent pyroptosis to alleviate sepsis. Biochemical pharmacology 2 41360226
2024 Higher expression of TSR2 aggravating hypertension via the PPAR signaling pathway. Aging 2 38814181

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