{ "paper": "headley-2026-inhibitory-rhythms", "claims": 29, "panels": { "total": [ "fig1a", "fig1b", "fig2a", "fig2b", "fig2c", "fig3a", "fig3b", "fig3c", "fig3d", "fig4a", "fig4b", "fig4f", "fig4g", "fig4h", "fig4i", "fig4c", "fig4d", "fig4e", "fig5a", "fig5b", "fig5c", "fig5s1a", "fig5s1b", "fig6", "fig6s1", "fig7a", "fig7b", "fig7c", "fig8a", "fig8b", "fig9a", "fig9b", "fig9c", "fig10a", "fig10b", "fig10c", "fig11a", "fig11b", "fig11c", "table1", "table2" ], "orphan": [ "fig1b", "fig2a", "fig2b", "fig2c", "fig3a", "fig3b", "fig3c", "fig3d", "fig4a", "fig4b", "fig4f", "fig4g", "fig4h", "fig4i", "fig4c", "fig4d", "fig4e", "fig5a", "fig5b", "fig5c", "fig5s1a", "fig5s1b", "fig6s1", "fig7a", "fig7b", "fig7c", "fig8a", "fig8b", "fig9a", "fig9b", "fig9c", "fig10a", "fig10b", "fig10c", "fig11a", "fig11b", "fig11c", "table1", "table2" ], "figure_level": [ "fig10", "fig2", "fig3", "fig4", "fig5", "fig7", "fig8", "fig9" ], "phantom": [], "pct": 4.9 }, "statistics": { "total": 0, "orphan": [], "pct": 100.0 }, "spans": { "segmented": 553, "obligations": 123, "textual": 430, "accounted": 24, "pct": 19.5, "orphans": [ { "uid": "results-003", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig1b" ] }, { "uid": "results-014", "section": "results", "text": "( B ) Examples of membrane potentials recorded simultaneously across the dendritic tree (in color) and soma (black) during naturalistic drive.", "stats": [], "panels": [ "fig1b" ] }, { "uid": "results-020", "section": "results", "text": "As a result, the basal and apical dendrites could generate Na + and NMDA spikes ( Figure 1B ; Goetz et al., 2021 ).", "stats": [], "panels": [ "fig1b" ] }, { "uid": "results-021", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig1c" ] }, { "uid": "results-023", "section": "results", "text": "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 et al., 2000 ).", "stats": [], "panels": [ "fig1c" ] }, { "uid": "results-025", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig1c" ] }, { "uid": "results-031", "section": "results", "text": "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 elicit action potentials.", "stats": [], "panels": [ "fig2a" ] }, { "uid": "results-036", "section": "results", "text": "( A ) Electrotonic distance between each dendritic compartment and the soma.", "stats": [], "panels": [ "fig2a" ] }, { "uid": "results-037", "section": "results", "text": "( B ) Dendritic compartments were grouped by their type (apical or basal) and electrotonic distance (percentile) from the soma.", "stats": [], "panels": [ "fig2b" ] }, { "uid": "results-040", "section": "results", "text": "( C ) Same format as B , but for NMDA spikes.", "stats": [], "panels": [ "fig2c" ] }, { "uid": "results-042", "section": "results", "text": "Dendritic compartments differed in their degree of passive electrical coupling to the soma (i.e. electrotonic distance; Figure 2A ).", "stats": [], "panels": [ "fig2a" ] }, { "uid": "results-052", "section": "results", "text": "Dendritic Na + spikes increased 2\u20133 ms prior to somatic action potentials in both basal and apical dendrites ( Figure 2B ).", "stats": [], "panels": [ "fig2b" ] }, { "uid": "results-055", "section": "results", "text": "The incidence of NMDA spikes increased ~25 ms prior to somatic action potentials, much earlier than seen with dendritic Na + spikes ( Figure 2C ).", "stats": [], "panels": [ "fig2c" ] }, { "uid": "results-059", "section": "results", "text": "This region is electrotonically close to the entire apical trunk, facilitating the propagation of Ca 2+ spikes ( Figure 3A ).", "stats": [], "panels": [ "fig3a" ] }, { "uid": "results-060", "section": "results", "text": "In our model, Ca 2+ spike occurrence increased within 20 ms of somatic action potentials ( Figure 3B ).", "stats": [], "panels": [ "fig3b" ] }, { "uid": "results-061", "section": "results", "text": "Furthermore, we found that NMDA spikes in the apical dendrites tended to precede Ca 2+ spikes ( Figure 3C ).", "stats": [], "panels": [ "fig3c" ] }, { "uid": "results-063", "section": "results", "text": "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.", "stats": [], "panels": [ "fig2c" ] }, { "uid": "results-065", "section": "results", "text": "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).", "stats": [], "panels": [ "fig3d" ] }, { "uid": "results-067", "section": "results", "text": "No such change was seen in basal dendrites ( Figure 3D , bottom).", "stats": [], "panels": [ "fig3d" ] }, { "uid": "results-069", "section": "results", "text": "( A ) Electrotonic distance between dendritic compartments and the apical nexus, where Ca 2+ spikes are generated.", "stats": [], "panels": [ "fig3a" ] }, { "uid": "results-070", "section": "results", "text": "( B ) Change in the incidence of Ca 2+ spikes at the nexus surrounding action potentials.", "stats": [], "panels": [ "fig3b" ] }, { "uid": "results-071", "section": "results", "text": "( C ) Percent change in NMDA spike presence in the apical dendrites centered on Ca 2+ spike initiation.", "stats": [], "panels": [ "fig3c" ] }, { "uid": "results-072", "section": "results", "text": "( D ) Percent change in NMDA spike coupling with action potentials during Ca 2+ spikes.", "stats": [], "panels": [ "fig3d" ] }, { "uid": "results-081", "section": "results", "text": "Both decreased the firing rate of the pyramidal cell from 5.5 Hz to less than 1 Hz ( Figure 4A ; control: 5.5\u00b10.85 Hz; distal: 0.20\u00b10.15 Hz; perisomatic: 0.70\u00b10.31 Hz; mean \u00b1 SD).", "stats": [], "panels": [ "fig4a" ] }, { "uid": "results-084", "section": "results", "text": "( A ) Action potential rate during periods with normal inhibitory tone (control), double rate on distal branches, or double rate on perisomatic.", "stats": [], "panels": [ "fig4a" ] }, { "uid": "results-086", "section": "results", "text": "( B ) Somatic excitability was measured by delivering current steps during the control, distal, and perisomatic inhibition states.", "stats": [], "panels": [ "fig4b" ] }, { "uid": "results-091", "section": "results", "text": "( C ) Impact of altered dendritic inhibition on rate of Na + spikes in apical and basal dendrites.", "stats": [], "panels": [ "fig4c" ] }, { "uid": "results-092", "section": "results", "text": "( D ) Same format as C , but for NMDA spikes.", "stats": [], "panels": [ "fig4d" ] }, { "uid": "results-093", "section": "results", "text": "( E ) Rate of Ca 2+ spikes in the apical dendrites.", "stats": [], "panels": [ "fig4e" ] }, { "uid": "results-096", "section": "results", "text": "( 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 perisomatic (blue) and distal (red) lags in inhibition from 0 to 500 ms. ( H ) Change in incidence of Ca 2+ spikes for distal (red, top) and proximal (blue, bottom) inhibition.", "stats": [], "panels": [ "fig4f", "fig4g", "fig4h" ] }, { "uid": "results-098", "section": "results", "text": "( I ) Same as ( H ) but for NMDA spikes.", "stats": [], "panels": [ "fig4h" ] }, { "uid": "results-100", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig4b" ] }, { "uid": "results-104", "section": "results", "text": "Although perisomatic inhibition produced the strongest subtractive effect, distal dendritic inhibition reduced firing rate the most ( Figure 4A ).", "stats": [], "panels": [ "fig4a" ] }, { "uid": "results-108", "section": "results", "text": "Perisomatic inhibition did not affect dendritic events compared to the control condition ( Figure 4C\u2013E ).", "stats": [], "panels": [ "fig4c", "fig4d", "fig4e" ] }, { "uid": "results-109", "section": "results", "text": "By contrast, dendritic inhibition decreased NMDA and Ca 2+ spikes ( Figure 4D and E ).", "stats": [], "panels": [ "fig4d", "fig4a", "fig4e" ] }, { "uid": "results-110", "section": "results", "text": "Na + spikes were relatively unaffected ( Figure 4C ).", "stats": [], "panels": [ "fig4c" ] }, { "uid": "results-119", "section": "results", "text": "To probe whether perisomatic or distal dendritic inhibition has distinct effects on E/I balance, we independently varied their lags ( Figure 4F\u2013I ).", "stats": [], "panels": [ "fig4f" ] }, { "uid": "results-121", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig4f" ] }, { "uid": "results-125", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig4g" ] }, { "uid": "results-126", "section": "results", "text": "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 ms. To better understand these effects, we calculated the cross-correlation (CC) between dendritic spikes and action potentials.", "stats": [], "panels": [ "fig4c" ] }, { "uid": "results-127", "section": "results", "text": "Increasing the lag decreased the coordination between Ca 2+ and somatic spikes ( Figure 4H ).", "stats": [], "panels": [ "fig4h" ] }, { "uid": "results-134", "section": "results", "text": "Thus, we emulated beta and gamma rhythmic input ( Figure 5A and F ).", "stats": [], "panels": [ "fig5a", "fig5d", "fig5f" ] }, { "uid": "results-135", "section": "results", "text": "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., 2018 ; Onorato et al., 2020 ; Adesnik, 2018 ; Murthy and Fetz, 1992 ).", "stats": [], "panels": [ "fig5b", "fig5a", "fig5d", "fig5g" ] }, { "uid": "results-140", "section": "results", "text": "( A ) Example data from the beta rhythmic inhibition simulation.", "stats": [], "panels": [ "fig5a" ] }, { "uid": "results-144", "section": "results", "text": "( B ) Action potential rate as a function of the phase of the beta rhythm.", "stats": [], "panels": [ "fig5b" ] }, { "uid": "results-146", "section": "results", "text": "( C ) Percent change in Ca 2+ spike presence at apical nexus by beta phase.", "stats": [], "panels": [ "fig5c" ] }, { "uid": "results-150", "section": "results", "text": "For all graphs, phase is given in radians with inhibition at a minimum for \u2013 \u03c0 and maximum at 0. Figure 5\u2014figure supplement 1. Phase-dependent effects on dendritic spikes of beta and gamma rhythmic inhibition delivered to opposite areas of the neuron.", "stats": [], "panels": [ "fig5s1" ] }, { "uid": "results-152", "section": "results", "text": "( A ) Action potential rate as a function of the phase of the beta rhythm.", "stats": [], "panels": [ "fig5a" ] }, { "uid": "results-154", "section": "results", "text": "( B ) Percent change in Ca 2+ spike presence at apical nexus by beta phase.", "stats": [], "panels": [ "fig5b" ] }, { "uid": "results-160", "section": "results", "text": "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\u2013E ).", "stats": [], "panels": [ "fig5c", "fig5d", "fig5e" ] }, { "uid": "results-162", "section": "results", "text": "In addition, the impact on Na + spikes was unexpected ( Figure 5E ), since delivery of the same inhibition tonically had little effect.", "stats": [], "panels": [ "fig5e" ] }, { "uid": "results-163", "section": "results", "text": "By contrast, gamma had virtually no effect on dendritic spikes ( Figure 5F\u2013H ).", "stats": [], "panels": [ "fig5f", "fig5g", "fig5h" ] }, { "uid": "results-167", "section": "results", "text": "During the peak phase, inhibition was greater than its mean rate, while during the trough phase, inhibition was lower (see Figure 5A and F ).", "stats": [], "panels": [ "fig5a", "fig5d", "fig5f" ] }, { "uid": "results-168", "section": "results", "text": "We found that the somatic action potential voltage threshold shifted lower during the \u2018trough\u2019 phase of gamma, when inhibition was at its weakest ( Figure 6A1 ) and without any change in the mean membrane voltage ( Figure 6A2 ).", "stats": [], "panels": [ "fig6a" ] }, { "uid": "results-171", "section": "results", "text": "During the \u2018peak\u2019 phase of beta, when inhibition was maximal, the threshold for evoking an action potential increased, which may reflect an \u2018off-path\u2019 shunting of excitatory current away from the soma and toward the dendrites ( Figure 6B1 ; Gidon and Segev, 2012 ).", "stats": [], "panels": [ "fig6b" ] }, { "uid": "results-172", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig6b" ] }, { "uid": "results-180", "section": "results", "text": "Figure 6\u2014figure supplement 1. Phase-dependent effects on somatic excitability of beta and gamma rhythmic inhibition delivered to opposite areas of the neuron.", "stats": [], "panels": [ "fig6s1" ] }, { "uid": "results-193", "section": "results", "text": "While phase modulation of firing rate was maintained with both rhythms, the overall level of spiking was dramatically reduced ( Figure 5\u2014figure supplement 1A and E ).", "stats": [], "panels": [ "fig5s1" ] }, { "uid": "results-194", "section": "results", "text": "Neither rhythm modulated Ca 2+ or NMDA spikes ( Figure 5\u2014figure supplement 1B, C, F, and G ).", "stats": [], "panels": [ "fig5s1" ] }, { "uid": "results-195", "section": "results", "text": "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, which last less than 1 ms, did show modulation by gamma rhythms delivered to the distal dendrites ( Figure 5\u2014figure supplement 1D and H ).", "stats": [], "panels": [ "fig5s1" ] }, { "uid": "results-197", "section": "results", "text": "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\u2014figure supplement 1A ).", "stats": [], "panels": [ "fig6s1" ] }, { "uid": "results-199", "section": "results", "text": "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\u2014figure supplement 1B ).", "stats": [], "panels": [ "fig6s1" ] }, { "uid": "results-206", "section": "results", "text": "Starting with distal dendrites, increasing inhibition frequency above 20 Hz diminished its entrainment of NMDA, Na + , and Ca 2+ spike onsets ( Figure 7A ).", "stats": [], "panels": [ "fig7a" ] }, { "uid": "results-211", "section": "results", "text": "( A ) Entrainment to an inhibitory rhythm delivered to the distal dendrites varied with its frequency.", "stats": [], "panels": [ "fig7a" ] }, { "uid": "results-214", "section": "results", "text": "( B ) Example voltage traces from dendritic compartments in either the distal basal or apical branches.", "stats": [], "panels": [ "fig7b" ] }, { "uid": "results-218", "section": "results", "text": "( 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.", "stats": [], "panels": [ "fig7c" ] }, { "uid": "results-222", "section": "results", "text": "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 peaks ( Figure 7B ).", "stats": [], "panels": [ "fig7b" ] }, { "uid": "results-223", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig7c" ] }, { "uid": "results-229", "section": "results", "text": "Lower frequency inhibition produced phase-dependent shifts in the mean membrane potential ( Figure 8A ).", "stats": [], "panels": [ "fig8a" ] }, { "uid": "results-232", "section": "results", "text": "By contrast, as frequency increased, the bias in momentary changes in the membrane potential diverged between peaks and troughs ( Figure 8B ).", "stats": [], "panels": [ "fig8b" ] }, { "uid": "results-239", "section": "results", "text": "( A ) The mean somatic membrane potential during either the trough or peak phase of the inhibitory rhythm.", "stats": [], "panels": [ "fig8a" ] }, { "uid": "results-240", "section": "results", "text": "( B ) Mean of the distribution of somatic membrane potential fluctuations as a function rhythm phase and frequency.", "stats": [], "panels": [ "fig8b" ] }, { "uid": "results-245", "section": "results", "text": "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).", "stats": [], "panels": [ "fig9a", "fig9d", "fig9f" ] }, { "uid": "results-247", "section": "results", "text": "( A ) Example data from the gamma rhythmic inhibition simulation.", "stats": [], "panels": [ "fig9a" ] }, { "uid": "results-251", "section": "results", "text": "( B ) Action potential rate as a function of the phase of the gamma rhythm.", "stats": [], "panels": [ "fig9b" ] }, { "uid": "results-253", "section": "results", "text": "( C ) Percent change in Ca 2+ spike presence at apical nexus by gamma phase.", "stats": [], "panels": [ "fig9c" ] }, { "uid": "results-259", "section": "results", "text": "Gamma bursts entrained spiking, with entrainment strongest during the middle of the burst ( Figure 9B ).", "stats": [], "panels": [ "fig9b" ] }, { "uid": "results-260", "section": "results", "text": "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 ).", "stats": [], "panels": [ "fig9c", "fig9d", "fig9e" ] }, { "uid": "results-261", "section": "results", "text": "Beta rhythms entrained somatic action potentials ( Figure 9G ), Ca 2+ spikes ( Figure 9H ), NMDA ( Figure 9I ), and Na + spikes ( Figure 9J ).", "stats": [], "panels": [ "fig9g", "fig9h" ] }, { "uid": "results-264", "section": "results", "text": "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 ( Iacaruso et al., 2017 ; Fu et al., 2012 ).", "stats": [], "panels": [ "fig10a" ] }, { "uid": "results-268", "section": "results", "text": "( A ) Schematic of the location for clustered excitatory synaptic inputs.", "stats": [], "panels": [ "fig10a" ] }, { "uid": "results-269", "section": "results", "text": "( 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 line) of the rhythm.", "stats": [], "panels": [ "fig10b" ] }, { "uid": "results-275", "section": "results", "text": "( C ) Summary of effects in panel B where the strength of each normalized cross-correlation was measured as its area under the curve.", "stats": [], "panels": [ "fig10c" ] }, { "uid": "results-281", "section": "results", "text": "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 phase, Figure 10B , top left).", "stats": [], "panels": [ "fig10b" ] }, { "uid": "results-282", "section": "results", "text": "Proximal inputs were either unaffected or moderately suppressed during the trough and suppressed during the peak ( Figure 10B , bottom left).", "stats": [], "panels": [ "fig10b" ] }, { "uid": "results-284", "section": "results", "text": "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).", "stats": [], "panels": [ "fig10b" ] }, { "uid": "results-285", "section": "results", "text": "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.", "stats": [], "panels": [ "fig10c" ] }, { "uid": "discussion-001", "section": "discussion", "text": "Discussion Arising from multiple interneuron subtypes, inhibition sculpts pyramidal neuron activity by acting at different membrane regions and distinct rhythmic frequencies ( Figure 11A ).", "stats": [], "panels": [ "fig11a" ] }, { "uid": "discussion-006", "section": "discussion", "text": "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 ).", "stats": [], "panels": [ "fig11b" ] }, { "uid": "discussion-007", "section": "discussion", "text": "In contrast, gamma modulated the threshold for action potential initiation, but only when delivered perisomatically ( Figure 11C ).", "stats": [], "panels": [ "fig11c" ] }, { "uid": "discussion-010", "section": "discussion", "text": "Figure 11. A summary schematic of the principal findings.", "stats": [], "panels": [ "fig11" ] }, { "uid": "discussion-011", "section": "discussion", "text": "( A ) The microcircuitry that was simulated in this study.", "stats": [], "panels": [ "fig11a" ] }, { "uid": "discussion-012", "section": "discussion", "text": "( B ) Beta rhythmic inhibition to the distal dendrites modulated dendritic spikes.", "stats": [], "panels": [ "fig11b" ] }, { "uid": "discussion-013", "section": "discussion", "text": "( C ) Gamma rhythmic inhibition to the perisomatic region modulated action potential initiation.", "stats": [], "panels": [ "fig11c" ] }, { "uid": "captions-052", "section": "captions", "text": "For all graphs, phase is given in radians with inhibition at a minimum for \u2013 \u03c0 and maximum at 0. [panels detected: a, b, c] === Figure 5s1 === Figure 5\u2014figure supplement 1. Phase-dependent effects on dendritic spikes of beta and gamma rhythmic inhibition delivered to opposite areas of the neuron.", "stats": [], "panels": [ "fig5s1" ] }, { "uid": "captions-066", "section": "captions", "text": "( B2 ) Same format as A2 , but for the beta rhythm. === Figure 6s1 === Figure 6\u2014figure supplement 1. Phase-dependent effects on somatic excitability of beta and gamma rhythmic inhibition delivered to opposite areas of the neuron.", "stats": [], "panels": [ "fig6s1" ] }, { "uid": "captions-112", "section": "captions", "text": "[panels detected: a, b, c] === Figure 11 === Figure 11. A summary schematic of the principal findings.", "stats": [], "panels": [ "fig11" ] }, { "uid": "tables-001", "section": "tables", "text": "Table 1. Inputs to layer 5 (L5) PN.", "stats": [], "panels": [ "table1" ] }, { "uid": "tables-007", "section": "tables", "text": "Divergence 2\u20138 2\u20138 ( 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\u00b10.22 0.53\u00b10.22 ( Br\u00e9maud et al., 2007 ) Inhibitory (perisomatic and somatic) Magnitude 162.5\u00b1103.1 pA 208.3\u00b158.7 pA ( Xiang et al., 2002 ) Firing rate 16.9\u00b114.3 Hz 16.9\u00b114.3 Hz ( Yu et al., 2019 ) Divergence 2.8\u00b11.9 2.8\u00b11.9 Number of synapses 406 406 ( Karimi et al., 2020 ) Release probability 0.88\u00b10.05 0.88\u00b10.05 ( Xiang et al., 2002 ) Inhibitory (basal) Magnitude 24.3\u00b118.4 pA 26.5\u00b11.6 pA ( Xiang et al., 2002 ) Firing rate 3.9\u00b14.9 Hz 3.9\u00b14.9 Hz ( Yu et al., 2019 ) Divergence 2.7\u00b11.6 2.7\u00b11.6 ( Tanaka et al., 2011 ; Thomson et al., 1996 ) Number of synapses 1023 1023 ( Karimi et al., 2020 ; Jadi et al., 2012 ) Release probability 0.72\u00b10.10 0.72\u00b10.10 ( Xiang et al., 2002 ) Inhibitory (apical) Magnitude 24.3\u00b133.1 pA 26.5\u00b11.6 pA ( Xiang et al., 2002 ) Firing rate 3.9\u00b14.9 Hz 3.9\u00b14.9 Hz ( Yu et al., 2019 ) Divergence 12\u00b13 12\u00b13 ( Silberberg and Markram, 2007 ; Vezoli et al., 2021 ) Number of synapses 1637 1637 ( Karimi et al., 2020 ) Release probability 0.30\u00b10.08 0.30\u00b10.08 ( Silberberg and Markram, 2007 ) Table 2. Short-term presynaptic plasticity.", "stats": [], "panels": [ "table2" ] } ] } }