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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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60 Own result · 17 Not checked yet · 9 Own interpretation · 4 Own method · 2 No basis given
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“We found that intersectional genetics can be used to obtain mouse lines where the expression of fluorescent proteins in specific morphological types is enriched.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Significance Statement
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“We found very little (2.2%) overlap of CR and Calb in the SST-cre line specially in deep layers”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
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In the paper
“In both intersectional lines the majority of tdTomatoexpressing cells coexpressed either CR or Calb”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
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- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
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In the paper
“The ascending axon of each morphological group exhibited a specific layer distribution of relative axonal length and proportion of axonal nodes”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“fanning-out Martinotti cells innervate L2/3 and the lower half of L1, and show adapting firing patterns”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
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In the paper
“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.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
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- Also stated in: § Morphological diversity of L5 SST interneurons · plain
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In the paper
“The proportion of tdTomato-expressing cells in L5b (ϳ67% of SST-INs) was larger than in L5a (ϳ44% of SST-INs”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
plaincorrelational- Asserted: passes — The authors put it forward as true.
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- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
plain- Asserted: passes — The authors put it forward as true.
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- Also stated in: § Electrophysiological properties of morphologically identified SST interneurons · plain
- It may state two separate claims as one. 22
- Probably not a substantial claim. 8
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
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In the paper
“non-Martinotti cells inhibit the dendrites of L4 neurons while avoiding L5 PCs”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plain- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
- Also stated in: § Morphological diversity of L5 SST interneurons · plain
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“L5 fanning-out Martinotti cells also inhibit nearby PCs but they target the dendrite mainly in L2/3”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Electrophysiological properties of morphologically identified SST interneurons
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“showing a high IR, a low AP threshold , and a high adaptation index”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Electrophysiological properties of morphologically identified SST interneurons
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
What the sources say
- Not checked yetcites Ma et al., 2006
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plain- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“Moreover the first three principal components, accounting for 93% of the variance, are mostly represented by features of the ascending axon.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plaincorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“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.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Morphological diversity of L5 SST interneurons
plain- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Electrophysiological properties of morphologically identified SST interneurons
plaincorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“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.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Electrophysiological properties of morphologically identified SST interneurons
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“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.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Electrophysiological properties of morphologically identified SST interneurons
plainn = 10- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“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.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Discussion
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It may state two separate claims as one.
- Rests on an assumption: The linear discriminant classifier built from firing properties accurately infers the morphological class of L5 SST-INs.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Here we show that L5 non-Martinotti cells specifically target L4 neurons and avoid L5 pyramidal cells.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
plaindirect- Asserted: passes — The authors put it forward as true.
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The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
- Rests on an assumption: Putative synaptic contacts identified morphologically correspond to functional synapses.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Indeed, neither calretinin nor calbindin are optimal molecular markers for fanning-out and T-shaped Martinotti respectively.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
plain- Asserted: passes — The authors put it forward as true.
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None of the sources cited for this claim have been checked yet.
In the paper
“SST-INs provide frequency-dependent feedback inhibition to pyramidal cells in supragranular and infragranular layers (Kapfer et al., 2007;Silberberg and Markram, 2007).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
What the sources say
- Not checked yetcites Kapfer et al., 2007
- Not checked yetcites Silberberg and Markram, 2007
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
What the sources say
- Not checked yetcites Li et al., 2013
- Not checked yetcites Hilscher et al., 2017
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
What the sources say
- Not checked yetcites Adesnik et al., 2012
- Not checked yetcites Veit et al., 2017
hedgedconditional- Asserted: passes — The authors put it forward as true.
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- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Long-range projecting nNos-expressing SST-INs have been described in L6 as well (He et al., 2016).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
What the sources say
- Not checked yetcites He et al., 2016
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
What the sources say
- Not checked yetcites Ma et al., 2006
- Not checked yetcites Xu et al., 2013
- Not checked yetcites Muñoz et al., 2017
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“However, L5 non-Martinotti tend to show a higher input resistance and seem to be less stuttering (Ma et al., 2006).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
What the sources say
- Not checked yetcites Ma et al., 2006
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
What the sources say
- Not checked yetcites Ma et al., 2006
- Not checked yetcites Muñoz et al., 2017
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The actions of different SST-INs subtypes are likely to have important computational roles in sensory perception during different behavioral states.”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: fails — It may be too vague to check against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“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).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
What the sources say
- Not checked yetcites Tan et al., 2008
- Not checked yetcites Hu and Agmon, 2016
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Indeed L4 non-Martinotti cells have been suggested to disinhibit L4 excitatory cells by primarily targeting L4 parvalbumin interneurons (Xu et al., 2013).”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of SST-INs subtypes
What the sources say
- Not checked yetcites Xu et al., 2013
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
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?
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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 citing paper
“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]
In the citing paper
“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 the citing paper
“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]
In the citing paper
“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]
In the citing paper
“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.
Rated major: it would touch a result — a value a result rests on, reported and recomputed p-values on opposite sides of the significance level, two passages that conflict with no sign which is right, or a cited work retracted before this paper was published.
In the paper
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Electrophysiological properties of morphologically identified SST interneurons
“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”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Diversity of cortical SST-INs
“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.”
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
“fanningout to pyramidal: 53.6%, 9 of 16”
§ Systems/Circuits Diversity and Connectivity of Layer 5 Somatostatin-Expressing Interneurons in the Mouse Barrel Cortex > Connectivity of morphological SST groups
Reported 53.6 · recomputed 56.25
Citations
Cites
GABAergic Interneurons in the Neocortex: From Cellular Properties to Circuits
Tremblay, R., Lee, S., & Rudy, B. (2016) · Neuron · doi:10.1016/j.neuron.2016.06.033 · Cited by 2,033
Self-citationGABAergic cell subtypes and their synaptic connections in rat frontal cortex
Kawaguchi, Y. (1997) · Cerebral Cortex · doi:10.1093/cercor/7.6.476 · Cited by 1,416 · cited 1×
“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×
“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).”
BackgroundDisynaptic Inhibition between Neocortical Pyramidal Cells Mediated by Martinotti Cells
Silberberg, G., & Markram, H. (2007) · Neuron · doi:10.1016/j.neuron.2007.02.012 · Cited by 768 · cited 1×
“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×
“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×
“An interesting approach might be to exploit the combination of patch-clamp recordings with single cell sequencing techniques to obtain more specific molecular markers in morphologically and/or electrophysiologically identified cells (Cadwell et al., 2016; Fuzik et al., 2016).”
BackgroundTargeting cells with single vectors using multiple-feature Boolean logic
Fenno, L. E., Mattis, J., Ramakrishnan, C., Hyun, M., Lee, S. Y., He, M., Tucciarone, J., Selimbeyoglu, A., Berndt, A., Grosenick, L., et al. (2014) · Nature Methods · doi:10.1038/nmeth.2996 · Cited by 557
Wang, Y., Toledo‐Rodriguez, M., Gupta, A., Wu, C., Silberberg, G., Luo, J., & Markram, H. (2004) · The Journal of Physiology · doi:10.1113/jphysiol.2004.073353 · Cited by 491 · cited 6×
“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×
“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×
“(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).”
BackgroundResultInfluential
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
“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
“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
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 299
“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
Metadata
- 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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