The Perineuronal ‘Safety’ Net? Perineuronal Net Abnormalities in Neurological Disorders
Open accessAbstract
Perineuronal nets (PNN) are extracellular matrix (ECM) assemblies that preferentially ensheath parvalbumin (PV) expressing interneurons. Converging evidence indicates that PV cells and PNN are impaired in a variety of neurological disorders. PNN development and maintenance is necessary for a number of processes within the CNS, including regulation of GABAergic cell function, protection of neurons from oxidative stress, and closure of developmental critical period plasticity windows. Understanding PNN functions may be essential for characterizing the mechanisms of altered cortical excitability observed in neurodegenerative and neurodevelopmental disorders. Indeed, PNN abnormalities have been observed in post-mortem brain tissues of patients with schizophrenia and Alzheimer's disease. There is impaired development of PNNs and enhanced activity of its key regulator matrix metalloproteinase-9 (MMP-9) in Fragile X Syndrome, a common genetic cause of autism. MMP-9, a protease that cleaves ECM, is differentially regulated in a number of these disorders. Despite this, few studies have addressed the interactions between PNN expression, MMP-9 activity and neuronal excitability. In this review, we highlight the current evidence for PNN abnormalities in CNS disorders associated with altered network function and MMP-9 levels, emphasizing the need for future work targeting PNNs in pathophysiology and therapeutic treatment of neurological disorders.
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Written from the paper’s full text.
TL;DR
This review synthesizes evidence that perineuronal nets (PNNs) around parvalbumin (PV) interneurons are abnormal in multiple neurological disorders, often alongside dysregulated MMP-9. It proposes that MMP-9-mediated PNN cleavage contributes to impaired PV cell function and cortical hyperexcitability, highlighting PNN/MMP-9 pathways as potential therapeutic targets.
Key findings
- PNNs preferentially ensheath PV interneurons and are implicated in regulating GABAergic cell function, protecting neurons from oxidative stress, and closing developmental critical period plasticity; PNN abnormalities are reported in schizophrenia, Alzheimer’s disease, epilepsy, CNS injury/stroke, hearing loss, and Fragile X syndrome (FXS).
- In FXS, Fmr1 knockout mice show delayed PNN development and increased MMP-9 activity in developing auditory cortex, mostly between P14 and P21; genetic reduction of MMP-9 normalized PV/PNN development and reversed auditory cortical neuron hyperresponsiveness. FXS is reported to occur in 1 in 4000 males and 1 in 8000 females.
- In schizophrenia, post-mortem brains show reduced WFA-labeled PNN cells and reduced CSPGs and PV expression in prefrontal cortex; elevated plasma MMP-9 has been reported, and minocycline trials are cited as showing overall improvement in negative symptoms and executive functioning.
- In Alzheimer’s disease, post-mortem hippocampus shows reduced PNN expression; PNN-ensheathed cells were more resistant to Aβ1-42 and iron-induced oxidative stress. Table 1 reports that in adult sensory cortices a majority of PV cells express PNNs, e.g., visual cortex ∼70% at 3–5 months, somatosensory layer 4 young adult ∼80%, and auditory cortex CBA young adult 50%.
Methods
- Narrative review; no new experimental data are presented. It synthesizes published human post-mortem, rodent model, in vitro, EEG/electrophysiology, and clinical trial evidence.
- Includes Table 1 compiling studies of PV cells enwrapped by PNNs across visual, somatosensory, and auditory cortices, with mouse strain, staining methods, and percentages by age and layer where available.
- Main analysis is qualitative integration around a working hypothesis linking MMP-9 activity, PNN cleavage/reorganization, PV cell dysfunction, and altered network excitability.
Limitations
- Stated in the paper: the precise role of PNNs in network function and development is not known; future studies should compare intrinsic and synaptic properties of the same cell types with and without PNNs; chABC likely has broader effects, so genetic removal of specific PNN components is needed; PNNs are heterogeneous and composition may be brain-region- and age-specific; mechanisms of PNN loss and restructuring in disease remain unclear.
- Evident from the text: as a narrative review, there is no systematic search strategy or quantitative meta-analysis; causal direction between MMP-9, PNN changes, and excitability is often inferred from correlational animal and human data; therapeutic suggestions such as minocycline or MMP-9 inhibitors are based largely on preliminary or indirect evidence.
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- Findingcitation
Studies of postmortem human tissues suggest developmental changes in lectican levels in human CNS.
“Studies of postmortem human tissues also suggest developmental changes in lectican levels in human CNS.”
Quote in the paperNot a substantial claim - Findingnone given
The expression of versican beta isoform increased 2-3 fold from embryonic day 14 until birth and then decreased in adulthood.
“In contrast, the expression of beta isoform increased 2–3 fold from embryonic day (E) 14 until birth and then decreased in adulthood.”
Quote in the paperNot a substantial claim - Findingcitation
A number of CNS diseases exhibit dysregulated expression of enzymes necessary for PNN cleavage and reorganization.
“In parallel, a number of these CNS diseases exhibit dysregulated expression of enzymes necessary for PNN cleavage and reorganization (Backstrom et al., 1996; Lorenzl et al., 2003; Rybakowski et al., 2009; Kim et al., 2009; Zhang et al., 2011; Yamamori et al., 2013; Liu et al., 2014; Rankin-Gee et al., 2015; Lepeta et al., 2017).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Findingnone given
There is enhanced activity of MMP-9 in Fragile X Syndrome.
“There is impaired development of PNNs and enhanced activity of its key regulator matrix metalloproteinase-9 (MMP-9) in Fragile X Syndrome, a common genetic cause of autism.”
Quote in the paperSubstantial claim - Findingcitation
High levels of brevican mRNA expression were detected in postmortem human cortex during early development until 8 years of age, and expression levels were lower in the adult human brain.
“For example, high levels of brevican mRNA expression were detected in postmortem human cortex during early development until 8 years of age, and the expression levels were lower in the adult human brain (Gary et al., 2000).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Gary et al., 2000 source
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- Findingcitation
Neurocan expression increased between E14 and postnatal day 3 and then decreased during early-late adolescence and into adulthood.
“Neurocan expression also increased between E14 and postnatal day (P) 3 and then decreased during early-late adolescence and into adulthood (Meyer-Puttlitz et al., 1996; Miller et al., 1996; Milev et al., 1998; Brückner et al., 2000).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Findingreasoning
MMP-9 is differentially regulated in a number of neurological disorders.
“MMP-9, a protease that cleaves ECM, is differentially regulated in a number of these disorders.”
Quote in the paperSubstantial claim - Findingnone given
PNN abnormalities have been observed in post-mortem brain tissues of patients with schizophrenia and Alzheimer's disease.
“Indeed, PNN abnormalities have been observed in post-mortem brain tissues of patients with schizophrenia and Alzheimer’s disease.”
Quote in the paperSubstantial claim - Findingcitation
A developmental upregulation of aggrecan and brevican, which peak in adulthood, was reported in rat brain.
“A developmental upregulation of aggrecan and brevican, which peak in adulthood, was reported in rat brain (Milev et al., 1998; Brückner et al., 2000; Popp et al., 2003).”
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The expression of versican alpha isoform was low during embryonic development, first and second postnatal weeks, and then increased steadily until around 3.5 months of age.
“The expression of alpha isoform was low during embryonic development, first and second postnatal weeks, and then increased steadily until around 3.5 months of age (Bignami et al., 1993b).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Bignami et al., 1993b
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- Findingnone given
There is impaired development of PNNs in Fragile X Syndrome.
“There is impaired development of PNNs and enhanced activity of its key regulator matrix metalloproteinase-9 (MMP-9) in Fragile X Syndrome, a common genetic cause of autism.”
Quote in the paperSubstantial claim - Findingcitation
PV, PNN, and MMP-9 interactions are important for driving adult plasticity.
“A recent study by Murase and colleagues demonstrated that PV, PNN, and MMP-9 interactions are important for driving adult plasticity (Murase et al., 2017).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Murase et al., 2017 source
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- Findingcitation
Astrocytes extend their processes into PNNs localized near the synapse.
“Astrocytes extend their processes into PNNs localized near the synapse (Brückner et al., 1993; Blümcke et al., 1995), suggesting a tetrapartite synaptic organization involving ECM, pre/post synaptic sites and astrocytes (Dityatev and Rusakov, 2011; Smith et al., 2015) whose synergistic functions remain unexplored.”
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Studies comparing PNNs detected with lectins or specific antibodies indicate substantial levels of molecular heterogeneity.
“Studies comparing PNNs detected with lectins or specific antibodies indicate substantial levels of molecular heterogeneity (Matthews et al., 2002; Irvine and Kwok, 2018), highlighting the need for better approaches which address PNN heterogeneity (Irvine and Kwok, 2018).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Matthews et al., 2002 source
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- cites Irvine and Kwok, 2018
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- Findingcitation
PV and PNN colocalization is correlated with critical period closure and reduced synaptic plasticity, particularly in sensory cortices.
“In fact, PV and PNN colocalization is correlated with critical period closure and reduced synaptic plasticity, particularly in sensory cortices (Nowicka et al., 2009; Beurdeley et al., 2012).”
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Auditory hypersensitivity is common in humans with FXS as measured by increased sound evoked responses, reduced habituation and expansion of brain regions activated by auditory stimuli.
“Auditory hypersensitivity is common in humans with FXS as measured by increased sound evoked responses, reduced habituation and expansion of brain regions activated by auditory stimuli (Rojas et al., 2001; Castrén et al., 2003; Van der Molen et al., 2012; Ethridge et al., 2016).”
Quote in the paperSubstantial claim0 of 3 sources checked - Findingnone given
In adult sensory cortices, a majority of PV cells express PNNs as seen with WFA staining.
“Interestingly, developing auditory cortex appears to exhibit fewer PNN and PV expressing cells than visual or somatosensory cortex, but in adult sensory cortices, a majority of PV cells express PNNs as seen with WFA staining.”
Quote in the paperNot a substantial claim - Findingcitation
Otx2 expression coincides with both PV and PNN expression in the cortex during critical period plasticity.
“Otx2 expression coincides with both PV and PNN expression in the cortex during critical period plasticity (Sugiyama et al., 2008).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Sugiyama et al., 2008 source
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- Findingcitation
Neurons in adult Fmr1 KO mouse auditory cortex show more variable response latencies.
“In vivo electrophysiological studies demonstrate that neurons in adult Fmr1 KO mouse auditory cortex (1) are hyperexcitable, exhibiting increased responses per stimulus, (2) show more variable response latencies, and (3) display broader frequency tuning curves (Rotschafer and Razak, 2013).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Rotschafer and Razak, 2013
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- Findingcitation
Neurons in adult Fmr1 KO mouse auditory cortex are hyperexcitable, exhibiting increased responses per stimulus.
“In vivo electrophysiological studies demonstrate that neurons in adult Fmr1 KO mouse auditory cortex (1) are hyperexcitable, exhibiting increased responses per stimulus, (2) show more variable response latencies, and (3) display broader frequency tuning curves (Rotschafer and Razak, 2013).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Rotschafer and Razak, 2013
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- Findingcitation
Mature Fmr1 KO mouse auditory cortex exhibits reduced habituation to repeated sound presentations.
“Recent work using EEG/ERP recordings shows that mature Fmr1 KO mouse auditory cortex exhibits reduced habituation to repeated sound presentations (Lovelace et al., 2016).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Lovelace et al., 2016 source
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- Findingnone given
Fmr1 KO mouse cortex shows increased gamma power in resting EEG.
“When resting EEG is analyzed for spectral power distribution, Fmr1 KO mouse cortex shows increased gamma power.”
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We found increased MMP-9 activity levels in the developing auditory cortex of Fmr1 KO mice.
“We found delayed PNN development and increased MMP-9 activity levels in the developing auditory cortex of Fmr1 KO mice (Wen et al., 2017), which may lead to impaired PV cell development and reduced inhibition.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Wen et al., 2017 source
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- Findingcitation
Minocycline administration in Fmr1 KO mice corrects ultrasonic vocalizations.
“Minocycline administration in Fmr1 KO mice prevents abnormal dendritic spine development in the hippocampus, improves anxiety-like behaviors (Bilousova et al., 2009; Dansie et al., 2013) and corrects ultrasonic vocalizations (Rotschafer et al., 2012).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Rotschafer et al., 2012 source
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- Findingcitation
PNNs are necessary for the uptake of Otx2 by PV cells, which promotes PV cell maturation.
“PNNs are necessary for the uptake of Otx2 by PV cells, which promotes PV cell maturation (Beurdeley et al., 2012).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Beurdeley et al., 2012 source
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- Findingcitation
Post-mortem brain tissues from schizophrenia patients exhibit reduced PV and/or GAD67 mRNA, particularly in prefrontal cortex.
“In situ hybridization and RT-PCR studies indicate that post-mortem brain tissues from schizophrenia patients exhibit reduced PV and/or GAD67 mRNA, particularly in prefrontal cortex (Akbarian et al., 1995; Volk et al., 2000; Hashimoto et al., 2003, 2008a,b; Mellios et al., 2009).”
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Minocycline administration in Fmr1 KO mice improves anxiety-like behaviors.
“Minocycline administration in Fmr1 KO mice prevents abnormal dendritic spine development in the hippocampus, improves anxiety-like behaviors (Bilousova et al., 2009; Dansie et al., 2013) and corrects ultrasonic vocalizations (Rotschafer et al., 2012).”
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The somatosensory cortex of adult FXS mice exhibit delayed PV cell maturation.
“The somatosensory cortex of adult FXS mice exhibit delayed PV cell maturation (Nomura et al., 2017).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Nomura et al., 2017 source
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- Findingcitation
In developing mice, PV interneurons receive reduced excitatory drive from neighboring pyramidal cells in layer 4 of somatosensory cortex.
“In developing mice, PV interneurons receive reduced excitatory drive from neighboring pyramidal cells in layer 4 of somatosensory cortex (Gibson et al., 2008).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Gibson et al., 2008 source
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- Findingcitation
Reduction of MMP-9 levels with minocycline in humans with FXS contributes to reversal of several FXS phenotypes, including behavioral deficits and enhanced auditory electrocortical activity in adults and/or adolescents with FXS.
“Reduction of MMP-9 levels with minocycline in humans with FXS (Dziembowska et al., 2013) contributes to reversal of several FXS phenotypes, including behavioral deficits and enhanced auditory electrocortical activity in adults and/or adolescents with FXS (Paribello et al., 2010; Leigh et al., 2013; Schneider et al., 2013).”
Quote in the paperSubstantial claim0 of 3 sources checked - Findingcitation
Minocycline administration in Fmr1 KO mice prevents abnormal dendritic spine development in the hippocampus.
“Minocycline administration in Fmr1 KO mice prevents abnormal dendritic spine development in the hippocampus, improves anxiety-like behaviors (Bilousova et al., 2009; Dansie et al., 2013) and corrects ultrasonic vocalizations (Rotschafer et al., 2012).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Findingcitation
Neurons in adult Fmr1 KO mouse auditory cortex display broader frequency tuning curves.
“In vivo electrophysiological studies demonstrate that neurons in adult Fmr1 KO mouse auditory cortex (1) are hyperexcitable, exhibiting increased responses per stimulus, (2) show more variable response latencies, and (3) display broader frequency tuning curves (Rotschafer and Razak, 2013).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Rotschafer and Razak, 2013
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- Findingcitation
We found delayed PNN development in the developing auditory cortex of Fmr1 KO mice.
“We found delayed PNN development and increased MMP-9 activity levels in the developing auditory cortex of Fmr1 KO mice (Wen et al., 2017), which may lead to impaired PV cell development and reduced inhibition.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Wen et al., 2017 source
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- Findingcitation
Analysis of post-mortem human brain tissues demonstrated reduced PV cell density and immunoreactivity at epileptic foci in the hippocampus and neocortex.
“Analysis of post-mortem human brain tissues demonstrated reduced PV cell density and immunoreactivity at epileptic foci in the hippocampus and neocortex (Sloviter et al., 1991; Ferrer et al., 1994; Andrioli et al., 2007).”
Quote in the paperSubstantial claim0 of 3 sources checked - Findingcitation
Network oscillatory activity that was increased by drug-induced changes in dopaminergic signaling in mouse prefrontal cortex was further enhanced following chABC-mediated degradation of PNNs.
“For example, network oscillatory activity that was increased by drug-induced changes in dopaminergic signaling in mouse prefrontal cortex was further enhanced following chABC-mediated degradation of PNNs (Steullet et al., 2014).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Steullet et al., 2014 source
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- Findingcitation
Brevican conditional knockdown in PV-expressing cells demonstrated that changes in the expression of specific lecticans alter synaptic input and firing frequency in PV cells.
“In addition, brevican conditional knockdown in PV-expressing cells demonstrated that changes in the expression of specific lecticans alter synaptic input and firing frequency in PV cells (Favuzzi et al., 2017).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Favuzzi et al., 2017 source
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- Findingcitation
In vitro studies in human neuroblastoma SH-SY5Y cells indicate that chondroitin sulfate application prior to exposure to oxidative-stress inducing agents, such as hydrogen peroxide, can prevent cell death and attenuate the production of reactive oxygen species.
“Specifically, in vitro studies conducted in human neuroblastoma SH-SY5Y cells indicate that chondroitin sulfate application prior to exposure to oxidative-stress inducing agents, such as hydrogen peroxide, can prevent cell death and attenuate the production of reactive oxygen species.”
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Genetic analysis of single nucleotide polymorphisms in schizophrenia patients demonstrates mutations in the gene encoding GCL, an enzyme important for production of cellular antioxidant GSH.
“Genetic analysis of single nucleotide polymorphisms in schizophrenia patients demonstrates mutations in the gene encoding GCL, an enzyme that is important for the production of the cellular antioxidant GSH (Tosic et al., 2006).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tosic et al., 2006 source
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- Findingcitation
Reduced PV and GAD67 protein levels are seen in both prefrontal and temporal cortex of post-mortem brain tissues from schizophrenia patients.
“Reduced PV and GAD67 protein levels are also seen in both prefrontal and temporal cortex of post-mortem brain tissues (Beasley and Reynolds, 1997; Curley et al., 2011; Glausier et al., 2014).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Findingcitation
A mouse model of schizophrenia exhibits reduced PNN levels in prefrontal cortex.
“A mouse model of schizophrenia also exhibits reduced PNN levels in prefrontal cortex (Cabungcal et al., 2014).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Cabungcal et al., 2014 source
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- Findingreasoning
Injection of a dopamine reuptake inhibitor to promote more oxidative stress via buildup of reactive oxygen species contributed to increased 8-oxo-dG signal and PV cell loss in GCL KO mice.
“During development, GCL KO mice had fewer PNN-expressing PV cells in anterior cingulate cortex, and the injection of a dopamine reuptake inhibitor to promote more oxidative stress via buildup of reactive oxygen species contributed to increased 8-oxo-dG signal and PV cell loss.”
Quote in the paperSubstantial claim - Findingcitation
Clinical trials using minocycline alone or as an adjunct to risperidone in patients with schizophrenia have demonstrated overall improvement in negative symptoms and executive functioning.
“Various clinical trials have been conducted using minocycline alone or as an adjunct to risperidone in patients with schizophrenia and have demonstrated overall improvement in negative symptoms and executive functioning (Miyaoka et al., 2008; Levkovitz et al., 2010; Khodaie-Ardakani et al., 2014).”
Quote in the paperSubstantial claim0 of 3 sources checked - Findingcitation
Increased gamma-band power was observed in baseline EEG recordings from humans with schizophrenia.
“Increased gamma-band power was observed in baseline EEG recordings from humans with schizophrenia (Gandal et al., 2012).”
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- Findingcitation
A functional polymorphism of the MMP-9 gene was discovered in human schizophrenia patients.
“A functional polymorphism of the MMP-9 gene was discovered in human schizophrenia patients (Rybakowski et al., 2009; Lepeta et al., 2017).”
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Elevated MMP-9 levels were shown in the plasma of schizophrenic patients.
“Additionally, Yamamori et al. (2013) showed elevated MMP-9 levels in the plasma of schizophrenic patients.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Yamamori et al. (2013) source
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- Findingcitation
Schizophrenia is associated with the loss of PNNs.
“Indeed, several studies indicate that schizophrenia is associated with the loss of PNNs.”
Quote in the paperSubstantial claim - Findingcitation
During development, GCL KO mice had fewer PNN-expressing PV cells in anterior cingulate cortex.
“During development, GCL KO mice had fewer PNN-expressing PV cells in anterior cingulate cortex, and the injection of a dopamine reuptake inhibitor to promote more oxidative stress via buildup of reactive oxygen species contributed to increased 8-oxo-dG signal and PV cell loss.”
Quote in the paperSubstantial claim - Findingnone given
In this mouse model, schizophrenia-like abnormalities are reported in the adult mouse prefrontal cortex as a result of neonatal lesion of ventral hippocampus.
“In this model, schizophrenia-like abnormalities are reported in the adult mouse prefrontal cortex as a result of neonatal lesion of ventral hippocampus.”
Quote in the paperNot a substantial claim - Findingcitation
Genomic sequencing data indicate that a number of mutations in the Scn1a gene contribute to a spectrum of different epileptic disorders affecting children, including de novo as well as missense, frame-shift, and nonsense mutations.
“Genomic sequencing data indicate that a number of mutations in the Scn1a gene contribute to a spectrum of different epileptic disorders affecting children, including de novo (Claes et al., 2001; Fujiwara et al., 2003) as well as missense, frame-shift, and nonsense mutations (Fujiwara et al., 2003).”
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PV KO mice are more likely to experience status epilepticus compared to controls following increased doses of pentylenetetrazol (PTZ), a GABAA receptor blocker.
“For example, PV KO mice are more likely to experience status epilepticus compared to controls following increased doses of pentylenetetrazol (PTZ), a GABAA receptor blocker (Schwaller et al., 2004).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Schwaller et al., 2004 source
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- Premisecitation
In the forebrain, PNNs are often found on parvalbumin-expressing GABAergic neurons.
“In the forebrain, PNNs are often found on PV expressing GABAergic neurons (Table 1).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Table 1
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- Premisereasoning
The complex of hyaluronan with ECM proteins forms the basis for PNN structure.
“This complex of hyaluronan with ECM proteins forms the basis for PNN structure.”
Quote in the paperNot a substantial claim - Interpretationcitation
PNNs are altered in various neurodevelopmental and neurodegenerative disorders associated with changes in brain activity.
“Recently, increasing evidence suggests that PNNs are altered in various neurodevelopmental and neurodegenerative disorders associated with changes in brain activity (De Luca and Papa, 2016; Sorg et al., 2016; Härtig et al., 2016).”
Quote in the paperSubstantial claim0 of 3 sources checked - Interpretationnone given
PV cells and PNNs are impaired in a variety of neurological disorders.
“Converging evidence indicates that PV cells and PNN are impaired in a variety of neurological disorders.”
Quote in the paperSubstantial claim - Premisecitation
Hyaluronan binds to other ECM molecules via N-terminal hyaluronan binding domains, including lecticans and link proteins such as cartilage link protein, Ctrl1, and brain-derived link proteins Bral1 and Bral2.
“Hyaluronan binds to other ECM molecules via N-terminal hyaluronan binding domains, including lecticans and link proteins such as cartilage link protein, Ctrl1, and brain-derived link proteins Bral1 and Bral2 (Frischknecht and Seidenbecher, 2008).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Frischknecht and Seidenbecher, 2008
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- Premisenone given
PNN development and maintenance is necessary for protection of neurons from oxidative stress.
“PNN development and maintenance is necessary for a number of processes within the CNS, including regulation of GABAergic cell function, protection of neurons from oxidative stress, and closure of developmental critical period plasticity windows.”
Quote in the paperNot a substantial claim - Interpretationcitation
Developmental regulation of ECM proteins that make up PNNs may underlie PNN formation, composition, and function.
“Developmental regulation of ECM proteins, which make up PNNs, may underlie PNN formation, composition, and function (Köppe et al., 1997).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Köppe et al., 1997 source
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- Interpretationcitation
Developmental changes in lectican expression levels suggest that PNNs themselves may also be developmentally regulated, underlying maturation of circuit function.
“These developmental changes in lectican expression levels suggest that the PNNs themselves may also be developmentally regulated, underlying maturation of circuit function (Köppe et al., 1997).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Köppe et al., 1997 source
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- Premisenone given
PNNs have been implicated in regulation of neural activity.
“Since then, PNNs have been implicated in a wide range of functions including neuroprotection, regulation of neural activity and experience-dependent plasticity.”
Quote in the paperNot a substantial claim - Premisenone given
PNN development and maintenance is necessary for regulation of GABAergic cell function.
“PNN development and maintenance is necessary for a number of processes within the CNS, including regulation of GABAergic cell function, protection of neurons from oxidative stress, and closure of developmental critical period plasticity windows.”
Quote in the paperSubstantial claim - Premisenone given
PNN development and maintenance is necessary for closure of developmental critical period plasticity windows.
“PNN development and maintenance is necessary for a number of processes within the CNS, including regulation of GABAergic cell function, protection of neurons from oxidative stress, and closure of developmental critical period plasticity windows.”
Quote in the paperSubstantial claim - Interpretationcitation
These auditory processing deficits may drive social, cognitive and language delays and deficits in FXS.
“These auditory processing deficits may drive social, cognitive and language delays and deficits in FXS (Rotschafer and Razak, 2014; Sinclair et al., 2017a,b).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Rotschafer and Razak, 2014
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- cites Sinclair et al., 2017a,b
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- Premisecitation
Aggrecan has been previously identified as an MMP-9 target.
“As aggrecan has been previously identified as an MMP-9 target (Mercuri et al., 2000; Madsen et al., 2010; Rankin-Gee et al., 2015), aggrecan-rich PNNs may be unstable in the presence of MMP-9, leading to abnormal development and neural excitability (Murase et al., 2017; Wen et al., 2017).”
Quote in the paperSubstantial claim0 of 3 sources checked - Interpretationcitation
PNNs may regulate PV cell functions.
“PNNs may regulate PV cell functions (Hensch, 2005; Cabungcal et al., 2013; Bernard and Prochiantz, 2016), as PNN reorganization is shown to result in abnormal CNS development and altered neural excitability (Pizzorusso et al., 2002; Dityatev et al., 2007; Liu et al., 2013; Balmer, 2016; Lensjø et al., 2017).”
Quote in the paperSubstantial claim0 of 3 sources checked - Interpretationreasoning
Abnormal interactions between PNNs, MMP-9 and PV activity may lead to neurological disorders.
“Conversely, abnormal interactions may lead to neurological disorders.”
Quote in the paperSubstantial claim - Premisecitation
Cortical PV cells contribute to synchronous oscillatory activity, particularly in the gamma range.
“Cortical PV cells contribute to synchronous oscillatory activity (Cardin et al., 2009; Sohal et al., 2009; Uhlhaas et al., 2009), particularly in the gamma range (∼30–100 Hz).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Interpretationreasoning
High levels of MMP-9 contribute to proteolytic cleavage of ECM, creating an extracellular environment that is permissive for synaptic plasticity.
“High levels of MMP-9 contribute to proteolytic cleavage of ECM, creating an extracellular environment that is permissive for synaptic plasticity.”
Quote in the paperSubstantial claim - Premisecitation
Other cell types, including putative excitatory neurons, also contain PNNs but are less numerous compared to GABAergic-PNN expressing cells.
“Other cell types, including putative excitatory neurons also contain PNNs but they are less numerous compared to GABAergic-PNN expressing cells (Wegner et al., 2003).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Wegner et al., 2003 source
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- Interpretationcitation
These findings suggest a link between PNNs, MMP-9 activity and PV activity that shapes normal brain development.
“These findings suggest a link between PNNs, MMP-9 activity and PV activity that shapes normal brain development.”
Quote in the paperSubstantial claim - Interpretationreasoning
Network excitability in FXS may also result from impaired synaptic scaling at PV cells due to PNN dysfunction.
“Network excitability in FXS may also result from impaired synaptic scaling at PV cells due to PNN dysfunction.”
Quote in the paperSubstantial claim - Interpretationreasoning
MMP-9 cleavage of PNNs may drive changes in PV expression and/or PV cell functions seen in FXS.
“In addition to regulating cortical development, MMP-9 cleavage of PNNs may drive changes in PV expression and/or PV cell functions seen in FXS.”
Quote in the paperSubstantial claim - Premisecitation
There is evidence for a role of PNNs in mediating synaptic scaling via Narp.
“There is evidence for a role of PNNs in mediating synaptic scaling via neuronal activity-regulated pentraxin, also known as Narp.”
Quote in the paperSubstantial claim - Premisecitation
Narp can interact with the N-terminal domain of AMPA receptors localized extracellularly and facilitate AMPA receptor clustering at both the pre- and post-synaptic cell in response to stimulation.
“Narp can interact with the N-terminal domain of AMPA receptors localized extracellularly and facilitate AMPA receptor clustering at both the pre- and post-synaptic cell in response to stimulation (O’Brien et al., 1999).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites O’Brien et al., 1999 source
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- Interpretationcitation
These data suggest that PNNs may drive PV cell maturation by directing transcription or translation of specific genes necessary for PV cell function.
“These data suggest that PNNs may drive PV cell maturation by directing transcription or translation of specific genes necessary for PV cell function.”
Quote in the paperSubstantial claim - Interpretationreasoning
In FXS, delayed PNN expression around PV cells during development may contribute to abnormal PV cell maturation.
“In FXS, delayed PNN expression around PV cells during development may contribute to abnormal PV cell maturation.”
Quote in the paperSubstantial claim - Interpretationcitation
These phenotypes in the mouse model are remarkably similar to those seen in humans with FXS.
“These phenotypes in the mouse model are remarkably similar to those seen in humans with FXS (Castrén et al., 2003; Ethridge et al., 2016; Wang et al., 2017).”
Quote in the paperSubstantial claim0 of 3 sources checked - Premisecitation
PV cell maturation is characterized by gradual increases in signaling, including faster firing rates and IPSC kinetics.
“PV cell maturation is characterized by gradual increases in signaling, including faster firing rates and IPSC kinetics (Lazarus and Huang, 2011).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Lazarus and Huang, 2011
Not checked in the free run.
- Interpretationreasoning
These findings suggest that MMP-9-mediated cleavage of PNNs during development may underlie sensory hypersensitivity in FXS.
“These findings suggest that MMP-9-mediated cleavage of PNNs during development may underlie sensory hypersensitivity in FXS.”
Quote in the paperSubstantial claim - Premisecitation
Recent studies have hinted at the potential role of MMP-9 in the manifestation of schizophrenia phenotypes.
“Recent studies have hinted at the potential role of MMP-9 in the manifestation of schizophrenia phenotypes.”
Quote in the paperNot a substantial claim - Premisecitation
Molecular mechanisms underlying seizure onset include impaired PV cell functions.
“The molecular mechanisms underlying seizure onset include impaired PV cell functions.”
Quote in the paperSubstantial claim - Interpretationreasoning
The neuroprotective effects of PNNs may be attributed to the CS-side chains of CSPGs.
“As chABC removes GAG side chains from CSPGs (Bradbury et al., 2002), these results suggest that the neuroprotective effects of PNNs may be attributed to the CS-side chains of CSPGs.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Bradbury et al., 2002 source
Not checked in the free run.
- Premisecitation
PV cells are particularly susceptible to oxidative stress given their fast cellular metabolism and increased mitochondrial function.
“PV cells are particularly susceptible to oxidative stress given their fast cellular metabolism and increased mitochondrial function (Hasenstaub et al., 2010; Kann et al., 2014; Inan et al., 2016).”
Quote in the paperSubstantial claim0 of 3 sources checked - Interpretationcitation
These findings lend support to the idea that PNN composition plays an important role in regulating PV cell function.
“These findings lend support to the idea that PNN composition plays an important role in regulating PV cell function.”
Quote in the paperSubstantial claim - Premisecitation
Effects of dysregulated dopamine signaling on prefrontal cortex oscillatory activity, which are suggested to drive schizophrenia phenotypes, can be also regulated by PNNs.
“The effects of dysregulated dopamine signaling on prefrontal cortex oscillatory activity, which are suggested to drive schizophrenia phenotypes, can be also regulated by PNNs.”
Quote in the paperSubstantial claim - Interpretationcitation
As PNNs regulate GABAergic cell function and oxidative stress levels, cortical abnormalities observed in schizophrenia may be attributed to PNN dysfunction.
“As PNNs regulate GABAergic cell function and oxidative stress levels, cortical abnormalities observed in schizophrenia may be attributed to PNN dysfunction (reviewed in Berretta, 2012; Berretta et al., 2015).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Cortical gamma band oscillations have been attributed to GABAergic cell activity, in particular PV interneurons.
“Cortical gamma band oscillations have been attributed to GABAergic cell activity, in particular PV interneurons (Sohal et al., 2009).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Sohal et al., 2009 source
Not checked in the free run.
- Interpretationreasoning
These data suggest altered PV cell function in schizophrenia.
“These data suggest altered PV cell function in schizophrenia, but why these cells are susceptible is unclear.”
Quote in the paperSubstantial claim - Interpretationcitation
Cells without PNNs are more susceptible to cell death.
“These results indicate that cells without PNNs are more susceptible to cell death (Morawski et al., 2004).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Morawski et al., 2004 source
Not checked in the free run.
- Interpretationcitation
Reduced PV cell spike output may contribute to seizures in mice with genetic knock-in of mutated human Scn1a, which results in subtle changes in Nav1.1 functionality.
“Recent studies demonstrate that reduced PV cell spike output may contribute to seizures in mice with genetic knock-in of mutated human Scn1a, which results in subtle changes in Nav1.1 functionality (Ogiwara et al., 2007; Hedrich et al., 2014).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Post-mortem brains of AD patients exhibit reduced expression of PNNs, particularly in the hippocampus.
“Post-mortem brains of AD patients exhibit reduced expression of PNNs, particularly in the hippocampus (Baig et al., 2005; Lendvai et al., 2013).”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationreasoning
The neuroprotective abilities of PNNs may stem from their ability to act as a barrier for oxidative stress-inducing molecules, such as metal ions.
“The neuroprotective abilities of PNNs may stem from their ability to act as a barrier for oxidative stress-inducing molecules, such as metal ions.”
Quote in the paperNot a substantial claim - Premisecitation
CSPGs present in PNNs appear to inhibit axonal growth.
“the CSPGs present in PNNs appear to inhibit axonal growth (Kwok et al., 2011).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Kwok et al., 2011 source
Not checked in the free run.
- Premisecitation
More cells died when pretreated with chABC in rodent cortical neuron experiments.
“More cells died when pretreated with chABC (Miyata et al., 2007).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Miyata et al., 2007 source
Not checked in the free run.
- Premisecitation
Cortical cells expressing PNNs exhibited accumulation of iron in post-fixed brain tissues obtained from both AD and normally aged controls.
“cortical cells expressing PNNs exhibited accumulation of iron in post-fixed brain tissues obtained from both AD and normally aged controls (Reinart et al., 2003).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Reinart et al., 2003 source
Not checked in the free run.
- Premisecitation
chABC treatment contributed to expanded receptive fields of uninjured primary afferents.
“but also contributed to expanded receptive fields of uninjured primary afferents (Bradbury et al., 2002; Massey et al., 2006).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Recent work has reported changes in PNN integrity in epilepsy.
“Recent work has reported changes in PNN integrity in epilepsy.”
Quote in the paperSubstantial claim - Interpretationcitation
Decrease in inhibitory cell function may underlie epilepsy-like activity following stroke.
“this decrease in inhibitory cell function may underlie epilepsy-like activity following stroke (Hobohm et al., 2005; Xie et al., 2014).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
In cortical areas surrounding the infarct and outside of the glial scar, WFA-expressing PNNs are completely degraded by day 7.
“in cortical areas surrounding the infarct and outside of the glial scar, WFA-expressing PNNs are completely degraded by day 7”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Hobohm et al., 2005 source
Not checked in the free run.
- Premisecitation
Neurons that exhibit buildup of tau protein and neurofibrillary tangles are more likely to lack PNNs.
“Neurons that exhibit buildup of tau protein and neurofibrillary tangles are more likely to lack PNNs (Brückner et al., 1999; Morawski et al., 2010).”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationcitation
MMP-9 may be responsible for ECM reorganization after CNS injury and stroke.
“MMP-9 may be responsible for this ECM reorganization”
Quote in the paperSubstantial claim0 of 3 sources checked - Premisecitation
Increased MMP-9 levels have been demonstrated following CNS injury and stroke in humans as well as animal models.
“a number of studies have demonstrated increased MMP-9 levels following CNS injury and stroke in humans as well as animal models (Zhang et al., 2011; Chaturvedi and Kaczmarek, 2014; Liu et al., 2014).”
Quote in the paperSubstantial claim0 of 3 sources checked - Premisecitation
Accumulation of lipofuscin, a byproduct of iron-catalyzed oxidative stress, was observed in cells without PNN.
“Accumulation of lipofuscin, a byproduct of iron-catalyzed oxidative stress, was also observed in cells without PNN (Morawski et al., 2004).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Morawski et al., 2004 source
Not checked in the free run.
- Premisecitation
RT-PCR analysis indicated upregulation of neurocan, which peaked on the third day following middle cerebral artery occlusion.
“RT-PCR analysis of the affected tissue indicated upregulation of neurocan, which peaked on the third day following middle cerebral artery occlusion”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Carmichael et al., 2005 source
Not checked in the free run.
- Premisecitation
Focal cortical stroke in rats contributes to upregulation of a number of CSPG-encoding genes particularly near the infarct and glial scar.
“focal cortical stroke in rats contributes to upregulation of a number of CSPG-encoding genes particularly near the infarct and glial scar.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Carmichael et al., 2005 source
Not checked in the free run.
- Premisecitation
CSPG degradation via application of chABC promoted regeneration of dorsal column axons after injury.
“CSPG degradation via application of chABC promoted regeneration of dorsal column axons after injury.”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
When sensory axons were transplanted into adult rats with dorsal column lesions, neurons were able to sprout and extend their processes to additional denervated areas but failed to penetrate the original lesion sites.
“When sensory axons were transplanted into adult rats with dorsal column lesions, neurons were able to sprout and extend their processes to additional denervated areas, but failed to penetrate the original lesion sites”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
In rat spinal cord injury models, increased concentrations of CSPGs and PNN expression were found at injury sites and surrounding secondary dorsal column relay neurons.
“In rat spinal cord injury models, increased concentrations of CSPGs and PNN expression were found at injury sites and surrounding secondary dorsal column relay neurons (Massey et al., 2008).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Massey et al., 2008 source
Not checked in the free run.
- Premisecitation
Similar degradation of WFA-expressing PNNs in the neocortex has also been observed following thromboembolic middle cerebral artery occlusion in rats, permanent middle artery occlusion in mice, and focal cerebral ischemia in sheep.
“Similar degradation of WFA-expressing PNNs in the neocortex has also been observed following thromboembolic middle cerebral artery occlusion in rats, permanent middle artery occlusion in mice, and focal cerebral ischemia in sheep (Härtig et al., 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Härtig et al., 2017 source
Not checked in the free run.
- Premisecitation
RT-PCR analysis indicated upregulation of versican and brevican, both of which peaked by day 28 following middle cerebral artery occlusion.
“as well as versican and brevican, both of which peaked by day 28 (Carmichael et al., 2005).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Carmichael et al., 2005 source
Not checked in the free run.
- Premisecitation
PNN-ensheathed rodent cortical neurons were more resistant to cell death when incubated with exogenous monomeric Aβ1-42 peptide for 48 h.
“PNN-ensheathed cells were more resistant to cell death when neurons were incubated with exogenous monomeric Aβ1-42 peptide for 48 h.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Miyata et al., 2007 source
Not checked in the free run.
- Premisecitation
chABC treatment resulted in an increase in terminals from initially injured afferents.
“This not only resulted in an increase in terminals from initially injured afferents”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationcitation
Perineuronal net loss in AD may contribute to altered E/I balance.
“Perineuronal net loss in AD may contribute to altered E/I balance (Palop et al., 2007; Sun et al., 2009)”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
MMP-9 is capable of cleaving Aβ in vitro.
“MMP-9 is capable of cleaving Aβ in vitro (Backstrom et al., 1996)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Backstrom et al., 1996 source
Not checked in the free run.
- Premisenone given
In several brain areas not all PV cells are ensheathed by PNNs as labeled with WFA.
“yet in several brain areas not all PV cells are ensheathed by PNNs as labeled with WFA (Table 1).”
Quote in the paperSubstantial claim - Premisecitation
PNNs containing aggrecan and brevican do not always overlap in mouse somatosensory cortex.
“PNNs containing aggrecan and brevican do not always overlap in mouse somatosensory cortex (Galtrey et al., 2008; Ueno et al., 2017b).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Galtrey et al., 2008 source
Not checked in the free run.
- cites Ueno et al., 2017b
Not checked in the free run.
- Interpretationreasoning
PNN loss is associated with a number of neurological disorders.
“Accumulating evidence suggests that PNN loss is associated with a number of neurological disorders.”
Quote in the paperSubstantial claim - Premisecitation
chABC-mediated degradation of PNNs around PV cells in brainstem and central auditory neurons results in reduced PV cell excitability and gain.
“chABC-mediated degradation of PNNs around PV cells in brainstem and central auditory neurons has similar effects, resulting in reduced PV cell excitability and gain (Balmer, 2016).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Balmer, 2016 source
Not checked in the free run.
- Premisecitation
In brainstem auditory nuclei of mice, genetic loss of brevican contributes to reduced sound-evoked firing rates.
“as well as reduced sound-evoked firing rates”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Blosa et al., 2015 source
Not checked in the free run.
- Premisecitation
In brainstem auditory nuclei of mice, genetic loss of brevican contributes to loss of brevican-expressing PNNs.
“In brainstem auditory nuclei of mice, genetic loss of brevican contributes to loss of brevican-expressing PNNs”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Blosa et al., 2015 source
Not checked in the free run.
- Interpretationcitation
Perineuronal net loss in AD may contribute to synaptic loss.
“synaptic loss (Scheibel and Tomiyasu, 1978; Ruiter and Uyling, 1987; Hoover et al., 2010)”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Scheibel and Tomiyasu, 1978
Not checked in the free run.
- cites Ruiter and Uyling, 1987
Not checked in the free run.
- cites Hoover et al., 2010 source
Not checked in the free run.
- Interpretationreasoning
Elevated MMP-9 levels can drive PNN loss and re-organization.
“As MMP-9 is capable of cleaving ECM, elevated MMP-9 levels can drive PNN loss and re-organization.”
Quote in the paperSubstantial claim- Rests on an assumption: MMP-9 is involved in PNN maintenance.
- Interpretationreasoning
Perineuronal nets may play a major role in PV neuron susceptibility and altered cortical processing in presbycusis.
“Perineuronal nets may play a major role in PV neuron susceptibility and altered cortical processing in presbycusis.”
Quote in the paperSubstantial claim - Premisecitation
In brainstem auditory nuclei of mice, genetic loss of brevican contributes to increased neuronal response thresholds.
“and increased neuronal response thresholds (Blosa et al., 2015).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Blosa et al., 2015 source
Not checked in the free run.
- Premisecitation
Genetic reduction of aggrecan resulted in reduced neuroprotective effects.
“Genetic reduction of aggrecan, which has the highest number of CS-side chains and presumably the largest negative charge, resulted in reduced neuroprotective effects (Suttkus et al., 2014).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Suttkus et al., 2014 source
Not checked in the free run.
- Premisecitation
PNN intensity is reduced in layers 1-4 of auditory cortex of adult CBA mice 1, 10, and 30 days post-noise exposure.
“PNN intensity is reduced in layers 1–4 of auditory cortex of adult CBA mice 1, 10, and 30 days post-noise exposure.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Nguyen et al., 2017 source
Not checked in the free run.
- Interpretationreasoning
PNNs may be necessary for neuronal excitability throughout the auditory pathway.
“These results indicate that PNNs may be necessary for neuronal excitability throughout the auditory pathway.”
Quote in the paperSubstantial claim - Premisecitation
Neuronal response properties susceptible to presbycusis include frequency selectivity, spontaneous activity levels and spectrotemporal selectivity.
“Neuronal response properties susceptible to presbycusis include frequency selectivity, spontaneous activity levels and spectrotemporal selectivity (Trujillo and Razak, 2013).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Trujillo and Razak, 2013
Not checked in the free run.
- Premisecitation
PNNs are heterogeneous structures.
“PNNs are heterogeneous structures (Ueno et al., 2017b; Yamada and Jinno, 2017; Irvine and Kwok, 2018)”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites Ueno et al., 2017b
Not checked in the free run.
- cites Yamada and Jinno, 2017 source
Not checked in the free run.
- cites Irvine and Kwok, 2018
Not checked in the free run.
- Interpretationreasoning
PNN composition may be brain region- and developmental age-specific.
“and PNN composition may be brain region- and developmental age-specific.”
Quote in the paperSubstantial claim - Premisenone given
Noise-induced hearing loss leads to increased excitability (gain) in the central auditory system.
“Noise-induced hearing loss leads to increased excitability (gain) in the central auditory system”
Quote in the paperSubstantial claim - Interpretationnone given
Reduced PNN intensity may be a result of increased cleavage or degradation of various PNN components, such as CSPGs, hyaluronan, or link proteins.
“Reduced PNN intensity may be a result of increased cleavage or degradation of various PNN components, such as CSPGs, hyaluronan, or link proteins.”
Quote in the paperNot a substantial claim - Premisecitation
Reduced GABAergic signaling has been demonstrated in the aged auditory system of rodents and humans.
“several studies have demonstrated reduced GABAergic signaling in the aged auditory system of rodents and humans (Caspary et al., 1990; Ouda et al., 2008; Martin del Campo et al., 2012; Gao et al., 2015).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisecitation
In layers 5-6, PNN intensity decreases 10 days post-noise exposure.
“In layers 5–6, PNN intensity decreases 10 days post-noise exposure (Nguyen et al., 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Nguyen et al., 2017 source
Not checked in the free run.
- Interpretationcitation
The CS-side chains of PNNs mediate its neuroprotective effects.
“These results lend further support to the idea that the CS-side chains of PNNs mediate its neuroprotective effects.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Suttkus et al., 2014 source
Not checked in the free run.
- Interpretationnone given
PNNs are necessary for PV cell function.
“Some of the work reviewed here indicates that PNNs are necessary for PV cell function”
Quote in the paperNot a substantial claim - Interpretationreasoning
PNN downregulation with age may make PV cells more susceptible to cell death.
“these data indicate that PNN downregulation with age may make PV cells more susceptible to cell death.”
Quote in the paperSubstantial claim - Interpretationcitation
Perineuronal net loss in AD may contribute to increased susceptibility to oxidative stress.
“and increased susceptibility to oxidative stress (Deibel et al., 1996; Varadarajan et al., 2000; Manczak et al., 2006; De Felice et al., 2007).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisenone given
Loss of PNNs around PV cells occurs following noise-induced and age-related hearing loss.
“Recent studies suggest that loss of PNNs around PV cells occurs following noise-induced and age-related hearing loss.”
Quote in the paperSubstantial claim - Premisecitation
PNNs are impaired following noise induced hearing loss, even within 1 day after noise exposure.
“PNNs are impaired following noise induced hearing loss, even within 1 day after noise exposure.”
Quote in the paperSubstantial claim - Premisecitation
Plasma levels of MMP-9 are elevated in AD patients.
“plasma levels of MMP-9 are elevated in AD patients (Backstrom et al., 1996; Lorenzl et al., 2003).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Co-treatment with MMP-9 inhibitors and NGF peptide following peripheral nerve injury can restore synaptic homeostasis, i.e., normalize glutamate/GABA ratio and expression of glial amino acid transporters.
“recent work in peripheral nerve injury models indicates that co-treatment with MMP-9 inhibitors and NGF peptide following insult can restore synaptic homeostasis, i.e., normalize glutamate/GABA ratio and expression of glial amino acid transporters (De Luca et al., 2016; Cirillo et al., 2016).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites De Luca et al., 2016
Not checked in the free run.
- cites Cirillo et al., 2016 source
Not checked in the free run.
- Premisecitation
MMP-9 upregulation in FXS can likely be attributed to loss of the Fmr1 gene.
“Unlike FXS, where MMP-9 upregulation can likely be attributed to loss of the Fmr1 gene (Janusz et al., 2013)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Janusz et al., 2013 source
Not checked in the free run.
- Premisecitation
Rats housed in enriched environments following ischemic damage exhibited faster neurological recovery compared to rats placed in standard housing.
“recent findings in rodent stroke models indicate that rats housed in enriched environments following ischemic damage exhibited faster neurological recovery compared to rats placed in standard housing (Madinier et al., 2014; Quattromani et al., 2018).”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationcitation
These findings provide important mechanistic insight for how MMP-9 dysregulation might arise in other CNS diseases.
“these findings provide important mechanistic insight for how MMP-9 dysregulation might arise in other CNS diseases.”
Quote in the paperNot a substantial claim - Premisecitation
Neuro-inflammation is evidenced in injury models, AD, epilepsy, and schizophrenia.
“there is evidence for neuro-inflammation in injury models, AD, epilepsy, and schizophrenia (Papa et al., 2014).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Papa et al., 2014 source
Not checked in the free run.
- Premisecitation
Absence of PNNs is associated with increased plasticity.
“As the absence of PNNs is associated with increased plasticity (Pizzorusso et al., 2002; Nowicka et al., 2009)”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Increased MMP-9 activity has been shown to degrade mature NGF.
“increased MMP-9 activity has been shown to degrade mature NGF (Bruno and Cuello, 2006).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Bruno and Cuello, 2006
Not checked in the free run.
- Premisecitation
Schizophrenia is associated with higher MMP-9/TIMP1 ratios compared to healthy controls.
“Schizophrenia is associated with higher MMP-9/TIMP1 ratios compared to healthy controls (Rahimi et al., 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Rahimi et al., 2017 source
Not checked in the free run.
- Premisecitation
Several ADAMTs are upregulated following spinal cord injury, stroke, and epilepsy.
“several ADAMTs are upregulated following spinal cord injury, stroke, and epilepsy (Lemarchant et al., 2013; Kim et al., 2016).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Premisecitation
Transcripts from MMP-9, 3, 7, and 14 are upregulated in rodent models of spinal cord injury.
“Quantitative PCR analysis indicates that transcripts from MMP-9, 3, 7, and 14 are upregulated in rodent models of spinal cord injury (Zhang et al., 2011).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Zhang et al., 2011 source
Not checked in the free run.
- Classified as probably not a substantial claim. 44
- May combine two claims. 55
Probably not a substantial claim: may not pass S3 (load-bearing).
FindingcitationBy week 4, tenascin expression is reduced to adult levels.
“By week 4, tenascin expression is reduced to adult levels (Brückner et al., 2000).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Brückner et al., 2000 source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
FindingcitationIn the mouse cortex, tenascin immunolabeling is first detected around E16 and tenascin levels increase after birth until the third postnatal week.
“In the mouse cortex, tenascin immunolabeling is first detected around E16 (Sheppard et al., 1991) and tenascin levels increase after birth until the third postnatal week (Brückner et al., 2000).”
Quote in the paperNot a substantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
FindingcitationSimilar to lecticans, tenascin and hyaluronan are also developmentally regulated.
“Similar to lecticans, tenascin and hyaluronan are also developmentally regulated.”
Quote in the paperNot a substantial claimMay combine two claims.
FindingcitationPeak MMP-9 expression is observed during postnatal development and is reduced during adulthood.
“Peak MMP-9 expression is observed during postnatal development and is reduced during adulthood (Bednarek et al., 2009; Reinhard et al., 2015; Wen et al., 2017).”
Quote in the paperNot a substantial claim0 of 3 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Findingnone givenWhile lecticans, tenascins, and hyaluronan are expressed during embryonic development, the classic net-like structure of PNNs is not typically seen until later during postnatal development.
“While lecticans, tenascins, and hyaluronan are expressed during embryonic development, the classic net-like structure of PNNs is not typically seen until later during postnatal development.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
FindingcitationHyaluronan is expressed during postnatal development and adulthood but peaks during embryonic development.
“Hyaluronan is expressed during postnatal development and adulthood but peaks during embryonic development (Bignami et al., 1993a).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Bignami et al., 1993a
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingnone givenDeveloping auditory cortex appears to exhibit fewer PNN and PV expressing cells than visual or somatosensory cortex.
“Interestingly, developing auditory cortex appears to exhibit fewer PNN and PV expressing cells than visual or somatosensory cortex, but in adult sensory cortices, a majority of PV cells express PNNs as seen with WFA staining.”
Quote in the paperNot a substantial claimMay combine two claims.
FindingcitationAuditory hypersensitivity is also evident in Fmr1 KO mice, which exhibit increased susceptibility to audiogenic seizures and abnormal auditory startle.
“Auditory hypersensitivity is also evident in Fmr1 knockout (KO) mice, a mouse model of FXS, which exhibit increased susceptibility to audiogenic seizures (Musumeci et al., 2000) and abnormal auditory startle (Frankland et al., 2004).”
Quote in the paperSubstantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
FindingcitationHyaluronan expression precedes the detection of both tenascin and lecticans, and is first observed near the notochord in E11-12 rat embryos.
“Hyaluronan expression precedes the detection of both tenascin and lecticans, and is first observed near the notochord in E11-12 rat embryos (Delpech and Delpech, 1984; Bignami et al., 1993a).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Delpech and Delpech, 1984
Not checked in the free run.
- cites Bignami et al., 1993a
Not checked in the free run.
May combine two claims.
FindingcitationIn adult mice, monocular deprivation followed by light exposure was associated with reduced WFA-expressing PNNs, lower aggrecan levels, and increased MMP-9 activity, as well as reduced PV cell output, increased firing of regular-spiking cells, and reduced gamma power.
“In this study, monocular deprivation followed by light exposure in adult mice was associated with reduced WFA-expressing PNNs, lower aggrecan levels, and increased MMP-9 activity, as well as reduced PV cell output, increased firing of regular-spiking cells, and reduced gamma power (Murase et al., 2017).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Murase et al., 2017 source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingown dataA large proportion of PNN-expressing PV cells is found within layer 4 of neocortex.
“A large proportion of PNN-expressing PV cells is found within layer 4 of neocortex (Table 1).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Table 1
Not checked in the free run.
May combine two claims.
FindingcitationOtx2 is localized to GABAergic cells, and is more likely found in PV cells that are enwrapped with PNNs, which contain a specific Otx2 binding site.
“Studies of rodent visual cortex development indicate that Otx2 is localized to GABAergic cells, and is more likely found in PV cells that are enwrapped with PNNs, which contain a specific Otx2 binding site (Beurdeley et al., 2012).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Beurdeley et al., 2012 source
Not checked in the free run.
May combine two claims.
Findingown dataThese phenotypes were mostly observed between P14 and P21, suggesting a window of abnormal development that can be targeted for therapeutic applications.
“These phenotypes were mostly observed between P14 and P21 suggesting a window of abnormal development that can be targeted for therapeutic applications.”
Quote in the paperSubstantial claimMay combine two claims.
FindingcitationRNA sequencing analysis of PV cells containing Otx2 indicates that Otx2 may be necessary for mitochondrial function, cellular respiration, and redox regulation.
“RNA sequencing analysis of PV cells containing Otx2 indicates that Otx2 may be necessary for mitochondrial function, cellular respiration, and redox regulation (Sakai et al., 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Sakai et al., 2017 source
Not checked in the free run.
May combine two claims.
FindingcitationThere is enhanced N1 amplitude and reduced gamma phase locking in auditory ERPs in Fmr1 KO mice.
“There is also enhanced N1 amplitude and reduced gamma phase locking in auditory ERPs (Lovelace et al., 2018).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Lovelace et al., 2018 source
Not checked in the free run.
May combine two claims.
Findingown dataAt P14, the responses of single neurons were similar across genotypes, while at P21, Fmr1 KO neurons show hyper-responsiveness.
“At P14, the responses of single neurons were similar across genotypes, while at P21, Fmr1 KO neurons show hyper-responsiveness.”
Quote in the paperSubstantial claimMay combine two claims.
FindingcitationGenetic reduction of MMP-9 levels in the Fmr1 KO mice normalized PV/PNN development and also reversed the hyperresponsiveness of auditory cortical neurons.
“Genetic reduction of MMP-9 levels in the Fmr1 KO mice normalized PV/PNN development and also reversed the hyperresponsiveness of auditory cortical neurons.”
Quote in the paperSubstantial claimMay combine two claims.
FindingcitationRedox dysregulation has been observed in schizophrenic patients, who have defective antioxidant systems and exhibit increased lipid peroxidation.
“Redox dysregulation has been observed in schizophrenic patients, who have defective antioxidant systems and exhibit increased lipid peroxidation (Do et al., 2000; Yao et al., 2006; Flatow et al., 2013).”
Quote in the paperSubstantial claim0 of 3 sources checkedMay combine two claims.
FindingcitationPost-mortem brains from patients with schizophrenia exhibit reduced density of WFA-labeled cells and reduced levels of CSPGs and PV expression in prefrontal cortex.
“Post-mortem brains from patients with schizophrenia exhibit reduced density of WFA-labeled cells (Mauney et al., 2013) and reduced levels of CSPGs and PV expression in prefrontal cortex (Enwright et al., 2016).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
FindingcitationGamma-band activity driven by rapid presentation of auditory stimuli and synchrony or phase-locking of electrocortical activity to these stimuli are reduced in schizophrenic patients compared to healthy controls.
“In contrast, gamma-band activity that is driven by rapid presentation of auditory stimuli and synchrony or phase-locking of electrocortical activity to these stimuli are reduced in schizophrenic patients compared to healthy controls (Kwon et al., 1999; Light et al., 2006).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Kwon et al., 1999
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- cites Light et al., 2006
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FindingcitationchABC-mediated removal of chondroitin sulfate side chains from PNNs reduced spiking of PV cells in adult rats and increased spiking variability.
“While this mechanism has not been confirmed, chABC-mediated removal of chondroitin sulfate side chains from PNNs reduced spiking of PV cells in adult rats (Balmer, 2016; Lensjø et al., 2017) and increased spiking variability (Lensjø et al., 2017).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
FindingcitationChondroitin sulfate promotes activation of protein kinase pathways including PKC and PI3K/Akt, and subsequent production of a neuronal antioxidant, heme oxygenase-1.
“Analysis of molecular mechanisms indicates that chondroitin sulfate promotes activation of protein kinase pathways including PKC and PI3K/Akt, and subsequent production of a neuronal antioxidant, heme oxygenase-1 (Cañas et al., 2007).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Cañas et al., 2007
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FindingcitationDrug-induced seizures led to increased expression of aggrecan cleavage products around PV cells and reduced expression of PNN components in rodent hippocampus, including Ctrl1 and hyaluronan synthase 3, both of which help to stabilize PNNs.
“Drug-induced seizures led to increased expression of aggrecan cleavage products around PV cells and reduced expression of PNN components in rodent hippocampus, including Ctrl1 and hyaluronan synthase 3, both of which help to stabilize PNNs (McRae and Porter, 2012; Rankin-Gee et al., 2015).”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationTenascins are fibrous, matricellular proteins that bind to the C-terminal domain of lecticans.
“Tenascins are fibrous, matricellular proteins, which bind to the C-terminal domain of lecticans (Aspberg et al., 1997; Day et al., 2004).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationPNNs are composed of chondroitin sulfate proteoglycans, tenascin, and link proteins, which facilitate binding to a hyaluronan backbone.
“PNNs are composed of CSPGs, tenascin, and link proteins, which facilitate binding to a hyaluronan backbone (Yamaguchi, 1999) (Figure 2).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Yamaguchi, 1999 source
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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisenone givenVersican consists of two different alpha and beta isoforms, which are differentially expressed during development.
“Versican consists of two different alpha and beta isoforms, which are differentially expressed during development.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
PremisecitationPerineuronal nets are specialized assemblies of extracellular matrix proteins that ensheath neurons and proximal dendrites.
“Perineuronal nets are specialized assemblies of ECM proteins which ensheath neurons and proximal dendrites (Celio et al., 1998).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Celio et al., 1998 source
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Premisenone givenChondroitin sulfate proteoglycans consist of a core protein and varying amounts of chondroitin sulfate chains covalently bound to a central region of the protein.
“Chondroitin sulfate proteoglycans consist of a core protein and varying amounts of chondroitin sulfate chains covalently bound to a central region of the protein.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenPNNs have been implicated in neuroprotection.
“Since then, PNNs have been implicated in a wide range of functions including neuroprotection, regulation of neural activity and experience-dependent plasticity.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
Premisenone givenMMP-9 is a protease that cleaves extracellular matrix.
“MMP-9, a protease that cleaves ECM, is differentially regulated in a number of these disorders.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationThe lectican family includes aggrecan, neurocan, versican, and brevican, which vary in core protein size and number of GAG side chains.
“These include aggrecan, neurocan, versican, and brevican, which vary in core protein size and the number of GAG side chains (Yamaguchi, 1999).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Yamaguchi, 1999 source
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InterpretationreasoningMMP-9 mediated cleavage and/or reorganization of PNN structures around PV-expressing cells contributes to abnormal PV cell development and cortical hyperexcitability.
“we propose a novel working hypothesis in which MMP-9 mediated cleavage and/or reorganization of PNN structures around PV-expressing cells contributes to abnormal PV cell development and cortical hyperexcitability (Figure 1).”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenPNNs have been implicated in experience-dependent plasticity.
“Since then, PNNs have been implicated in a wide range of functions including neuroprotection, regulation of neural activity and experience-dependent plasticity.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenThe functional impact of developmental changes in PNN lectican composition is unknown.
“However, the functional impact of developmental changes in PNN lectican composition is unknown.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based).
Premisenone givenFew studies have addressed the interactions between PNN expression, MMP-9 activity and neuronal excitability.
“Despite this, few studies have addressed the interactions between PNN expression, MMP-9 activity and neuronal excitability.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationHyaluronan is an unsulfated GAG that is synthesized at the cell surface by membrane-bound enzymes called HAS and protrudes through the plasma membrane into the extracellular space.
“Hyaluronan is an unsulfated GAG that is synthesized at the cell surface by the membrane-bound enzymes called HAS and protrudes through the plasma membrane into the extracellular space (Fraser et al., 1997).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Fraser et al., 1997 source
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PremisecitationChondroitin sulfate side chains are a type of GAG polysaccharide consisting of disaccharide blocks of N-acetylgalactosamine and glucuronic acid.
“These chondroitin sulfate side chains are a type of GAG polysaccharide consisting of disaccharide blocks of N-acetylgalactosamine and glucuronic acid (Sherman and Back, 2007).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Sherman and Back, 2007 source
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PremisecitationThe role of PNNs in neural and network excitability is only beginning to be understood.
“However, the role of PNNs in neural and network excitability is only beginning to be understood (Balmer, 2016; Murase et al., 2017; Wen et al., 2017).”
Quote in the paperNot a substantial claim0 of 3 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenThe function of lecticans in CNS development remains unclear.
“Although lecticans are developmentally regulated, their function in CNS development remains unclear.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisenone givenThe lectican family of CSPGs is most prominent in PNNs of the CNS.
“Various CSPGs exist in the ECM throughout the body, but the lectican family of CSPGs is most prominent in PNNs of the CNS.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S2 (checkable).
InterpretationreasoningSystematic studies of the interplay between MMP-9, PNN and PV cells may be key to understanding mechanisms of multiple CNS disorders at the cellular and circuit levels.
“we bring together evidence from studies of schizophrenia, AD, and epilepsy to suggest that systematic studies of the interplay between MMP-9, PNN and PV cells may be key to understanding mechanisms of multiple CNS disorders at the cellular and circuit levels.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationFXS is a leading genetic cause of autism, occurring in 1 in 4000 males and 1 in 8000 females.
“FXS is a leading genetic cause of autism, occurring in 1 in 4000 males and 1 in 8000 females (Murray et al., 1997).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Murray et al., 1997 source
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Premisenone givenParvalbumin cells are a major class of GABAergic, fast-spiking inhibitory interneurons in the neocortex.
“Parvalbumin cells are a major class of GABAergic, fast-spiking inhibitory interneurons in the neocortex.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
InterpretationcitationBoth neurons and glial cells are involved in the assembly of PNN around neurons.
“It is unclear how these molecules are assembled to form PNN around neurons, but some studies suggest that both neurons and glial cells are involved in the process (Miyata et al., 2007; Pyka et al., 2011).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
InterpretationcitationAggrecan-rich PNNs may be unstable in the presence of MMP-9, leading to abnormal development and neural excitability.
“As aggrecan has been previously identified as an MMP-9 target (Mercuri et al., 2000; Madsen et al., 2010; Rankin-Gee et al., 2015), aggrecan-rich PNNs may be unstable in the presence of MMP-9, leading to abnormal development and neural excitability (Murase et al., 2017; Wen et al., 2017).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationThis leads to loss of FMRP, which is necessary for regulation of protein synthesis.
“This leads to loss of Fragile X Mental Retardation protein (FMRP), which is necessary for regulation of protein synthesis (Verkerk et al., 1991).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Verkerk et al., 1991 source
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Premisenone givenThe most common cause of FXS is an expansion of CGG trinucleotide repeats in the 5′ region of the Fmr1 gene that results in its hypermethylation and inactivation.
“The most common cause of FXS is an expansion of CGG trinucleotide repeats in 5′ region of the Fragile X mental retardation 1 (Fmr1) gene that results in its hypermethylation and inactivation.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
InterpretationcitationThere is a tetrapartite synaptic organization involving ECM, pre/post synaptic sites and astrocytes.
“Astrocytes extend their processes into PNNs localized near the synapse (Brückner et al., 1993; Blümcke et al., 1995), suggesting a tetrapartite synaptic organization involving ECM, pre/post synaptic sites and astrocytes (Dityatev and Rusakov, 2011; Smith et al., 2015) whose synergistic functions remain unexplored.”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Dityatev and Rusakov, 2011
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- cites Smith et al., 2015 source
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Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIt is unclear how these molecules are assembled to form PNN around neurons.
“It is unclear how these molecules are assembled to form PNN around neurons, but some studies suggest that both neurons and glial cells are involved in the process (Miyata et al., 2007; Pyka et al., 2011).”
Quote in the paperNot a substantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationIndividuals with FXS suffer from social and communication deficits, sensory hypersensitivity, and anxiety disorders.
“Individuals with this neurodevelopmental disorder suffer from social and communication deficits, sensory hypersensitivity, and anxiety disorders (Abbeduto and Hagerman, 1997; Rogers et al., 2001).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Abbeduto and Hagerman, 1997
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- cites Rogers et al., 2001
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Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationLike PNNs, PV expression is developmentally regulated and experience-dependent.
“Like PNNs, PV expression is developmentally regulated and experience-dependent (Hensch, 2005).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Hensch, 2005 source
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PremisecitationMMP-9 is a zinc-dependent endopeptidase expressed in various cells within the CNS which is capable of cleaving ECM proteins, including collagen and laminin.
“MMP-9 is a zinc-dependent endopeptidase expressed in various cells within the CNS which is capable of cleaving ECM proteins, including collagen and laminin (reviewed in Reinhard et al., 2015).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Reinhard et al., 2015 source
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PremisecitationOne mechanism that has been proposed to explain some symptoms of schizophrenia is a reduction in GABAergic signaling in specific cortical regions.
“One mechanism that has been proposed to explain some symptoms of schizophrenia is a reduction in GABAergic signaling in specific cortical regions.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationDelayed PNN development and increased MMP-9 activity may lead to impaired PV cell development and reduced inhibition.
“We found delayed PNN development and increased MMP-9 activity levels in the developing auditory cortex of Fmr1 KO mice (Wen et al., 2017), which may lead to impaired PV cell development and reduced inhibition.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationcitationDuring development, sensory stimuli can activate Narp expression which promotes PV interneuron excitation, leading to feed-forward inhibition.
“This suggests that during development, sensory stimuli can activate Narp expression which promotes PV interneuron excitation, leading to feed-forward inhibition (O’Brien et al., 1999; Gu et al., 2013).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites O’Brien et al., 1999 source
Not checked in the free run.
- cites Gu et al., 2013
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May combine two claims.
InterpretationcitationOtx2 may play a role in PV cell maturation and refinement of cortical inhibition.
“An orthodenticle homeoprotein, Otx2, may play a role in PV cell maturation and refinement of cortical inhibition.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningThese results indicate that there is altered E/I balance in auditory cortex, which may underlie auditory hypersensitivity phenotypes.
“These results indicate that there is altered E/I balance in auditory cortex, which may underlie auditory hypersensitivity phenotypes.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningIn FXS, delayed development of PNNs may contribute to abnormal Narp expression, impairing the ability of PV cells to scale cortical networks, resulting in cortical hyperexcitability.
“In FXS, delayed development of PNNs may contribute to abnormal Narp expression, impairing the ability of PV cells to scale cortical networks, resulting in cortical hyperexcitability.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationcitationPerineuronal net dysfunction in schizophrenia may drive changes in GABAergic cell function leading to network dysfunction, as normal PNN expression regulates excitability of PV cells.
“Perineuronal net dysfunction in schizophrenia may also drive changes in GABAergic cell function leading to network dysfunction, as normal PNN expression regulates excitability of PV cells (Liu et al., 2013; Balmer, 2016).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
InterpretationcitationPNNs may protect PV cells from oxidative stress, and thus serve as a 'safety net' to support the rapid firing properties of PV cells.
“Recent studies show that PNNs may protect PV cells from oxidative stress (Cabungcal et al., 2013; Morishita et al., 2015), and thus serve as a ‘safety net’ to support the rapid firing properties of PV cells.”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationScn1a is a gene that encodes Nav1.1, a sodium channel necessary for PV cell firing.
“Scn1a is a gene that encodes Nav1.1, a sodium channel necessary for PV cell firing (Ogiwara et al., 2007; Hedrich et al., 2014).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
InterpretationcitationMinocycline can reduce plasma levels of MMP-9 and inhibit MMP-9 activity, suggesting a possible role of MMP-9 inhibition in the beneficial effects of minocycline in schizophrenia.
“Beside its antibiotic activity, minocycline can reduce plasma levels of MMP-9 (Dziembowska et al., 2013) and inhibit MMP-9 activity (Lee et al., 2006), suggesting a possible role of MMP-9 inhibition in the beneficial effects of minocycline in schizophrenia.”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationFailure of transplanted sensory axons to penetrate original lesion sites led to denervation of upstream pathways and inactivation of parts of somatosensory cortex.
“which lead to denervation of upstream pathways and inactivation of parts of somatosensory cortex (Davies et al., 1999; Kaas et al., 2008).”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationMetal ions can react with free radicals generated by cellular metabolism to generate even more free radicals, resulting in lipid peroxidation, oxidative stress, and cell death.
“The molecules can react with free radicals generated by cellular metabolism to generate even more free radicals, resulting in lipid peroxidation, oxidative stress, and cell death (Stohs and Bagchi, 1995).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Stohs and Bagchi, 1995 source
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PremisecitationIn cortical areas surrounding the infarct and outside of the glial scar, CSPG immunoreactivity is down regulated although it remains detectable up to 14 days post-injury.
“CSPG immunoreactivity is down regulated although they remain detectable up to 14 days post-injury (Hobohm et al., 2005).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Hobohm et al., 2005 source
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PremisecitationReduced PV-immunoreactivity and PV cell function have been observed following stroke.
“reduced PV-immunoreactivity and PV cell function have also been observed following stroke”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
InterpretationcitationAlzheimer's disease may lead to hippocampal and cortical hyperexcitability as well as increased seizure susceptibility in both AD humans and rodent models.
“This may lead to hippocampal and cortical hyperexcitability as well as increased seizure susceptibility in both AD humans and rodent models (Palop et al., 2007; Roberson et al., 2011).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Palop et al., 2007 source
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- cites Roberson et al., 2011
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Classified as probably not a substantial claim. May combine two claims.
InterpretationcitationAs PNNs regulate PV cell activity and protect PV cells from oxidative stress, PNNs can be potentially targeted to control seizures.
“As PNNs regulate PV cell activity (Balmer, 2016; Lensjø et al., 2017) and protect PV cells from oxidative stress (Cabungcal et al., 2013), PNNs can be potentially targeted to control seizures.”
Quote in the paperSubstantial claim0 of 3 sources checkedMay combine two claims.
PremisecitationThe number of PNN expressing cells is reduced in aging mouse auditory cortex with a particular impact on PV/PNN co-localized cells.
“The number of PNN expressing cells is reduced in aging mouse auditory cortex with a particular impact on PV/PNN co-localized cells (Brewton et al., 2016).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Brewton et al., 2016 source
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PremisecitationOverexposure to loud sounds can cause hair cell damage.
“Overexposure to loud sounds can cause hair cell damage (Puel et al., 1998)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Puel et al., 1998 source
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PremisecitationMinocycline administration may reverse both schizophrenia and FXS phenotypes, perhaps due to its ability to lower MMP-9 levels or activity.
“Recent studies indicate that minocycline administration may reverse both schizophrenia and FXS phenotypes, perhaps due to its ability to lower MMP-9 levels or activity (Dziembowska et al., 2013).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Dziembowska et al., 2013 source
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PremisecitationGABAergic signaling is involved in shaping frequency selectivity, spontaneous activity levels and spectrotemporal selectivity, with fast-spiking PV cells shaping both spectral and temporal selectivity.
“GABAergic signaling is involved in shaping these properties, with fast-spiking PV cells shaping both spectral and temporal selectivity (Moore and Wehr, 2012).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Moore and Wehr, 2012 source
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InterpretationreasoningActivity-dependent synaptogenesis and the development of stable neuronal networks may be a result of normal PNN interactions with inhibitory interneurons, in particular fast spiking PV cells.
“activity-dependent synaptogenesis and the development of stable neuronal networks may be a result of normal PNN interactions with inhibitory interneurons, in particular fast spiking PV cells.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based).
Premisenone givenPNNs are implicated in experience-dependent plasticity during development.
“PNNs are also implicated in experience-dependent plasticity during development.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationAge-related hearing loss (presbycusis) is associated with difficulty interpreting speech and slower auditory processing.
“Age-related hearing loss, also known as presbycusis, is associated with difficulty interpreting speech and slower auditory processing (Gates and Mills, 2005)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Gates and Mills, 2005
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InterpretationreasoningAltered PNN formation may contribute to abnormal GABAergic signaling and impaired temporal processing with age, leading to deficits in speech comprehension.
“Altered PNN formation may contribute to abnormal GABAergic signaling and impaired temporal processing with age, leading to deficits in speech comprehension.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S4 (based).
Interpretationnone givenThe use of chABC likely has broader effects beyond addressing PNN function.
“While the use of chABC has been useful in addressing PNN function, this enzyme likely has broader effects.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationOverexposure to loud sounds can cause hearing loss and reduced afferent input in the auditory system.
“hearing loss and reduced afferent input in the auditory system (Syka, 2002).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Syka, 2002 source
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May combine two claims.
PremisecitationIn the rodent hippocampus, aggrecan-expressing PNNs overlap with WFA labeling in stratum oriens, but stratum pyramidale exhibits aggrecan-rich PNNs, which are not WFA-positive.
“in the rodent hippocampus, aggrecan-expressing PNNs overlap with WFA labeling in stratum oriens, but stratum pyramidale exhibits aggrecan-rich PNNs, which are not WFA-positive (Yamada and Jinno, 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Yamada and Jinno, 2017
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PremisecitationIron-induced oxidative stress resulted in apoptotic cell death in cells without PNNs, while neurons with PNNs were more likely to survive.
“While iron-induced oxidative stress resulted in apoptotic cell death in cells without PNNs, neurons with PNNs were more likely to survive (Suttkus et al., 2012, 2014).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Suttkus et al., 2012, 2014
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InterpretationreasoningElevated MMP-9 levels may contribute to both PNN degradation and altered levels of Aβ in AD.
“elevated MMP-9 levels may contribute to both PNN degradation and altered levels of Aβ in AD.”
Quote in the paperSubstantial claim- Rests on an assumption: MMP-9 is involved in PNN maintenance.
May combine two claims.
PremisecitationTenascin-R expressing PNNs are not observed in rodent PFC, but are prominent in other cortical areas.
“Tenascin-R expressing PNNs are not observed in rodent PFC, but are prominent in other cortical areas (Ueno et al., 2017a).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Ueno et al., 2017a
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Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationTIMP levels may be indicators for risk of stroke mortality.
“and may be indicators for risk of stroke mortality (Hansson et al., 2011).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Hansson et al., 2011 source
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PremisecitationTIMP levels are increased in postmortem brains of AD patients.
“TIMP levels are increased in postmortem brains of AD patients (Peress et al., 1995)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Peress et al., 1995 source
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Interpretationnone givenThe ADAMTs family of aggrecanases may also cause PNN cleavage and reorganization.
“the ADAMTs family of aggrecanases may also cause PNN cleavage and reorganization.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningAbsence of PNNs may allow for axonal outgrowth and synaptogenesis, particularly in surviving cortical areas, which exhibit reduced CSPG and PNN expression and are therefore growth permissive.
“this may allow for axonal outgrowth and synaptogenesis, particularly in surviving cortical areas, which exhibit reduced CSPG and PNN expression and are therefore growth permissive (2014).”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationTIMPs form 1:1 complexes with MMPs in order to inhibit MMP activity.
“tissue inhibitors of MMPs, known as TIMPs, which form 1:1 complexes with MMPs in order to inhibit MMP activity (Gomis-Rüth et al., 1997; Brew and Nagase, 2010).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Gomis-Rüth et al., 1997 source
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- cites Brew and Nagase, 2010
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Classified as probably not a substantial claim.
InterpretationcitationNeuro-inflammation may be crucial in understanding MMP-9 dysregulation in CNS disorders.
“these may be crucial in understanding MMP-9 dysregulation in CNS disorders”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Papa et al., 2014 source
Not checked in the free run.
The paper relies on this without support.
Re-establishing neuronal plasticity is key for re-innervation and recovery after CNS injury and stroke.
“In CNS injury and stroke, re-establishing neuronal plasticity appears to be key for re-innervation and recovery.”
The paper relies on this without support.
MMP-9 is involved in PNN maintenance.
“As MMP-9 is also involved in PNN maintenance”
The paper relies on this without support.
Neurotrophins are necessary for neuronal growth and plasticity.
“Neurotrophins are necessary for neuronal growth and plasticity”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Perineuronal nets (PNN) regulate synaptic plasticity and rewiring of the network of interneurons.
“Importantly, PNN regulate the synaptic plasticity (Kochlamazashvili et al., 2010) and rewiring the network (Pollock et al., 2014) of the interneurons (Wen et al., 2018).”
From Disrupted structural connectivity in ArcAβ mouse model of Aβ amyloidosis (2020) · cites this paper as Wen et al., 2018
Microglia release matrix metalloproteinases or other ECM-degrading proteases that act on perineuronal net components in acute injuries such as stroke.
“microglia undergo activation and subsequently release matrix metalloproteinases (e.g. MMP9) or other ECM-degrading proteases that act on PNN components [33].”
From Microglia facilitate loss of perineuronal nets in the Alzheimer's disease brain (2020) · cites this paper as [33]
Cites
Oxidative mechanisms in the toxicity of metal ions
Stohs, S. J., & Bagchi, D. (1995) · Free Radical Biology and Medicine · doi:10.1016/0891-5849(94)00159-h · Cited by 4,219 · see citing works
Verkerk, A. J. M. H., Pieretti, M., Sutcliffe, J. S., Fu, Y.-H., Kuhl, D. P. A., Pizzuti, A., Reiner, O., Richards, S., Victoria, M. F., Zhang, F., et al. (1991) · Cell · doi:10.1016/0092-8674(91)90397-h · Cited by 3,481 · see citing works
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Microglia facilitate loss of perineuronal nets in the Alzheimer's disease brain
Crapser, J. D., Spangenberg, E. E., Barahona, R. A., Arreola, M. A., Hohsfield, L. A., & Green, K. N. (2020) · EBioMedicine · doi:10.1016/j.ebiom.2020.102919 · Cited by 280 · see citing works
MMPs in learning and memory and neuropsychiatric disorders
Beroun, A., Mitra, S., Michaluk, P., Pijet, B., Stefaniuk, M., & Kaczmarek, L. (2019) · Cellular and Molecular Life Sciences · doi:10.1007/s00018-019-03180-8 · Cited by 207 · see citing works
Yu, Z., Chen, N., Hu, D., Chen, W., Yuan, Y., Meng, S., Zhang, W., Lu, L., Han, Y., & Shi, J. (2020) · Frontiers in Molecular Neuroscience · doi:10.3389/fnmol.2020.00004 · Cited by 52 · see citing works
Reinhard, S. M., Abundez-Toledo, M., Espinoza, K., & Razak, K. A. (2019) · Hearing Research · doi:10.1016/j.heares.2019.107781 · Cited by 15 · see citing works
Self-citationDisrupted structural connectivity in ArcAβ mouse model of Aβ amyloidosis
Al-Amin, M. M., Grandjean, J., Klohs, J., & Kim, J. (2020) · doi:10.1101/2020.04.27.064634 · Cited by 2 · see citing works
“Importantly, PNN regulate the synaptic plasticity (Kochlamazashvili et al., 2010) and rewiring the network (Pollock et al., 2014) of the interneurons (Wen et al., 2018).”
Background
- Authors
- Wen, T. H., Binder, D. K., Ethell, I. M., & Razak, K. A.
- Affiliations
- University of California, Riverside (US)
- Corresponding author
- Ethell, Iryna M.
- Razak, Khaleel A.
- Venue
- Frontiers in Molecular Neuroscience
- Publisher
- Frontiers Media SA
- Published
- 2018-08-03
- Volume
- 11
- Pages
- 270-270
- ISSN
- 1662-5099 (ISSN-L)
- PMC
- PMC6085424
- OpenAlex
- W2886787141
- Cited by
- 170
- References
- 209
- Citations by year
- 2026: 15 · 2025: 21 · 2024: 16 · 2023: 26 · 2022: 27 · 2021: 27 · 2020: 24 · 2019: 12 · 2018: 2
- FWCI
- 2.10
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Frontiers in Molecular Neuroscience (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- eScholarship (California Digital Library) (submittedVersion)
- eScholarship (California Digital Library) (submittedVersion, cc-by)
- Funders
- National Institutes of Health (U54 HD082008)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetics and Neurodevelopmental Disorders
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cancer Research › Protease and Inhibitor Mechanisms
- Keywords
- perineuronal nets, extracellular matrix, parvalbumin interneurons, MMP-9, fragile X syndrome, autism spectrum disorder, schizophrenia, Alzheimer's disease, cortical excitability, critical period plasticity, GABAergic inhibition, oxidative stress, neurodevelopmental disorders, neurodegenerative diseases
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-06.
Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-06.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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