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stale · v3 provisional awaiting approval

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for Spatially targeted inhibitory rhythms differentially affect neuronal integration · this layer across all papers · json

Provisional

This layer needs a corpus-scope decision that has not been ruled on yet, so what it produces would change if the decision changed. It waits on claim-format, relation-vocab.

Awaiting approval

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Out of date

These inputs changed after this ran:

  • claims/headley-2026-inhibitory-rhythms/ca-spikes-couple-20ms-before-ap.md
  • claims/headley-2026-inhibitory-rhythms/gamma-perisomatic-no-dendritic-spike-change.md
  • claims/headley-2026-inhibitory-rhythms/perisomatic-inhib-subtractive-divisive.md

What it produced99 spans.orphans

Read from coverage/headley-2026-inhibitory-rhythms.json · 33 KB. claims 29panels.pct 4.9statistics.total 0statistics.pct 100spans.segmented 553spans.obligations 123spans.textual 430spans.accounted 24spans.pct 19.5

# uidsectiontextstatspanels
1 results-003 results In brief, this model featured a multicompartmental dendritic tree that produced dendritic Na + , NMDA, and Ca 2+ spikes, along with somatic action potentials that could backpropagate ( Figure 1B ). [] ["fig1b"]
2 results-014 results ( B ) Examples of membrane potentials recorded simultaneously across the dendritic tree (in color) and soma (black) during naturalistic drive. [] ["fig1b"]
3 results-020 results As a result, the basal and apical dendrites could generate Na + and NMDA spikes ( Figure 1B ; Goetz et al., 2021 ). [] ["fig1b"]
4 results-021 results Dendritic Na + spikes were regenerative events lasting less than 1 ms that were not preceded by somatic action potentials ( Figure 1C1 ; Golding and Spruston, 1998 ). [] ["fig1c"]
5 results-023 results NMDA spikes occur when adjacent NMDA-bearing synapses were synergistically recruited by a combination of glutamatergic activation and local depolarization ( Figure 1C2 ; Larkum et al., 2009 ; Schiller… [] ["fig1c"]
6 results-025 results Ca 2+ spikes are depolarizations generated at the nexus of the apical trunk upon activation of voltage-gated Ca 2+ channels ( Figure 1C3 ; Schiller et al., 1997 ; Larkum and Zhu, 2002 ). [] ["fig1c"]
7 results-031 results L5 pyramidal neurons have a substantially longer apical trunk, which increases the electrotonic distance of their apical tuft from the soma ( Figure 2A ) and diminishes the ability of tuft synapses to… [] ["fig2a"]
8 results-036 results ( A ) Electrotonic distance between each dendritic compartment and the soma. [] ["fig2a"]
9 results-037 results ( B ) Dendritic compartments were grouped by their type (apical or basal) and electrotonic distance (percentile) from the soma. [] ["fig2b"]
10 results-040 results ( C ) Same format as B , but for NMDA spikes. [] ["fig2c"]
11 results-042 results Dendritic compartments differed in their degree of passive electrical coupling to the soma (i.e. electrotonic distance; Figure 2A ). [] ["fig2a"]
12 results-052 results Dendritic Na + spikes increased 2–3 ms prior to somatic action potentials in both basal and apical dendrites ( Figure 2B ). [] ["fig2b"]
13 results-055 results The incidence of NMDA spikes increased ~25 ms prior to somatic action potentials, much earlier than seen with dendritic Na + spikes ( Figure 2C ). [] ["fig2c"]
14 results-059 results This region is electrotonically close to the entire apical trunk, facilitating the propagation of Ca 2+ spikes ( Figure 3A ). [] ["fig3a"]
15 results-060 results In our model, Ca 2+ spike occurrence increased within 20 ms of somatic action potentials ( Figure 3B ). [] ["fig3b"]
16 results-061 results Furthermore, we found that NMDA spikes in the apical dendrites tended to precede Ca 2+ spikes ( Figure 3C ). [] ["fig3c"]
17 results-063 results Since NMDA spikes in the apical tuft normally have a weak relationship to somatic spiking ( Figure 2C ), they may elicit somatic spiking indirectly by driving Ca 2+ spikes. [] ["fig2c"]
18 results-065 results To test this, we measured how a Ca 2+ spike changed the spike-triggered average between apical tuft NMDA spikes and action potentials ( Figure 3D , top). [] ["fig3d"]
19 results-067 results No such change was seen in basal dendrites ( Figure 3D , bottom). [] ["fig3d"]
20 results-069 results ( A ) Electrotonic distance between dendritic compartments and the apical nexus, where Ca 2+ spikes are generated. [] ["fig3a"]
21 results-070 results ( B ) Change in the incidence of Ca 2+ spikes at the nexus surrounding action potentials. [] ["fig3b"]
22 results-071 results ( C ) Percent change in NMDA spike presence in the apical dendrites centered on Ca 2+ spike initiation. [] ["fig3c"]
23 results-072 results ( D ) Percent change in NMDA spike coupling with action potentials during Ca 2+ spikes. [] ["fig3d"]
24 results-081 results Both decreased the firing rate of the pyramidal cell from 5.5 Hz to less than 1 Hz ( Figure 4A ; control: 5.5±0.85 Hz; distal: 0.20±0.15 Hz; perisomatic: 0.70±0.31 Hz; mean ± SD). [] ["fig4a"]
25 results-084 results ( A ) Action potential rate during periods with normal inhibitory tone (control), double rate on distal branches, or double rate on perisomatic. [] ["fig4a"]
26 results-086 results ( B ) Somatic excitability was measured by delivering current steps during the control, distal, and perisomatic inhibition states. [] ["fig4b"]
27 results-091 results ( C ) Impact of altered dendritic inhibition on rate of Na + spikes in apical and basal dendrites. [] ["fig4c"]
28 results-092 results ( D ) Same format as C , but for NMDA spikes. [] ["fig4d"]
29 results-093 results ( E ) Rate of Ca 2+ spikes in the apical dendrites. [] ["fig4e"]
30 results-096 results ( F ) Examples of membrane potential recorded in control (top), and both distal (middle) and proximal (bottom) inhibition lagged by 500 ms. ( G ) Change in firing rate for control (black dot) and for … [] ["fig4f","fig4g","fig4h"]
31 results-098 results ( I ) Same as ( H ) but for NMDA spikes. [] ["fig4h"]
32 results-100 results A series of current pulses were injected into the soma to measure the relationship between firing rate and injected current (f-I curve), which captures the gain function of the neuron ( Figure 4B ). [] ["fig4b"]
33 results-104 results Although perisomatic inhibition produced the strongest subtractive effect, distal dendritic inhibition reduced firing rate the most ( Figure 4A ). [] ["fig4a"]
34 results-108 results Perisomatic inhibition did not affect dendritic events compared to the control condition ( Figure 4C–E ). [] ["fig4c","fig4d","fig4e"]
35 results-109 results By contrast, dendritic inhibition decreased NMDA and Ca 2+ spikes ( Figure 4D and E ). [] ["fig4d","fig4a","fig4e"]
36 results-110 results Na + spikes were relatively unaffected ( Figure 4C ). [] ["fig4c"]
37 results-119 results To probe whether perisomatic or distal dendritic inhibition has distinct effects on E/I balance, we independently varied their lags ( Figure 4F–I ). [] ["fig4f"]
38 results-121 results Increasing either of their lags by 500 ms produced obvious differences in the emission of dendritic spikes and their coordination with action potentials ( Figure 4F ). [] ["fig4f"]
39 results-125 results We systematically characterized these lag effects for the following spiking events modulated by tonic changes in inhibition: action potentials, Ca 2+ , and NMDA spikes ( Figure 4G ). [] ["fig4g"]
40 results-126 results Increasing the lag of perisomatic inhibition lowered action potential firing, while for distal dendritic inhibition, the firing rate decreased out to a lag of 125 ms and then returned to normal at 500… [] ["fig4c"]
41 results-127 results Increasing the lag decreased the coordination between Ca 2+ and somatic spikes ( Figure 4H ). [] ["fig4h"]
42 results-134 results Thus, we emulated beta and gamma rhythmic input ( Figure 5A and F ). [] ["fig5a","fig5d","fig5f"]
43 results-135 results Depths of modulation were set to similarly entrain action potentials ( Figure 5B and G ) and were comparable to spontaneous and optogenetically induced gamma and beta bursts seen in vivo ( Amir et al.… [] ["fig5b","fig5a","fig5d","fig5g"]
44 results-140 results ( A ) Example data from the beta rhythmic inhibition simulation. [] ["fig5a"]
45 results-144 results ( B ) Action potential rate as a function of the phase of the beta rhythm. [] ["fig5b"]
46 results-146 results ( C ) Percent change in Ca 2+ spike presence at apical nexus by beta phase. [] ["fig5c"]
47 results-150 results For all graphs, phase is given in radians with inhibition at a minimum for – π and maximum at 0. Figure 5—figure supplement 1. Phase-dependent effects on dendritic spikes of beta and gamma rhythmic in… [] ["fig5s1"]
48 results-152 results ( A ) Action potential rate as a function of the phase of the beta rhythm. [] ["fig5a"]
49 results-154 results ( B ) Percent change in Ca 2+ spike presence at apical nexus by beta phase. [] ["fig5b"]
50 results-160 results The phase of beta modulated the occurrence of Ca 2+ , NMDA, and Na + spikes, with each showing an ~75% depth of modulation with respect to their mean level ( Figure 5C–E ). [] ["fig5c","fig5d","fig5e"]
51 results-162 results In addition, the impact on Na + spikes was unexpected ( Figure 5E ), since delivery of the same inhibition tonically had little effect. [] ["fig5e"]
52 results-163 results By contrast, gamma had virtually no effect on dendritic spikes ( Figure 5F–H ). [] ["fig5f","fig5g","fig5h"]
53 results-167 results During the peak phase, inhibition was greater than its mean rate, while during the trough phase, inhibition was lower (see Figure 5A and F ). [] ["fig5a","fig5d","fig5f"]
54 results-168 results We found that the somatic action potential voltage threshold shifted lower during the ‘trough’ phase of gamma, when inhibition was at its weakest ( Figure 6A1 ) and without any change in the mean memb… [] ["fig6a"]
55 results-171 results During the ‘peak’ phase of beta, when inhibition was maximal, the threshold for evoking an action potential increased, which may reflect an ‘off-path’ shunting of excitatory current away from the soma… [] ["fig6b"]
56 results-172 results Additionally, there was a decrease in membrane voltage during the peak phase, which may correspond to decreased excitation arising from the suppression of dendritic spikes ( Figure 6B2 ). [] ["fig6b"]
57 results-180 results Figure 6—figure supplement 1. Phase-dependent effects on somatic excitability of beta and gamma rhythmic inhibition delivered to opposite areas of the neuron. [] ["fig6s1"]
58 results-193 results While phase modulation of firing rate was maintained with both rhythms, the overall level of spiking was dramatically reduced ( Figure 5—figure supplement 1A and E ). [] ["fig5s1"]
59 results-194 results Neither rhythm modulated Ca 2+ or NMDA spikes ( Figure 5—figure supplement 1B, C, F, and G ). [] ["fig5s1"]
60 results-195 results It is likely that the slow timescale of Ca 2+ and NMDA spikes, ~50 ms, is not optimal for the fast periodicity of the gamma rhythm, which cycles every ~15 ms. In agreement with this, Na + spikes, whic… [] ["fig5s1"]
61 results-197 results Gamma rhythmic inhibition on the dendrites had minimal or no impact on action potential threshold, but did shift the somatic membrane potential more negative ( Figure 6—figure supplement 1A ). [] ["fig6s1"]
62 results-199 results By contrast, delivering beta rhythmic inhibition to the soma raised the action potential threshold and hyperpolarized the membrane potential during the peak phase ( Figure 6—figure supplement 1B ). [] ["fig6s1"]
63 results-206 results Starting with distal dendrites, increasing inhibition frequency above 20 Hz diminished its entrainment of NMDA, Na + , and Ca 2+ spike onsets ( Figure 7A ). [] ["fig7a"]
64 results-211 results ( A ) Entrainment to an inhibitory rhythm delivered to the distal dendrites varied with its frequency. [] ["fig7a"]
65 results-214 results ( B ) Example voltage traces from dendritic compartments in either the distal basal or apical branches. [] ["fig7b"]
66 results-218 results ( C ) Percent change from the mean in the rate of dendritic spike onsets (red gradient) and offsets (blue gradient) as a function of rhythm frequency and phase. [] ["fig7c"]
67 results-222 results Indeed, examination of voltage traces in the dendrites during beta rhythmic inhibition revealed that NMDA and Ca 2+ spike onsets tended to occur during the trough, while offsets happened during the pe… [] ["fig7b"]
68 results-223 results To quantify this, we plotted the percent change in the probability of dendritic spike onsets and offsets with respect to both the phase and frequency of the inhibitory rhythm ( Figure 7C ). [] ["fig7c"]
69 results-229 results Lower frequency inhibition produced phase-dependent shifts in the mean membrane potential ( Figure 8A ). [] ["fig8a"]
70 results-232 results By contrast, as frequency increased, the bias in momentary changes in the membrane potential diverged between peaks and troughs ( Figure 8B ). [] ["fig8b"]
71 results-239 results ( A ) The mean somatic membrane potential during either the trough or peak phase of the inhibitory rhythm. [] ["fig8a"]
72 results-240 results ( B ) Mean of the distribution of somatic membrane potential fluctuations as a function rhythm phase and frequency. [] ["fig8b"]
73 results-245 results Gamma and beta bursts were delivered to the same model with mean depth of modulation like the tonic case ( Figure 9A and F ; see Methods for details). [] ["fig9a","fig9d","fig9f"]
74 results-247 results ( A ) Example data from the gamma rhythmic inhibition simulation. [] ["fig9a"]
75 results-251 results ( B ) Action potential rate as a function of the phase of the gamma rhythm. [] ["fig9b"]
76 results-253 results ( C ) Percent change in Ca 2+ spike presence at apical nexus by gamma phase. [] ["fig9c"]
77 results-259 results Gamma bursts entrained spiking, with entrainment strongest during the middle of the burst ( Figure 9B ). [] ["fig9b"]
78 results-260 results As with the tonically imposed rhythm, there was none or minimal modulation of Ca 2+ ( Figure 9C ), NMDA ( Figure 9D ), and Na + spikes ( Figure 9E ). [] ["fig9c","fig9d","fig9e"]
79 results-261 results Beta rhythms entrained somatic action potentials ( Figure 9G ), Ca 2+ spikes ( Figure 9H ), NMDA ( Figure 9I ), and Na + spikes ( Figure 9J ). [] ["fig9g","fig9h"]
80 results-264 results To examine this further, we added patches of concentrated excitatory synaptic inputs onto either the distal or proximal dendrites ( Figure 10A ), with densities similar to functional clusters in vivo … [] ["fig10a"]
81 results-268 results ( A ) Schematic of the location for clustered excitatory synaptic inputs. [] ["fig10a"]
82 results-269 results ( B ) Normalized cross-correlation between synaptic drive onto a clustered input and spiking at the soma, stratified by whether the presynaptic spike arrived during the peak (red line) or trough (blue… [] ["fig10b"]
83 results-275 results ( C ) Summary of effects in panel B where the strength of each normalized cross-correlation was measured as its area under the curve. [] ["fig10c"]
84 results-281 results Relative to the arhythmic Poisson inhibition case, beta rhythms enhanced the transmission of distal inputs when inhibition was low (trough phase) and suppressed them when inhibition was high (peak pha… [] ["fig10b"]
85 results-282 results Proximal inputs were either unaffected or moderately suppressed during the trough and suppressed during the peak ( Figure 10B , bottom left). [] ["fig10b"]
86 results-284 results It barely affected or moderately suppressed distal inputs ( Figure 10B , top right), while proximal inputs were enhanced during the trough and suppressed during the peak ( Figure 10B , bottom right). [] ["fig10b"]
87 results-285 results Summarizing these results ( Figure 10C ), we found that somatic spiking driven by clustered proximal synapses was bidirectionally modulated by gamma rhythms and suppressed by beta. [] ["fig10c"]
88 discussion-001 discussion Discussion Arising from multiple interneuron subtypes, inhibition sculpts pyramidal neuron activity by acting at different membrane regions and distinct rhythmic frequencies ( Figure 11A ). [] ["fig11a"]
89 discussion-006 discussion Beta rhythmic inhibition entrained dendritic spikes, focusing them into the phase when inhibition was at a minimum, but only when delivered to the distal dendrites ( Figure 11B ). [] ["fig11b"]
90 discussion-007 discussion In contrast, gamma modulated the threshold for action potential initiation, but only when delivered perisomatically ( Figure 11C ). [] ["fig11c"]
91 discussion-010 discussion Figure 11. A summary schematic of the principal findings. [] ["fig11"]
92 discussion-011 discussion ( A ) The microcircuitry that was simulated in this study. [] ["fig11a"]
93 discussion-012 discussion ( B ) Beta rhythmic inhibition to the distal dendrites modulated dendritic spikes. [] ["fig11b"]
94 discussion-013 discussion ( C ) Gamma rhythmic inhibition to the perisomatic region modulated action potential initiation. [] ["fig11c"]
95 captions-052 captions For all graphs, phase is given in radians with inhibition at a minimum for – π and maximum at 0. [panels detected: a, b, c] === Figure 5s1 === Figure 5—figure supplement 1. Phase-dependent effects on … [] ["fig5s1"]
96 captions-066 captions ( B2 ) Same format as A2 , but for the beta rhythm. === Figure 6s1 === Figure 6—figure supplement 1. Phase-dependent effects on somatic excitability of beta and gamma rhythmic inhibition delivered to … [] ["fig6s1"]
97 captions-112 captions [panels detected: a, b, c] === Figure 11 === Figure 11. A summary schematic of the principal findings. [] ["fig11"]
98 tables-001 tables Table 1. Inputs to layer 5 (L5) PN. [] ["table1"]
99 tables-007 tables Divergence 2–8 2–8 ( Markram et al., 1997 ; Reimann et al., 2015 ; Deuchars et al., 1994 ) Number of synapses 16070 16070 ( Karimi et al., 2020 ) Release probability 0.53±0.22 0.53±0.22 ( Brémaud et a… [] ["table2"]

Artifacts

Versions

From the run ledger. There is no changelog beside it to keep in step.

  1. v3 · 2026-09-13 · scripts/pipeline.py run

    ran via scripts/pipeline.py

    cd extract && python3 -m elife_extract.cli coverage --paper headley-2026-inhibitory-rhythms --json ../coverage/headley-2026-inhibitory-rhythms.json

  2. v2 · 2026-09-12 · scripts/pipeline.py run

    re-run after prepare v2

    cd extract && python3 -m elife_extract.cli coverage --paper headley-2026-inhibitory-rhythms --json ../coverage/headley-2026-inhibitory-rhythms.json

  3. v1 · 2026-09-11 · scripts/pipeline.py run

    coverage for the nine papers that had none

    cd extract && python3 -m elife_extract.cli coverage --paper headley-2026-inhibitory-rhythms --json ../coverage/headley-2026-inhibitory-rhythms.json

This layer across the corpus

Across the corpus

4 blocked upstream · 6 stale·a paper links to its own cell, where this layer's output for it is rendered

PaperStateVersionLast runOutputCell
A three-dimensional immunofluorescence atlas of the …ran via scripts/pipeline.pyblocked upstreamv32026-09-13artiushin-2026-spider-atlas.jsonjson
Distinct representational properties of cues and con…ran via scripts/pipeline.pystalev42026-09-13bouyeure-2026-fear-rsa.jsonjson
Computational modelling identifies key determinants …ran via scripts/pipeline.pystalev32026-09-13ejdrup-2026-dopamine.jsonjson
Contributions of insula and superior temporal sulcus…rebuild after edge completion passstalev72026-09-13gadeke-2026-guilt-insula.jsonjson
Spatially targeted inhibitory rhythms differentially…ran via scripts/pipeline.pystalev32026-09-13headley-2026-inhibitory-rhythms.jsonjson
Feedback of peripheral saccade targets to early fove…ran via scripts/pipeline.pyblocked upstreamv32026-09-13kammer-2026-foveal-feedback.jsonjson
iGABASnFR2 is an improved genetically encoded protei…ran via scripts/pipeline.pyblocked upstreamv52026-09-13kolb-2026-igabasnfr2.jsonjson
A deep learning pipeline for mapping in situ network…ran via scripts/pipeline.pyblocked upstreamv52026-09-13rozak-2026-neurovascular-dl.jsonjson
Self-association enhances early attentional selectio…ran via scripts/pipeline.pystalev32026-09-13scheller-2026-self-prioritization.jsonjson
Impaired excitability of fast-spiking neurons in a n…ran via scripts/pipeline.pystalev52026-09-13wengert-2026-kcnc1.jsonjson

Inputs and outputs

Produces
  • coverage/{paper}.json

One per paper — the table above links each one that exists.

Views
  • document — declared, and this artifact is not the shape this view needs
  • table — rendered above, over the 128 spans.orphans in the artifact

Running it

The command comes from the declaration, so this text and what actually runs cannot diverge. pipeline.py run also runs the unmet dependencies first. 

python3 scripts/pipeline.py run <paper> coverage

Underneath, that runs cd extract && python3 -m claim_graphs.cli coverage --paper {paper} --json ../coverage/{paper}.json.