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Phase 3: Read intentExplainer14 min

Event-related potentials and oscillations

Two ways the brain's electrical activity carries information: a stereotyped waveform locked to an event, and a rhythm whose power or phase changes. Averaging, time-frequency analysis, and the difference between evoked and induced.

AssumesWhat the spectrum didProject B: The P300 spellerSpineDecoding / signal processing / MLComputational neuroscience

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The P300 speller worked by averaging: the same waveform, locked to each flash, grew while the noise shrank. The motor imagery classifier worked by band power: a rhythm got weaker over one hemisphere. Those are the two languages of EEG. An is a change in voltage locked in time to an event. An oscillation is a rhythm whose power or phase changes. They are analyzed differently, they can hide each other, and understanding both is the foundation of every decoder in this phase.

AveragingSimulated signal
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Figure 1. An ERP emerging from averaging. The average keeps what is the same on every trial and cancels what is not.

Averaging keeps what is locked

Add fifty epochs, each aligned to its stimulus, and divide by fifty. Anything with the same shape and timing on every trial survives. Anything with random timing, or random sign, shrinks toward zero at a rate of one over the square root of the number of trials. The survivor is the ERP. Its named bumps are called components: P1, N1, P2 (early sensory), N170 (faces), P300 (rare targets), N400 (semantic surprise), the ERN (errors), each with a typical latency, polarity, and scalp location.

Averaging assumes the response is the same every time. It is not, quite; latency jitters from trial to trial, which smears the average and makes fast components look smaller. That is a limitation to know about, not a reason to stop averaging.

Oscillations change in power and phase

Alpha does not have a fixed shape locked to anything; it waxes and wanes on its own schedule. Averaging the raw signal across trials cancels it, because its phase differs on every trial. To see it, compute power (square the band-passed signal, or take the spectrogram) on each trial first, then average the power. Now a rhythm that reliably gets weaker after a movement cue shows up as a dip in mu power, the , even though the raw average shows nothing.

So: average the signal to see ERPs; average the power to see oscillatory changes. Doing the wrong one shows nothing and leads people to conclude there is nothing.

Predict before you look

After a stimulus, alpha power over visual cortex drops by half for a second, with random phase on each trial. What does the ERP (the average of the raw signal) show?

Almost nothing. Random phase means alpha averages to zero whether its power is high or low. The change is only visible in the average of power, or in a time-frequency plot. This is the definition of an induced response: present in power, absent in the average. An evoked response is one that is phase-locked and survives averaging.

Time-frequency analysis

The or the , applied per trial and averaged, gives a picture with time along the bottom, frequency up the side, and colour for power relative to baseline. ERD appears as a blue patch in the mu band during movement. A rebound of beta after movement appears red. Gamma bursts during attention appear, weakly, and often turn out to be muscle. The trade the spectrum page described holds: short windows for time, long for frequency, and wavelets adjust the window to the frequency automatically.

Phase

Beyond power, oscillations have phase, and phase carries information. Inter-trial phase coherence measures whether a rhythm’s phase is consistent across trials at each time; a strong value right after a stimulus means the stimulus reset the rhythm. Phase-locking between two channels measures whether two regions oscillate in step, which is the most common (and most over-interpreted) measure of “connectivity.” And an ERP can itself be thought of as a phase reset of ongoing rhythms; the debate over whether that is what ERPs are has run for twenty years.

Where decoders live

P300 spellers decode an ERP: the classifier looks at the average-like shape of each epoch. Motor imagery decodes oscillations: the classifier looks at band power over channels. SSVEP decodes a driven oscillation: the classifier looks for a frequency. Knowing which language your paradigm speaks tells you what features to extract and what preprocessing will destroy the signal. High-pass at 1 Hz kills slow ERPs; averaging raw signal kills induced responses.

Deep dive Latency jitter and single-trial ERPs 3 min

If the P300 arrives at 350 ms on one trial and 420 on the next, the average is a broader, lower bump than any single trial’s. Methods exist to estimate the latency per trial (template matching, Woody filtering) and realign before averaging, at the risk of aligning noise. For BCIs, jitter is why classifiers that look at the whole 0 to 800 ms window beat any single-time-point measure.

Deep dive The gamma problem 3 min

Scalp “gamma” (above 30 Hz) is small, and muscle activity from the scalp, jaw, and eyes is large in exactly that band. Micro-saccades, tiny eye movements, produce a burst of muscle activity that looks like a gamma response around 200 ms after a stimulus. A run of high-profile scalp gamma findings turned out to be this. Treat scalp gamma with suspicion, and never report it without an eye-movement control.

Deep dive Connectivity claims 3 min

Two channels showing phase-locking at 10 Hz does not mean two brain regions are communicating. Volume conduction puts the same source on both electrodes with zero lag, which is perfect phase-locking of nothing. Measures that ignore zero-lag (imaginary coherence, phase-lag index) help. Source-space analysis helps more. A large fraction of published EEG connectivity is volume conduction.

Recall
What is the difference between an evoked and an induced response, and how do you see each?
Evoked responses are phase-locked to the event and survive averaging the raw signal (ERPs). Induced responses are changes in oscillatory power with random phase, invisible in the raw average, visible when power is computed per trial and then averaged.
Recall
Why does averaging the raw signal hide event-related desynchronization?
The mu rhythm's phase is random across trials, so it averages to zero regardless of its power; the power decrease only shows in the average of per-trial power.
Recall
Why is scalp gamma treated with suspicion?
Muscle activity from micro-saccades and the jaw is large in the same frequency range and time-locks to stimuli, producing fake gamma responses; several published findings were this.
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