← Home

Perturbations: gamma gates rates not just mean inhibition

exp042 · 2 June 2026 · pdf

Abstract

exp025′s rate gap admits a cheap reading: PING fires less because the I-loop delivers more inhibition. This entry forecloses that reading and identifies what about the I-stream is doing the suppressing: qualitatively (rhythm vs mean), and quantitatively (which temporal precision is required). Scaffolded by ar009 §Leg 1 item 2.

Methods

Pure inference on the trained exp025 PING baseline (seed 42, 𝜃𝑢= off). For each batch the I-population spike tensor 𝒔base𝐼{0,1}𝑇×𝐵×𝑁𝐼 is recorded from a baseline forward pass, then an override tensor replaces it in a second pass via the exp037 hidden-perturbation hook. The E-population experiences only the override I-stream through 𝑊𝐼𝐸; the readout consumes the perturbed E spikes.

Every perturbation only moves spikes in time (or, for Poisson, redraws at the matched count) — none adds or removes them — so the mean per-cell I rate is matched to baseline by construction: exactly for phase-shuffle, and to within ≈ 3% for the jitter families across the range where each result is read. The one exception is cycle-coherent jitter at the largest 𝜎: a Gaussian block offset with 𝜎=100 ms displaces part of each burst past the ends of the fixed presentation window, where it is clamped and lost, so the realised I rate falls to 40.5 Hz (24% below the 53.2 Hz baseline). Realised I is therefore plotted on every sweep, and the strict same-mean-inhibition comparison (the compound figure below) is anchored at 𝜎=14 ms, where realised I is still within 3% of baseline on both arms.

Five perturbation families:

  1. Baseline: no override; trained PING dynamics.
  2. Cycle-coherent jitter: partition the trial into blocks of length 1/𝑓𝛾 (≈ 22.8 ms at the trained operating point from exp041). For each (trial, block), draw a single Gaussian offset Δ𝒩︀(0,𝜎2) and shift every I-spike in that block by Δ. Within-burst cross-cell synchrony is preserved exactly; only the placement of each burst is perturbed. Sweep 𝜎{0,1,3,7,14,21,28,42,60,100} ms.
  3. Per-I-cell jitter: draw an independent Gaussian offset Δ𝑏,𝑛,𝑘𝒩︀(0,𝜎2) for every I-spike and shift each spike by its own offset. Destroys within-burst cross-cell synchrony while keeping the mean per-cell rate; the within-burst counterpart of cycle-coherent jitter. Sweep 𝜎{0,0.5,1,2,5,9,14,21,50} ms.
  4. Phase-shuffle: per-trial permutation 𝜋𝑏 of the time axis applied to all I-cells together: 𝒔shuf[𝑡,𝑏,𝑛]𝐼=𝒔base[𝜋𝑏(𝑡),𝑏,𝑛]𝐼. Preserves cross-cell co-firing within a timestep; destroys all phase structure.
  5. Rate-matched Poisson: per-(trial, cell) Bernoulli with 𝑝=count𝑏,𝑛/𝑇. Destroys both temporal and cross-cell structure; tests the 𝑔𝑖 variance limit.

Results

A two-by-two panel; both top rasters use the same jitter magnitude, sigma 14 ms, differing only in the kind of jitter. Top row: two single-trial rasters of trained PING (E spikes black, I spikes red). Top left, per-I-cell jitter, where the I-bursts have dissolved into continuous asynchronous firing and E is silent. Top right, cycle-coherent jitter, where the I-bursts stay sharp but are displaced and E firing appears in the gaps; both panels have near-identical realised I rates. Bottom row: two twin-axis line plots of hidden E rate (black diamonds) and accuracy (red squares) versus jitter sigma, each with a grey realised mean I-rate trace. Bottom left (per-cell) E and accuracy fall to near zero while realised I stays flat near 53 Hz; bottom right (cycle-coherent) E rate rises and accuracy stays high, while the grey realised-I trace holds flat then droops at the largest sigma.
Figure 1: Two inference-time perturbations of the trained-PING I-stream, both holding the mean per-cell I rate fixed at ≈ 53.2 Hz, push the E rate in opposite directions, which a mean-inhibition account cannot produce. Both columns use the same jitter magnitude, σ = 14 ms — only the kind of jitter differs. Left column, smear the bursts: per-I-cell jitter (realised I 52.5 Hz) scatters the spikes within each burst, destroying synchrony while leaving the mean untouched; the burst dissolves into a continuous shunt, the E rate falls to zero, and accuracy collapses toward chance (9.9% at the Poisson limit). Right column, move the bursts: cycle-coherent jitter (realised I 51.7 Hz — within 3% of the left column) displaces each gamma burst bodily but keeps its within-burst synchrony; the I-stream opens gaps and the E rate rises from 9.1 Hz at baseline to 35.8 Hz, accuracy holding near 84.9%. Same jitter magnitude, same mean inhibition (both ≈ 53.2 Hz), opposite outcome: what gates the E rate is the timing of inhibition, the rhythm, not its average level. The cycle-coherent rise continues past the full phase-shuffle level to 66.3 Hz by σ = 100 ms (bottom-right sweep), but there the finite trial window truncates the most-displaced bursts and realised I falls 24%, so the strict rate-matched reading is taken at σ = 14 ms; see Methods.

Per-I-cell jitter

Twin-axis line plot: hidden E rate (black diamonds, left axis) and test accuracy (red squares, right axis) versus per-I-cell jitter sigma in milliseconds on a symlog axis. Both fall steeply from baseline at small sigma; E rate is essentially zero by 5 ms and accuracy reaches chance by 9 ms. A grey line shows the realised mean I rate, flat near 53 Hz across the whole sweep and dipping only slightly to about 49 Hz by 50 ms.
Figure 2: Per-I-cell jitter sweep, three seeds. Each spike receives an independent Gaussian offset; mean per-cell I rate is preserved exactly. E rate (black diamonds, left axis) falls monotonically from baseline (9.1 Hz), already more than halved to 4.3 Hz by 𝜎=0.5 ms and essentially zero (0 Hz) by 𝜎5 ms, below 𝜏GABA=6 ms. Accuracy (red squares, right axis) holds at ≈ 89.2% up to 𝜎=0.5 ms, then collapses through 83.4% (σ = 1), 65.4% (σ = 2) and 17.7% (σ = 5), bottoming at chance (10.6%) by 𝜎=9 ms. The grey trace is the realised mean I rate, held flat near 53.2 Hz across the sweep: the E collapse happens under matched inhibition. The 𝜎 asymptote is the rate-matched Poisson regime: E silent, accuracy at chance.
Five stacked single-trial rasters of trained PING under per-I-cell jitter at sigma 0, 1, 5, 9 and 50 ms; E spikes black, I spikes red. Crisp vertical I-bursts at sigma 0 smear at sigma 1 and dissolve into continuous asynchronous I firing by sigma 5, while E firing goes silent.
Figure 3: Single trial replayed at five per-cell jitter levels. At 𝜎=0 the I-bursts are crisp vertical bands. At 𝜎=1 ms the bursts visibly smear into a few-ms-wide cluster and E firing already collapses to the low single digits (per-trial E annotated on each panel; sweep mean 1.7 Hz). At 𝜎5 ms the I-stream looks indistinguishable from a continuous low-variance shunt, and E is silenced. Per-cell jitter doesn’t release E; it destroys the bursty structure that gave E its recovery troughs in the first place.

Cycle-coherent jitter

Twin-axis line plot on a symlog sigma axis: hidden E rate (black diamonds, left axis) rises monotonically from about 9 Hz to 66 Hz as cycle-coherent jitter sigma grows, while test accuracy (red squares, right axis) declines gently from about 91 to 82 percent. A grey line shows the realised mean I rate: flat near 53 Hz up to about sigma 14 ms, then drooping to about 40 Hz by sigma 100 ms.
Figure 4: E rate (black diamonds) and accuracy (red squares) vs cycle-coherent jitter 𝜎, three seeds. As 𝜎 grows the displaced bursts open wider gaps and the E rate climbs from baseline (9.1 Hz) past the full phase-shuffle level (25.9 Hz, the reference with within-burst structure destroyed) to 66.3 Hz by 𝜎=100 ms, with the sharpest rise near the predicted transition timescale 𝜎=1/𝑓𝛾 ≈ 22.8 ms; accuracy declines only gently, holding near 81.7%. The grey trace is the realised mean I rate: it holds within 3% of baseline through 𝜎14 ms — where the E rate has already risen to 35.8 Hz — then droops to 40.5 Hz (24% below baseline) by 𝜎=100 ms, as the finite trial window truncates the most-displaced bursts. The strict same-mean-inhibition comparison is read at the smaller 𝜎, where the rate is matched and the E rise is already unambiguous.
Five stacked single-trial rasters of trained PING under cycle-coherent jitter at sigma 0, 7, 14, 28 and 100 ms; E spikes black, I spikes red. The red I-bursts stay sharp and vertical at every sigma but shift position, while E firing (black) grows denser and fills the widening gaps as sigma increases.
Figure 5: Single trial replayed at five jitter levels (𝜎=0,7,14,28,100 ms; seed 42, MNIST digit 0 sample 0). Per-trial E rate annotated on each panel. The I-bands stay vertical and crisp at every 𝜎: within-burst synchrony is preserved exactly. What changes is where each burst lands: at larger 𝜎 the bursts are displaced bodily from their phase-locked positions, opening longer gaps in the I-stream that E fires through, and the E rate climbs accordingly.

Next steps

Toroidal (wrapped) jitter, to extend the rate-matched range and disentangle release from truncation. The strict same-mean-inhibition claim is now anchored at 𝜎=14 ms, where realised I holds within 3% of baseline on both arms and the E rate has already risen to 35.8 Hz — the qualitative result stands on rate-matched ground. Beyond 𝜎30 ms, though, the cycle-coherent E-rate rise and the realised-I droop become confounded: some of the extra E firing is genuine gap-opening, and some is simply less inhibition delivered, because the finite window clamps and loses the most-displaced bursts. Wrapping each block offset modulo the trial length would restore exact spike-count preservation at every 𝜎 and separate the two, at the cost of re-injecting a wrapped burst at the opposite trial edge — a phase artifact of its own, so the wrapped sweep is a robustness check, not a replacement. The prediction: if the wrapped E rate still climbs past the phase-shuffle level (25.9 Hz), the release at large 𝜎 is real; if it flattens there, part of the 66.3 Hz overshoot at 𝜎=100 ms was the truncation. Either way the anchored rhythm-vs-mean conclusion is unaffected.