Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex
Open accessAbstract
Inhibitory interneurons represent 10–15% of the neurons in the somatosensory cortex, and their activity powerfully shapes sensory processing. Three major groups of GABAergic interneurons have been defined according to developmental, molecular, morphological, electrophysiological, and synaptic features. Dendritic-targeting somatostatin-expressing interneurons (SST-INs) have been shown to display diverse morphological, electrophysiological, and molecular properties and activity patterns in vivo. However, the correlation between these properties and SST-IN subtype is unclear. In this study, we aimed to correlate the morphological diversity of layer 5 (L5) SST-INs with their electrophysiological and molecular diversity in mice of either sex. Our morphological analysis demonstrated the existence of three subtypes of L5 SST-INs with distinct electrophysiological properties: T-shaped Martinotti cells innervate L1, and are low-threshold spiking; fanning-out Martinotti cells innervate L2/3 and the lower half of L1, and show adapting firing patterns; non-Martinotti cells innervate L4, and show a quasi-fast spiking firing pattern. We estimated the proportion of each subtype in L5 and found that T-shaped Martinotti, fanning-out Martinotti, and Non-Martinotti cells represent ∼10, ∼50, and ∼40% of L5 SST-INs, respectively. Last, we examined the connectivity between the three SST-IN subtypes and L5 pyramidal cells (PCs). We found that L5 T-shaped Martinotti cells inhibit the L1 apical tuft of nearby PCs; L5 fanning-out Martinotti cells also inhibit nearby PCs but they target the dendrite mainly in L2/3. On the other hand, non-Martinotti cells inhibit the dendrites of L4 neurons while avoiding L5 PCs. Our data suggest that morphologically distinct SST-INs gate different excitatory inputs in the barrel cortex. SIGNIFICANCE STATEMENT Morphologically diverse layer 5 SST-INs show different patterns of activity in behaving animals. However, little is known about the abundance and connectivity of each morphological type and the correlation between morphological subtype and spiking properties. We demonstrate a correlation between the morphological and electrophysiological diversity of layer 5 SST-INs. Based on these findings we built a classifier to infer the abundance of each morphological subtype. Last, using paired recordings combined with morphological analysis, we investigated the connectivity of each morphological subtype. Our data suggest that, by targeting different cell types and cellular compartments, morphologically diverse SST-INs might gate different excitatory inputs in the mouse barrel cortex.
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TL;DR
Layer 5 somatostatin-expressing interneurons (SST-INs) in mouse barrel cortex comprise three morphologically distinct subtypes—T-shaped Martinotti, fanning-out Martinotti, and non-Martinotti—with correlated firing patterns and different postsynaptic targets. These subtypes are estimated to make up roughly 10%, 50%, and 40% of L5 SST-INs, respectively, and they connect to distinct dendritic compartments or cell types.
Key findings
- Morphology correlated with firing pattern: T-shaped Martinotti cells target L1 (50 ± 3% of ascending axonal length and 56 ± 4% of nodes in L1) and show low-threshold spiking with rebound spiking in 7 of 8 cells; fanning-out Martinotti cells target L2/3 (65.8 ± 6% of ascending axon and 71 ± 8% of nodes) and show adapting firing; non-Martinotti cells target L4 (87.6 ± 4% of ascending axon and 92 ± 3% of nodes) and show stuttering in all 8 cells, with fast biphasic AHP in 7 of 8.
- A linear discriminant classifier estimated that in the SST-cre line, fanning-out Martinotti cells represent 50.8% of L5 SST-INs, T-shaped Martinotti cells 13.8%, and non-Martinotti cells 35.4% (abstract rounds these to ~10%, ~50%, and ~40%). The SST/CR line was enriched for fanning-out Martinotti cells (88.9%), while the SST/Calb line was enriched for T-shaped Martinotti cells (61.5%).
- Both Martinotti subtypes innervated L5 pyramidal cells with similar probabilities (fanning-out: 53.6%, 9 of 16; T-shaped: 40%, 4 of 10), but their apical dendritic contact distributions differed significantly (p = 0.033): fanning-out synapses were mainly in L2/3, 150–300 µm below the pia, whereas T-shaped contacts showed a bimodal distribution with a peak in L4 (~450 µm below pia) and a second peak in L1/L2-3 border.
- Non-Martinotti cells did not connect to L5 pyramidal cells (0 of 13) but showed high connection probability with L4 spiny stellate cells (83.3%, 5 of 6), with putative contacts located on dendrites.
Methods
- Whole-cell patch-clamp recordings in acute barrel cortex slices from P20–P40 mice of either sex, using SST-cre, SST/CR, and SST/Calb transgenic lines; biocytin filling followed by confocal reconstruction.
- Reconstructed 30 SST-INs and analyzed 11 electrophysiological parameters in 29 morphologically identified cells; used principal component analysis, hierarchical clustering, and a linear discriminant classifier to classify 76 additional neurons.
- Performed dual and triple paired recordings (39 pairs) between L5 SST-INs and nearby L5 pyramidal cells or L4 spiny stellate cells, combined with morphological analysis of putative synaptic contacts.
Limitations
- Stated: slicing artifacts can affect reconstructions, so the authors applied quality criteria and unsupervised clustering; current molecular markers are insufficient, as SST/CR labels only part of the fanning-out population and SST/Calb labels a heterogeneous population.
- Evident: sample sizes are small for some subtypes (30 reconstructed cells, 29 electrophysiologically analyzed, 39 paired recordings), and the classifier was trained on 20 of 29 cells and made two errors in a 21-cell control group.
- Evident: all data come from in vitro slices, so connectivity and firing properties may not fully reflect in vivo activity or circuit behavior.
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Claims the paper makes
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- Findingown data
Intersectional genetics can be used to obtain mouse lines where the expression of fluorescent proteins in specific morphological types is enriched.
“We found that intersectional genetics can be used to obtain mouse lines where the expression of fluorescent proteins in specific morphological types is enriched.”
Quote in the paperSubstantial claim - Findingown data
Very little (2.2%) overlap of CR and Calb exists in the SST-cre line, especially in deep layers.
“We found very little (2.2%) overlap of CR and Calb in the SST-cre line specially in deep layers”
Quote in the paperSubstantial claim - Findingown data
In both intersectional lines, the majority of tdTomato-expressing cells coexpressed either CR or Calb.
“In both intersectional lines the majority of tdTomatoexpressing cells coexpressed either CR or Calb”
Quote in the paperSubstantial claim - Findingown data
T-shaped Martinotti cells represent approximately 10% (13.8% in the SST-cre line) of L5 SST-INs.
“T-shaped Martinotti, fanning-out Martinotti, and Non-Martinotti cells represent ϳ10, ϳ50, and ϳ40% of L5 SST-INs, respectively.”
- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
“fanning-out Martinotti cells represent 50.8% of labeled cells in L5, T-Shaped cells represent 13.8% of labeled cells, and non-Martinotti represent 35.4% of labeled cells”
Quote in the paperSubstantial claim - Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
- Findingown data
The ascending axon of each morphological group exhibited a specific layer distribution of relative axonal length and proportion of axonal nodes.
“The ascending axon of each morphological group exhibited a specific layer distribution of relative axonal length and proportion of axonal nodes”
Quote in the paperSubstantial claim
- Findingown data
Fanning-out Martinotti cells innervate L2/3 and the lower half of L1.
“fanning-out Martinotti cells innervate L2/3 and the lower half of L1, and show adapting firing patterns”
Quote in the paperSubstantial claim - Findingown data
Non-Martinotti cells innervate L4.
“non-Martinotti cells innervate L4, and show a quasi-fast spiking firing pattern”
- Also stated in: § Morphological diversity of L5 SST interneurons · plain
“Non-Martinotti cells are characterized by their specific innervation of L4.”
Quote in the paperSubstantial claim - Also stated in: § Morphological diversity of L5 SST interneurons · plain
- Findingown data
In the SST/Calb mouse line, the proportion of tdTomato-expressing cells in L5b (~67% of SST-INs) is larger than in L5a (~44% of SST-INs).
“The proportion of tdTomato-expressing cells in L5b (ϳ67% of SST-INs) was larger than in L5a (ϳ44% of SST-INs”
Quote in the paperSubstantial claim - Findingown data
T-shaped Martinotti cells are low-threshold spiking.
“T-shaped Martinotti cells innervate L1, and are low-threshold spiking”
- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
“7 of 8 T-shaped cells showed rebound spiking and a LTS firing pattern”
Quote in the paperSubstantial claim - Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
- Findingown data
Non-Martinotti cells represent approximately 40% (35.4% in the SST-cre line) of L5 SST-INs.
“T-shaped Martinotti, fanning-out Martinotti, and Non-Martinotti cells represent ϳ10, ϳ50, and ϳ40% of L5 SST-INs, respectively.”
- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
“fanning-out Martinotti cells represent 50.8% of labeled cells in L5, T-Shaped cells represent 13.8% of labeled cells, and non-Martinotti represent 35.4% of labeled cells”
Quote in the paperSubstantial claim - Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
- Findingown data
Principal component analysis of 16 morphological axonal parameters divided the dataset into three morphological groups.
“A principal component analysis of 16 morphological axonal parameters of the reconstructed cells showed that the whole dataset could be divided in three morphological groups in the principal component space”
Quote in the paperSubstantial claim - Findingown data
T-shaped Martinotti cells innervate L1.
“T-shaped Martinotti cells innervate L1, and are low-threshold spiking”
- Also stated in: § Morphological diversity of L5 SST interneurons · plain
“T-shaped Martinotti cells specifically target L1, where they have 50 Ϯ 3% of their ascending axonal length, and 56 Ϯ 4% of their ascending axonal nodes”
Quote in the paperSubstantial claim - Also stated in: § Morphological diversity of L5 SST interneurons · plain
- Findingown data
Fanning-out Martinotti cells represent approximately 50% (50.8% in the SST-cre line) of L5 SST-INs.
“T-shaped Martinotti, fanning-out Martinotti, and Non-Martinotti cells represent ϳ10, ϳ50, and ϳ40% of L5 SST-INs, respectively.”
- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
“fanning-out Martinotti cells represent 50.8% of labeled cells in L5, T-Shaped cells represent 13.8% of labeled cells, and non-Martinotti represent 35.4% of labeled cells”
Quote in the paperSubstantial claim - Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
- Findingown data
L5 T-shaped Martinotti cells inhibit the L1 apical tuft of nearby pyramidal cells.
“We found that L5 T-shaped Martinotti cells inhibit the L1 apical tuft of nearby PCs”
Quote in the paperSubstantial claim - Findingown data
The proportion of YFP cells expressing CR and Calb was higher in L2/3 (15.9%).
“However the proportion of YFP cells expressing CR and Calb was higher in L2/3 (15.9%”
Quote in the paperNot a substantial claim - Findingown data
Fanning-out Martinotti cells show adapting firing patterns.
“fanning-out Martinotti cells innervate L2/3 and the lower half of L1, and show adapting firing patterns”
- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
“The majority of fanning-out Martinotti cells showed an adapting firing pattern characterized by trains of APs that after an initial adaptation showed regular ISIs”
Quote in the paperSubstantial claim - Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
- Findingown data
Non-Martinotti cells show a quasi-fast spiking firing pattern.
“non-Martinotti cells innervate L4, and show a quasi-fast spiking firing pattern”
- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · hedged
“Non-Martinotti cells showed several features of the quasi-fast spiking phenotype such as a lower IR, a shorter AHP duration , and a higher F max compared with Martinotti cells”
Quote in the paperSubstantial claim - Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · hedged
- Findingown data
Three subtypes of L5 SST-INs exist with distinct electrophysiological properties.
“Our morphological analysis demonstrated the existence of three subtypes of L5 SST-INs with distinct electrophysiological properties: T-shaped Martinotti cells innervate L1, and are low-threshold spiking; fanning-out Martinotti cells innervate L2/3 and the lower half of L1, and show adapting firing patterns; non-Martinotti cells innervate L4, and show a quasi-fast spiking firing pattern.”
- Also stated in: § Morphological diversity of L5 SST interneurons · plain
“A principal component analysis of 16 morphological axonal parameters of the reconstructed cells showed that the whole dataset could be divided in three morphological groups in the principal component space”
Quote in the paperSubstantial claim - Also stated in: § Morphological diversity of L5 SST interneurons · plain
- Findingown data
In vitro, the same morphological diversity of SST-INs as observed in vivo by Muñoz et al. (2017) was found.
“We found in vitro the same morphological diversity observed by Mun ˜oz et al. (2017) in vivo and were able to correlate morphological subtypes to the diverse firing patterns observed in the SST-INs population (Ma et al., 2006).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Muñoz et al., 2017
Not checked in the free run.
- cites Ma et al., 2006
Not checked in the free run.
- Findingown data
There is no significant difference in the IPSC charge evoked by Martinotti cells onto pyramidal cells (fanning-out Martinotti: 965.6 ± 204.5 nC; T-shaped Martinotti: 1232.6 ± 262.3 nC; p = 0.48, two-sample t test).
“We did not find any significant difference in the IPSC charge evoked by Martinotti cells onto pyramidal cells (fanning-out Martinotti: 965.6 Ϯ 204.5 nC; T-shaped Martinotti: 1232.6 Ϯ 262.3 nC; p ϭ 0.48, two-sample t test).”
Quote in the paperSubstantial claim - Findingown data
7 of 8 non-Martinotti cells showed a fast biphasic AHP.
“7 of 8 cells showed a fast biphasic AHP”
Quote in the paperNot a substantial claim - Findingown data
Fanning-out cells had a significantly higher Fmax than T-shaped Martinotti cells.
“Fanning-out cells had a significantly higher F max , a more depolarized V threshold (Fig. 6C), and a smaller adaptation index compared with T-shaped Martinotti cells”
Quote in the paperSubstantial claim - Findingown data
Only 2 of 14 fanning-out cells showed rebound spiking.
“Only 2 of 14 fanning-out cells showed rebound spiking”
Quote in the paperNot a substantial claim - Findingown data
The rising phase of the action potential was faster in non-Martinotti cells than in Martinotti cells.
“the rising phase of the AP was faster than that of Martinotti cells”
Quote in the paperNot a substantial claim - Findingown data
Non-Martinotti cells are not connected to L5 pyramidal cells (0 of 13).
“Our dual and triple recordings showed that these cells are not connected to L5 pyramidal cells (0 of 13).”
Quote in the paperSubstantial claim - Findingown data
Fanning-out and T-shaped Martinotti interneurons differed in their electrophysiological properties.
“We found that fanning-out and T-shaped Martinotti interneurons differed in their electrophysiological properties”
Quote in the paperSubstantial claim - Findingown data
T-shaped Martinotti cells showed the lowest maximal firing frequency (Fmax).
“T-shaped Martinotti showed the lowest F max”
Quote in the paperNot a substantial claim - Findingown data
The distribution of contact points on the apical dendrites of L5 pyramidal neurons is significantly different between the two types of Martinotti cells (p = 0.033, two-sample Kolmogorov-Smirnov test).
“We measured the distance from the pia of the contact points located on the apical dendrites of L5 pyramidal neurons, and found that the distribution of contact points was significantly different between the two types of Martinotti cells ( p ϭ 0.033, two-sample Kolmogorov-Smirnov test; Fig. 7 F, G).”
Quote in the paperSubstantial claim- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
- Findingown data
These morphological subtypes target different layers of the cortical column.
“These morphological subtypes target different layers of the cortical column (Fig. 4) and are active during different behavioral states (Mun ˜oz et al., 2017).”
Quote in the paperSubstantial claim - Findingown data
Fanning-out Martinotti cells had most of their synapses located in L2/3, between 150 and 300 μm below the pia.
“Indeed while fanning-out Martinotti cells had most of their synapses located in L2/3, between 150 and 300 m below the pia (Fig. 7 A, F )”
Quote in the paperSubstantial claim- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
- Findingown data
The morphological diversity of SST-INs in L5 correlates with their spike properties and repetitive firing characteristics.
“In the present study we show that the morphological diversity of SST-INs in L5 correlates with their spike properties and repetitive firing characteristics.”
Quote in the paperSubstantial claim - Findingown data
7 of 8 T-shaped cells showed rebound spiking.
“7 of 8 T-shaped cells showed rebound spiking and a LTS firing pattern”
Quote in the paperSubstantial claim - Findingown data
All non-Martinotti cells showed stuttering (n=8).
“We found that all the non-Martinotti showed stuttering (n ϭ 8)”
Quote in the paperSubstantial claim - Findingown data
A substantial proportion (37.7%) of L5 SST-INs was classified as non-Martinotti cells.
“Moreover we found that a substantial proportion (37.7%) of L5 SST-INs was classified as non-Martinotti cells.”
Quote in the paperSubstantial claim- Rests on an assumption: The linear discriminant classifier built from firing properties accurately infers the morphological class of L5 SST-INs.
- Findingown data
Fanning-out Martinotti cells showed intermediate values in most electrophysiological parameters compared with the other two morphological groups.
“In summary, fanning-out Martinotti cells showed intermediate values in most electrophysiological parameters compared with the other two morphological groups.”
Quote in the paperSubstantial claim - Findingcitation
Non-Martinotti cells showed a 'quasi-fast spiking' phenotype, similar to that described in X94 cells.
“We found that non-Martinotti cells showed a “quasi-fast spiking” phenotype, similar to that described in X94 cells (Ma et al., 2006).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ma et al., 2006
Not checked in the free run.
- Findingown data
T-shaped Martinotti cells showed a bimodal distribution of synapses with a peak in L4 (~450 μm below the pia) and a second peak in L1 and at the border between L1 and L2/3.
“T-shaped Martinotti cells showed a bimodal distribution with a peak in L4 (ϳ450 m below the pia) and a second peak in L1 and at the border between L1 and L2/3 (Fig. 7C,G).”
Quote in the paperSubstantial claim- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
- Findingown data
Non-Martinotti cells have a high connection probability with L4 spiny stellate cells (83.3%, 5 of 6).
“Indeed we found a high connection probability of non-Martinotti cells with L4 spiny stellate cells (83.3%, 5 of 6).”
Quote in the paperSubstantial claim - Findingown data
Fanning-out Martinotti cells have 18 ± 4% of their axon reaching the deeper half of L1, where it branches.
“A smaller proportion (18 Ϯ 4%) of the axon of these cells can reach the deeper half of L1, where it branches”
Quote in the paperNot a substantial claim - Findingcitation
In the barrel cortex, L5 contains at least three morphological subtypes of SST-INs.
“In the barrel cortex, L5 contains at least three morphological subtypes of SST-INs (Mun ˜oz et al., 2017;present study).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Muñoz et al., 2017
Not checked in the free run.
- Findingown data
Both types of Martinotti cells innervate L5 pyramidal neurons with similar connection probabilities (fanning-out to pyramidal: 53.6%, 9 of 16; T-shaped to pyramidal: 40%, 4 of 10).
“We found that both types of Martinotti cells innervate L5 pyramidal neurons with similar connection probabilities (fanningout to pyramidal: 53.6%, 9 of 16; T-shaped to pyramidal: 40%, 4 of 10; Fig. 7A-D).”
Quote in the paperSubstantial claim - Findingreasoning
The SST/Calb line labels a population that is still very heterogeneous.
“The SST/CR mouse line labels only part of the fanning-out population in L5, and the SST/Calb labels a population that is still very heterogeneous.”
Quote in the paperSubstantial claim - Findingown data
The SST/CR mouse line labels only part of the fanning-out population in L5.
“The SST/CR mouse line labels only part of the fanning-out population in L5, and the SST/Calb labels a population that is still very heterogeneous.”
Quote in the paperSubstantial claim - Findingown data
No significant correlation was found between the age of the animal and the electrophysiological properties analyzed.
“We did not find any significant correlation between the age of the animal and the electrophysiological properties analyzed here.”
Quote in the paperNot a substantial claim - Findingown data
Fanning-out cells had a more depolarized Vthreshold than T-shaped Martinotti cells.
“Fanning-out cells had a significantly higher F max , a more depolarized V threshold (Fig. 6C), and a smaller adaptation index compared with T-shaped Martinotti cells”
Quote in the paperNot a substantial claim - Findingown data
All putative contact points between non-Martinotti cells and L4 spiny stellate cells were located on dendrites.
“All the putative contact points between non-Martinotti cells and L4 spiny stellate cells were located on dendrites (Fig. 8C,G).”
Quote in the paperNot a substantial claim- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
- Findingown data
Non-Martinotti cells have a very small amount of axon in L1 (0.2 ± 0.2%) that does not branch in this layer.
“Non-Martinotti cells are also characterized by having a very small amount of axon in L1 (0.2 Ϯ 0.2%) that does not branch in this layer.”
Quote in the paperNot a substantial claim - Findingown data
The majority of fanning-out Martinotti cells showed an adapting firing pattern.
“The majority of fanning-out Martinotti cells showed an adapting firing pattern characterized by trains of APs that after an initial adaptation showed regular ISIs”
Quote in the paperNot a substantial claim - Findingown data
Fanning-out cells had a smaller adaptation index than T-shaped Martinotti cells.
“Fanning-out cells had a significantly higher F max , a more depolarized V threshold (Fig. 6C), and a smaller adaptation index compared with T-shaped Martinotti cells”
Quote in the paperNot a substantial claim - Interpretationown data
Morphologically distinct SST-INs gate different excitatory inputs in the barrel cortex.
“Our data suggest that morphologically distinct SST-INs gate different excitatory inputs in the barrel cortex.”
- Also stated in: § Significance Statement · hedged
“Our data suggest that, by targeting different cell types and cellular compartments, morphologically diverse SST-INs might gate different excitatory inputs in the mouse barrel cortex.”
Quote in the paperSubstantial claim - Also stated in: § Significance Statement · hedged
- Interpretationown data
Martinotti and non-Martinotti cells belong to distinct inhibitory circuits within L5.
“These studies showed that Martinotti and non-Martinotti cells belong to distinct inhibitory circuits within L5.”
Quote in the paperSubstantial claim - Interpretationown data
Approximately half of the cells labeled in the SST-CR line are also labeled in the SST-Calb mouse line.
“The amount of YFP IR cells expressing both CR and Calb over the YFP-CR IR population was Ͼ50% throughout layers, indicating that approximately half of the cells labeled in the SST-CR line are also labeled in the SST-Calb mouse line”
Quote in the paperNot a substantial claim - Study design and conductown data
In the SST-cre mouse line, 95.6% of td-Tomato cells coexpress SST.
“SST-cre, where 95.6% of td-Tomato cells were found to coexpress SST”
Quote in the paperSubstantial claim - Study design and conductown data
The linear discriminant classifier performed correctly for 19 of 21 control cells using six electrophysiological parameters.
“The classifier performed correctly for 19 of 21 cells using just six parameters.”
Quote in the paperSubstantial claim - Premisecitation
L2/3 SST-INs show adapting firing patterns resembling that of L5 fanning-out Martinotti.
“Indeed L2/3 SST-INs show adapting firing patterns resembling that of L5 fanning-out Martinotti (Halabisky et al., 2006;Ma et al., 2006;Xu et al., 2013).”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites Halabisky et al., 2006
Not checked in the free run.
- cites Ma et al., 2006
Not checked in the free run.
- cites Xu et al., 2013
Not checked in the free run.
- Interpretationown data
The low Fmax of T-shaped Martinotti cells is likely due to their tendency to reach depolarization block early during the f-I curve.
“likely due to their tendency to reach depolarization block early during the f-I curve”
Quote in the paperNot a substantial claim - Study design and conductown data
For all connections analyzed and presented in this study, there was clear morphological evidence that the apical dendrite of the pyramidal cells was intact and passed through the axonal domain of the SST-INs.
“We point out that for all the connections analyzed and presented in this study we had clear morphological evidence showing the apical dendrite of the pyramidal cells was intact and passed through the axonal domain of the SST-INs.”
Quote in the paperSubstantial claim - Interpretationown data
Slicing artifacts can be excluded as an explanation of the observed lack of connectivity of non-Martinotti cells and L5 pyramidal cells.
“Thus, we can exclude slicing artifacts as an explanation of the observed lack of connectivity of non-Martinotti cells and L5 pyramidal cells (Fig. 8 A, C).”
Quote in the paperSubstantial claim - Interpretationcitation
The electrophysiological properties that define the three groups of SST-INs described in L5 seem to be differentially distributed across layers.
“The electrophysiological properties that define the three groups of SST-INs described in L5 seem to be differentially distributed across layers.”
Quote in the paperNot a substantial claim - Premisecitation
These morphological subtypes are active during different behavioral states.
“These morphological subtypes target different layers of the cortical column (Fig. 4) and are active during different behavioral states (Mun ˜oz et al., 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Muñoz et al., 2017
Not checked in the free run.
- Study design and conductreasoning
The performance of the classifier was reliable.
“Given that no other similar error was found in the test group we concluded that the performance of the classifier was reliable.”
Quote in the paperSubstantial claim - Premisecitation
L2/3 SST-INs are Martinotti cells targeting L1.
“L2/3 SST-INs have been shown to be Martinotti cells targeting L1 (Ma et al., 2006;Dumitriu et al., 2007;Xu et al., 2013;Mun ˜oz et al., 2017).”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites Ma et al., 2006
Not checked in the free run.
- cites Dumitriu et al., 2007
Not checked in the free run.
- cites Xu et al., 2013
Not checked in the free run.
- Interpretationown data
Morphologically distinct/diverse L5 SST-INs belong to/participate in different microcircuits in the mouse barrel cortex.
“Last, using paired recordings we show that morphologically distinct SST-INs belong to different microcircuits in the barrel cortex.”
- Also stated in: § Connectivity of SST-INs subtypes · plain
“In conclusion, we provide evidence that morphologically diverse L5 SST-INs participate in different microcircuits in the mouse barrel cortex.”
Quote in the paperSubstantial claim - Also stated in: § Connectivity of SST-INs subtypes · plain
- Interpretationcitation
T-shaped Martinotti cells seem to be the sole SST-IN subtype to provide dendritic inhibition to the apical dendrite of L5 PCs in L4.
“Interestingly, T-shaped Martinotti cells seem to be the sole SST-IN subtype to provide dendritic inhibition to the apical dendrite of L5 PCs in L4, where these cells receive sensory inputs from the ventral posteromedial thalamic nucleus (Petreanu et al., 2009).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Petreanu et al., 2009
Not checked in the free run.
- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
- May combine two claims. 22
- Classified as probably not a substantial claim. 8
May combine two claims.
Findingown dataIn the SST/CR intersectional mouse line, approximately 47% of SST-INs in L2/3, 14% in L5a, and 4% in L5b express tdTomato.
“in the SST/CR intersectional mouse line ϳ47% of SST-INs in L2/3, 14% of SST-INs in L5a, and 4% of SST-INs in L5b express tdTomato”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataNon-Martinotti cells inhibit the dendrites of L4 neurons while avoiding L5 pyramidal cells.
“non-Martinotti cells inhibit the dendrites of L4 neurons while avoiding L5 PCs”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThirty neurons were morphologically reconstructed and analyzed: 8 non-Martinotti, 13 fanning-out Martinotti, and 9 T-shaped Martinotti cells.
“We morphologically reconstructed and analyzed a total of 30 neurons: 19 in the SST-cre, 5 in the SST/CR, and 6 in the SST/ Calb mouse lines.”
- Also stated in: § Morphological diversity of L5 SST interneurons · plain
“8 Non-Martinotti, 13 fanning-out Martinotti, and 9 T-shaped Martinotti cells”
Quote in the paperSubstantial claim- Also stated in: § Morphological diversity of L5 SST interneurons · plain
May combine two claims.
Findingown dataL5 fanning-out Martinotti cells inhibit nearby pyramidal cells but target the dendrite mainly in L2/3.
“L5 fanning-out Martinotti cells also inhibit nearby PCs but they target the dendrite mainly in L2/3”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe SST/CR mouse line targets mainly fanning-out Martinotti cells (88.9% of labeled cells), with none classified as T-shaped and 11.1% as non-Martinotti.
“The SST/CR mouse line targets mainly fanning-out Martinotti cells, that represented 88.9% of labeled cells, whereas none of the cells was classified as T-shaped Martinotti, and non-Martinotti cells 11.1% of labeled cells”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataT-shaped cells showed a high input resistance, a low AP threshold, and a high adaptation index.
“showing a high IR, a low AP threshold , and a high adaptation index”
Quote in the paperSubstantial claimMay combine two claims.
FindingcitationWithin the Martinotti group, fanning-out Martinotti cells showed an adapting type of firing pattern reminiscent of the GIN SST-INs, and T-shaped Martinotti cells showed an LTS firing pattern similar to that observed in some X98 cells.
“Within the Martinotti group, fanning-out Martinotti cells showed an adapting type of firing pattern reminiscent of the GIN SST-INs, and T-shaped Martinotti cells showed an LTS firing pattern similar to that observed in some X98 cells (Ma et al., 2006).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Ma et al., 2006
Not checked in the free run.
May combine two claims.
Findingown dataNon-Martinotti cells had 87.6 ± 4% of their ascending axon and 92 ± 3% of their axonal nodes in L4.
“cells belonging to this morphological group had 87.6 Ϯ 4% of their ascending axon, and 92 Ϯ 3% of their axonal nodes in L4”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataT-shaped Martinotti cells have 50 ± 3% of their ascending axonal length and 56 ± 4% of their ascending axonal nodes in L1.
“T-shaped Martinotti cells specifically target L1, where they have 50 Ϯ 3% of their ascending axonal length, and 56 Ϯ 4% of their ascending axonal nodes”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataThe first three principal components account for 93% of the variance and are mostly represented by features of the ascending axon.
“Moreover the first three principal components, accounting for 93% of the variance, are mostly represented by features of the ascending axon.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataFanning-out Martinotti cells have 65.8 ± 6% of their ascending axon and 71 ± 8% of their nodes in L2/3.
“Fanning-out Martinotti cells are characterized by targeting mainly L2/3, where these cells have 65.8 Ϯ 6% of their ascending axon, and 71 Ϯ 8% of their nodes.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataNon-Martinotti cells showed a lower input resistance, a shorter AHP duration, and a higher Fmax compared with Martinotti cells.
“Non-Martinotti cells showed several features of the quasi-fast spiking phenotype such as a lower IR, a shorter AHP duration , and a higher F max compared with Martinotti cells”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe SST/Calb mouse line showed enrichment for T-shaped Martinotti neurons (61.5% of labeled cells compared with 13.1% in the SST-Cre line), with fanning-out Martinotti cells at 30.8% and non-Martinotti cells at 7.7%.
“The SST/Calb mouse line showed some enrichment for T-shaped Martinotti neurons that represented 61.5% of labeled cells (compared with 13.1% in the SST-Cre line), whereas fanning-out Martinotti cells represented 30.8% of labeled cells, and non-Martinotti cells represented only 7.7% of labeled cells.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataIn the control group, the classifier correctly identified all non-Martinotti cells (n=10), 7 of 8 fanning-out Martinotti cells, and 2 of 3 T-shaped cells.
“In the control group the classifier correctly identified all the non-Martinotti cells (n ϭ 10), 7 of 8 fanning-out Martinotti cells, and 2 of 3 T-shaped cells.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataThe SST/CR line is highly enriched in fanning-out Martinotti cells, while the SST/Calb line is enriched in T-shaped Martinotti cells.
“We found that the SST/CR line is highly enriched in fanning-out Martinotti cells, while the SST/Calb line is enriched in T-shaped Martinotti cells.”
Quote in the paperSubstantial claim- Rests on an assumption: The linear discriminant classifier built from firing properties accurately infers the morphological class of L5 SST-INs.
May combine two claims.
Findingown dataL5 non-Martinotti cells specifically target L4 neurons and avoid L5 pyramidal cells.
“Here we show that L5 non-Martinotti cells specifically target L4 neurons and avoid L5 pyramidal cells.”
Quote in the paperSubstantial claimMay combine two claims.
FindingreasoningT-shaped Martinotti form synapses onto tufts in L1, whereas fanning-out Martinotti target the apical dendrites in L2/3.
“As expected from their axonal distribution, T-shaped Martinotti form synapses onto tufts in L1, whereas fanning-out Martinotti target the apical dendrites in L2/3 (Figs. 4, 7).”
Quote in the paperSubstantial claim- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
May combine two claims.
InterpretationreasoningNeither calretinin nor calbindin are optimal molecular markers for fanning-out and T-shaped Martinotti respectively.
“Indeed, neither calretinin nor calbindin are optimal molecular markers for fanning-out and T-shaped Martinotti respectively.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationSST-INs provide frequency-dependent feedback inhibition to pyramidal cells in supragranular and infragranular layers.
“SST-INs provide frequency-dependent feedback inhibition to pyramidal cells in supragranular and infragranular layers (Kapfer et al., 2007;Silberberg and Markram, 2007).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Kapfer et al., 2007
Not checked in the free run.
- cites Silberberg and Markram, 2007
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIn the neocortical L5, computational and experimental work has shown that Martinotti cells can synchronize pyramidal cells through distal inhibition.
“In the neocortical L5, computational and experimental work has shown that Martinotti cells can synchronize pyramidal cells through distal inhibition (Li et al., 2013;Hilscher et al., 2017).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Li et al., 2013
Not checked in the free run.
- cites Hilscher et al., 2017
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationThis form of inhibition is thought to underlie surround inhibition and gamma-band synchronization in L2/3 of the visual cortex.
“This form of inhibition is thought to underlie surround inhibition, and gammaband synchronization in L2/3 of the visual cortex (Adesnik et al., 2012;Veit et al., 2017).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Adesnik et al., 2012
Not checked in the free run.
- cites Veit et al., 2017
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationLong-range projecting nNos-expressing SST-INs have been described in L6.
“Long-range projecting nNos-expressing SST-INs have been described in L6 as well (He et al., 2016).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites He et al., 2016
Not checked in the free run.
May combine two claims.
PremisecitationL4 SST-INs are quasi-fast spiking L4 targeting non-Martinotti cells.
“On the other hand, L4 SST-INs have been shown to be quasi-fast spiking L4 targeting non-Martinotti cells (Ma et al., 2006;Xu et al., 2013;Mun ˜oz et al., 2017).”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites Ma et al., 2006
Not checked in the free run.
- cites Xu et al., 2013
Not checked in the free run.
- cites Muñoz et al., 2017
Not checked in the free run.
May combine two claims.
PremisecitationL5 non-Martinotti cells tend to show a higher input resistance and seem to be less stuttering.
“However, L5 non-Martinotti tend to show a higher input resistance and seem to be less stuttering (Ma et al., 2006).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Ma et al., 2006
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
InterpretationcitationIt is likely that one or both types of Martinotti cells are also present in L6.
“It is likely that one or both types of Martinotti cells are also present in this layer (Ma et al., 2006;Mun ˜oz et al., 2017).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Ma et al., 2006
Not checked in the free run.
- cites Muñoz et al., 2017
Not checked in the free run.
Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing), S4 (based).
Interpretationnone givenThe actions of different SST-INs subtypes are likely to have important computational roles in sensory perception during different behavioral states.
“The actions of different SST-INs subtypes are likely to have important computational roles in sensory perception during different behavioral states.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationL5 non-Martinotti cells receive thalamic input from the ventral posteromedial nucleus and this input can shift thalamic feedforward inhibition from perisomatic to dendritic.
“Using the X94 mouse line it was shown that L5 non-Martinotti cells receive thalamic input from the ventral posteromedial nucleus and that this input can shift thalamic feedforward inhibition from perisomatic to dendritic (Tan et al., 2008;Hu and Agmon, 2016).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Tan et al., 2008
Not checked in the free run.
- cites Hu and Agmon, 2016
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationL4 non-Martinotti cells have been suggested to disinhibit L4 excitatory cells by primarily targeting L4 parvalbumin interneurons.
“Indeed L4 non-Martinotti cells have been suggested to disinhibit L4 excitatory cells by primarily targeting L4 parvalbumin interneurons (Xu et al., 2013).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Xu et al., 2013
Not checked in the free run.
The paper relies on this without support.
The correlation between morphological, electrophysiological, and molecular properties and SST-IN subtype is unclear.
“However, the correlation between these properties and SST-IN subtype is unclear.”
“However, the relationship between the morphological and electrophysiological diversity remains poorly understood.”
The paper relies on this without support.
Little is known about the specific postsynaptic partners of different morphological subtypes of L5 SST-INs.
“However, little is known about the specific postsynaptic partners of different morphological subtypes of L5 SST-INs.”
The paper relies on this without support.
The output connectivity of non-Martinotti cells has not been studied.
“Moreover, the output connectivity of non-Martinotti cells has not been studied.”
The paper relies on this without support.
It is unknown whether both types of Martinotti cells are connected to L5 PCs and participate in feedback inhibition.
“but it is unknown whether both types of Martinotti cells are connected to L5 PCs and participate in this type of inhibition”
The paper relies on this without support.
The linear discriminant classifier built from firing properties accurately infers the morphological class of L5 SST-INs.
“We used the firing properties of morphologically identified cells to build a linear discriminant classifier to infer the proportion of cells belonging to each morphological class in L5.”
The paper relies on this without support.
Putative synaptic contacts identified morphologically correspond to functional synapses.
“We also quantified the putative synaptic contacts in connected pairs.”
“All the putative contact points between non-Martinotti cells and L4 spiny stellate cells were located on dendrites (Fig. 8C,G).”
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?
Martinotti cells are particularly abundant in layer 5 and common in layers 2 and 3, where they represent approximately 60% of SST+ neurons.
“Martinotti cells are particularly abundant in layer 5, but they are also common in layers 2 and 3. These cells represent approximately 60% of the SST+ neurons in these layers”
From Development and Functional Diversification of Cortical Interneurons (2018) · cites this paper as Nigro et al., 2018
SST+ non-Martinotti cells are found throughout layers 2 to 6, lack axons in layer 1, and have lower resting membrane potential and higher firing frequency than Martinotti cells but lower than PV+ basket cells.
“SST+ non-Martinotti cells are found throughout layers 2 to 6 and characteristically lack axons in layer 1. They exhibit lower resting membrane potential and higher firing frequency than Martinotti cells but lower than PV+ basket cells”
From Development and Functional Diversification of Cortical Interneurons (2018) · cites this paper as Nigro et al., 2018
Three previously described morphological types of L5 Pvalb cells, as well as two morphological types of L5 Martinotti cells, corresponded to different t-types.
“We also showed that three previously described morphological types of L5 Pvalb cells5, as well as two morphological types of L5 Martinotti cells29,30, corresponded to different t-types.”
From Phenotypic variation of transcriptomic cell types in mouse motor cortex (2021) · cites this paper as [30]
In the Sst subclass, morpho-electric properties varied continuously across the transcriptomic landscape, with neighbouring t-types consistently showing similar morphologies and similar rebound values.
“We observed the same in the Sst subclass, which is known to be transcriptomically4 and morpho-electrically29,30,33 diverse in L5. Here we also found that morpho-electric properties varied continuously across the transcriptomic landscape, with neighbouring t-types consistently showing similar morphologies and similar rebound values (Fig. 4b).”
From Phenotypic variation of transcriptomic cell types in mouse motor cortex (2021) · cites this paper as [30]
Recent evidence suggests that different morphological phenotypes correspond to different synaptic connectivity profiles.
“Recent evidence also suggests that different morphological phenotypes correspond to differential synaptic connectivity profiles (Nigro et al., 2018; Naka et al., 2019; Scala et al., 2019).”
From Integrated Morphoelectric and Transcriptomic Classification of Cortical GABAergic Cells (2020) · cites this paper as Nigro et al., 2018
In V1 L4, Martinotti cells did not show the heterogeneity reported for L5 Martinotti cells; none showed rebound firing after stimulation offset.
“Previous literature suggested that Martinotti cells in L5 form two groups with different morphology and/or electrophysiology29,42, but we did not observe such heterogeneity in L4: in particular, none of the MCs showed a rebound firing after stimulation offset42.”
From Layer 4 of mouse neocortex differs in cell types and circuit organization between sensory areas (2019) · cites this paper as [42]
All non-fast-spiking SOM+ cells in V1 L5 and the majority of SOM+ cells in S1 L5 are known to be Martinotti cells.
“Morphologically, all non-fast-spiking SOM+ cells in V1 L529,47 and the majority of SOM+ cells in S1 L522,29,42,47 are known to be MCs.”
From Layer 4 of mouse neocortex differs in cell types and circuit organization between sensory areas (2019) · cites this paper as [42]
A subclass of deep-layer SST-expressing low-threshold spiking (LTS) interneurons shows increased excitability in adult jSI mice compared with group-housed mice.
“We thus screened sub-types of mPFC interneurons expressing somatostatin (SST)12 and parvalbumin11, and found that a subclass of deep-layer SST-expressing low-threshold spiking (LTS) interneurons showed increased excitability in adult jSI mice compared with GH mice (Extended Data Fig. 8).”
From A prefrontal–paraventricular thalamus circuit requires juvenile social experience to regulate adult sociability in mice (2020) · cites this paper as [12]
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.
L5 subtype proportions differ between Results and Discussion
MajorThe Results report 50.8% fanning-out, 13.8% T-shaped, and 35.4% non-Martinotti cells in the SST-cre line. The Discussion reports 49.2%, 13.1%, and 37.7%, respectively, for the same L5 population; these differences exceed rounding.
“fanning-out Martinotti cells represent 50.8% of labeled cells in L5, T-Shaped cells represent 13.8% of labeled cells, and non-Martinotti represent 35.4% of labeled cells”
“Specifically, 49.2% were classified as fanning-out and 13.1% were classified as T-shaped.Moreover we found that a substantial proportion (37.7%) of L5 SST-INs was classified as non-Martinotti cells.”
Fanning-out connection probability does not match the pair count
MinorNine connected pairs out of 16 give 56.25%, not 53.6%.
Reported 53.6 · recomputed 56.25
“fanningout to pyramidal: 53.6%, 9 of 16”
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Self-citationGABAergic cell subtypes and their synaptic connections in rat frontal cortex
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“Electrophysiological properties of morphologically identified SST interneurons Layer 5 SST-INs have been previously shown to have heterogeneous spiking properties in slice recordings (Kawaguchi and Kubota, 1997; Wang et al., 2004; Ma et al., 2006).”
BackgroundThe subcellular organization of neocortical excitatory connections
Petreanu, L., Mao, T., Sternson, S. M., & Svoboda, K. (2009) · Nature · doi:10.1038/nature07709 · Cited by 1,047 · cited 1× · see citing works
“A wide morphological diversity has been described in other GABAergic interneuron groups, such as VIP-INs (bipolar and multipolar) and PV-INs (basket, chandelier, translaminar; DeFelipe et al., 1989; Buchanan et al., 2012; Bortone et al., 2014; Prönneke et al., 2015; He et al., 2016).”
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“Using the X94 mouse line it was shown that L5 non-Martinotti cells receive thalamic input from the ventral posteromedial nucleus and that this input can shift thalamic feedforward inhibition from perisomatic to dendritic (Tan et al., 2008; Hu and Agmon, 2016).”
MethodA neural circuit for spatial summation in visual cortex
Adesnik, H., Bruns, W., Taniguchi, H., Huang, Z. J., & Scanziani, M. (2012) · Nature · doi:10.1038/nature11526 · Cited by 702 · cited 1× · see citing works
“This form of inhibition is thought to underlie surround inhibition, and gammaband synchronization in L2/3 of the visual cortex (Adesnik et al., 2012; Veit et al., 2017).”
BackgroundElectrophysiological, transcriptomic and morphologic profiling of single neurons using Patch-seq
Cadwell, C. R., Palasantza, A., Jiang, X., Berens, P., Deng, Q., Yilmaz, M., Reimer, J., Shen, S., Bethge, M., Tolias, K. F., et al. (2016) · Nature Biotechnology · doi:10.1038/nbt.3445 · Cited by 581 · cited 1× · see citing works
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BackgroundTargeting cells with single vectors using multiple-feature Boolean logic
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“Indeed L2/3 SST-INs show adapting firing patterns resembling that of L5 fanning-out Martinotti (Halabisky et al., 2006; Ma et al., 2006; Xu et al., 2013).”
BackgroundInfluentialTermination of afferent axons in macaque striate cortex
Blasdel, G. G., & Lund, J. S. (1983) · Journal of Neuroscience · doi:10.1523/jneurosci.03-07-01389.1983 · Cited by 452 · cited 1× · see citing works
“Axonal nodes represent branching points and have been previously used as a measure of axonal targeting (Blasdel and Lund, 1983; Florence and Casagrande, 1987; Freund et al., 1989; Lee et al., 2005; Yamada et al., 2010).”
MethodDistinct Subtypes of Somatostatin-Containing Neocortical Interneurons Revealed in Transgenic Mice
Ma, Y., Hu, H., Berrebi, A. S., Mathers, P. H., & Agmon, A. (2006) · The Journal of Neuroscience · doi:10.1523/jneurosci.0661-06.2006 · Cited by 432 · cited 21× · see citing works
“(2017) in vivo and were able to correlate morphological subtypes to the diverse firing patterns observed in the SST-INs population (Ma et al., 2006).”
BackgroundResultInfluentialXu, X., Roby, K. D., & Callaway, E. M. (2010) · Journal of Comparative Neurology · doi:10.1002/cne.22229 · Cited by 429 · cited 1× · see citing works
“Axonal nodes represent branching points and have been previously used as a measure of axonal targeting (Blasdel and Lund, 1983; Florence and Casagrande, 1987; Freund et al., 1989; Lee et al., 2005; Yamada et al., 2010).”
MethodSupralinear increase of recurrent inhibition during sparse activity in the somatosensory cortex
Kapfer, C., Glickfeld, L. L., Atallah, B. V., & Scanziani, M. (2007) · Nature Neuroscience · doi:10.1038/nn1909 · Cited by 427 · cited 1× · see citing works
“SST-INs provide frequency-dependent feedback inhibition to pyramidal cells in supragranular and infragranular layers (Kapfer et al., 2007; Silberberg and Markram, 2007).”
BackgroundTranscriptional Architecture of Synaptic Communication Delineates GABAergic Neuron Identity
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“Interestingly, T-shaped Martinotti cells seem to be the sole SST-IN subtype to provide dendritic inhibition to the apical dendrite of L5 PCs in L4, where these cells receive sensory inputs from the ventral posteromedial thalamic nucleus (Petreanu et al., 2009).”
BackgroundFuzik, J., Zeisel, A., Máté, Z., Calvigioni, D., Yanagawa, Y., Szabó, G., Linnarsson, S., & Harkany, T. (2016) · Nature Biotechnology · doi:10.1038/nbt.3443 · Cited by 386 · cited 1× · see citing works
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BackgroundDendritic encoding of sensory stimuli controlled by deep cortical interneurons
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Neocortical Somatostatin-Expressing GABAergic Interneurons Disinhibit the Thalamorecipient Layer 4
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“Dendritic targeting GABAergic interneurons expressing the neuropeptide somatostatin (SST-INs) represent 30% of GABAergic interneurons (Xu et al., 2010; Tremblay et al., 2016).”
BackgroundSelf-citationCortical gamma band synchronization through somatostatin interneurons
Veit, J., Hakim, R., Jadi, M. P., B, T. J. S. A., & Adesnik, H. (2017) · Nature Neuroscience · doi:10.1038/nn.4562 · Cited by 364 · cited 5× · see citing works
“SST-INs are a very diverse group of inhibitory cells that show differential expression of molecular markers, different morphologies and electrophysiological properties (Wang et al., 2004; Ma et al., 2006).”
BackgroundInfluentialStrategies and Tools for Combinatorial Targeting of GABAergic Neurons in Mouse Cerebral Cortex
He, M., Tucciarone, J., Lee, S., Nigro, M. J., Kim, Y., Levine, J. M., Kelly, S. M., Krugikov, I., Wu, P., Chen, Y., et al. (2016) · Neuron · doi:10.1016/j.neuron.2016.08.021 · Cited by 339 · see citing works
Self-citationVisualization of chandelier cell axons by parvalbumin immunoreactivity in monkey cerebral cortex.
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“A wide morphological diversity has been described in other GABAergic interneuron groups, such as VIP-INs (bipolar and multipolar) and PV-INs (basket, chandelier, translaminar; DeFelipe et al., 1989; Buchanan et al., 2012; Bortone et al., 2014; Prönneke et al., 2015; He et al., 2016).”
BackgroundLayer-specific modulation of neocortical dendritic inhibition during active wakefulness
Muñoz, W., Tremblay, R., Levenstein, D., & Rudy, B. (2017) · Science · doi:10.1126/science.aag2599 · Cited by 305 · cited 1× · see citing works
“Indeed, T-shaped and fanning-out Martinotti cells may control Ca 2 electrogenesis in different dendritic compartments, differentially contributing to sensory perception (Murayama et al., 2009).”
BackgroundSelf-citationTranslaminar inhibitory cells recruited by layer 6 corticothalamic neurons suppress visual cortex
Bortone, D. S., Olsen, S. R., & Scanziani, M. (2014) · Neuron · doi:10.1016/j.neuron.2014.02.021 · Cited by 300 · cited 1× · see citing works
“A wide morphological diversity has been described in other GABAergic interneuron groups, such as VIP-INs (bipolar and multipolar) and PV-INs (basket, chandelier, translaminar; DeFelipe et al., 1989; Buchanan et al., 2012; Bortone et al., 2014; Prönneke et al., 2015; He et al., 2016).”
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“Axonal nodes represent branching points and have been previously used as a measure of axonal targeting (Blasdel and Lund, 1983; Florence and Casagrande, 1987; Freund et al., 1989; Lee et al., 2005; Yamada et al., 2010).”
MethodRobust but delayed thalamocortical activation of dendritic-targeting inhibitory interneurons
Tan, Z., Hu, H., Huang, Z. J., & Agmon, A. (2008) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0710628105 · Cited by 155 · cited 4× · see citing works
“SST-INs are a very heterogeneous group of cortical GABAergic neurons that vary in morphological, electrophysiological, and molecular features (Tremblay et al., 2016).”
BackgroundInfluentialTarget-specific expression of presynaptic NMDA receptors in neocortical microcircuits
Buchanan, K. A., Blackman, A. V., Moreau, A. W., Elgar, D., Costa, R. P., Lalanne, T., Tudor Jones, A. A., Oyrer, J., & Sjöström, P. J. (2012) · Neuron · doi:10.1016/j.neuron.2012.06.017 · Cited by 144 · cited 1× · see citing works
“A wide morphological diversity has been described in other GABAergic interneuron groups, such as VIP-INs (bipolar and multipolar) and PV-INs (basket, chandelier, translaminar; DeFelipe et al., 1989; Buchanan et al., 2012; Bortone et al., 2014; Prönneke et al., 2015; He et al., 2016).”
BackgroundChrna2-Martinotti cells synchronize layer 5 type A pyramidal cells via rebound excitation
Hilscher, M. M., Leão, R. N., Edwards, S. J., Leão, K. E., & Kullander, K. (2017) · PLoS Biology · doi:10.1371/journal.pbio.2001392 · Cited by 118 · cited 1× · see citing works
“In the neocortical L5, computational and experimental work has shown that Martinotti cells can synchronize pyramidal cells through distal inhibition (Li et al., 2013; Hilscher et al., 2017).”
MethodHalabisky, B., Shen, F., Huguenard, J. R., & Prince, D. A. (2006) · Journal of Neurophysiology · doi:10.1152/jn.01079.2005 · Cited by 111 · cited 2× · see citing works
“L5 SST-INs also show diverse firing properties when depolarized by current injection (Halabisky et al., 2006; Ma et al., 2006; Tremblay et al., 2016).”
BackgroundExuberant thalamocortical axon arborization in cortex-specific NMDAR1 knockout mice
Lee, L., Iwasato, T., Itohara, S., et al. (2005) · The Journal of Comparative Neurology · doi:10.1002/cne.20481 · Cited by 106 · cited 1× · see citing works
“Axonal nodes represent branching points and have been previously used as a measure of axonal targeting (Blasdel and Lund, 1983; Florence and Casagrande, 1987; Freund et al., 1989; Lee et al., 2005; Yamada et al., 2010).”
MethodDumitriu, D., Cossart, R., Huang, Z. J., & Yuste, R. (2006) · Cerebral Cortex · doi:10.1093/cercor/bhj126 · Cited by 103 · see citing works
Muñoz, W., Tremblay, R., & Rudy, B. (2014) · Cell Reports · doi:10.1016/j.celrep.2014.11.042 · Cited by 65 · cited 3× · see citing works
“Recently this morphological diversity was confirmed in vivo (Muñoz et al., 2014, 2017).”
BackgroundSelf-citationOrganization of individual afferent axons in layer IV of striate cortex in a primate
Florence, S. L., & Casagrande, V. A. (1987) · Journal of Neuroscience · doi:10.1523/jneurosci.07-12-03850.1987 · Cited by 57 · cited 1× · see citing works
“Axonal nodes represent branching points and have been previously used as a measure of axonal targeting (Blasdel and Lund, 1983; Florence and Casagrande, 1987; Freund et al., 1989; Lee et al., 2005; Yamada et al., 2010).”
MethodRole of pre- and postsynaptic activity in thalamocortical axon branching
Yamada, A., Uesaka, N., Hayano, Y., Tabata, T., Kano, M., & Yamamoto, N. (2010) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0900613107 · Cited by 53 · cited 1× · see citing works
“Axonal nodes represent branching points and have been previously used as a measure of axonal targeting (Blasdel and Lund, 1983; Florence and Casagrande, 1987; Freund et al., 1989; Lee et al., 2005; Yamada et al., 2010).”
MethodHu, H., & Agmon, A. (2016) · Journal of Neuroscience · doi:10.1523/jneurosci.0739-16.2016 · Cited by 52 · see citing works
Li, X., Morita, K., Robinson, H. P. C., & SMALL, M. P. (2013) · Journal of Neurophysiology · doi:10.1152/jn.00397.2012 · Cited by 21 · cited 1× · see citing works
“In the neocortical L5, computational and experimental work has shown that Martinotti cells can synchronize pyramidal cells through distal inhibition (Li et al., 2013; Hilscher et al., 2017).”
Method
Flagged first, then by citations
Research works printed but not linked: 2
- Martinotti, 1889, Contributo allo studio della corteccia cerebrale, ed all'origine centrale dei nervi, Ann Freniatr Sci Affini, 1, 14
- Ramon y Cajal, 1891, Sur la structure de l'ecorce cerebrale de quelques mammiferes, Cellule, 7, 3
Cited by
Sorted by citations
Development and Functional Diversification of Cortical Interneurons
Lim, L., Mi, D., Llorca, A., & Marín, O. (2018) · Neuron · doi:10.1016/j.neuron.2018.10.009 · Cited by 714 · see citing works
“Consistently, MGE/POA progenitor cells take on properties of the LGE and CGE in mice lacking Nkx2-1 (Butt et al., 2008; Sussel et al., 1999).”
InfluentialIntegrated Morphoelectric and Transcriptomic Classification of Cortical GABAergic Cells
Gouwens, N. W., Sorensen, S. A., Baftizadeh, F., Budzillo, A., Lee, B. R., Jarsky, T., Alfiler, L., Baker, K., Barkan, E., Berry, K., et al. (2020) · Cell · doi:10.1016/j.cell.2020.09.057 · Cited by 538 · see citing works
“described L2/3 Martinotti and L5/6 fanning-out Martinotti cells (Nigro et al., 2018; Munõz et al., 2017).”
Phenotypic variation of transcriptomic cell types in mouse motor cortex
Scala, F., Kobak, D., Bernabucci, M., Bernaerts, Y., Cadwell, C. R., Castro, J. R., Hartmanis, L., Jiang, X., Laturnus, S., Miranda, E., et al. (2021) · Nature · doi:10.1038/s41586-020-2907-3 · Cited by 398 · see citing works
Burst-dependent synaptic plasticity can coordinate learning in hierarchical circuits
Payeur, A., Guerguiev, J., Zenke, F., Richards, B. A., & Naud, R. (2021) · Nature Neuroscience · doi:10.1038/s41593-021-00857-x · Cited by 286 · see citing works
“We modeled somatostatin-positive interneurons [46, 71, 92] using the adaptive exponential integrate-and-fire (AdEx) model [68]:”
MethodLayer 4 of mouse neocortex differs in cell types and circuit organization between sensory areas
Scala, F., Kobak, D., Shan, S., Bernaerts, Y., Laturnus, S., Cadwell, C. R., Hartmanis, L., Froudarakis, E., Castro, J. R., Tan, Z. H., et al. (2019) · Nature Communications · doi:10.1038/s41467-019-12058-z · Cited by 193 · see citing works
Neuronal circuits in barrel cortex for whisker sensory perception
Staiger, J. F., & Petersen, C. C. H. (2021) · Physiological Reviews · doi:10.1152/physrev.00019.2019 · Cited by 165 · see citing works
Single-cell transcriptomic evidence for dense intracortical neuropeptide networks
Smith, S. J., Sümbül, U., Graybuck, L. T., Collman, F., Seshamani, S., Gala, R., Gliko, O., Elabbady, L., Miller, J. A., Bakken, T. E., et al. (2019) · eLife · doi:10.7554/elife.47889 · Cited by 156 · see citing works
Yamamuro, K., Bicks, L. K., Leventhal, M. B., Kato, D., Im, S., Flanigan, M. E., Garkun, Y., Norman, K. J., Caro, K., Sadahiro, M., et al. (2020) · Nature Neuroscience · doi:10.1038/s41593-020-0695-6 · Cited by 169 · see citing works
Cortical Somatostatin Interneuron Subtypes Form Cell-Type Specific Circuits
Wu, S. J., Sevier, E., Dwivedi, D., Saldi, G.-A., Hairston, A., Yu, S. Q., Abbott, L., Choi, D. H., Sherer, M., Qiu, Y., et al. (2022) · Neuron · doi:10.1016/j.neuron.2023.05.032 · Cited by 130 · see citing works
Self-citationComplementary networks of cortical somatostatin interneurons enforce layer specific control
Naka, A., Veit, J., Shababo, B., Chance, R. K., Risso, D., Stafford, D., Snyder, B., Egladyous, A., Chu, D., Sridharan, S., et al. (2019) · eLife · doi:10.7554/elife.43696 · Cited by 117 · see citing works
Diversity and Function of Somatostatin-Expressing Interneurons in the Cerebral Cortex
Riedemann, T. (2019) · International Journal of Molecular Sciences · doi:10.3390/ijms20122952 · Cited by 94 · see citing works
“In layer 2/3 and in layers 5 and 6, pyramidal neurons represent the preferred postsynaptic target of SOM+ INs [59,75,76,127,128].”
BackgroundNaskar, S., Qi, J., Pereira, F., Gerfen, C. R., & Lee, S. (2021) · Cell Reports · doi:10.1016/j.celrep.2021.108774 · Cited by 72 · see citing works
“SST INs are diverse not only in their molecular and genetic profiles but also in their axonal morphology and intrinsic electrophysiological properties (Jiang et al., 2015; Ma et al., 2006; Muñoz et al., 2017; Naka et al., 2019; Nigro et al., 2018; Xu et al., 2013).”
Cortical disinhibitory circuits: cell types, connectivity and function
Kullander, K., & Topolnik, L. (2021) · Trends in Neurosciences · doi:10.1016/j.tins.2021.04.009 · Cited by 83 · see citing works
Genetic mosaicism in the human brain: from lineage tracing to neuropsychiatric disorders
Bizzotto, S., & Walsh, C. A. (2022) · Nature reviews. Neuroscience · doi:10.1038/s41583-022-00572-x · Cited by 84 · see citing works
Hage, T. A., Bosma-Moody, A., Baker, C. A., Kratz, M. B., Campagnola, L., Jarsky, T., Zeng, H., & Murphy, G. J. (2022) · eLife · doi:10.7554/elife.71103 · Cited by 84 · see citing works
“How, if at all, does the pattern of vertical inhibitory input to L2/3 PCs resemble that of vertical excitatory input to the same cells? The diverse axonal projection patterns of interneurons and reports of translaminar synaptic inhibition (Kapfer et al., 2007; Silberberg and Markram, 2007; Nigro et al., 2018; Gouwens…”
BackgroundRen, C., Peng, K., Yang, R., Liu, W., Liu, C., & Komiyama, T. (2022) · Neuron · doi:10.1016/j.neuron.2022.04.031 · Cited by 74 · see citing works
Head Movements Control the Activity of Primary Visual Cortex in a Luminance-Dependent Manner
Bouvier, G., Senzai, Y., & Scanziani, M. (2020) · Neuron · doi:10.1016/j.neuron.2020.07.004 · Cited by 64 · see citing works
Nguyen, R., Venkatesan, S., Binko, M. A., Bang, J. Y., Cajanding, J. D., Briggs, C., Sargin, D., Imayoshi, I., Lambe, E. K., & Kim, J. C. (2020) · Journal of Neuroscience · doi:10.1523/jneurosci.1919-19.2020 · Cited by 72 · see citing works
“The interneurons targeted include the lowthreshold spiking and Regular spiking subgroups, broadly consistent with the somatostatin (SST) and VIP subclasses (Ma et al., 2006; Prönneke et al., 2015; Batista-Brito et al., 2017; Nigro et al., 2018).”
BackgroundCell-type-specific inhibitory circuitry from a connectomic census of mouse visual cortexPreprint
Schneider-Mizell, C. M., Bodor, Á. L., Brittain, D., Buchanan, J., Bumbarger, D. J., Elabbady, L., Gamlin, C. R., Kapner, D., Kinn, S., Mahalingam, G., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.01.23.525290 · Cited by 70 · see citing works
Mechanisms underlying the response of mouse cortical networks to optogenetic manipulation
Mahrach, A., Chen, G., Li, N., van Vreeswijk, C., & Hansel, D. (2020) · eLife · doi:10.7554/elife.49967 · Cited by 68 · see citing works
“PCs in layer 2/3 project to layer 5 and receive feedback from the latter ( Hooks et al., 2013 ; Naka and Adesnik, 2016 ).”
BackgroundJoffe, M. E., Maksymetz, J., Luchsinger, J. R., Dogra, S., Ferranti, A. S., Luessen, D. J., Gallinger, I. M., Xiang, Z., Branthwaite, H. E., Melugin, P. R., et al. (2022) · Neuron · doi:10.1016/j.neuron.2021.12.027 · Cited by 69 · see citing works
“One of the major types of SST-IN is the Martinotti cell, which can be functionally characterized by low-threshold spiking activity as well as cellular anatomy, morphology, and function (Nigro et al., 2018).”
BackgroundParallel social information processing circuits are differentially impacted in autism
Lewis, E. M., Stein-O’Brien, G. L., Patino, A. V., Nardou, R., Grossman, C. D., Brown, M., Bangamwabo, B., Ndiaye, N., Giovinazzo, D., Dardani, I., et al. (2020) · Neuron · doi:10.1016/j.neuron.2020.10.002 · Cited by 72 · see citing works
Exciting Complexity: The Role of Motor Circuit Elements in ALS Pathophysiology
Gunes, Z. I., Kan, V. W. Y., Ye, X., & Liebscher, S. (2020) · Frontiers in Neuroscience · doi:10.3389/fnins.2020.00573 · Cited by 67 · see citing works
The Organization and Development of Cortical Interneuron Presynaptic Circuits is Area Specific
Pouchelon, G., Dwivedi, D., Bollmann, Y., Agba, C. K., Xu, Q., Mirow, A. M., Kim, S., Qiu, Y., Sevier, E., Ritola, K., et al. (2021) · Cell Reports · doi:10.1016/j.celrep.2021.109993 · Cited by 52 · see citing works
“40% of non-Mar-tinotti cells in S1 (Nigro et al., 2018).”
The covariance environment defines cellular niches for spatial inference
Haviv, D., Remšík, J., Gatie, M. I., Snopkowski, C., Takizawa, M. T., Pereira, N., Bashkin, J., Jovanovich, S. B., Nawy, T., Chaligné, R., et al. (2024) · Nature Biotechnology · doi:10.1038/s41587-024-02193-4 · Cited by 69 · see citing works
Joffe, M. E., Winder, D. G., & Conn, P. J. (2020) · Neuropharmacology · doi:10.1016/j.neuropharm.2020.108126 · Cited by 52 · see citing works
Malik, R., Pai, E. L.-L., Rubin, A. N., Stafford, A. M., Angara, K., Minasi, P., Rubenstein, J. L. R., Sohal, V. S., & Vogt, D. L. (2019) · Nature Communications · doi:10.1038/s41467-019-12962-4 · Cited by 55 · see citing works
Cone, J. J., Scantlen, M. D., Histed, M. H., & Maunsell, J. H. R. (2019) · eNeuro · doi:10.1523/eneuro.0337-18.2019 · Cited by 49 · see citing works
“, 2017) and diverse electrophysiological and morphologic properties within a cortical layer (Nigro et al., 2018).”
BackgroundInterneuron development and dysfunction
Yang, J., Yang, X. W., & Tang, K. (2021) · FEBS Journal · doi:10.1111/febs.15872 · Cited by 52 · see citing works
Layer 6b controls brain state via apical dendrites and the higher-order thalamocortical system
Zolnik, T. A., Bronec, A., Ross, A., Staab, M., Sachdev, R. N. S., Molnár, Z., Eickholt, B. J., & Larkum, M. E. (2023) · Neuron · doi:10.1016/j.neuron.2023.11.021 · Cited by 51 · see citing works
Inhibition in the auditory cortex
Studer, F., & Barkat, T. R. (2021) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2021.11.021 · Cited by 52 · see citing works
“One study found increased spontaneous and sound-evoked activity of L1 inhibitory neurons during movement, which potentially explains the decreased activity observed in L2/3 excitatory neurons (Zhou et al., 2014).”
BackgroundInfluentialCell-type-specific propagation of visual flicker
Schneider, M., Τζάνου, Α., Uran, C., & Vinck, M. A. (2023) · Cell Reports · doi:10.1016/j.celrep.2023.112492 · Cited by 56 · see citing works
Huang, A. Y., Li, P., Rodin, R. E., Kim, S. N., Dou, Y., Kenny, C. J., Akula, S. K., Hodge, R. D., Bakken, T. E., Miller, J. A., et al. (2020) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2006163117 · Cited by 50 · see citing works
“Previous studies cataloging interneurons in mouse and human have suggested that MGE-derived inhibitory neuron subtypes (SST+ and PVALB+) are enriched in infragranular cortical layers, while CGE-derived interneuron subtypes (LAMP5/PAX6+, VIP+) tend to occupy upper cortical layers preferentially (5, 42, 43), and thus ou…”
BackgroundAn enhancer-AAV toolbox to target and manipulate distinct interneuron subtypes
Furlanis, E., Dai, M., Garcia, B. L., Tran, T. K., Vergara, J., Pereira, A., Gorissen, B. L., Wills, S., Vlachos, A., Hairston, A., et al. (2024) · Neuron · doi:10.1016/j.neuron.2025.05.002 · Cited by 66 · see citing works
Somatostatin and Somatostatin-Containing Interneurons—From Plasticity to Pathology
Liguz‐Lecznar, M., Dobrzański, G., & Kossut, M. M. (2022) · Biomolecules · doi:10.3390/biom12020312 · Cited by 46 · see citing works
“L5 non-Martinotti interneurons account for approximately one-third of all L5 SOM-INs and regulate the activity of the excitatory cells of L4, but not of L5 [54].”
BackgroundPhenotypic variation within and across transcriptomic cell types in mouse motor cortex
(2020) · doi:10.1101/2020.02.03.929158 · Cited by 397 · see citing works
“Sst Myh8_1, Sst Etv1, and Sst Pvalb Etv1 showed ‘T-shaped’ Martinotti morphologies (Muñoz et al., 2017; Nigro et al., 2018) and strong rebound firing.”
BackgroundInfluentialSomatostatin-Positive Interneurons Contribute to Seizures in SCN8A Epileptic Encephalopathy
Wengert, E. R., Miralles, R. M., Wedgwood, K. C. A., Wagley, P. K., Strohm, S. M., Panchal, P. S., Idrissi, A. M., Wenker, I. C., Thompson, J. A., Gaykema, R. P., et al. (2021) · Journal of Neuroscience · doi:10.1523/jneurosci.0718-21.2021 · Cited by 48 · see citing works
Origin, Development, and Synaptogenesis of Cortical Interneurons
Llorca, A., & Deogracias, R. (2022) · Frontiers in Neuroscience · doi:10.3389/fnins.2022.929469 · Cited by 40 · see citing works
(2020) · doi:10.1101/2020.02.03.932244 · Cited by 537 · see citing works
“As mentioned, this met-type reflects the well-described Martinotti-type neurons, and includes cells that resemble the recently described L2/3 Martinotti and L5/6 fanning-out Martinotti cells (Muñ oz et al., 2017; Nigro et al., 2018).”
MethodFisher, J., Verhagen, M. G., Long, Z., Moissidis, M., Yan, Y., He, C., Wang, J., Micoli, E., Alastruey, C. M., Moors, R., et al. (2023) · Neuron · doi:10.1016/j.neuron.2023.11.013 · Cited by 36 · see citing works
“…transcriptional profile in the developmental dataset aligns well with Sst-MET-6, which corresponds to deep-layer alpha-2 T-shaped MCs, 23 MC3 matches several subtypes of fanning-out MCs (Sst-MET-3, Sst-MET-4, and Sst-MET-5) that occupy different layers in the adult cortex 24,25 (Figures 3A and 3B).”
Regulation of Recurrent Inhibition by Asynchronous Glutamate Release in Neocortex
Deng, S., Li, J., He, Q., Zhang, X., Zhu, J., Li, L., Mi, Z., Yang, X., Jiang, M., Dong, Q., et al. (2019) · Neuron · doi:10.1016/j.neuron.2019.10.038 · Cited by 37 · see citing works
Almási, Z., Dávid, C., Witte, M., & Staiger, J. F. (2019) · Frontiers in Neuroanatomy · doi:10.3389/fnana.2019.00045 · Cited by 37 · see citing works
Yang, J., Serrano, P., Yin, X., Sun, X., Lin, Y., & Chen, S. X. (2022) · Neuron · doi:10.1016/j.neuron.2022.08.018 · Cited by 36 · see citing works
Stimulus novelty uncovers coding diversity in survey of visual cortexPreprint
Garrett, M., Groblewski, P. A., Piet, A. T., Ollerenshaw, D. R., Najafi, F., Yavorska, I., Amster, A., Bennett, C., Buice, M. A., Caldejon, S., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.02.14.528085 · Cited by 29 · see citing works
Guthman, E. M., Garcia, J. D., Ma, M., Chu, P., Baca, S. M., Smith, K. R., Restrepo, D., & Huntsman, M. M. (2020) · eLife · doi:10.7554/elife.50601 · Cited by 32 · see citing works
“Further, in piriform and neocortex, FS Sst+ INs show clear biophysical and circuit connectivity differences from other Sst+ IN subtypes (Large et al., 2016; Ma et al., 2006; Naka et al., 2019; Nigro et al., 2018).”
BackgroundInfluentialThe hippocampus associated GABAergic neural network impairment in early-stage of Alzheimer's disease
Tang, Y., Yan, Y., Mao, J., Ni, J., & Qing, H. (2023) · Ageing Research Reviews · doi:10.1016/j.arr.2023.101865 · Cited by 34 · see citing works
Genetically Defined Subtypes of Somatostatin-Containing Cortical Interneurons
Hostetler, R. E., Hu, H., & Agmon, A. (2023) · eNeuro · doi:10.1523/eneuro.0204-23.2023 · Cited by 30 · see citing works
“We therefore resorted to an intersectional strategy (He et al., 2016; Kim et al., 2017; Nigro et al., 2018; Wu et al., 2022), but chose the following novel approach: we crossed each of the four Cre line with the Sst-Flp line and the combinatorial RC::FLTG reporter (Plummer et al.”
InfluentialMultimodal cortical neuronal cell type classification
Mao, X., & Staiger, J. F. (2024) · Pflügers Archiv - European Journal of Physiology · doi:10.1007/s00424-024-02923-2 · Cited by 34 · see citing works
“Layer 2/3 VIP cells possess axonal arborizations that extend downward to the layer 6/white matter boundary [22], with Vip-Rspo1-Itga4 displaying regular spiking and Vip-Ptprt-Pkp2 (mostly associated with Chat + cells) displaying irregular spiking [9, 10].”
BackgroundTranslaminar recurrence from layer 5 suppresses superficial cortical layers
Onodera, K., & Kato, H. (2022) · Nature Communications · doi:10.1038/s41467-022-30349-w · Cited by 25 · see citing works
Adolescent binge drinking leads to long-lasting changes in cortical microcircuits in mice
Sicher, A. R., Starnes, W. D., Griffith, K. R., Dao, N. C., Smith, G. C., Brockway, D. F., & Crowley, N. A. (2023) · Neuropharmacology · doi:10.1016/j.neuropharm.2023.109561 · Cited by 27 · see citing works
Connectomic reconstruction of a cortical columnPreprint
Sievers, M., Motta, A., Schmidt, M., Yener, Y., Loomba, S., Song, K., Bruett, J., & Helmstaedter, M. N. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.03.22.586254 · Cited by 31 · see citing works
A postnatal molecular switch drives activity-dependent maturation of parvalbumin interneurons
Moissidis, M., Abbasova, L., Selten, M. M., Alis, R., Bernard, C., Domínguez-Canterla, Y., Oozeer, F., Qin, S., Kelly, A., Mòdol, L., et al. (2025) · Cell · doi:10.1016/j.cell.2025.06.029 · Cited by 34 · see citing works
Wang, H., Flores, R. J., Yarur, H. E., Limoges, A., Bravo-Rivera, H., Casello, S. M., Loomba, N., Enríquez-Traba, J., Arenivar, M. A., Wang, Q., et al. (2024) · Neuron · doi:10.1016/j.neuron.2024.03.015 · Cited by 25 · see citing works
Distinct roles of PV and Sst interneurons in visually induced gamma oscillations
Onorato, I., Τζάνου, Α., Schneider, M., Uran, C., Broggini, A. C., & Vinck, M. A. (2025) · Cell Reports · doi:10.1016/j.celrep.2025.115385 · Cited by 36 · see citing works
Li, M., Cabrera-García, D., Salling, M. C., Au, E., Yang, G., & Harrison, N. L. (2021) · Neuropharmacology · doi:10.1016/j.neuropharm.2021.108501 · Cited by 24 · see citing works
Targeting prefrontal cortex GABAergic microcircuits for the treatment of alcohol use disorder
Fish, K. N., & Joffe, M. E. (2022) · Frontiers in Synaptic Neuroscience · doi:10.3389/fnsyn.2022.936911 · Cited by 20 · see citing works
“Most SST-INs are lowthreshold spiking cells, characterized by high input resistance, depolarized resting potential, low rheobase, and occasional firing upon hyperpolarization rebound (Kawaguchi and Kubota, 1997, 1998; Marlin and Carter, 2014; McGarry and Carter, 2016; Tremblay et al., 2016; Nigro et al., 2018; Joffe e…”
BackgroundConnectomics of predicted Sst transcriptomic types in mouse visual cortex
Gamlin, C. R., Schneider-Mizell, C. M., Mallory, M., Elabbady, L., Gouwens, N. W., Williams, G., Mukora, A., Dalley, R. A., Bodor, Á. L., Brittain, D., et al. (2025) · Nature · doi:10.1038/s41586-025-08805-6 · Cited by 31 · see citing works
“Previous functional studies have also shown that MCs can connect broadly in a ‘blanket of inhibition’ 25–27 or inhibit a subset of excitatory neuron targets within and across cortical layers 28–32 .”
InfluentialJuxtacellular opto-tagging of hippocampal CA1 neurons in freely moving mice
Ding, L., Balsamo, G., Chen, H., Blanco-Hernández, E., Zouridis, I. S., Naumann, R. K., Preston‐Ferrer, P., & Burgalossi, A. (2022) · eLife · doi:10.7554/elife.71720 · Cited by 20 · see citing works
Neocortical somatostatin neuron diversity in cognition and learning
Park, E., Mosso, M. B., & Barth, A. L. (2025) · Trends in Neurosciences · doi:10.1016/j.tins.2024.12.004 · Cited by 20 · see citing works
“Experiments resolving SST subtypes have demonstrated differences in intrinsic properties [28], laminar location [30,31], and target speci fi city [25,32] between subsets of Martinotti neurons, highlighting the advantage of this approach for understanding SST function.”
Regulatory Elements Inserted into AAVs Confer Preferential Activity in Cortical Interneurons
Rubin, A. N., Malik, R., Cho, K. K. A., Lim, K. J., Lindtner, S., Schwartz, S. E. R., Vogt, D. L., Sohal, V. S., & Rubenstein, J. L. R. (2020) · eNeuro · doi:10.1523/eneuro.0211-20.2020 · Cited by 18 · see citing works
“Second, the Cre-driver lines are not able to distinguish between additional subclasses within the PV, SST, and VIP CIN subtypes, which have distinct physiological properties and molecular markers (Nigro et al., 2018).”
BackgroundSamaran, J., Gabriel Peyré, A., & Cantini, L. (2024) · Nature Communications · doi:10.1038/s41467-024-51382-x · Cited by 22 · see citing works
“Furthermore, Somatostatin-expressing neurons (Sst), which are known to be morphologically diverse 82 , seem to be organized according to their layer of origin, with layer 2/3, layer 5, and layer 6 moving from left to right in the last UMAP plot of Fig.”
Zhou, X., Mansori, I., Fischer, T., Witte, M., & Staiger, J. F. (2020) · Journal of Comparative Neurology · doi:10.1002/cne.24756 · Cited by 18 · see citing works
“In layer V, GIN cells are mainly represented by fanning-out Martinotti cells, which innervate most extensively layer II/III, and to a lesser extent layer I (Ma et al., 2006; Nigro et al., 2018).”
BackgroundLimoni, G., Murthy, S. S., Jabaudon, D., Dayer, A. G., & Niquille, M. (2021) · Cell Reports · doi:10.1016/j.celrep.2020.108644 · Cited by 21 · see citing works
Synaptic zinc potentiates AMPA receptor function in mouse auditory cortex
Bender, P. T. R., McCollum, M., Boyd-Pratt, H., Mendelson, B. Z., & Anderson, C. T. (2023) · Cell Reports · doi:10.1016/j.celrep.2023.112932 · Cited by 18 · see citing works
Hwang, Y.-S., Maclachlan, C., Blanc, J., Dubois, A., Petersen, C. C. H., Knott, G. W., & Lee, S. (2020) · Cerebral Cortex · doi:10.1093/cercor/bhaa378 · Cited by 16 · see citing works
“In the mouse somatosensory cortex, SST+ neurons show various physiological features that can be used to further classify the sub-classes of SST+ neurons (Munoz et al. 2017; Nigro et al. 2018).”
BackgroundTranscranial Magnetic Stimulation and Neocortical Neurons: The Micro-Macro Connection
Tian, D., & Izumi, S. (2022) · Frontiers in Neuroscience · doi:10.3389/fnins.2022.866245 · Cited by 18 · see citing works
Hernáth, F., Schlett, K., & Szűcs, A. L. (2019) · Scientific Reports · doi:10.1038/s41598-019-49197-8 · Cited by 16 · see citing works
Inhibitory and disinhibitory VIP IN-mediated circuits in neocortexPreprint
Dellal, S. S., Zurita, H., Kruglikov, I., Valero, M., Abad-Perez, P., Geron, E., Meng, J. H., Prönneke, A., Hanson, J. L., Mir, E., et al. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.02.26.640383 · Cited by 23 · see citing works
Self-citationCortical Hyperexcitability in the Driver’s Seat in ALS
Gunes, Z. I., Kan, V. W. Y., Jiang, S., Logunov, E., Ye, X., & Liebscher, S. (2022) · Clinical and Translational Neuroscience · doi:10.3390/ctn6010005 · Cited by 17 · see citing works
“Activity levels and information processing in M1 is regulated by a diverse set of local and remote inhibitory neurons, which constitute ~20% of the cortical neuronal population [92,93].”
BackgroundOpioid modulation of prefrontal cortex cells and circuits
Cole, R. H., Moussawi, K., & Joffe, M. E. (2024) · Neuropharmacology · doi:10.1016/j.neuropharm.2024.109891 · Cited by 16 · see citing works
Cattaneo, S., Ripamonti, M., Bedogni, F., Sessa, A., & Taverna, S. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/566984 · Cited by 14 · see citing works
“…described in cortical and hippocampal SST-INs (Oliva et al., 2000; Taverna et al., 2005; Ma et al., 2006; Xu et al., 2006; Yekhlef et al., 2015; Nigro et al., 2018), and distinct from those of other major cell types such as spiny projection neurons (SPNs, (Taverna et al., 2008; Kreitzer,…”
BackgroundThe synaptic architecture of layer 5 thick tufted excitatory neurons in mouse visual cortex
Bodor, A. L., Schneider-Mizell, C. M., Zhang, C., Elabbady, L., Mallen, A., Bergeson, A., Brittain, D., Buchanan, J., Bumbarger, D. J., Dalley, R. A., et al. (2025) · Nature Neuroscience · doi:10.1038/s41593-025-02004-2 · Cited by 17 · see citing works
Hilscher, M. M., Mikulovic, S., Perry, S. E., Lundberg, S., & Kullander, K. (2023) · Pharmacological Research · doi:10.1016/j.phrs.2023.106895 · Cited by 15 · see citing works
“…cells with “ T-shaped ” morphology (prominent innervation of layer 1) was suppressed during whisking whereas the spiking activity of layer 5 Martinotti cells with “ fanning-out ” morphology (prominent innervation of layer 2/3, with less ascending axon reaching layer 1) was increased [140,141].”
Agmon, A., & Barth, A. L. (2024) · Frontiers in Neural Circuits · doi:10.3389/fncir.2024.1436915 · Cited by 12 · see citing works
Transcriptomic cell-type specificity of local cortical circuits
Patiño, M., Rossa, M., Lagos, W. N., Patne, N. S., & Callaway, E. M. (2024) · Neuron · doi:10.1016/j.neuron.2024.09.003 · Cited by 17 · see citing works
Wu, S. J., Dai, M., Yang, S.-P., McCann, C., Qiu, Y., Kumar, V., Marrero, G. J., Tsyporin, J., Huang, S., Shin, D., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.07.20.604399 · Cited by 17 · see citing works
László, Z. I., Bercsényi, K., Mayer, M., Lefkovics, K., Szabó, G., Katona, I., & Lele, Z. (2019) · Cerebral Cortex · doi:10.1093/cercor/bhz168 · Cited by 13 · see citing works
Transcriptional maintenance of cortical somatostatin interneuron subtype identity during migration
Munguba, H., Nikouei, K., Hochgerner, H., Oberst, P., Kouznetsova, A., Ryge, J., Muñoz‐Manchado, A. B., Close, J., Batista‐Brito, R., Linnarsson, S., et al. (2023) · Neuron · doi:10.1016/j.neuron.2023.07.018 · Cited by 19 · see citing works
Sensitization of Cutaneous Primary Afferents in Bone Cancer Revealed by In Vivo Calcium Imaging
de Clauser, L., Luiz, A. P., Santana‐Varela, S., Wood, J. N., & Sikandar, S. (2020) · Cancers · doi:10.3390/cancers12123491 · Cited by 12 · see citing works
“Where indicated, homozygous Pirt-GCaMP3 and Rosa-flox-stop tdTomato mice were crossed with homozygous Calb1-cre [69] or Tmem233-cre mice.”
BackgroundIbrahim, L. A., Wamsley, B., Alghamdi, N., Yusuf, N., Sevier, E., Hairston, A., Sherer, M., Jaglin, X. H., Xu, Q., Guo, L., et al. (2023) · eLife · doi:10.7554/elife.86842 · Cited by 14 · see citing works
Cell-type-specific enhancement of deviance detection by synaptic zinc in the mouse auditory cortex
McCollum, M., Manning, A., Bender, P. T. R., Mendelson, B. Z., & Anderson, C. T. (2024) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2405615121 · Cited by 11 · see citing works
“In addition, whereas the organizing principles of inhibitory circuits in superficial cortical layers are relatively well understood ( 83 – 93 ), the translaminar activation of layer 5 GABAergic interneurons during auditory processing is less well studied.”
Mapping of morpho-electric features to molecular identity of cortical inhibitory neurons
Roussel, Y., Verasztó, C., Rodarie, D., Damart, T., Reimann, M., Ramaswamy, S., Markram, H., & Keller, D. X. (2023) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1010058 · Cited by 11 · see citing works
Gamlin, C. R., Schneider-Mizell, C. M., Mallory, M., Elabbady, L., Gouwens, N. W., Williams, G., Mukora, A., Dalley, R. A., Bodor, Á. L., Brittain, D., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.03.22.533857 · Cited by 11 · see citing works
Genetic approaches to elucidating cortical and hippocampal GABAergic interneuron diversity
Machold, R. P., & Rudy, B. (2024) · Frontiers in Cellular Neuroscience · doi:10.3389/fncel.2024.1414955 · Cited by 11 · see citing works
“A number of recent studies have begun to illuminate the diversity of SST MC, non-MC, and OLM IN populations using intersectional genetic approaches (He et al., 2016; Nigro et al., 2018; Gouwens et al., 2020; Wu et al., 2023; Chamberland et al., 2024).”
BackgroundMethodSelf-citationFrequency-dependent gating of feedforward inhibition in thalamofrontal synapses
Lee, J., Choi, J. H., & Rah, J. (2020) · Molecular Brain · doi:10.1186/s13041-020-00608-2 · Cited by 10 · see citing works
Park, E., Kuljis, D. A., Swindell, R. A., Ray, A., Zhu, M., Christian, J. A., & Barth, A. L. (2025) · Cell Reports · doi:10.1016/j.celrep.2025.115606 · Cited by 9 · see citing works
Bershteyn, M., Zhou, H., Fuentealba, L. C., Chen, C., Subramanyam, G., Cherkowsky, D., Sevilla, E. S., Hampel, P., Salvatierra, J., Sezan, M., et al. (2025) · Neuron · doi:10.1016/j.neuron.2025.06.010 · Cited by 13 · see citing works
Selective Inhibitory Circuit Dysfunction after Chronic Frontal Lobe Contusion
Nolan, A., Sohal, V. S., & Rosi, S. (2022) · Journal of Neuroscience · doi:10.1523/jneurosci.0097-22.2022 · Cited by 11 · see citing works
Nogueira, I., Lima, T. Z., Malfatti, T., & Leão, K. E. (2023) · Frontiers in Aging Neuroscience · doi:10.3389/fnagi.2023.1152497 · Cited by 8 · see citing works
“A larger sample of Chrna2+ MCs may allow for further subdivision of membrane properties and cell morphology (Nigro et al., 2018) since the genetic profiles of Chrna2+ MCs show several possible subclassifications based on clustering of transcriptomic data (Allen brain Atlas, Transcriptomics Explorer).”
BackgroundFriendlyClearMap: an optimized toolkit for mouse brain mapping and analysis
Negwer, M., Bosch, B., Bormann, M., Hesen, R., Lütje, L., Aarts, L., Rossing, C., Kasri, N. N., & Schubert, D. (2022) · GigaScience · doi:10.1093/gigascience/giad035 · Cited by 5 · see citing works
Transcriptional maintenance of cortical somatostatin interneuron subtype identity during migrationPreprint
Munguba, H., Nikouei, K., Hochgerner, H., Oberst, P., Kouznetsova, A., Ryge, J., Batista‐Brito, R., Muñoz‐Manchado, A. B., Close, J., Linnarsson, S., et al. (2019) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/593285 · Cited by 9 · see citing works
Huang, A. Y., Li, P., Rodin, R. E., et al. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.04.19.046904 · Cited by 9 · see citing works
“suggested that MGE-derived inhibitory neuron subtypes (SST+ and PVALB+) are enriched in 320 infragranular cortical layers, while CGE-derived interneuron subtypes (LAMP5/PAX6+, VIP+) 321 tend to occupy upper cortical layers preferentially (5, 42, 43) and thus our mapping of PRDD-seq 322”
BackgroundDistinct roles of PV and Sst interneurons in visually-induced gamma oscillationsPreprint
Onorato, I., Τζάνου, Α., Schneider, M., et al. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.04.08.535291 · Cited by 9 · see citing works
“We found that the Sst interneurons included both Nw and Bw cells, consistent with previous reports (Mu˜noz et al., 2014; Kim et al., 2016; Nigro et al., 2018; Kvitsiani et al., 2013), (Fig.”
InfluentialPyramidal neurons proportionately alter cortical interneuron subtypes
Wu, S. J., Dai, M., Yang, S.-P., McCann, C., Qiu, Y., Kumar, V., Marrero, G. J., Tsyporin, J., Huang, S., Shin, D., et al. (2026) · Nature · doi:10.1038/s41586-025-09996-8 · Cited by 26 · see citing works
Cell-Type–Specific Effects of Somatostatin on Synaptic Transmission in the Anterior Cingulate Cortex
Riedemann, T., & Sutor, B. (2024) · Journal of Neuroscience · doi:10.1523/jneurosci.0598-23.2024 · Cited by 9 · see citing works
Cortical somatostatin interneuron subtypes form cell-type specific circuitsPreprint
Wu, S. J., Sevier, E., Saldi, G.-A., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.09.29.510081 · Cited by 8 · see citing works
“8,10 Another SST subtype that resides in L5a is the SST-Etv1 subtype, which resembles a fanning-out Martinotti shape and can be partially targeted using an Etv1 CreER ;Sst FlpO intersectional strategy.”
InfluentialSelf-citation
- Authors
- Nigro, M. J., Hashikawa-Yamasaki, Y., & Rudy, B.
- Affiliations
- Neuroscience Institute (IT)
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2018-01-11
- Volume
- 38
- Issue
- 7
- Pages
- 1622-1633
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC5815450
- OpenAlex
- W2782653800
- Semantic Scholar
- f587144c9781b3912ec6840b8be4b2b7202ba19d
- Cited by
- 149
- References
- 34
- Citations by year
- 2026: 7 · 2025: 14 · 2024: 24 · 2023: 29 · 2022: 23 · 2021: 13 · 2020: 22 · 2019: 15 · 2018: 2
- FWCI
- 5.92
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Journal of Neuroscience (publishedVersion, cc-by-nc-sa)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health (5r01ns030989-22)
- National Institute of Neurological Disorders and Stroke (R01NS107257, R01-NS030989, P01NS074972, R01NS110079, R21NS093987)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neuropharmacology Research
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- layer 5, barrel cortex, Martinotti cells, dendritic inhibition, somatosensory cortex
- MeSH terms
- Animals, Dendrites, Female, Interneurons, Male, Neural Inhibition, Neural Pathways, Neurons, Somatosensory Cortex, Somatostatin, Pyramidal Cells, Mice, Electrophysiological Phenomena, In Vitro Techniques
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