Affinage

SLC6A5

Sodium- and chloride-dependent glycine transporter 2 · UniProt Q9Y345

Round 2 corrected
Length
797 aa
Mass
87.4 kDa
Annotated
2026-04-28
130 papers in source corpus 35 papers cited in narrative 34 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

SLC6A5 (GlyT2) is a presynaptic Na⁺/Cl⁻-dependent glycine transporter that concentrates glycine into glycinergic nerve terminals to sustain vesicular glycine loading and inhibitory neurotransmission in the spinal cord, brainstem, and cerebellum (PMID:7861131, PMID:9387864, PMID:17554001, PMID:18815261). Transport depends on residues in transmembrane domains that coordinate Na⁺ and glycine binding (e.g., Tyr-289, Asp-471), N-glycosylation for ER-to-surface trafficking assisted by calnexin, and association with cholesterol-rich membrane rafts (PMID:10788509, PMID:22132725, PMID:11036075, PMID:23650557, PMID:18266927). Surface expression is dynamically regulated by syntaxin 1A–mediated exocytic delivery, clathrin-dependent endocytosis driven by PKC-stimulated ubiquitination of a C-terminal lysine cluster (K751/K773/K787/K791) by E3 ligases LNX1/LNX2, and recycling through Rab11-positive endosomes, with Na⁺/K⁺-ATPase, PMCA, and NCX1 forming a functional raft-resident complex that sustains local ion homeostasis required for transport (PMID:11278707, PMID:21910806, PMID:23484054, PMID:31628376, PMID:19374720, PMID:23986260, PMID:25315779). Loss-of-function mutations in SLC6A5 cause hereditary hyperekplexia (startle disease) in humans by impairing glycine uptake and presynaptic glycine recycling (PMID:16751771, PMID:22700964).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 1995 High

    Establishing where GlyT2 acts: immunolocalization to axonal varicosities in spinal cord, brainstem, and cerebellum—matching glycine receptor distribution—positioned GlyT2 as the presynaptic glycine reuptake system at inhibitory synapses.

    Evidence Immunocytochemistry with antibodies against recombinant GlyT2 domains in mouse brain

    PMID:7582108 PMID:7861131

    Open questions at the time
    • Subcellular resolution to synaptic boutons not achieved
    • Glial expression possibility not addressed
  2. 1997 High

    Demonstrating GlyT2 is the glycine concentrator: functional uptake assays showed GlyT2 expression directly correlates with intracellular glycine accumulation, establishing its role in loading glycinergic terminals with substrate.

    Evidence Double-immunofluorescence and glycine uptake in spinal neurons and GlyT2-transfected COS cells

    PMID:9387864

    Open questions at the time
    • Coupling to vesicular filling not yet demonstrated
    • Stoichiometry of Na⁺/Cl⁻/glycine cotransport not established
  3. 1998 High

    Molecular characterization of human GlyT2 revealed a 797-residue transporter with Km ~108 µM for glycine, sarcosine-insensitive, mapping to 11p15.1-15.2, and alternative splicing yielding functionally distinct isoforms (uptake vs. exchange modes).

    Evidence Expression cloning in CHO cells with kinetic analysis; RACE analysis and transport assays of splice variants in COS cells

    PMID:9509996 PMID:9845349

    Open questions at the time
    • Physiological relevance of the exchange-mode isoform (GLYT2b) unclear
    • 3 Na⁺ stoichiometry not yet formally demonstrated
  4. 2000 High

    Structure–function dissection identified key determinants of transport: Tyr-289 in TM III is essential for ion coupling and substrate permeation, four N-glycosylation sites (N345/N355/N360/N366) are required for surface delivery and activity, and syntaxin 1A interaction regulates membrane expression.

    Evidence Site-directed mutagenesis with electrophysiology and uptake assays in HEK-293 and COS cells; co-immunoprecipitation of syntaxin 1A in COS cells and rat brain with functional consequence

    PMID:10722844 PMID:10788509 PMID:11036075

    Open questions at the time
    • High-resolution structural basis for ion/glycine coordination not available
    • Whether syntaxin 1A interaction is direct or scaffolded unclear
  5. 2001 High

    Resolving how GlyT2 reaches the surface: syntaxin 1A mediates SNARE-dependent exocytic delivery of GlyT2 from intracellular vesicles to the plasma membrane, as BoNT/C blockade prevents surface insertion without affecting internalization, and immunogold EM placed GlyT2 on small synaptic-like vesicles.

    Evidence Synaptosome stimulation, BoNT/C neurotoxin treatment, immunogold electron microscopy in brain-derived preparations

    PMID:11278707

    Open questions at the time
    • Identity of the vesicle population (recycling vs. biosynthetic) carrying GlyT2 not resolved
    • Whether SNARE-dependent insertion is activity-regulated in vivo unknown
  6. 2006 High

    SLC6A5 was established as a human disease gene: multiple loss-of-function mutations cause hereditary hyperekplexia by impairing glycine uptake and/or subcellular localization, linking presynaptic glycine recycling failure to startle disease.

    Evidence Sequencing of hyperekplexia patient cohorts with heterologous functional validation of missense, nonsense, and frameshift mutations

    PMID:16751771 PMID:16884688

    Open questions at the time
    • Genotype–phenotype severity correlations incomplete
    • Whether partial loss-of-function alleles cause milder phenotypes not established
  7. 2007 High

    The physiological rationale for GlyT2 was clarified: GlyT2's inability to operate in reverse (unlike GlyT1) enables it to maintain cytosolic glycine at concentrations sufficient for vesicular loading by VIAAT, directly coupling plasma membrane uptake to quantal glycinergic release.

    Evidence Reconstitution in neuroendocrine cells co-expressing VIAAT and plasmalemmal transporters with double-sniffer patch clamp

    PMID:17554001

    Open questions at the time
    • Structural basis for the irreversibility of GlyT2 transport not known
    • Whether VIAAT and GlyT2 physically interact not tested
  8. 2008 High

    GlyT2-mediated glycine refilling was shown to be rate-limiting for sustained glycinergic transmission: pharmacological GlyT2 blockade in spinal neurons caused a switch from glycinergic to GABAergic phenotype, revealing two kinetically distinct vesicle pools with different refilling requirements; membrane raft association was demonstrated as essential for transport activity.

    Evidence Electrophysiology in GlyT2-eGFP transgenic neurons with GlyT2 inhibitors; detergent-resistant membrane fractionation and cholesterol depletion in synaptosomes

    PMID:18266927 PMID:18815261

    Open questions at the time
    • Molecular mechanism of raft-dependent transport enhancement unknown
    • Whether glycine-to-GABA switch occurs in vivo not shown
  9. 2008 High

    PKC was identified as a central regulator of GlyT2 surface levels: PKC activation promotes GlyT2 internalization and redistribution from raft to non-raft domains, with K422 identified as a regulatory determinant; this introduced ubiquitination-dependent trafficking control.

    Evidence PMA treatment with surface biotinylation, raft fractionation, and K422E mutagenesis in HEK cells and brainstem neurons

    PMID:18341477

    Open questions at the time
    • Whether K422 is directly ubiquitinated or indirectly involved not resolved
    • In vivo PKC activating signal identity unknown
  10. 2009 High

    GlyT2 recycling was mapped to Rab11-positive endosomes: dominant-negative Rab11 impaired both GlyT2 trafficking and glycine transport, establishing the slow recycling pathway as the intracellular itinerary for internalized transporter.

    Evidence Biochemical fractionation, immunogold EM, dominant-negative Rab11 co-expression with transport assay in rat brainstem

    PMID:19374720

    Open questions at the time
    • Sorting signals directing GlyT2 into Rab11 compartments not identified
    • Whether Rab4-dependent fast recycling also contributes not tested
  11. 2011 High

    The endocytic route was defined: GlyT2 internalization is primarily clathrin-mediated, with constitutive endocytosis occurring from raft domains and PKC-stimulated endocytosis following redistribution to non-raft membrane.

    Evidence Clathrin pathway inhibitors, dominant-negative mutants, siRNA knockdown with internalization assays in HEK cells and neurons

    PMID:21910806

    Open questions at the time
    • Adaptor proteins linking GlyT2 to clathrin not identified
    • Relative contribution of ubiquitin-dependent vs. -independent endocytosis not quantified
  12. 2012 High

    Expanded genotype–function mapping revealed diverse pathogenic mechanisms in hyperekplexia: 20 SLC6A5 variants disrupted Cl⁻ binding, Na⁺ affinity, cation-π interactions, or extracellular loop conformation; the dominant Y705C mutation in TM11 impaired secretory pathway maturation via aberrant disulfide interactions.

    Evidence Large patient cohort sequencing with functional validation of 16 mutations; electrophysiology, cysteine labeling, and secretory pathway analysis for Y705C

    PMID:22700964 PMID:22753417

    Open questions at the time
    • No high-resolution structure to map all mutation sites
    • Therapeutic rescue strategies for trafficking-defective mutants not explored
  13. 2013 High

    The GlyT2 interactome was expanded: Na⁺/K⁺-ATPase was identified as a direct partner that preferentially associates with raft-resident active GlyT2 and regulates its endocytosis across species; C-terminal ubiquitination at K751/K773/K787/K791 was shown to drive constitutive endocytosis and degradation; calnexin was identified as an ER chaperone for GlyT2 biogenesis via both glycan-dependent and lectin-independent modes.

    Evidence Mass spectrometry interactomics with reciprocal co-IP, ouabain treatment in neurons/zebrafish/rats; systematic lysine mutagenesis with ubiquitination and turnover assays; CNX knockdown/overexpression with transport and surface expression assays

    PMID:23484054 PMID:23650557 PMID:23986260

    Open questions at the time
    • Direct binding interface between NKA and GlyT2 not mapped
    • Whether calnexin interaction is specific to GlyT2 among SLC6 family unknown
  14. 2014 Medium

    A raft-resident functional complex of GlyT2 with PMCA2/3 and NCX1 was identified, suggesting coordinated local Na⁺/Ca²⁺ handling at glycinergic terminals; GSK3β was shown to differentially stimulate GlyT2 activity while inhibiting GlyT1.

    Evidence Co-immunoprecipitation and pharmacological inhibition of PMCA/NCX with transport assays; co-expression in COS-7 and oocytes with phosphorylation and transport readouts

    PMID:25301276 PMID:25315779

    Open questions at the time
    • Whether PMCA/NCX/GlyT2 complex exists in vivo at synapses not confirmed by super-resolution imaging
    • GSK3β phosphorylation site(s) on GlyT2 not mapped
    • Functional significance of GSK3β regulation in vivo unknown
  15. 2019 High

    The E3 ubiquitin ligases responsible for GlyT2 ubiquitination were identified: LNX1 and LNX2 ubiquitinate the same C-terminal lysine cluster via their RING domains; genetic deletion of LNX2 in spinal neurons increased GlyT2 expression, and LNX2 is required for PKC-mediated transport regulation.

    Evidence Unbiased E3 ligase screening, co-IP, RING-domain mutagenesis, LNX2 knockout neurons with PKC activation

    PMID:31628376

    Open questions at the time
    • Whether LNX1 and LNX2 are redundant or specialized in vivo not determined
    • Upstream signals activating LNX2 beyond PKC not identified

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the high-resolution structure of GlyT2 in multiple conformational states, the structural basis for its inability to operate in reverse, whether GlyT2 and VIAAT are physically coupled for vesicular glycine refilling, and therapeutic strategies to rescue trafficking-defective hyperekplexia mutants.
  • No experimental GlyT2 structure available
  • Physical coupling between GlyT2 and VIAAT not tested
  • Pharmacological chaperone rescue of misfolded mutants not explored

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005215 transporter activity 5 GO:0140104 molecular carrier activity 2
Localization
GO:0005886 plasma membrane 6 GO:0005768 endosome 2 GO:0031410 cytoplasmic vesicle 2
Pathway
R-HSA-382551 Transport of small molecules 5 R-HSA-9609507 Protein localization 5 R-HSA-112316 Neuronal System 4 R-HSA-5653656 Vesicle-mediated transport 3
Partners
ATP1A1ATP2B2ATP2B3CANXLNX1LNX2SLC8A1STX1A

Evidence

Reading pass · 34 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1995 GLYT2 (SLC6A5) is expressed predominantly in neurons of the spinal cord, brainstem, and cerebellum, with immunoreactivity localized to axonal processes with varicosities but not cell bodies, correlating with the distribution of glycine receptors and suggesting a role in terminating glycine neurotransmission at inhibitory synapses. Western blot, immunocytochemistry with polyclonal antibodies against recombinant N-terminus and loop fusion proteins in mouse brain Journal of neurochemistry High 7582108 7861131
1997 GLYT2 is the active concentrator of glycine in glycinergic neurons: GLYT2 expression in cultured spinal neurons and GLYT2-transfected COS cells directly correlates with intracellular glycine accumulation, establishing that GLYT2-mediated uptake is responsible for the high glycine content of glycinergic neurons and that GLYT2 can be used as a reliable marker for glycinergic neurons. Double-immunofluorescence co-localization, glycine uptake assays in cultured spinal neurons and GLYT2-transfected COS cells Brain research. Molecular brain research High 9387864
1998 Human GlyT2 encodes a 797 amino acid Na+/Cl−-dependent glycine transporter with an apparent Km of 108 μM for glycine, insensitive to sarcosine, and the gene maps to chromosome 11p15.1-15.2. Expression cloning, stable expression in CHO cells, glycine uptake assays, radiation hybrid mapping FEBS letters High 9845349
1998 Two alternative 5' isoforms of rat GLYT2 (GLYT2a and GLYT2b) are generated by alternative exon usage; GLYT2a actively accumulates glycine in transfected COS cells whereas GLYT2b only exchanges (releases) glycine, indicating functionally distinct transport modes. RACE analysis, genomic DNA analysis, transport assays in transfected COS cells Neuroscience letters Medium 9509996
2000 Tyrosine 289 in transmembrane domain III of GLYT2a is critical for ion coupling and glycine transport: mutations Y289W, Y289F, and Y289S abolish or dramatically reduce transport activity, alter Na+ affinity and cooperativity, reduce Na+ selectivity, and decrease Cl− dependence, indicating that TM III forms part of the ion and substrate permeation pathway. Site-directed mutagenesis, electrophysiology (steady-state currents), glycine uptake assays in HEK-293 cells The Journal of biological chemistry High 10788509
2000 N-glycosylation at four sites (Asn-345, Asn-355, Asn-360, Asn-366) in the large extracellular loop of GLYT2 is required for transport activity, proper intracellular trafficking to the plasma membrane in COS cells, and apical sorting in polarized MDCK cells; enzymatic deglycosylation reduces transport activity by 35-40%; apical localization occurs via a glycolipid raft-independent pathway. Site-directed mutagenesis of glycosylation sites, enzymatic deglycosylation, transport assays, biotinylation assays, transfection in COS and MDCK cells The Journal of biological chemistry High 11036075
2000 Syntaxin 1A physically interacts with both GLYT1 and GLYT2 in COS cells and rat brain tissue (co-immunoprecipitation), and co-transfection of syntaxin 1A inhibits glycine transport by ~40%; this inhibition is reversed by Munc18; syntaxin 1A interaction partially removes the transporters from the plasma membrane as shown by biotinylation. Co-immunoprecipitation in COS cells and rat brain, glycine transport assays, cell-surface biotinylation FEBS letters High 10722844
2000 Ethanol selectively and acutely inhibits GLYT2a but not GLYT1b in a non-competitive (allosteric) manner, with a cut-off at alcohols with four carbons suggesting a specific binding site; chronic ethanol treatment decreases GLYT2a activity and surface expression while slightly increasing GLYT1b function. Glycine uptake assays in stably transfected HEK-293 cells, n-alkanol inhibition series British journal of pharmacology Medium 10683205
2001 Syntaxin 1 mediates GLYT2 trafficking to the plasma membrane via a SNARE-dependent mechanism: under conditions stimulating vesicular glycine release (exocytosis) in synaptosomes, GLYT2 traffics first to the plasma membrane and then is internalized; inactivation of syntaxin 1 with BoNT/C prevents GLYT2 from reaching the plasma membrane but does not block internalization. Immunogold labeling shows GLYT2 is present in small synaptic-like vesicles. Synaptosome stimulation, BoNT/C neurotoxin treatment, surface expression assays, immunogold electron microscopy The Journal of biological chemistry High 11278707
2004 Calpain (Ca2+-dependent protease) cleaves the extended N-terminal cytoplasmic region of GlyT2 at two sites (after Met-156 and Gly-164) in spinal cord synaptosomes and cultured neurons; the resulting ~70 kDa N-terminally truncated GlyT2 retains full transport activity upon expression in HEK293 cells, suggesting calpain-mediated proteolysis may regulate GlyT2 trafficking or function. Calpain activation in synaptosomes and cultured neurons, protein sequence analysis of cleavage sites, expression of truncated GlyT2 in HEK293 cells with transport assay Journal of neurochemistry Medium 14675166
2006 Missense, nonsense, and frameshift mutations in SLC6A5 (encoding GlyT2) cause hereditary hyperekplexia; these mutations result in defective subcellular GlyT2 localization, decreased glycine uptake, or both; selected mutations affect predicted glycine and Na+ binding sites, establishing SLC6A5 as a presynaptic disease gene. Human genetics (sequencing), heterologous expression with glycine uptake assays, subcellular localization studies Nature genetics High 16751771
2006 Truncating mutations in the human GlyT2 gene (SLC6A5) cause complete loss of glycine transport function, confirming SLC6A5 as a disease gene in human hyperekplexia. DNA sequencing of patients, heterologous expression and glycine uptake assays Biochemical and biophysical research communications High 16884688
2007 GlyT2 (but not GlyT1) maintains the high cytosolic glycine concentration required for efficient vesicular loading by VIAAT: in a neuroendocrine cell reconstitution system, GlyT2 more effectively than GlyT1 supports glycinergic quantal release because GlyT2 cannot operate in reverse mode, thereby maintaining cytosolic glycine for vesicle refilling. Reconstitution in neuroendocrine cells co-expressing VIAAT and plasmalemmal transporters; double-sniffer patch-clamp technique measuring quantal glycine/GABA release The Journal of neuroscience High 17554001
2007 The C-terminal PDZ-ligand motif of GlyT2 is required for efficient synaptic localization in neurons: inactivation of this motif does not impair transport function in HEK293T cells but reduces co-localization with synaptic markers by up to 50% in hippocampal neurons, indicating the PDZ motif is important for trafficking to and/or stabilization at synaptic sites. Site-directed mutagenesis, transfection of hippocampal neurons, immunofluorescence co-localization with synaptic markers Molecular and cellular neurosciences Medium 17851090
2007 GLYT2 is expressed in a subset of GFAP-positive astrocyte-derived gliosomes in mouse spinal cord and contributes to glycine-evoked GABA release from GABAergic synaptosomes, demonstrating that GLYT2 localization is not exclusively neuronal and that it functionally contributes to GABAergic signaling via glycine transport. Purified synaptosome and gliosome preparations, pharmacological blockade with selective GLYT1/GLYT2 inhibitors, confocal microscopy co-labeling with GAT1, GFAP, MAP2 Neurochemistry international Medium 17597258
2008 GlyT2-mediated glycine uptake is directly coupled to the refilling of recycling synaptic vesicles: GlyT2 inhibition in cultured spinal neurons causes a switch from glycinergic to GABAergic inhibitory phenotype through cytosolic glycine depletion, fully reversible by glycine resupply; high-frequency stimulation reveals two kinetically distinct phases of vesicle release with different requirements for GlyT2-mediated glycine refilling. Whole-cell and synaptic electrophysiology in GlyT2-eGFP transgenic mouse spinal cord neurons; glycine transport current recording; pharmacological GlyT2 inhibition The Journal of neuroscience High 18815261
2008 GLYT2 is associated with membrane rafts (Triton X-100-insoluble fractions) in brainstem terminals and neurons; cholesterol and sphingomyelin depletion with methyl-β-cyclodextrin disrupts raft association and impairs transport activity, indicating that the lipid raft environment is required for optimal GLYT2 function. Detergent-resistant membrane fractionation, methyl-β-cyclodextrin treatment, confocal microscopy, electron microscopy on purified raft fractions, transport assays in synaptosomes and neurons Journal of neurochemistry High 18266927
2008 Protein kinase C (PKC) activation by PMA inhibits GLYT2 transport by increasing internalization rate and accumulating GLYT2 in intracellular compartments; PMA also redistributes GLYT2 from raft to non-raft membrane domains; a regulatory lysine mutant K422E is resistant to PMA-induced trafficking modulation; Rac1 signaling also participates in PMA action on GLYT2. PMA treatment, monensin trafficking assay, surface biotinylation, raft fractionation, mutagenesis (K422E), siRNA knockdown in HEK cells, brainstem neurons and synaptosomes The Biochemical journal High 18341477
2009 GLYT2 resides intracellularly in Rab11-positive recycling endosomes and synaptophysin-containing vesicles in rat brainstem; coexpression of dominant-negative Rab11 impairs GLYT2 trafficking and glycine transport, establishing that GlyT2 recycles through Rab11-positive endosomal compartments. Biochemical fractionation, immunofluorescence and immunogold electron microscopy, dominant-negative Rab11 co-expression with transport assay Traffic (Copenhagen, Denmark) High 19374720
2011 GlyT2 endocytosis is mediated primarily by the clathrin pathway; PKC activation promotes redistribution of GLYT2 from raft to non-raft membrane domains and increases ubiquitinated GLYT2 endocytosis; GLYT2 is constitutively internalized from cell-surface lipid rafts while remaining raft-associated in recycling structures. Pharmacological inhibitors of clathrin pathway, dominant-negative mutants, siRNA knockdown, raft fractionation, internalization assays in HEK cells and neurons Traffic (Copenhagen, Denmark) High 21910806
2012 Twenty SLC6A5 sequence variants were identified in hyperekplexia patients, all defective in glycine transport; pathogenic mechanisms include protein truncation (R439X most common), splice site mutations, missense mutations affecting Cl− binding, extracellular loop 4 conformational changes, and cation-π interactions; mutation A275T causes voltage-sensitive decrease in glycine transport through lower Na+ affinity. DNA sequencing of 93 hyperekplexia patients, glycine uptake assays for 16 novel mutations, electrophysiology (A275T) The Journal of biological chemistry High 22700964
2012 The dominant hyperekplexia mutation Y705C in TM11 of GlyT2 introduces a cysteine that interacts with the Cys-311–Cys-320 pair in external loop 2, impairing transporter maturation through the secretory pathway, reducing surface expression, and inhibiting transport; Y705C also shows altered H+ and Zn2+ sensitivity of glycine transport. Molecular modeling, electrophysiology, [3H]glycine transport assays, cell surface expression assays, cysteine labeling, secretory pathway analysis The Journal of biological chemistry High 22753417
2012 A MusD retrotransposon insertion in mouse Slc6a5 abolishes GlyT2 protein expression (null allele), causing handling-induced spasms from day 5 and death by two weeks; loss of GlyT2 accelerates NMJ synapse elimination and the embryonic-to-adult AChR subunit switch, consistent with increased motor neuron activity; heterozygous mice show repetitive grooming and hyperactivity. Spontaneous mouse mutant characterization, Western blot, neuromuscular junction analysis, behavioral phenotyping PloS one High 22272310
2012 Asp471 in the external vestibule of GLYT2 controls cation access and Na+ coupling: substitution of Asp471 reduces Na+ affinity and cooperativity, increases sodium leakage and uncoupled ion movements through GLYT2, and alters Na+/Li+-induced conformational changes; this residue is distinct from the homologous Asp295 in GLYT1, explaining the different lithium sensitivity of the two transporters. Homology modeling, molecular dynamics simulation, site-directed mutagenesis, electrophysiology, [3H]glycine transport assays, cation accessibility assays The Biochemical journal High 22132725
2013 Na+/K+-ATPase (NKA) is a direct interacting partner of GlyT2 in the CNS: identified by mass spectrometry interactomics, confirmed by reciprocal co-immunoprecipitation and immunocytochemistry; NKA preferentially interacts with the raft-associated active pool of GlyT2; ouabain-mediated NKA inhibition modulates GlyT2 endocytosis and total expression in neurons, in zebrafish embryos, and in adult rats, indicating an evolutionarily conserved regulatory mechanism. High-throughput mass spectrometry, reciprocal co-immunoprecipitation, immunocytochemistry, ouabain treatment in neurons, in vivo zebrafish and rat models The Journal of neuroscience High 23986260
2013 Ubiquitination of a C-terminal four-lysine cluster (K751, K773, K787, K791) in GlyT2 is required for constitutive endocytosis, sorting into the slow recycling pathway, and transporter turnover; increased ubiquitination by PKC activation accelerates GlyT2 degradation, while decreased ubiquitination increases stability; UCHL1, as major regulator of neuronal ubiquitin homeostasis, indirectly modulates GlyT2 turnover. Mutagenesis of lysine cluster, ubiquitination assays, endocytosis assays, turnover/half-life measurements, PKC activation, siRNA UCHL1 knockdown PloS one High 23484054
2013 Calnexin (CNX) assists GlyT2 biogenesis in the ER: CNX binds transiently to an under-glycosylated GlyT2 precursor via both glycan-dependent and polypeptide-based (lectin-independent) interactions; CNX knockdown reduces GlyT2 accumulation and transport activity while CNX overexpression enhances them; CNX can stabilize non-glycosylated GlyT2 conformational states and rescue transport but not surface expression of fully non-glycosylated mutants; GlyT2 is degraded in lysosomes. siRNA-mediated CNX knockdown, CNX overexpression, pharmacological inhibition of glycosylation, transport assays, surface expression assays, lysosomal inhibitor treatments PloS one High 23650557
2013 GlyT2 activation by glycine uptake can stimulate GABA release from cerebellar nerve terminals via two mechanisms: (1) reversal of GAT1 GABA transporter driven by Na+ influx through GlyT2 and subsequent mitochondrial Na+/Ca2+ exchange; (2) Ca2+-dependent anion channels permeable to GABA; ~20% of evoked GABA release is Ca2+-dependent via NCX reversal. [3H]GABA release assays from purified cerebellar nerve endings, pharmacological dissection with selective GlyT2, GAT1 and NCX inhibitors, Ca2+ manipulation Journal of neuroscience research Medium 24273061
2014 GlyT2 forms a functional complex with plasma membrane Ca2+-ATPase (PMCA) isoforms 2 and 3 and Na+/Ca2+ exchanger 1 (NCX1) in lipid raft subdomains of the presynaptic terminal; endogenous PMCA and NCX activities are required for GlyT2 activity; this complex is proposed to help control local Na+ increases generated by GlyT2 cotransport. Co-immunoprecipitation, lipid raft fractionation, pharmacological inhibition of PMCA and NCX, GlyT2 transport assays The Journal of biological chemistry Medium 25315779
2014 Glycogen synthase kinase 3β (GSK3β) differentially modulates GlyT1 and GlyT2: GSK3β co-expression inhibits GlyT1 and stimulates GlyT2 activity/surface expression in COS-7 cells and Xenopus oocytes; GSK3β increases 32Pi incorporation into GlyT1 and decreases it in GlyT2; pharmacological inhibition of endogenous GSK3β in brainstem/spinal cord neurons produces opposite modulation of the two transporters. Co-expression in COS-7 cells and Xenopus oocytes, transport assays, phosphorylation (32Pi) assays, GSK3β inhibitors (LiCl, TDZD-8) in neuronal cultures Neuropharmacology Medium 25301276
2015 BDNF acting on truncated TrkB-T1 receptors promotes internalization of GlyT2 (and GlyT1) in astrocytes via a Rho-GTPase-dependent, dynamin/clathrin-dependent endocytosis mechanism; BDNF decreases GlyT2 Vmax without changing Km, consistent with transporter internalization rather than kinetic inhibition. Primary astrocyte cultures, [3H]glycine uptake assays, dynasore treatment, siRNA TrkB-T knockdown, Rho-GTPase inhibitor (toxin B), immunofluorescence of GlyT endosomes Glia Medium 26200505
2017 P2X purinergic receptor stimulation upregulates GlyT2 surface expression and transport activity by reducing transporter ubiquitination; this effect requires co-stimulation of homomeric P2X3 and P2X2 receptors, or heteromeric P2X2/3 receptors; P2X3 receptor activation modulates glycinergic neurotransmission through GlyT2 regulation. Pharmacological P2X receptor agonists/antagonists, siRNA knockdown of P2X subtypes, dorsal root ganglion enrichment, transport assays, surface expression, ubiquitination assays, glycinergic current recording Neuropharmacology Medium 28734869
2019 E3 ubiquitin ligases LNX1 and LNX2 ubiquitinate a C-terminal lysine cluster (K751, K773, K787, K791) in GlyT2 via their RING-finger domains, regulating GlyT2 expression and transport activity; genetic deletion of endogenous LNX2 in spinal cord neurons increases GlyT2 expression; LNX2 is required for PKC-mediated control of GlyT2 transport. Unbiased E3 ligase screening, co-immunoprecipitation, ubiquitination assays, RING-domain mutagenesis, LNX2 knockout neurons, PKC activation experiments Scientific reports High 31628376
2021 Molecular dynamics simulations of GlyT2 in a complex neuronal membrane reveal a distinct lipid fingerprint: GlyT2 shows specific enrichment and depletion of lipid species (including cholesterol contacts) compared to other SLC6 transporters (dDAT, hDAT, hSERT), with the lipid environment influencing local membrane biophysical properties near the transporter. Molecular dynamics simulation in complex neuronal membrane, lipid fingerprint analysis BBA advances Low 37082011

Source papers

Stage 0 corpus · 130 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
1991 Expression cloning of a cocaine- and antidepressant-sensitive human noradrenaline transporter. Nature 819 2008212
2009 Candidate gene studies of ADHD: a meta-analytic review. Human genetics 713 19506906
2011 Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in bioinformatics 656 21873635
2000 Orthostatic intolerance and tachycardia associated with norepinephrine-transporter deficiency. The New England journal of medicine 416 10684912
2005 Diversification of transcriptional modulation: large-scale identification and characterization of putative alternative promoters of human genes. Genome research 409 16344560
1999 Net1, a Sir2-associated nucleolar protein required for rDNA silencing and nucleolar integrity. Cell 335 10219245
2012 Adolescent impulsivity phenotypes characterized by distinct brain networks. Nature neuroscience 313 22544311
2008 Genome-wide association scan of quantitative traits for attention deficit hyperactivity disorder identifies novel associations and confirms candidate gene associations. American journal of medical genetics. Part B, Neuropsychiatric genetics : the official publication of the International Society of Psychiatric Genetics 310 18821565
1994 Stable expression of biogenic amine transporters reveals differences in inhibitor sensitivity, kinetics, and ion dependence. The Journal of biological chemistry 279 8125921
2001 Functional interaction between monoamine plasma membrane transporters and the synaptic PDZ domain-containing protein PICK1. Neuron 260 11343649
2001 The role of neuronal and extraneuronal plasma membrane transporters in the inactivation of peripheral catecholamines. Pharmacology & therapeutics 252 11707293
1995 Regional distribution and developmental variation of the glycine transporters GLYT1 and GLYT2 in the rat CNS. The European journal of neuroscience 243 7582108
2010 Poor replication of candidate genes for major depressive disorder using genome-wide association data. Molecular psychiatry 220 20351714
2006 Mutations in the gene encoding GlyT2 (SLC6A5) define a presynaptic component of human startle disease. Nature genetics 179 16751771
2009 Genetic predictors of response to antidepressants in the GENDEP project. The pharmacogenomics journal 171 19365399
2009 Gene-centric association signals for lipids and apolipoproteins identified via the HumanCVD BeadChip. American journal of human genetics 164 19913121
2005 Support for association between ADHD and two candidate genes: NET1 and DRD1. American journal of medical genetics. Part B, Neuropsychiatric genetics : the official publication of the International Society of Psychiatric Genetics 159 15717291
2004 Association of distinct alpha(2) adrenoceptor and serotonin transporter polymorphisms with constipation and somatic symptoms in functional gastrointestinal disorders. Gut 154 15138209
1998 Acute regulation of norepinephrine transport: II. PKC-modulated surface expression of human norepinephrine transporter proteins. The Journal of pharmacology and experimental therapeutics 141 9808705
2009 Sequence variations of ABCB1, SLC6A2, SLC6A3, SLC6A4, CREB1, CRHR1 and NTRK2: association with major depression and antidepressant response in Mexican-Americans. Molecular psychiatry 136 19844206
2009 Genetical genomic determinants of alcohol consumption in rats and humans. BMC biology 135 19874574
2004 Prediction of antidepressant response to milnacipran by norepinephrine transporter gene polymorphisms. The American journal of psychiatry 132 15337646
1995 Localization of glycine neurotransmitter transporter (GLYT2) reveals correlation with the distribution of glycine receptor. Journal of neurochemistry 131 7861131
2003 A regulated interaction of syntaxin 1A with the antidepressant-sensitive norepinephrine transporter establishes catecholamine clearance capacity. The Journal of neuroscience : the official journal of the Society for Neuroscience 129 12629174
1997 The glycine transporter GLYT2 is a reliable marker for glycine-immunoreactive neurons. Brain research. Molecular brain research 129 9387864
2007 Toward a confocal subcellular atlas of the human proteome. Molecular & cellular proteomics : MCP 114 18029348
2006 A polymorphism in the norepinephrine transporter gene alters promoter activity and is associated with attention-deficit hyperactivity disorder. Proceedings of the National Academy of Sciences of the United States of America 111 17146058
2001 Net1 stimulates RNA polymerase I transcription and regulates nucleolar structure independently of controlling mitotic exit. Molecular cell 111 11511359
2011 Structure-based discovery of prescription drugs that interact with the norepinephrine transporter, NET. Proceedings of the National Academy of Sciences of the United States of America 104 21885739
2006 Genetic polymorphisms in monoamine neurotransmitter systems show only weak association with acute post-surgical pain in humans. Molecular pain 102 16848906
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2011 Differential regulation of the two RhoA-specific GEF isoforms Net1/Net1A by TGF-β and miR-24: role in epithelial-to-mesenchymal transition. Oncogene 95 21986943
2008 Genome-wide association study of response to methylphenidate in 187 children with attention-deficit/hyperactivity disorder. American journal of medical genetics. Part B, Neuropsychiatric genetics : the official publication of the International Society of Psychiatric Genetics 93 18821564
2001 The activity of guanine exchange factor NET1 is essential for transforming growth factor-beta-mediated stress fiber formation. The Journal of biological chemistry 93 11278519
2001 Calcium- and syntaxin 1-mediated trafficking of the neuronal glycine transporter GLYT2. The Journal of biological chemistry 93 11278707
2003 A mutation in the human norepinephrine transporter gene (SLC6A2) associated with orthostatic intolerance disrupts surface expression of mutant and wild-type transporters. The Journal of neuroscience : the official journal of the Society for Neuroscience 90 12805287
2000 The role of N-glycosylation in transport to the plasma membrane and sorting of the neuronal glycine transporter GLYT2. The Journal of biological chemistry 88 11036075
2008 NET1-mediated RhoA activation facilitates lysophosphatidic acid-induced cell migration and invasion in gastric cancer. British journal of cancer 87 18827818
2008 The glycine transporter GlyT2 controls the dynamics of synaptic vesicle refilling in inhibitory spinal cord neurons. The Journal of neuroscience : the official journal of the Society for Neuroscience 86 18815261
2004 Norepinephrine transporter (NET), serotonin transporter (SERT), vesicular monoamine transporter (VMAT2) and organic cation transporters (OCT1, 2 and EMT) in human placenta from pre-eclamptic and normotensive pregnancies. Placenta 86 15135235
2011 The small GTPase RhoA localizes to the nucleus and is activated by Net1 and DNA damage signals. PloS one 85 21390328
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2012 Interaction of antidepressants with the serotonin and norepinephrine transporters: mutational studies of the S1 substrate binding pocket. The Journal of biological chemistry 79 23086945
1996 Isolation of a novel oncogene, NET1, from neuroepithelioma cells by expression cDNA cloning. Oncogene 79 8649828
2007 The transporters GlyT2 and VIAAT cooperate to determine the vesicular glycinergic phenotype. The Journal of neuroscience : the official journal of the Society for Neuroscience 73 17554001
2000 Characterization of the interactions between the glycine transporters GLYT1 and GLYT2 and the SNARE protein syntaxin 1A. FEBS letters 70 10722844
2012 Mutations in the GlyT2 gene (SLC6A5) are a second major cause of startle disease. The Journal of biological chemistry 68 22700964
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2006 Net1 and Myeov: computationally identified mediators of gastric cancer. British journal of cancer 66 16552434
1998 Molecular cloning and functional expression of the human glycine transporter GlyT2 and chromosomal localisation of the gene in the human genome. FEBS letters 66 9845349
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2001 Characterization of the Net1 cell cycle-dependent regulator of the Cdc14 phosphatase from budding yeast. The Journal of biological chemistry 64 11274204
2006 Mutations within the human GLYT2 (SLC6A5) gene associated with hyperekplexia. Biochemical and biophysical research communications 63 16884688
2002 Cdc5 influences phosphorylation of Net1 and disassembly of the RENT complex. BMC molecular biology 63 11960554
2011 The nuclear guanine nucleotide exchange factors Ect2 and Net1 regulate RhoB-mediated cell death after DNA damage. PloS one 62 21373644
2011 Direct effect of morphine on breast cancer cell function in vitro: role of the NET1 gene. British journal of anaesthesia 60 21857017
2004 Overall distribution of GLYT2 mRNA-containing versus GAD67 mRNA-containing neurons and colocalization of both mRNAs in midbrain, pons, and cerebellum in rats. Neuroscience research 60 15140559
2002 Mass spectrometry-based methods for phosphorylation site mapping of hyperphosphorylated proteins applied to Net1, a regulator of exit from mitosis in yeast. Molecular & cellular proteomics : MCP 60 12096118
1999 Developmental expression of the glycine transporter GLYT2 in the auditory system of rats suggests involvement in synapse maturation. The Journal of comparative neurology 60 10440707
2008 A bacterial cytotoxin identifies the RhoA exchange factor Net1 as a key effector in the response to DNA damage. PloS one 58 18509476
2002 Budding yeast Cdc5 phosphorylates Net1 and assists Cdc14 release from the nucleolus. Biochemical and biophysical research communications 57 12056824
2014 Propofol and bupivacaine in breast cancer cell function in vitro - role of the NET1 gene. Anticancer research 56 24596379
2013 Regulation of focal adhesion kinase activation, breast cancer cell motility, and amoeboid invasion by the RhoA guanine nucleotide exchange factor Net1. Molecular and cellular biology 52 23689132
1998 Characterization of the 5' region of the rat brain glycine transporter GLYT2 gene: identification of a novel isoform. Neuroscience letters 52 9509996
2007 Functional expression of release-regulating glycine transporters GLYT1 on GABAergic neurons and GLYT2 on astrocytes in mouse spinal cord. Neurochemistry international 48 17597258
2010 Smad3 regulates Rho signaling via NET1 in the transforming growth factor-beta-induced epithelial-mesenchymal transition of human retinal pigment epithelial cells. The Journal of biological chemistry 45 20547485
2007 The nuclear RhoA exchange factor Net1 interacts with proteins of the Dlg family, affects their localization, and influences their tumor suppressor activity. Molecular and cellular biology 42 17938206
2019 LncRNA CTC-497E21.4 promotes the progression of gastric cancer via modulating miR-22/NET1 axis through RhoA signaling pathway. Gastric cancer : official journal of the International Gastric Cancer Association and the Japanese Gastric Cancer Association 41 31451992
2015 BDNF, via truncated TrkB receptor, modulates GlyT1 and GlyT2 in astrocytes. Glia 41 26200505
2010 GlyT2+ neurons in the lateral cerebellar nucleus. Cerebellum (London, England) 39 19826891
2011 Startle disease in Irish wolfhounds associated with a microdeletion in the glycine transporter GlyT2 gene. Neurobiology of disease 38 21420493
2012 A novel dominant hyperekplexia mutation Y705C alters trafficking and biochemical properties of the presynaptic glycine transporter GlyT2. The Journal of biological chemistry 37 22753417
2009 Coexpression of alpha6beta4 integrin and guanine nucleotide exchange factor Net1 identifies node-positive breast cancer patients at high risk for distant metastasis. Cancer epidemiology, biomarkers & prevention : a publication of the American Association for Cancer Research, cosponsored by the American Society of Preventive Oncology 37 19124484
2021 Simultaneous binding of Guidance Cues NET1 and RGM blocks extracellular NEO1 signaling. Cell 36 33740419
2011 Endocytosis of the neuronal glycine transporter GLYT2: role of membrane rafts and protein kinase C-dependent ubiquitination. Traffic (Copenhagen, Denmark) 36 21910806
2022 Circ_0000745 promotes acute lymphoblastic leukemia progression through mediating miR-494-3p/NET1 axis. Hematology (Amsterdam, Netherlands) 35 34957935
2007 Mapping of glutathione and its precursor amino acids reveals a role for GLYT2 in glycine uptake in the lens core. Investigative ophthalmology & visual science 35 17962467
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2013 Na+/K+-ATPase is a new interacting partner for the neuronal glycine transporter GlyT2 that downregulates its expression in vitro and in vivo. The Journal of neuroscience : the official journal of the Society for Neuroscience 33 23986260
2021 Baicalin suppresses lung cancer growth phenotypes via miR-340-5p/NET1 axis. Bioengineered 32 33955315
2016 The guanine nucleotide exchange factor Net1 facilitates the specification of dorsal cell fates in zebrafish embryos by promoting maternal β-catenin activation. Cell research 32 27910850
2013 GlyT1 and GlyT2 in brain astrocytes: expression, distribution and function. Brain structure & function 31 23529192
2010 Blockade of glycine transporter (GlyT) 2, but not GlyT1, ameliorates dynamic and static mechanical allodynia in mice with herpetic or postherpetic pain. Journal of pharmacological sciences 31 20173309
2008 Expression and clinical significance of NET-1 and PCNA in hepatocellular carcinoma. Medical oncology (Northwood, London, England) 31 18214716
2008 Trafficking properties and activity regulation of the neuronal glycine transporter GLYT2 by protein kinase C. The Biochemical journal 31 18341477
2008 The neuronal glycine transporter GLYT2 associates with membrane rafts: functional modulation by lipid environment. Journal of neurochemistry 30 18266927
2014 miR-22 regulates expression of oncogenic neuro-epithelial transforming gene 1, NET1. The FEBS journal 29 25041463
2009 Subcellular localization of the neuronal glycine transporter GLYT2 in brainstem. Traffic (Copenhagen, Denmark) 29 19374720
2007 The C-terminal PDZ-ligand motif of the neuronal glycine transporter GlyT2 is required for efficient synaptic localization. Molecular and cellular neurosciences 29 17851090
2012 Glycine transporter type 2 (GlyT2) inhibitor ameliorates bladder overactivity and nociceptive behavior in rats. European urology 26 22341128
2009 Interaction of the RhoA exchange factor Net1 with discs large homolog 1 protects it from proteasome-mediated degradation and potentiates Net1 activity. The Journal of biological chemistry 26 19586902
2018 miR‑22‑3p enhances multi‑chemoresistance by targeting NET1 in bladder cancer cells. Oncology reports 25 29620229
2014 Presynaptic control of glycine transporter 2 (GlyT2) by physical and functional association with plasma membrane Ca2+-ATPase (PMCA) and Na+-Ca2+ exchanger (NCX). The Journal of biological chemistry 25 25315779
2023 AAV-Net1 facilitates the trans-differentiation of supporting cells into hair cells in the murine cochlea. Cellular and molecular life sciences : CMLS 24 36917323
2013 Constitutive endocytosis and turnover of the neuronal glycine transporter GlyT2 is dependent on ubiquitination of a C-terminal lysine cluster. PloS one 24 23484054
2024 SP1-induced circ_0017552 modulates colon cancer cell proliferation and apoptosis via up-regulation of NET1. Cancer genetics 23 38810361
2000 Differential effects of ethanol on glycine uptake mediated by the recombinant GLYT1 and GLYT2 glycine transporters. British journal of pharmacology 23 10683205
2018 Contributions of the RhoA guanine nucleotide exchange factor Net1 to polyoma middle T antigen-mediated mammary gland tumorigenesis and metastasis. Breast cancer research : BCR 22 29769144
2013 Inhibition of hepatocellular carcinoma growth and angiogenesis by dual silencing of NET-1 and VEGF. Journal of molecular histology 22 23636606
2011 A functional and transcriptomic analysis of NET1 bioactivity in gastric cancer. BMC cancer 22 21284875
2008 Association study of polymorphisms in the neutral amino acid transporter genes SLC1A4, SLC1A5 and the glycine transporter genes SLC6A5, SLC6A9 with schizophrenia. BMC psychiatry 22 18638388
2000 Transmembrane domain III plays an important role in ion binding and permeation in the glycine transporter GLYT2. The Journal of biological chemistry 22 10788509
2019 NET1 Enhances Proliferation and Chemoresistance in Acute Lymphoblastic Leukemia Cells. Oncology research 21 31046876
2013 Distribution of glycinergic neurons in the brain of glycine transporter-2 transgenic Tg(glyt2:Gfp) adult zebrafish: relationship to brain-spinal descending systems. The Journal of comparative neurology 20 22736487
2004 Calpain-mediated proteolytic cleavage of the neuronal glycine transporter, GlyT2. Journal of neurochemistry 20 14675166
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2011 Meiotic nuclear divisions in budding yeast require PP2A(Cdc55)-mediated antagonism of Net1 phosphorylation by Cdk. The Journal of cell biology 19 21690311
2022 Synergistic Control of Transmitter Turnover at Glycinergic Synapses by GlyT1, GlyT2, and ASC-1. International journal of molecular sciences 18 35269698
2014 The RhoGEF Net1 is required for normal mammary gland development. Molecular endocrinology (Baltimore, Md.) 17 25321414
2013 Rho GTPase-independent regulation of mitotic progression by the RhoGEF Net1. Molecular biology of the cell 16 23864709
2012 An aspartate residue in the external vestibule of GLYT2 (glycine transporter 2) controls cation access and transport coupling. The Biochemical journal 16 22132725
2012 A MusD retrotransposon insertion in the mouse Slc6a5 gene causes alterations in neuromuscular junction maturation and behavioral phenotypes. PloS one 16 22272310
2012 Neuroepithelial transforming gene 1 (Net1) binds to caspase activation and recruitment domain (CARD)- and membrane-associated guanylate kinase-like domain-containing (CARMA) proteins and regulates nuclear factor κB activation. The Journal of biological chemistry 16 22343628
2019 E3 ubiquitin ligases LNX1 and LNX2 are major regulators of the presynaptic glycine transporter GlyT2. Scientific reports 15 31628376
2017 P2X receptors up-regulate the cell-surface expression of the neuronal glycine transporter GlyT2. Neuropharmacology 15 28734869
2021 NET1 promotes HCC growth and metastasis in vitro and in vivo via activating the Akt signaling pathway. Aging 14 33839702
2020 Hog1 activation delays mitotic exit via phosphorylation of Net1. Proceedings of the National Academy of Sciences of the United States of America 14 32265285
2013 Calnexin-assisted biogenesis of the neuronal glycine transporter 2 (GlyT2). PloS one 14 23650557
2019 Evaluation of therapeutic effect of targeting nanobubbles conjugated with NET-1 siRNA by shear wave elastography: an in vivo study of hepatocellular carcinoma bearing mice model. Drug delivery 13 31544556
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2014 Glycine transporters GlyT1 and GlyT2 are differentially modulated by glycogen synthase kinase 3β. Neuropharmacology 13 25301276
2013 A new function for glycine GlyT2 transporters: Stimulation of γ-aminobutyric acid release from cerebellar nerve terminals through GAT1 transporter reversal and Ca(2+)-dependent anion channels. Journal of neuroscience research 13 24273061
2012 The effect of NET-1 on the proliferation, migration and endocytosis of the SMMC-7721 HCC cell line. Oncology reports 13 22378020
2021 Investigating the lipid fingerprint of SLC6 neurotransmitter transporters: a comparison of dDAT, hDAT, hSERT, and GlyT2. BBA advances 12 37082011
2019 The C-terminal region of Net1 is an activator of RNA polymerase I transcription with conserved features from yeast to human. PLoS genetics 12 30802237
2019 Cdc14 activation requires coordinated Cdk1-dependent phosphorylation of Net1 and PP2A-Cdc55 at anaphase onset. Cellular and molecular life sciences : CMLS 12 30927017
2011 RhoC and guanine nucleotide exchange factor Net1 in androgen-unresponsive mouse mammary carcinoma SC-4 cells and human prostate cancer after short-term endocrine therapy. The Prostate 12 22095553
2007 The nucleolar Net1/Cfi1-related protein Dnt1 antagonizes the septation initiation network in fission yeast. Molecular biology of the cell 12 17538026