Theta nested Gamma

A 5 Hz afferent-rate envelope compared with constant excitation of the same mean.

Abstract

Independent excitatory afferents drive a recurrent excitatory–inhibitory circuit. We compare a 5 Hz sinusoidal rate envelope ranging from 0.2 to 3.8 Hz per afferent with constant 2 Hz input. The connectivity, weights and initial state are paired across conditions. Over 200–800 ms, mean E firing was 16.76 versus 15.82 Hz, and amplitude-weighted phase concentration was 0.526 versus 0.077. This single-seed comparison does not establish an independent gamma oscillator.

Question

Does a theta-frequency envelope on otherwise independent afferent excitation organise fast population bursting? We extend the circuit in Susin and Destexhe (2021) with periodic input and a constant-input comparison. The circuit is a modified conductance-based LIF version of the published PING network.[1]

Both populations receive excitatory afferents. From the start of the simulation, with no burn-in, the theta condition follows

λ(t)=λ0+Asin⁡(2πfθt).(1)\lambda(t)=\lambda_0+A\sin(2\pi f_\theta t). \tag{1}

Here tt is time in seconds from initialization, λ(t)\lambda(t) is the rate of each afferent in spikes/s, λ0=2\lambda_0=2 spikes/s is the mean, A=1.8A=1.8 spikes/s is the modulation amplitude, and fθ=5f_\theta=5 Hz is the envelope frequency. The rate ranges from 0.2 to 3.8 Hz and starts at its rising mean crossing. The comparison remains at 2 Hz. The two conditions therefore have the same prescribed mean over complete cycles; their realised spike counts and neuronal firing rates need not match.

Results

Recorded activity

Figure 2: Select the 5 Hz theta-modulated or constant-input recording. Each replays the complete one-second simulation from initialization, with no burn-in. A: Private inputs and population spikes are shown for 160 E-targeted and 40 I-targeted channels and their receiving neurons. Each channel aggregates 400 independent excitatory afferents. Blue denotes E-targeted channels and E neurons; rose denotes I-targeted channels and I neurons. Both private inputs are excitatory. Displayed recurrent edges exist in the simulated circuit. Pulses travel over the 1.5 ms synaptic delay; highlights last 3 ms for visibility. B: Pistons show full-population mean excitatory and inhibitory conductances onto E neurons and mean E/I voltages at the same timestep. Both conditions share conductance and voltage scales. C: Prescribed per-afferent rates, with a moving time cursor; E- and I-targeted envelopes coincide. The selector changes the recorded condition, while speed changes playback only. At 1×, one recorded second plays over 25 seconds. Positions and pulse appearance are display choices.

Paired comparison

We measured activity from 0.2 to 0.8 seconds, spanning three complete theta cycles and excluding 200 ms at each end from measurement. The full simulation, including the initial transient, is shown; no burn-in was simulated. Mean E firing rates were 16.76 Hz with theta input and 15.82 Hz with constant input; mean I rates were 19.67 and 18.29 Hz, respectively.

Paired afferent-rate traces, full E/I rasters, and unsmoothed population spike counts, all spanning one second for theta-modulated and constant input.

Figure 3: Fresh simulations with theta-modulated input on the left and constant input on the right. A/E: prescribed per-afferent rates, black solid for E-targeted and red dashed for I-targeted inputs; the two traces coincide. B/F: all 1,600 E neurons in black below 400 I neurons in red over 5 theta periods. D/H: unsmoothed population spike counts in 1 ms bins. Every panel spans the full 0–1,000 ms recording. E contains four times as many cells as I. Axes share scales across conditions. Time starts at initialization with no burn-in; both conditions use seed 8.

MeasurementTheta inputConstant input
E spectral peak within 30–90 Hz44 Hz40 Hz
I spectral peak within 30–90 Hz44 Hz40 Hz
E amplitude-weighted phase concentration0.5260.077
I amplitude-weighted phase concentration0.5270.079
Table 1.

Descriptive measurements from one paired seed. Spectral peaks are raw frequency-bin maxima in the specified search interval, with 2 Hz bin spacing. Phase concentration measures how strongly the 30–90 Hz amplitude envelope is concentrated at the prescribed theta phase; the same reference clock is assigned to the constant condition. Values range from zero to one. No significance test or surrogate correction was applied.

These observations characterise this circuit under the specified input envelopes. They do not establish a separate gamma oscillator: sharp theta-locked bursts can contribute harmonics to the filtered band, and a maximum at a search boundary is not evidence of an isolated spectral peak. Repeated seeds, spectral-background checks and feedback interventions would be needed for a stronger mechanistic claim.

Methods

  1. Pair the input conditions. Reuse connectivity, weights and the initial state; generate target-specific Poisson counts with the same random streams. Only the prescribed source-rate envelope differs. Parameters are listed in the appendix.
  2. Measure population rates and spectra. Count all spikes in 1 ms bins and divide by population size and bin duration. Use 200–800 ms for rate and spectrum summaries. Welch spectra use a Hann window, 500 ms segments and 50% overlap; report the raw maximum within 30–90 Hz, with 2 Hz bin spacing.
  3. Measure phase concentration. Apply a fourth-order Butterworth 30–90 Hz bandpass in second-order sections forward and backward over the full recording. Take the Hilbert-transform magnitude as the amplitude envelope, then crop to the measurement interval. Concentration is the magnitude of the amplitude-weighted mean unit phase vector at the prescribed 5 Hz phase. The constant condition uses the same clock. This is descriptive and has no surrogate correction.

Network diagrams

Structural diagram of 1,600 excitatory and 400 inhibitory neurons, all four recurrent pathways, and private excitatory afferents to both populations.

Figure A1: Shared circuit architecture for both conditions. Each E or I neuron receives its own aggregate excitatory input channel, representing 400 independent Poisson afferents. Recurrent excitation uses AMPA synapses and inhibition uses GABA synapses. Only the afferent-rate envelope differs between conditions; the connectivity and weights are identical. Loops on the E and I boxes represent connections between neurons within each population; individual neurons do not connect to themselves.

Parameters

ParameterRecorded value
Network
Native parameter snapshotUnavailable
Input channelsUnavailable
PopulationsUnavailable
Membrane modelUnavailable
Neurons
Membrane capacitanceUnavailable
Leak conductanceUnavailable
Resting potentialUnavailable
Reset potentialUnavailable
ThresholdUnavailable
Refractory periodUnavailable
Membrane time constantUnavailable
Synapses and weights
Native synapse and weight parametersUnavailable
Input and simulation
External afferents per neuron400
Baseline E source rate2 Hz
Baseline I source rate2 Hz
Theta envelope frequency5 Hz
Envelope amplitude per afferent1.8 Hz
Analysis start200 ms
Analysis end800 ms
Seed8
Timestep0.1 ms
Duration1000 ms
Burn-in0 ms
Analysis
amplitudefourth-order Butterworth 30–90 Hz SOS forward-backward filter; Hilbert magnitude; computed on full recording, then cropped away from endpoints
analysis interval ms[200,800]
concentrationabsolute amplitude-weighted mean phase vector; descriptive, no surrogate significance test
limitationsone paired seed; rate matching is prescribed input expectation, not realized count or output-rate matching; harmonics of theta-locked bursts can contribute to bandpassed amplitude; no independent gamma mechanism established
phaseprescribed 5 Hz sine phase, radians; same reference phase assigned to constant comparison
phase profilemean gamma-band amplitude within 18 equal prescribed-phase bins
spectrumWelch of 1 ms per-neuron population rates; Hann; 500 ms segments; 50% overlap; raw bin maximum in 30–90 Hz; 2 Hz bin spacing
Table A1. Model parameters follow the selected presentation to its original compute's native snapshot; input and protocol settings come from that compute's recorded configuration. Paired neuron values follow E / I order. Missing snapshots or fields are unavailable. Historical figures retain their original evidence; current recipe defaults are never substituted.

Dataset

Upstream: —

Downstream: —

RunDateDurationSizeEvidenceStorageRan onVerification
exp130-r026-present1s6.3 MBCurrentR2LocalLocal: absent; R2: catalog-verified
exp130-r025-analyse<1s431.4 KBCurrentR2LocalLocal: absent; R2: catalog-verified
exp130-r024-compute14s5.1 MBCurrentR2LocalLocal: absent; R2: catalog-verified

References

  1. Susin E, Destexhe A (2021). Integration, coincidence detection and resonance in networks of spiking neurons expressing Gamma oscillations and asynchronous states. PLOS Computational Biology, 17(9), e1009416.