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Relevance to Autism

Transgenic mice overexpressing grik4 in the forebrain displayed social impairment, enhanced anxiety, and depressive states, accompanied by altered synaptic transmission, in Aller et al., 2015. Furthermore, a novel de novo probably damaging missense variant in GRIK4 was identified in an ASD proband from the Autism Sequencing Consortium (De Rubeis et al., 2014).

Molecular Function

This gene encodes a protein that belongs to the glutamate-gated ionic channel family. Glutamate functions as the major excitatory neurotransmitter in the central nervous system through activation of ligand-gated ion channels and G protein-coupled membrane receptors. The protein encoded by this gene forms functional heteromeric kainate-preferring ionic channels with the subunits encoded by related gene family members.

External Links

        

References

Type
Title
Type of Disorder
Associated Disorders
Author, Year
Primary
Increased Dosage of High-Affinity Kainate Receptor Gene grik4 Alters Synaptic Transmission and Reproduces Autism Spectrum Disorders Features.
Support
Genomic diagnosis for children with intellectual disability and/or developmental delay.
ASD, ID, epilepsy/seizures
Support
Synaptic, transcriptional and chromatin genes disrupted in autism.
ASD
Support
Inherited and De Novo Genetic Risk for Autism Impacts Shared Networks.
ASD

Rare

Variant ID
Variant Type
Allele Change
Residue Change
Inheritance Pattern
Inheritance Association
Family Type
Author, Year
 GEN775R001 
 missense_variant 
 c.398G>T 
 p.Arg133Ile 
 De novo 
  
  
 GEN775R002 
 missense_variant;inframe_deletion 
 c.[1856G>A];[2713_2727delCTGGCGCAGGAGGCCinsGCT] 
 p.[Arg619His];[Leu905_Glu908del] 
  
  
  
 GEN775R003 
 frameshift_variant 
 delC 
  
 Familial 
 Paternal 
 Multiplex 

Common

No Common Variants Available
Chromosome
CNV Locus
CNV Type
# of studies
Animal Model
11
Duplication
 1
 
11
Duplication
 1
 
11
Duplication
 1
 
11
Deletion-Duplication
 15
 
11
Duplication
 1
 
11
Deletion
 5
 

Model Summary

Overexpression of Grik4 in the forebrain of mice causes social impairment and increase in anxiety as well as changes in synaptic transmission in the hippocampal trisynaptic circuit.

References

Type
Title
Author, Year
Additional
Genetic ablation of the GluK4 kainate receptor subunit causes anxiolytic and antidepressant-like behavior in mice.
Additional
The GluK4 kainate receptor subunit regulates memory, mood, and excitotoxic neurodegeneration.
Additional
Increased Dosage of High-Affinity Kainate Receptor Gene grik4 Alters Synaptic Transmission and Reproduces Autism Spectrum Disorders Features.
Additional
Increased Grik4 Gene Dosage Causes Imbalanced Circuit Output and Human Disease-Related Behaviors.

M_Grik4_3_KO_HM

Model Type: Genetic
Model Genotype: Homozygous
Mutation: Exon 14 of Grik4 gene was targeted with a loxP site targeted upstream of exon14, and a floxed neo cassette was inserted downstream.The founder lines containing the targeted gene were crossed with transgenic mice expressing cre recombinase under the promoter for protamine 1, which leads to recombination and excision of exon 14 from all tissues being activated in the male germ line.
Allele Type:
Strain of Origin:
Genetic Background: C57BL/6*129SVE
ES Cell Line:
Mutant ES Cell Line:
Model Source:

M_Grik4_4_KO_HT

Model Type: Genetic
Model Genotype: Heterozygous
Mutation: Exon 14 of Grik4 gene was targeted with flanking loxP sites in mouse ES cells. There founder lines containing the targeted gene were crossed with transgenic mice expressing cre recombinase under the promoter for protamine 1, which leads to recombination and excision of exon 4 from all tissues being activated in the male germ line.
Allele Type:
Strain of Origin:
Genetic Background: C57BL/6*129SVE
ES Cell Line:
Mutant ES Cell Line:
Model Source:

M_Grik4_5_KO_Grik5_DM

Model Type: Genetic
Model Genotype: Homozygous
Mutation: Grik4 homozygous null mice were crossed with Grik5 homozygous null mice to give rise to the double KO mice.
Allele Type:
Strain of Origin:
Genetic Background: C57BL/6*129SVE
ES Cell Line:
Mutant ES Cell Line:
Model Source:

M_Grik4_2_KO_HM

Model Type: Genetic
Model Genotype: Homozygous
Mutation: The targeting construct had loxP sites flanking exon 16 of Grik4 gene. Mice positive for this construct were crossed with mice homozygous for a transgene encoding Cre recombinase under the EIIa promoter than causes exon 16 excision via recombination in the early embryonic stages.
Allele Type: Targeted(knockout)
Strain of Origin:
Genetic Background: C57BL/6*129SVE
ES Cell Line:
Mutant ES Cell Line:
Model Source:

M_Grik4_2_KO_HM_Kainate

Model Type: Pharmaceutical intervention
Model Genotype: Homozygous
Mutation: Kainate (0.15 nmol, in PBS) was stereotaxically injected in the hippocampi of the Grik4 KO mice.
Allele Type: Targeted(knockout)
Strain of Origin:
Genetic Background: C57BL/6*129SVE
ES Cell Line:
Mutant ES Cell Line:
Model Source:

M_Grik4_1_TG

Model Type: Genetic
Model Genotype: Transgenic
Mutation: A myc-tagged Grik4 construct (from rat) was developed under the control of a CamKII promoter. This construct was injected into the pronucleus of fertilized eggs to generate the founder lines. Two of these founder lines with similar expression of Grik4 (over expression because of an additional transgene) were used in all the analyses.
Allele Type: Transgenic
Strain of Origin:
Genetic Background: C57BL/6
ES Cell Line:
Mutant ES Cell Line:
Model Source:

M_GRIK4_6_KO_HM

Model Type: Genetic
Model Genotype: Homozygous
Mutation: A targeting construct containing two lox P sites flanking exon 14 of the Grik4 gene was generated and injected into embryonic mouse cells (Fernandes et al., 2009; PMID 19778510).
Allele Type: Knockout
Strain of Origin:
Genetic Background: C57BL/6
ES Cell Line:
Mutant ES Cell Line:
Model Source: Dr. A. Contractor (Northwestern University)

M_Grik4_3_KO_HM

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
Synaptic plasticity: hippocampal LTP1
Decreased
Description: CA3 pyramidal neurons show signficantly impaired mossy fiber LTP, in slices obtained from Grik4 null mice
 Whole-cell patch clamp: CA3 pyramidal neurons
 P14-28
Anxiety1
Decreased
Description: Grik4 null mice have significantly reduced anxiety compared to wild type controls in several tasks including elevated zero maze. The latency to restart feeding is also shorter in the novelty induced suppressed feeding paradigm and the number of marbles buried in a brightly lit novel cage is also lesser than the ones buried by wild type controls. Over all authors take the results of these tests to indicate reduction in anxiety
Exp Paradigm: Marble-burying test
 Marble-burying test
 Adult
Depression1
Decreased
Description: Grik4 null mice display a decreased tendency for depression as they actually have an increased intake of sucrose compared to wild type littermate controls in the sucrose intake test, as well as decreased imobility and increased activity in the forced swim test
Exp Paradigm: Sucrose preference test
 Sucrose preference test
 Adult
Anxiety1
Decreased
Description: Grik4 null mice have significantly reduced anxiety compared to wild type controls in several tasks including elevated zero maze. The latency to restart feeding is also shorter in the novelty induced suppressed feeding paradigm and the number of marbles buried in a brightly lit novel cage is also lesser than the ones buried by wild type controls. Over all authors take the results of these tests to indicate reduction in anxiety
Exp Paradigm:  Elevated zero maze test
 Elevated zero maze test
 Adult
Depression1
Decreased
Description: Grik4 null mice display a decreased tendency for depression as they actually have an increased intake of sucrose compared to wild type littermate controls in the sucrose intake test, as well as decreased imobility and increased activity in the forced swim test
Exp Paradigm:  Forced swim test
 Forced swim test
 Adult
Anxiety1
Decreased
Description: Grik4 null mice have significantly reduced anxiety compared to wild type controls in several tasks including elevated zero maze. The latency to restart feeding is also shorter in the novelty induced suppressed feeding paradigm and the number of marbles buried in a brightly lit novel cage is also lesser than the ones buried by wild type controls. Over all authors take the results of these tests to indicate reduction in anxiety
Exp Paradigm: Novelty-suppressed feeding paradigm
 Novelty-induced hypophagia
 Adult
Exploratory activity1
 No Change
 Open field test: conducted in dimly lit room
 Adult
Spatial working memory1
 No Change
 Y-maze test
 Adult
General locomotor activity1
 No Change
 Open field test: conducted in dark
 Adult
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Homeostasis, Immune response, Maternal behavior, Molecular profile, Neuroanatomy / Ultrastructure / Cytoarchitecture, Repetitive behavior, Seizure, Sensory, Social behavior

M_Grik4_4_KO_HT

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
Anxiety1
 No Change
 Marble-burying test
 Adult
Exploratory activity1
 No Change
 Open field test: conducted in dimly lit room
 Adult
Spatial working memory1
 No Change
 Y-maze test
 Adult
General locomotor activity1
 No Change
 Open field test: conducted in dark
 Adult
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Homeostasis, Immune response, Maternal behavior, Molecular profile, Neuroanatomy / Ultrastructure / Cytoarchitecture, Neurophysiology, Repetitive behavior, Seizure, Sensory, Social behavior

M_Grik4_5_KO_Grik5_DM

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
Synaptic plasticity: hippocampal LTP1
Decreased
Description: CA3 pyramidal neurons show significantly impaired mossy fiber LTP, in slices obtained from Grik4/Grink5 double KO mice
 Whole-cell patch clamp: CA3 pyramidal neurons
 P14-28
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Emotion, Homeostasis, Immune response, Learning & memory, Maternal behavior, Molecular profile, Motor phenotype, Neuroanatomy / Ultrastructure / Cytoarchitecture, Repetitive behavior, Seizure, Sensory, Social behavior

M_Grik4_2_KO_HM

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
Hyperactivity1
Increased
Description: Grik4 KO mice have increased distance traveled in the open field test, indicating increased hyperactivity in a novel environment
 Open field test: conducted in dark
 2-3 months
Startle response: acoustic stimulus1
Decreased
Description: Grik4 KO mice have impaired startle response at 110 dB of sound ( authors note no difference with WT at 80, 90 and 100 dB)
 Acoustic startle reflex test
 2-3 months
Sensorimotor gating1
Decreased
Description: Grik4 KO mice have significantly reduced prepulse inhibition compared to WT controls
 Prepulse inhibition
 2-3 months
Spatial reference memory1
Decreased
Description: Grik4 KO mice have reduced spatial reference memory in the probe trial of the MWM as they do not preferentially spend more time in the target quadrant compared to the other three quadrants, unlike WT controls
 Morris water maze test: probe trial
 2-3 months
Spatial learning1
Decreased
Description: Grik4 KO mice have reduced spatial learning in the hidden platform phase of MWM determined by increased path length relative to WT controls in the task
 Morris water maze test: hidden platform test
 2-3 months
Swimming ability1
 No Change
 Morris water maze test
 2-3 months
Hearing1
 No Change
 Auditory brainstem response test
 2-3 months
Vision1
 No Change
 Morris water maze test
 2-3 months
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Emotion, Homeostasis, Immune response, Maternal behavior, Molecular profile, Neuroanatomy / Ultrastructure / Cytoarchitecture, Neurophysiology, Repetitive behavior, Seizure, Social behavior

M_Grik4_2_KO_HM_Kainate

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
Apoptosis: brain cells1
Decreased
Description: Kainate injected Grik4 KO mice had reduced cell death in the CA3 region (indicating a protective role by Grik4 KO, to excitotoxicity), compared to WT mice injected with Kainate
 Stereology
 2-3 months
Apoptosis: brain cells1
 No Change
 Stereology
 2-3 months
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Emotion, Homeostasis, Immune response, Learning & memory, Maternal behavior, Molecular profile, Motor phenotype, Neuroanatomy / Ultrastructure / Cytoarchitecture, Repetitive behavior, Seizure, Sensory, Social behavior

M_Grik4_1_TG

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
General locomotor activity: Ambulatory activity2
Decreased
Description: Mutants showed a decrease in total distance traveled compared to controls.
 Open field test
 8-12 weeks
General locomotor activity1
Decreased
Description: Grik4-overexp mice have reduced locomotor activity, measured as distance traveled in elevated plus maze, the open field and the light-dark explorationt est
Exp Paradigm: Light-dark exploration test
 Light-dark exploration test: Light-dark exploration test
 2-3 months
General locomotor activity1
Decreased
Description: Grik4-overexp mice have reduced locomotor activity, measured as distance traveled in elevated plus maze, the open field and the light-dark explorationt est
Exp Paradigm:  Open field test
 Open field test
 2-3 months
General locomotor activity1
Decreased
Description: Grik4-overexp mice have reduced locomotor activity, measured as distance traveled in elevated plus maze, the open field and the light-dark explorationt est
Exp Paradigm:  Elevated plus maze test
 Elevated plus maze test
 2-3 months
Miniature post synaptic current amplitude: AMPAR/NMDAR ratio2
Increased
Description: Mutants showed an increase in the ratio of AMPAR/NMDAR-mediated EPSCs compared to controls.
 Whole-cell patch clamp
 P17-21
Presynaptic function: paired-pulse facilitation1
Decreased
Description: In contrast to mEPSCs, paired-pulse facilitation is dampened in the mossy fiber-CA3 synapses
 Paired-pulse ratio
 P18-20
Spontaneous post synaptic events: excitatory currents1
Increased
Description: Grik4-overexp mice have increased frequency of spontaneous excitatory postsynaptic currents in the CA3 pyramidal cells
 Whole-cell patch clamp
 P18-20
Miniature post synaptic current amplitude: excitatory2
Increased
Description: Mutants showed an increase in the amplitude of AMPAR-mediated EPSCs compared to controls. Mutants also showed an increase in the amplitude of KainateR-mediated EPSCs compared to controls. Moreover, mutants showed an increase in the number of neurons in which KainateR-mediated ERSCs could be resolved (13/32 in mutants compared to 10/41 in controls).
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current frequency: excitatory2
Increased
Description: Mutants showed an increase in the frequency of miniature post synaptic currents in regular spiking cells compared to controls.
Exp Paradigm: Regular spiking cells
 Whole-cell patch clamp
 P17-21
Spontaneous post synaptic event amplitude: excitatory currents1
Increased
Description: Grik4-overexp mice have increased amplitude of kainate receptor mediated spontaneous synaptic transmission in the CA3 pyramidal cells
 Whole-cell patch clamp
 P18-20
Decay kinetics of miniature post synaptic currents1
Increased
Description: There is faster decay of the postsynaptic currents in Grik4-overexp CA3 pyramidal cells
 Whole-cell patch clamp
 P18-20
Miniature post synaptic current amplitude: excitatory2
Increased
Description: Mutants showed an increase in the amplitude of miniature post synaptic currents in regular spiking cells compared to controls.
Exp Paradigm: Regular spiking cells
 Whole-cell patch clamp
 P17-21
Neurotransmitter release: quantal parameters2
Increased
Description: Mutants showed an increase in quantal content compared to controls.
 Whole-cell patch clamp
 P17-21
Spontaneous post synaptic event frequency: inhibitory currents2
Increased
Description: Mutants showed an increase in the frequency of spontaneous inhibitory post synaptic currents in late spiking cells compared to controls.
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current amplitude: excitatory2
Decreased
Description: Mutants showed a decrease in the amplitude of miniature post synaptic currents in late spiking cells compared to controls.
Exp Paradigm: Late spiking cells
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current frequency: excitatory2
Increased
Description: Mutants showed an increase in the frequency of miniature post synaptic currents compared to controls.
 Whole-cell patch clamp
 P17-21
Rectification of ion channels2
Decreased
Description: Mutants showed a decrease in rectification index compared to controls.
 Whole-cell patch clamp
 P13-21
Decay kinetics of evoked post synaptic currents2
Increased
Description: Mutants showed an increase in the decay kinetics of KainateR-mediated EPSCs compared to controls.
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current amplitude: excitatory2
Increased
Description: Mutants showed an increase in the amplitude of miniature post synaptic currents compared to controls.
 Whole-cell patch clamp
 P17-21
Neuronal activation following behavioral stimulation: c-Fos levels2
Increased
Description: Mutants showed an increas in c-Fos-positive cells and c-Fos immunoreactivity compared to controls.
Exp Paradigm: Centromedial amygdala
 Immunohistochemistry
 P17-21
Spontaneous post synaptic event frequency: excitatory currents2
Increased
Description: Mutants showed an increase in the frequency of spontaneous excitatory post synaptic currents in regular spiking cells compared to controls.
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current amplitude: excitatory1
Increased
Description: Grik4-overexp mice have increased amplitude of AMPA receptor mediated mEPSCs in the CA3 pyramidal neurons, evoked by activation of mossy fibers
 Whole-cell patch clamp: CA3 pyramidal neurons
 P18-20
Miniature post synaptic current frequency: excitatory2
Decreased
Description: Mutants showed a decrease in the frequency of miniature post synaptic currents in late spiking cells compared to controls.
Exp Paradigm: Late spiking cells
 Whole-cell patch clamp
 P17-21
Presynaptic function: paired-pulse facilitation2
Decreased
Description: Mutants showed a decrease in the paired-pulse ratio of evoked responses compared to controls.
 Whole-cell patch clamp
 P17-21
Synaptic plasticity2
Decreased
Description: Mutants showed a decrease in frequency facilitation compared to controls.
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current amplitude: excitatory2
Increased
Description: Mutants showed an increase in the amplitude of AMPAR-mediated evoked post synaptic currents compared to controls.
 Whole-cell patch clamp
 P17-21
Neuronal activation following behavioral stimulation: c-Fos levels2
Increased
Description: Mutants showed an increas in c-Fos-positive cells and c-Fos immunoreactivity compared to controls.
Exp Paradigm: Basolateral amygdala
 Immunohistochemistry
 P17-21
Spontaneous post synaptic event frequency: excitatory currents2
Decreased
Description: Mutants showed a decrease in the frequency of spontaneous excitatory post synaptic currents in late spiking cells compared to controls.
 Whole-cell patch clamp
 P17-21
Local field potential1
Increased
Description: Grik4-overexp mice have increased field EPSP in the CA1 region of the hippocampal trisynaptic circuit, measured in vivo with intracranial electrodes
 In vivo local field potential (LFP) recordings: CA1 region
 2-3 months
Miniature post synaptic current frequency: excitatory1
Increased
Description: Grik4-overexp mice have increased frequency of AMPA receptor mediated mEPSCs in the CA3 pyramidal neurons, evoked by activation of mossy fibers
 Whole-cell patch clamp: CA3 pyramidal neurons
 P18-20
Social memory2
Decreased
Description: Mutants showed a decrease in preference for a novel mouse compared to controls.
 Three-chamber social approach test
 8-12 weeks
Social memory1
Decreased
Description: Grik4-overexp mice show no preference to a novel target mouse over a familiar one and spend the same amount of time with both targets, unlike wild type controls
 Three-chamber social approach test
 2-3 months
Social withdrawal1
Increased
Description: Grik4-overexp mcie show an increased tendency to spend time at the corners of the cage containing the familiar mouse ( like withdrawal or non-engaging), authors note that this is not seen in the wild type controls at all
 
 2-3 months
Depression1
Increased
Description: Grik4-overexp mice show reduced intake of sucrose indicating increased anhedonia or depression and also have a larger index of immobility in the forced swim test again indicating increase in depression
Exp Paradigm: Sucrose preference test
 Sucrose preference test
 2-3 months
Anxiety1
Increased
Description: Grik4-overexp mice show increase in anxiety as they spend lesser time in the center of the open field chamber and the open arms of the elevated plus maze compared to wild type controls and more time in the dark in the light-dark exploration tests
Exp Paradigm:  Open field test
 Open field test
 2-3 months
Depression1
Increased
Description: Grik4-overexp mice show reduced intake of sucrose indicating increased anhedonia or depression and also have a larger index of immobility in the forced swim test again indicating increase in depression
Exp Paradigm:  Forced swim test
 Forced swim test
 2-3 months
Anxiety1
Increased
Description: Grik4-overexp mice show increase in anxiety as they spend lesser time in the center of the open field chamber and the open arms of the elevated plus maze compared to wild type controls and more time in the dark in the light-dark exploration tests
Exp Paradigm:  Elevated plus maze test
 Elevated plus maze test
 2-3 months
Anxiety2
Increased
Description: Mutants showed a decrease in time spent in the open arm of elevated plus maze compared to controls.
 Elevated plus maze test
 8-12 weeks
Depression2
Increased
Description: Mutants showed an increase in immobility time in swimming test compared to controls.
 Forced swim test
 8-12 weeks
Anxiety1
Increased
Description: Grik4-overexp mice show increase in anxiety as they spend lesser time in the center of the open field chamber and the open arms of the elevated plus maze compared to wild type controls and more time in the dark in the light-dark exploration tests
Exp Paradigm: Light-dark exploration test
 Light-dark exploration test: Light-dark exploration test
 2-3 months
Targeted expression1
Increased
Description: Grik4-overexp mice show increased expression of Grik4, in the hippocampus, neocortex and striatum
Exp Paradigm: Immunohistochemistry
 Immunohistochemistry
 > 4 weeks
Targeted expression1
Increased
Description: Grik4-overexp mice show increased expression of Grik4, in the hippocampus, neocortex and striatum
Exp Paradigm:  Western blot
 Western blot
 > 4 weeks
Spatial working memory1
 No Change
 Y-maze test
 2-3 months
Motor coordination and balance1
 No Change
 Accelerating rotarod test
 2-3 months
Neuroreceptor levels: glutamate receptors: AMPA receptors1
 No Change
 Western blot
 > 4 weeks
Neuronal activation following behavioral stimulation: c-Fos levels2
 No Change
 Immunohistochemistry
 P17-21
Spontaneous post synaptic event amplitude: excitatory currents2
 No Change
 Whole-cell patch clamp
 P17-21
Spontaneous post synaptic event amplitude: excitatory currents2
 No Change
 Whole-cell patch clamp
 P17-21
Spontaneous post synaptic event amplitude: inhibitory currents2
 No Change
 Whole-cell patch clamp
 P17-21
Spontaneous post synaptic event amplitude: inhibitory currents2
 No Change
 Whole-cell patch clamp
 P17-21
Spontaneous post synaptic event frequency: inhibitory currents2
 No Change
 Whole-cell patch clamp
 P17-21
Social approach1
 No Change
 Three-chamber social approach test
 2-3 months
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Homeostasis, Immune response, Learning & memory, Maternal behavior, Molecular profile, Neuroanatomy / Ultrastructure / Cytoarchitecture, Repetitive behavior, Seizure, Sensory

M_GRIK4_6_KO_HM

Category
Entity
Quantity
Experimental Paradigm
Age at Testing
General locomotor activity: Ambulatory activity1
Increased
Description: Mutants showed an increase in total distance traveled compared to controls.
 Open field test
 8-12 weeks
Miniature post synaptic current frequency: excitatory1
Increased
Description: Mutants showed an increase in the amplitude of miniature post synaptic currents in late spiking cells compared to controls.
Exp Paradigm: Late spiking cells
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current frequency: excitatory1
Decreased
Description: Mutants showed a decrease in the amplitude of miniature post synaptic currents in regular spiking cells compared to controls.
Exp Paradigm: Regular spiking cells
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current amplitude: excitatory1
Decreased
Description: Mutants showed a decrease in the amplitude of miniature post synaptic currents compared to controls.
 Whole-cell patch clamp
 P17-21
Presynaptic function: paired-pulse facilitation1
Increased
Description: Mutants showed an increase in the paired-pulse ratio of evoked responses compared to controls.
 Whole-cell patch clamp
 P17-21
Miniature post synaptic current amplitude: excitatory1
Decreased
Description: Mutants showed a decrease in the amplitude of AMPAR-mediated evoked post synaptic currents compared to controls.
 Whole-cell patch clamp
 P17-21
Depression1
Decreased
Description: Mutants showed a decrease in immobility time in swimming test compared to controls.
 Forced swim test
 8-12 weeks
Anxiety1
 No Change
 Elevated plus maze test
 8-12 weeks
Presynaptic function: paired-pulse facilitation1
 No Change
 Whole-cell patch clamp
 P17-21
Social memory1
 No Change
 Three-chamber social approach test
 8-12 weeks
 Not Reported: Circadian sleep/wake cycle, Communications, Developmental profile, Homeostasis, Immune response, Learning & memory, Maternal behavior, Molecular profile, Neuroanatomy / Ultrastructure / Cytoarchitecture, Repetitive behavior, Seizure, Sensory

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