{
 "paper": "headley-2026-inhibitory-rhythms",
 "model": "supplied:runs/headley-2026-inhibitory-rhythms/plain-claim.answer.json",
 "prompt": "extract/prompts/plain-claim.md",
 "claims": [
  {
   "slug": "alt-distal-inhibition-raises-somatic-threshold",
   "role": "hypothesis",
   "panel": "",
   "plain": "Distal dendritic inhibition cuts firing the same way perisomatic inhibition does, by raising the spike threshold rather than by suppressing dendritic spikes."
  },
  {
   "slug": "alt-perisomatic-and-distal-share-mechanism",
   "role": "hypothesis",
   "panel": "",
   "plain": "Perisomatic and distal inhibition work through one shared mechanism, so perisomatic inhibition also controls dendritic spiking."
  },
  {
   "slug": "alt-phase-modulation-requires-buildup",
   "role": "hypothesis",
   "panel": "",
   "plain": "Phase-dependent control of dendritic spikes and spike timing needs many cycles of a rhythm to build up."
  },
  {
   "slug": "beta-bidirectional-dendritic-control",
   "role": "empirical",
   "panel": "fig5, fig7",
   "plain": "Beta-rhythm inhibition on distal dendrites both raises dendritic spiking in its troughs and suppresses it in its peaks."
  },
  {
   "slug": "beta-gates-distal-apical-inputs",
   "role": "empirical",
   "panel": "fig10",
   "plain": "Clustered apical inputs reach the soma when they arrive in the trough of a distal beta rhythm and are blocked at its peak."
  },
  {
   "slug": "beta-optimal-distal-dendritic-entrainment",
   "role": "empirical",
   "panel": "fig7",
   "plain": "Across a sweep of rhythm frequencies, beta near 20 Hz locks distal dendritic Ca²⁺ and NMDA spikes to phase most strongly."
  },
  {
   "slug": "burst-effects-emerge-first-cycles",
   "role": "empirical",
   "panel": "fig9",
   "plain": "Phase-dependent control of dendritic spikes and spike timing appears within the first few cycles of a beta or gamma burst."
  },
  {
   "slug": "ca-spikes-couple-20ms-before-ap",
   "role": "empirical",
   "panel": "fig2, fig3",
   "plain": "Ca²⁺ spikes in apical tuft dendrites lead somatic action potentials by about 20 ms, most tightly in the most distal compartments."
  },
  {
   "slug": "distal-inhib-drops-firing-02hz",
   "role": "empirical",
   "panel": "fig4, fig5",
   "plain": "Doubling distal dendritic inhibition nearly silences the neuron by shutting down dendritic Ca²⁺ and NMDA spikes, not by raising spike threshold."
  },
  {
   "slug": "ei-lag-sensitivity-firing-rate",
   "role": "empirical",
   "panel": "fig4",
   "plain": "Changing the delay between excitation and inhibition barely moves the firing rate but changes which dendrites drive the spikes."
  },
  {
   "slug": "gamma-gates-proximal-basal-inputs",
   "role": "empirical",
   "panel": "fig10",
   "plain": "Gamma-rhythm inhibition near the soma gates clustered proximal and basal inputs by phase while leaving distal apical inputs largely alone."
  },
  {
   "slug": "gamma-optimal-perisomatic-ap-modulation",
   "role": "empirical",
   "panel": "fig8",
   "plain": "Across a sweep of rhythm frequencies, gamma modulates the somatic spike threshold most strongly when inhibition sits near the soma."
  },
  {
   "slug": "gamma-perisomatic-no-dendritic-spike-change",
   "role": "empirical",
   "panel": "fig5",
   "plain": "Gamma rhythms near the soma shift the spike threshold by phase without changing how often dendritic spikes occur."
  },
  {
   "slug": "hypothesis-distinct-compartmental-roles",
   "role": "hypothesis",
   "panel": "hypothesis",
   "plain": "Perisomatic inhibition controls the soma's spike output, while distal dendritic inhibition controls dendritic spikes and their timing."
  },
  {
   "slug": "hypothesis-frequency-compartment-matching",
   "role": "hypothesis",
   "panel": "hypothesis",
   "plain": "A rhythm controls a compartment best when its cycle matches the local spike timescale — gamma at the soma, beta at distal dendrites."
  },
  {
   "slug": "interprets-pv-gamma-sst-beta-associations",
   "role": "literature-context",
   "panel": "fig10 synthesis / discussion",
   "plain": "Earlier work found that PV+ interneurons target the soma and accompany gamma, while SST+ interneurons target distal dendrites and beta."
  },
  {
   "slug": "l5-model-single-cell-scope",
   "role": "scope",
   "panel": "fig1A",
   "plain": "Every result comes from a compartmental model of one layer 5 pyramidal neuron, with no network or population effects simulated."
  },
  {
   "slug": "na-spikes-couple-2to3ms-before-ap",
   "role": "empirical",
   "panel": "fig2, fig3",
   "plain": "Na+ dendritic spikes in proximal compartments lead somatic action potentials by 2–3 ms, with coupling weakening farther from the soma."
  },
  {
   "slug": "naturalistic-drive-parameterization",
   "role": "scope",
   "panel": "fig1A (inset)",
   "plain": "Synaptic drive uses published experimental parameters and yields a typical in vivo firing rate, with no sensitivity analysis over those choices."
  },
  {
   "slug": "nmda-spikes-couple-25ms-before-ap",
   "role": "empirical",
   "panel": "fig2, fig3",
   "plain": "NMDA spikes lead somatic action potentials by about 25 ms, longer than Na+ spikes because their kinetics are slower."
  },
  {
   "slug": "perisomatic-inhib-drops-firing-07hz",
   "role": "empirical",
   "panel": "fig4, fig5",
   "plain": "Doubling perisomatic inhibition nearly silences the neuron by raising the spike threshold, while dendritic spikes carry on."
  },
  {
   "slug": "perisomatic-inhib-subtractive-divisive",
   "role": "empirical",
   "panel": "fig5, fig6",
   "plain": "Perisomatic inhibition lowers both the neuron's baseline firing and the slope of its input-output curve, compressing its dynamic range."
  },
  {
   "slug": "prediction-beta-optimal-distal",
   "role": "prediction",
   "panel": "prediction",
   "plain": "Distal inhibition should work best at beta, whose cycle matches the roughly 20–25 ms lead of dendritic spikes over somatic spikes."
  },
  {
   "slug": "prediction-distal-dendritic-spike-mechanism",
   "role": "prediction",
   "panel": "prediction",
   "plain": "Doubling distal dendritic inhibition should cut firing by suppressing apical Ca²⁺ and NMDA spikes, leaving the spike threshold alone."
  },
  {
   "slug": "prediction-gamma-optimal-perisomatic",
   "role": "prediction",
   "panel": "prediction",
   "plain": "Perisomatic inhibition should work best at gamma, whose cycles match the 2–3 ms lead of proximal Na+ spikes over somatic spikes."
  },
  {
   "slug": "prediction-orthogonal-input-gating",
   "role": "prediction",
   "panel": "prediction",
   "plain": "Beta at the distal dendrites should gate apical inputs and gamma at the soma should gate basal inputs, each working independently."
  },
  {
   "slug": "prediction-perisomatic-input-output-shaping",
   "role": "prediction",
   "panel": "prediction",
   "plain": "Perisomatic inhibition should both shift and flatten the somatic input-output curve rather than simply moving its operating point."
  },
  {
   "slug": "prediction-perisomatic-threshold-mechanism",
   "role": "prediction",
   "panel": "prediction",
   "plain": "Doubling perisomatic inhibition should cut firing by raising the somatic spike threshold, leaving dendritic Ca²⁺ and NMDA spikes intact."
  },
  {
   "slug": "pv-gamma-sst-beta-correspondence",
   "role": "interpretation",
   "panel": "fig10 (synthesis / discussion)",
   "plain": "The model explains why PV+ cells pair with gamma and SST+ cells with beta: each rhythm is optimal where that cell type makes its synapses."
  }
 ]
}