The DOI matches its Crossref record; 52 of the 52 references it cites were found in a registry or an index; none of the 58 sources its claims cite is checked for support yet.
Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons
Open accessAbstract
UNLABELLED: Long-term potentiation of excitatory synapses on pyramidal neurons in the stratum radiatum rarely occurs in hippocampal area CA2. Here, we present evidence that perineuronal nets (PNNs), a specialized extracellular matrix typically localized around inhibitory neurons, also surround mouse CA2 pyramidal neurons and envelop their excitatory synapses. CA2 pyramidal neurons express mRNA transcripts for the major PNN component aggrecan, identifying these neurons as a novel source for PNNs in the hippocampus. We also found that disruption of PNNs allows synaptic potentiation of normally plasticity-resistant excitatory CA2 synapses; thus, PNNs play a role in restricting synaptic plasticity in area CA2. Finally, we found that postnatal development of PNNs on CA2 pyramidal neurons is modified by early-life enrichment, suggesting that the development of circuits containing CA2 excitatory synapses are sensitive to manipulations of the rearing environment. SIGNIFICANCE STATEMENT: Perineuronal nets (PNNs) are thought to play a major role in restricting synaptic plasticity during postnatal development, and are altered in several models of neurodevelopmental disorders, such as schizophrenia and Rett syndrome. Although PNNs have been predominantly studied in association with inhibitory neurons throughout the brain, we describe a dense expression of PNNs around excitatory pyramidal neurons in hippocampal area CA2. We also provide insight into a previously unrecognized role for PNNs in restricting plasticity at excitatory synapses and raise the possibility of an early critical period of hippocampal plasticity that may ultimately reveal a key mechanism underlying learning and memory impairments of PNN-associated neurodevelopmental disorders.
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46 Own result · 39 Not checked yet · 11 Own interpretation · 4 Own method
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“We also found that disruption of PNNs allows synaptic potentiation of normally plasticity-resistant excitatory CA2 synapses; thus, PNNs play a role in restricting synaptic plasticity in area CA2.”
- Also stated in: § PNNs suppress synaptic plasticity in area CA2 · plain
“However, a 2 h incubation with ChABC enabled LTP induction of excitatory synaptic responses in the CA2 SR (140 Ϯ 0.2% baseline, n ϭ 10), reaching levels comparable to that induced at CA1 synapses ( p ϭ 0.02, two-tailed unpaired t test compared with untreated slices).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Abstract
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- Also stated in: § PNNs suppress synaptic plasticity in area CA2 · plain
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“CA2 pyramidal neurons express mRNA transcripts for the major PNN component aggrecan, identifying these neurons as a novel source for PNNs in the hippocampus.”
- Also stated in: § PNNs localize to excitatory synapses on CA2 pyramidal neurons · plain
“We found that CA2 neurons identified by Purkinje cell protein 4 (PCP4 ) expression, but not CA1 and CA3 neurons, expressed aggrecan mRNA (Fig. 1c).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Abstract
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- Also stated in: § PNNs localize to excitatory synapses on CA2 pyramidal neurons · plain
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“Finally, we found that postnatal development of PNNs on CA2 pyramidal neurons is modified by early-life enrichment, suggesting that the development of circuits containing CA2 excitatory synapses are sensitive to manipulations of the rearing environment.”
- Also stated in: § PNN development is regulated by experience · plain
“We found that WFA staining intensity in the CA2 SP was significantly greater in EE-exposed animals than those reared in standard housing at PN 21 and PN 45 ( p ϭ 0.047 and p ϭ 0.001 respec-tively; two-way ANOVA, Bonferroni's post hoc test for pairwise comparison), but not at PN 14, the earliest age tested (Fig. 3b; p Ͼ 0.05).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: WFA staining intensity reliably measures the amount or development of perineuronal nets in CA2.
- Also stated in: § PNN development is regulated by experience · plain
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“In contrast, staining for PNNs in the neighboring CA1 SP and SR layers was sparse (Fig. 2c).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs localize to excitatory synapses on CA2 pyramidal neurons
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“CA2 neuron input resistance was significantly decreased after 0.05 U/ml ChABC treatment (n ϭ 11 per group, *p Ͻ 0.0043, two-tailed unpaired t test).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
plaindirectn = 11 ·- Asserted: passes — The authors put it forward as true.
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“We show here that PNNs in area CA2 are upregulated by early-life exposure to EE.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
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“Our findings that PNN degradation enables LTP in area CA2”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
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“Furthermore, in the presence of bicuculline, we found no significant differences in basal excitatory synaptic transmission (Fig. 4d), paired-pulse facilitation (Fig. 4e), or AMPAR/NMDAR ratio (Fig. 4f ).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
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“WFA staining intensity was similarly significantly greater in the CA2 SR at PN 21 and PN 45 ( p ϭ 0.015 and p ϭ 0.001 respectively; two-way ANOVA, Bonferroni's post hoc test for pairwise comparison; Fig. 3c).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNN development is regulated by experience
plaincorrelational- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: WFA staining intensity reliably measures the amount or development of perineuronal nets in CA2.
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In the paper
“Intrinsic properties did not change in a way that could explain this newly uncovered plasticity in area CA2 ( p Ͼ 0.05, two-way ANOVA, Bonferroni's post hoc test for pair-wise comparison; Fig. 4c; Tables 1, 2).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
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- It may state two separate claims as one. 27
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- Probably not a substantial claim. 24
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Here, we present evidence that perineuronal nets (PNNs), a specialized extracellular matrix typically localized around inhibitory neurons, also surround mouse CA2 pyramidal neurons and envelop their excitatory synapses.”
- Also stated in: § PNNs localize to excitatory synapses on CA2 pyramidal neurons · plain
“Three different PNN markers, WFA, the chondroitin sulfate proteoglycans aggrecan and neurocan, labeled select inhibitory interneurons throughout the hippocampus (Fig. 1a), but also intensely labeled EGFP-expressing CA2 pyramidal neurons (Fig. 1b).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Abstract
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It may state two separate claims as one.
- Also stated in: § PNNs localize to excitatory synapses on CA2 pyramidal neurons · plain
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Three different PNN markers, WFA, the chondroitin sulfate proteoglycans aggrecan and neurocan, labeled select inhibitory interneurons throughout the hippocampus (Fig. 1a), but also intensely labeled EGFP-expressing CA2 pyramidal neurons (Fig. 1b).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs localize to excitatory synapses on CA2 pyramidal neurons
boostedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“However, a 2 h incubation with ChABC enabled LTP induction of excitatory synaptic responses in the CA2 SR (140 Ϯ 0.2% baseline, n ϭ 10), reaching levels comparable to that induced at CA1 synapses ( p ϭ 0.02, two-tailed unpaired t test compared with untreated slices).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
plaindirectn = 10 ·- Asserted: passes — The authors put it forward as true.
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In the paper
“Significant effects were not detected in other properties, including resting membrane potential, membrane capacitance, and decay time.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“Similar results were obtained when experiments on ChABC-treated or control-treated slices were performed in the presence of the GABA A -receptor blocker bicuculline (significant difference between ChABC-treated and untreated slices, p ϭ 0.022 at 10 min, two-tailed unpaired t test compared with untreated slices, n ϭ 8 and 4 respectively).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
plaindirectn = 8 ·- Asserted: passes — The authors put it forward as true.
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In the paper
“As reported previously for untreated slices, synaptic responses recorded from CA2 neurons were not potentiated following an LTP”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
What the sources say
- Not checked yetcites Zhao et al., 2007
plainn = 10- Asserted: passes — The authors put it forward as true.
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In the paper
“We found that WFA staining intensity in the CA2 SP was significantly greater in EE-exposed animals than those reared in standard housing at PN 21 and PN 45 ( p ϭ 0.047 and p ϭ 0.001 respec-tively; two-way ANOVA, Bonferroni's post hoc test for pairwise comparison), but not at PN 14, the earliest age tested (Fig. 3b; p Ͼ 0.05).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNN development is regulated by experience
plain- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
- Rests on an assumption: WFA staining intensity reliably measures the amount or development of perineuronal nets in CA2.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Normalized WFA fluorescence intensity was significantly greater in CA2 SP of EE mice compared with control at PN 21 (N ϭ 8 and 7 for control and EE respectively) and PN 45 (N ϭ 6 and 9 for control and EE respectively; Bonferroni's post hoc test for pairwise comparison after two-way ANOVA,*p Ͻ 0.05, ***p ϭ 0.0008).No significant difference was observed at PN 14”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
plaincorrelationaln = 9- Asserted: passes — The authors put it forward as true.
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In the paper
“PNNs increase during postnatal development and are increased with experience.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
plaindirectn = 7- Asserted: passes — The authors put it forward as true.
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In the paper
“Electron micrographs showing WFA staining along dendritic spines of CA2 pyramidal neurons in the SR (top, red arrowheads), and area CA2 somata in the SP (bottom), and sparse labeling of WFA in area CA1 SP and SR (bottom; scale bar, 2 m).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
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In the paper
“Anti-aggrecan (top), WFA (middle), and neurocan (bottom), indicated in green, label EGFP-expressing CA2 pyramidal neurons and their proximal neurites in red”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
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In the paper
“PNNs first appear in the brain around postnatal day (PN) 14 in the mouse, and gradually increase until they are fully expressed in adulthood, often tracking the end of critical windows of synaptic plasticity (Hensch, 2004;Horii-Hayashi et al., 2015).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Introduction
What the sources say
- Not checked yetcites Hensch, 2004
- Not checked yetcites Horii-Hayashi et al., 2015
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“presence and absence of PNNs in the visual cortex and the closure and reopening of ocular dominance plasticity, strongly supporting a role for PNNs in limiting experience-dependent plasticity.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Significance Statement
What the sources say
- Not checked yetcites Pizzorusso et al. (2002)
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In the paper
“Similarly, decreasing activity via dark rearing delays and attenuates PNN development around inhibitory neurons in the visual cortex (Pizzorusso et al., 2002).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Significance Statement
What the sources say
- Not checked yetcites Pizzorusso et al., 2002
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“CA2 neurons receive excitatory synapses from the dentate gyrus, entorhinal cortex, and area CA3, and it is these synapses from CA3 in CA2 stratum radiatum (SR) that are highly resistant to plasticity (Zhao et al., 2007;Chevaleyre and Siegelbaum, 2010;Kohara et al., 2014).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Significance Statement
What the sources say
- Not checked yetcites Zhao et al., 2007
- Not checked yetcites Chevaleyre and Siegelbaum, 2010
- Not checked yetcites Kohara et al., 2014
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“Interestingly, the onset of PNN expression requires normal early experience in several brain regions, including motor, visual, and somatosensory systems, and similar manipulations performed in adulthood were without effect (Kalb and Hockfield, 1988;Guimara ˜es et al., 1990;Kind et al., 1995;Lander et al., 1997;McRae et al., 2007).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Introduction
What the sources say
- Not checked yetcites Kalb and Hockfield, 1988
- Not checked yetcites Guimara ˜es et al., 1990
- Not checked yetcites Kind et al., 1995
plaindirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Degradation of PNNs around inhibitory neurons in culture (mainly PV-positive interneurons) with the bacterial enzyme chondroitinase (ChABC) increases interneuron excitability without affecting the number or distribution of perisomatic GABAergic presynaptic terminals (Dityatev et al., 2007), suggesting that PNNs function to regulate inhibitory neuron activity.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Significance Statement
What the sources say
- Not checked yetcites Dityatev et al., 2007
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Hippocampal area CA2, positioned between areas CA3 and CA1, has recently been appreciated as an important module of the hippocampus that is molecularly distinct from its neighboring areas (Caruana et al., 2012;Dudek et al., 2016).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Significance Statement
What the sources say
- Not checked yetcites Caruana et al., 2012
- Not checked yetcites Dudek et al., 2016
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Although components of the extracellular matrix (ECM) have been implicated in promoting synaptic plasticity, perineuronal nets (PNNs), a specialized form of ECM typically found around inhibitory neurons, are thought to inhibit plasticity (Dityatev et al., 2010).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Introduction
What the sources say
- Not checked yetcites Dityatev et al., 2010
hedgedconditional- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“All procedures were approved by National Institute of Environmental Health Sciences, University of Alabama at Birmingham, and University of North Carolina Animal Care and Use Committees and were in accordance with the National Institutes of Health guidelines for care and use of animals.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Animals.
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“ECM, restrict plasticity at the normally plasticity-resistant synapses in CA2 SR.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
plaindirect- Asserted: passes — The authors put it forward as true.
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The quote only partly states this claim.
- Rests on an assumption: The LTP facilitation seen after ChABC in CA2 comes from perineuronal net breakdown, not from other chondroitinase-sensitive extracellular matrix components.
In the paper
“adult mice exposed to enrichment had reduced PNN density around cerebellar neurons (Foscarin et al., 2011)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Foscarin et al., 2011
plaindirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“PNN proteoglycans are highly negatively charged and may function to buffer and restrict diffusion of cations like calcium (Bru ¨ckner et al., 1993).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Bru ¨ckner et al., 1993
hedgedconditional- Asserted: passes — The authors put it forward as true.
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In the paper
“other studies have reported a decrease in PNNs after exposure to enrichment in adulthood in several brain regions”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Sale et al., 2007
- Not checked yetcites Foscarin et al., 2011
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Both receptors are known to be coupled to the”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
boostedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“These findings are consistent with a previous report showing that ChABC treatment had no effect on basal synaptic transmission and pairedpulse facilitation in area CA1.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
What the sources say
- Not checked yetcites Bukalo et al., 2001
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“amblyopic rats exposed to EE in adulthood had reduced PNNs in the visual cortex (Sale et al., 2007)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Sale et al., 2007
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“PNN degradation can reopen critical windows of plasticity in a number of brain regions”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Pizzorusso et al., 2002
- Not checked yetcites Gogolla et al., 2009
- Not checked yetcites Romberg et al., 2013
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
In the paper
“neurons in area CA2 have long been noted as being remarkably resistant to cell death from both seizure and ischemic insult (Sloviter et al., 1991)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Sloviter et al., 1991
boosted- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
In the paper
“which has been implicated in the signaling pathway suppressing LTP in area CA2 (Lee et al., 2010)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Lee et al., 2010
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“For each experiment, fluorescence quantification was repeated and analyzed with the experimenter blinded to condition.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Electron microscopy.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“Series and input resistances were monitored by measuring the response to a 10 mV step at each sweep and cells were included for analysis if Ͻ25% change in series and input resistance was detected.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Electron microscopy.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“However, in contrast to our work in area CA2, PNN degradation resulted in an attenuation of LTP in area CA1 (Bukalo et al., 2001).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > PNNs suppress synaptic plasticity in area CA2
What the sources say
- Not checked yetcites Bukalo et al., 2001
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“indicating that CA2 synapses have the cellular machinery for LTP, but that calcium is limited in the postsynaptic neurons (Simons et al., 2009)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Simons et al., 2009
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Raising external calcium levels to 4 mM is insufficient to induce LTP in area CA2, but the very high concentration of 10 mM allows for the induction of LTP”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Simons et al., 2009
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“PNNs have also been linked to neuroprotective functions (Morawski et al., 2004;Cabungcal et al., 2013)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Morawski et al., 2004
- Not checked yetcites Cabungcal et al., 2013
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“CA2 neurons express high levels of the protein regulator of G-protein signaling 14 (RGS 14 )”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Lee et al., 2010
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“PNN degradation modestly disrupts LTP in area CA1 (Bukalo et al., 2001)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Bukalo et al., 2001
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“This study was repeated in a separate cohort of animals and the data combined by normalizing each mean gray value-the value for the PN 45 enriched cohort-for each study.”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Electron microscopy.
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Developmental regulation of PNNs in area CA2 may therefore represent a therapeutic target for PNNassociated developmental disorders, such as schizophrenia and Rett syndrome”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Belichenko et al., 1997
- Not checked yetcites Mauney et al., 2013
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: fails — It may be too vague to check against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Considering that the development of PNNs in hippocampal area CA2 pyramidal neurons parallels the expression timeline of PNNs located in brain regions noted for their critical periods for plasticity”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“recent reports implicate area CA2 in social aggression and social recognition memory (Hitti and Siegelbaum, 2014;Pagani et al., 2015;Smith et al., 2016)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Hitti and Siegelbaum, 2014
- Not checked yetcites Pagani et al., 2015
- Not checked yetcites Smith et al., 2016
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“early postnatal enrichment was found to rescue PNN development in dark-reared animals and promote critical period maturation (Bartoletti et al., 2004)”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
What the sources say
- Not checked yetcites Bartoletti et al., 2004
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“our data raise the intriguing possibility of an early critical period for synaptic plasticity in area CA2”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
The paper relies on this without support.
The LTP facilitation seen after ChABC in CA2 comes from perineuronal net breakdown, not from other chondroitinase-sensitive extracellular matrix components.
“We also found that disruption of PNNs allows synaptic potentiation of normally plasticity-resistant excitatory CA2 synapses; thus, PNNs play a role in restricting synaptic plasticity in area CA2.”
The paper relies on this without support.
WFA staining intensity reliably measures the amount or development of perineuronal nets in CA2.
“WFA staining intensity was quantified using measures of pixel luminescence value on ImageJ software (National Institutes of Health) using a region of interest (ROI) contoured around either CA2 pyramidal neurons in stratum pyramidale (SP) or CA2 apical dendrites using GFP fluorescence in a dorsal hippocampal section.”
The paper relies on this without support.
Aggrecan mRNA in CA2 pyramidal cells means those cells synthesize aggrecan protein.
“Localization of aggrecan mRNA in the pyramidal cell layer of area CA2 indicates that CA2 pyramidal neurons synthesize at least one major component of the PNN matrix, identifying such neurons as a novel source of PNNs in the hippocampus.”
The paper relies on this without support.
Perineuronal nets are mainly found on inhibitory neurons in the neocortex.
“Because PNNs are predominantly associated with inhibitory neurons in the neocortex”
The paper relies on this without support.
ChABC's effect on synaptic plasticity is caused by PNN degradation rather than by other actions of the enzyme.
“Plasticity of EPSC amplitudes is enhanced in CA2 neurons treated with the PNN-degrading enzyme ChABC, compared with untreated controls.”
The paper relies on this without support.
The markers WFA, aggrecan, and neurocan specifically label perineuronal nets.
“PNN markers surround CA2 pyramidal neurons and dendrites.”
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?
The source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“All the social memory associated regions are highly populated with PNN-surrounded PV neurons [46, 66, 130, 131].”
From The extracellular matrix and perineuronal nets in memory (2022) · cites this paper as [66]
Experimental — accuracy not yet measuredThe cited source says what the claim attributes to it.
In the citing paper
“For example, in the CA2, an area associated with social memory and which usually does not exhibit LTP, PNN depletion enables LTP [66].”
From The extracellular matrix and perineuronal nets in memory (2022) · cites this paper as [66]
The cited source says what the claim attributes to it.
In the citing paper
“While PNNs predominantly enwrap PV inhibitory neurons in most brain areas, they surround excitatory neurons in the CA2 [66, 71].”
From The extracellular matrix and perineuronal nets in memory (2022) · cites this paper as [66]
The cited source says what the claim attributes to it.
In the citing paper
“PNNs are usually associated with restriction of synaptic plasticity on inhibitory PV neurons, but in CA2, the PNNs also suppress LTP in excitatory synapses on pyramidal neurons [66].”
From The extracellular matrix and perineuronal nets in memory (2022) · cites this paper as [66]
The cited source says what the claim attributes to it.
In the citing paper
“PNNs are usually associated with restriction of synaptic plasticity on inhibitory PV neurons, but in CA2, the PNNs also suppress LTP in excitatory synapses on pyramidal neurons [66].”
From The extracellular matrix and perineuronal nets in memory (2022) · cites this paper as [66]
In the citing paper
“it has recently been reported that PNNs are highly enriched in the CA2 region of the murine hippocampus33.”
From Elimination of the four extracellular matrix molecules tenascin-C, tenascin-R, brevican and neurocan alters the ratio of excitatory and inhibitory synapses (2019) · cites this paper as [33]
In the citing paper
“it has recently been demonstrated that pyramidal neurons in the CA2 region are a major source of perineuronal nets in the hippocampus and positive for WFA and aggrecan33.”
From Elimination of the four extracellular matrix molecules tenascin-C, tenascin-R, brevican and neurocan alters the ratio of excitatory and inhibitory synapses (2019) · cites this paper as [33]
In the citing paper
“A recent study demonstrated that especially the CA2 excitatory pyramidal neurons are surrounded by PNNs.”
From Elimination of the four extracellular matrix molecules tenascin-C, tenascin-R, brevican and neurocan alters the ratio of excitatory and inhibitory synapses (2019) · cites this paper as [33]
In the citing paper
“When mice were kept in an enriched environment, the PNN expression was enhanced, indicating a correlation between PNN development and environmental stimuli within the juvenile brain33.”
From Elimination of the four extracellular matrix molecules tenascin-C, tenascin-R, brevican and neurocan alters the ratio of excitatory and inhibitory synapses (2019) · cites this paper as [33]
In the citing paper
“A prominent PNN-positive area was found in the CA2 region of the hippocampus33.”
From Elimination of the four extracellular matrix molecules tenascin-C, tenascin-R, brevican and neurocan alters the ratio of excitatory and inhibitory synapses (2019) · cites this paper as [33]
In the citing paper
“In the CA2 subregion, however, PNNs also enwrap excitatory pyramidal cells (Carstens et al., 2016).”
From Adult neurogenesis, glia, and the extracellular matrix (2019) · cites this paper as Carstens et al., 2016
In the citing paper
“but increases them in the CA2 region of the hippocampus (33)”
From Perineuronal nets, inhibitory interneurons and anxiety-related ventral hippocampal neuronal oscillations are altered by early life adversity (2019) · cites this paper as [33]
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
“A two-way ANOVA for the two conditions at different ages indicated significant main effects of age (F (2,33) ϭ 32.33), condition (F (2,33) ϭ 11.72), and interaction (F (2,33) ϭ 3.338).”
§ Perineuronal Nets Suppress Plasticity of Excitatory Synapses on CA2 Pyramidal Neurons > Discussion
“A two-way ANOVA for the two conditions at different ages indicated significant main effects of age (F (2,33) ϭ 69.55), condition (F (1,33) ϭ 11.93), and interaction (F (2,33) ϭ 3.940).”
Citations
Cites
Imaging Neuronal Subsets in Transgenic Mice Expressing Multiple Spectral Variants of GFP
Feng, G., Mellor, R. H., Bernstein, M., Keller-Peck, C., Nguyen, Q. T., Wallace, M., Nerbonne, J. M., Lichtman, J. W., & Sanes, J. R. (2000) · Neuron · doi:10.1016/s0896-6273(00)00084-2 · Cited by 3,042
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The hippocampal CA2 region is essential for social memory
Hitti, F. L., & Siegelbaum, S. A. (2014) · Nature · doi:10.1038/nature13028 · Cited by 976
Perineuronal Nets Protect Fear Memories from Erasure
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The dual role of the extracellular matrix in synaptic plasticity and homeostasis
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Brain extracellular matrix affects AMPA receptor lateral mobility and short-term synaptic plasticity
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Cell type–specific genetic and optogenetic tools reveal hippocampal CA2 circuits
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Perineuronal nets protect fast-spiking interneurons against oxidative stress
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Cited by
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Items 101–200 of 271
Lehner, A., Hoffmann, L., Rampp, S., Coras, R., Paulsen, F. P., Frischknecht, R., Hamer, H. M., Walther, K., Brandner, S., Hofer, W., et al. (2024) · Epilepsia Open · doi:10.1002/epi4.12963 · Cited by 23
Structural Changes Observed in the Piriform Cortex in a Rat Model of Pre-motor Parkinson’s Disease
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Stöber, T. M., Lehr, A. B., Hafting, T., Kumar, A., & Fyhn, M. (2020) · Hippocampus · doi:10.1002/hipo.23256 · Cited by 20
Benoy, A., Ibrahim, M. Z. B., Behnisch, T., & Sajikumar, S. (2021) · Journal of Neuroscience · doi:10.1523/jneurosci.2885-20.2021 · Cited by 20
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Self-citationSex and region-specific effects of high fat diet on PNNs in obesity susceptible rats
Dingess, P. M., Zhang, Z., Sorg, B. A., Ferrario, C. R., & Brown, T. E. (2020) · Physiology & Behavior · doi:10.1016/j.physbeh.2020.112963 · Cited by 24
Development of the hippocampal CA2 region and the emergence of social recognition
Diethorn, E. J., & Gould, E. (2023) · Developmental Neurobiology · doi:10.1002/dneu.22919 · Cited by 23
Carstens, K. E., Gloss, B. R., Alexander, G. M., & Dudek, S. M. (2020) · European Journal of Neuroscience · doi:10.1111/ejn.15050 · Cited by 19
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Role of A1 receptor-activated GIRK channels in the suppression of hippocampal seizure activity
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Parvalbumin interneuron-derived tissue-type plasminogen activator shapes perineuronal net structure
Lépine, M., Douceau, S., Devienne, G. F., Prunotto, P., Lenoir, S., Regnauld, C., Pouettre, E., Piquet, J., Lebouvier, L., Hommet, Y., et al. (2022) · BMC Biology · doi:10.1186/s12915-022-01419-8 · Cited by 21
Hippocampal area CA2: interneuron disfunction during pathological states
Piskorowski, R. A., & Chevaleyre, V. (2023) · Frontiers in Neural Circuits · doi:10.3389/fncir.2023.1181032 · Cited by 22
Perineuronal Nets Alterations Contribute to Stress-Induced Anxiety-Like Behavior
Li, X., Ren, D., Luo, B., Liu, Z., Li, N.-Y., Zhou, T. G., & Fei, E. (2023) · Molecular Neurobiology · doi:10.1007/s12035-023-03596-1 · Cited by 20
Grødem, S., Thompson, E. H., Røe, M. B., Vatne, G. H., Nymoen, I., Buccino, A. P., Otterstad, T., Hafting, T., Fyhn, M., & Lensjø, K. K. (2025) · Molecular Psychiatry · doi:10.1038/s41380-025-02894-5 · Cited by 21
Building a pathway to recovery: Targeting ECM remodeling in CNS injuries
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Nonneuronal contributions to synaptic function
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Alteration of hippocampal CA2 plasticity and social memory in adult rats impacted by juvenile stress
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Do Perineuronal Nets Stabilize the Engram of a Synaptic Circuit?
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Metadata
- Authors
- Carstens, K. E., Phillips, M. L., Pozzo-Miller, L., Weinberg, R. J., & Dudek, S. M.
- Affiliations
- University of North Carolina at Chapel Hill (US)
- Civitan International (US)
- National Institutes of Health (US)
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2016-06-08
- Volume
- 36
- Issue
- 23
- Pages
- 6312-6320
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC4899529
- OpenAlex
- W2410908538
- Semantic Scholar
- e25a3569c55d49e43bcaa10c5b535c9d4b82ebca
- Cited by
- 271
- References
- 52
- Citations by year
- 2026: 23 · 2025: 39 · 2024: 40 · 2023: 25 · 2022: 21 · 2021: 23 · 2020: 27 · 2019: 23 · 2018: 21 · 2017: 22 · 2016: 6
- FWCI
- 13.46
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Journal of Neuroscience (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Carolina Digital Repository (University of North Carolina at Chapel Hill) (submittedVersion)
- UNC Libraries
- Funders
- National Institutes of Health
- National Institute of Neurological Disorders and Stroke (R01NS039444, R01NS065027)
- National Institute of Environmental Health Sciences (ES100221, Z01ES100221)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Life Sciences › Neuroscience › Developmental Neuroscience › Neurogenesis and neuroplasticity mechanisms
- Keywords
- perineuronal nets, excitatory synapses, synaptic plasticity, hippocampal CA2, pyramidal neurons, aggrecan, extracellular matrix, long-term potentiation, postnatal development, rearing conditions, neurodevelopmental disorders, schizophrenia, Rett syndrome, learning and memory
- MeSH terms
- Animals, Animals, Newborn, Extracellular Matrix, Female, Glutamate Decarboxylase, Male, Membrane Proteins, Mice, Inbred C57BL, Mice, Transgenic, Nerve Net, Nerve Tissue Proteins, Extracellular Matrix Proteins, Pyramidal Cells, Gene Expression Regulation, Developmental, Excitatory Amino Acid Agents, Excitatory Postsynaptic Potentials, Satellite Cells, Perineuronal, Green Fluorescent Proteins, Mice, CA2 Region, Hippocampal
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