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.
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The P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. Related: Event-related potential (ERP)Event-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. , Oddball paradigmOddball paradigm An experiment in which a rare target is mixed in among frequent standards; the rare one produces a P300. . Glossary entry speller worked by averaging: the same waveform, locked to each flash, grew while the noise shrank. The motor imageryMotor imagery Imagining a movement without making it, which changes rhythms over motor cortex in a way a classifier can sometimes detect. Related: Sensorimotor rhythm (mu)Sensorimotor rhythm (mu) An 8 to 13 Hz rhythm over motor cortex that weakens when you move, or imagine moving, the corresponding body part. , Event-related desynchronization (ERD)Event-related desynchronization (ERD) The drop in rhythmic power over a cortical area when it becomes active, such as mu weakening when you imagine moving your hand. , BCI inefficiencyBCI inefficiency The finding that around one in five people cannot control a motor-imagery BCI even after training; once called "BCI illiteracy," a label the field is retiring. . Glossary entry classifierClassifier An algorithm that assigns each trial a label, such as left hand or right hand, after being trained on labelled examples. Related: Linear discriminant analysis (LDA)Linear discriminant analysis (LDA) A simple, robust classifier that draws a straight boundary between classes; the workhorse of EEG decoding for two decades. , Support vector machine (SVM)Support vector machine (SVM) A classifier that finds the boundary with the widest margin between classes; strong on small datasets. , Cross-validationCross-validation Testing a model on data it never saw during training, repeatedly, so the reported accuracy means something. . Glossary entry worked by band power: a rhythm got weaker over one hemisphere. Those are the two languages of EEGElectroencephalography (EEG) Recording the brain's electrical activity from electrodes on the scalp, a few tens of microvolts of blurred, summed cortical activity. Related: Electrocorticography (ECoG)Electrocorticography (ECoG) Recording from electrodes placed on the surface of the brain under the skull, usually during epilepsy surgery evaluation; cleaner and sharper than EEG. , Magnetoencephalography (MEG)Magnetoencephalography (MEG) Recording the tiny magnetic fields produced by cortical currents with superconducting sensors in a shielded room; less blurred by the skull than EEG, and far more expensive. , Volume conductionVolume conduction The spreading of currents from a source through the conductive tissues of the head, which is why one source shows up on many electrodes and why scalp EEG is blurry. , Microvolt (µV)Microvolt (µV) One millionth of a volt. Scalp EEG is tens of microvolts; a AA battery is 1.5 million of them. . Glossary entry. An event-related potentialEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller., N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right., Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice., Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials.. Glossary entry 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 decoderDecoder The algorithm that maps neural signals to intended actions, such as a classifier or a Kalman filter. Related: ClassifierClassifier An algorithm that assigns each trial a label, such as left hand or right hand, after being trained on labelled examples. , Kalman filterKalman filter An estimator that blends a prediction from a motion model with a noisy measurement, weighting each by how much it trusts it; the decoder behind most intracortical cursor control. , Brain-computer interface (BCI)Brain-computer interface (BCI) A system that turns measured brain activity into control of something outside the body, from a cursor to a wheelchair to a speech synthesizer. . Glossary entry in this phase.
Averaging keeps what is locked
Add fifty epochsEpoch A fixed-length slice of the recording locked to an event, such as one second starting 200 milliseconds before each stimulus. Related: Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. , Baseline correctionBaseline correction Subtracting the mean of the pre-stimulus interval from each epoch so trials line up. . Glossary entry, 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 ERPEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. , N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. , Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. , Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. . Glossary entry. Its named bumps are called components: P1, N1, P2 (early sensory), N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. Related: Event-related potential (ERP)Event-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. . Glossary entry (faces), P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. Related: Event-related potential (ERP)Event-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. , Oddball paradigmOddball paradigm An experiment in which a rare target is mixed in among frequent standards; the rare one produces a P300. . Glossary entry (rare targets), N400 (semantic surprise), the ERNError-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. Related: Event-related potential (ERP)Event-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. . Glossary entry (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 spectrogramSpectrogram A spectrum computed over sliding windows and plotted against time, showing how frequency content changes. Related: Wavelet transformWavelet transform A time-frequency analysis that uses short wiggles at high frequencies and long ones at low, matching the trade-off to the frequency. , SpectrumSpectrum A plot of how much of a signal lives at each frequency. . Glossary entry) 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 event-related desynchronizationEvent-related desynchronization (ERD) The drop in rhythmic power over a cortical area when it becomes active, such as mu weakening when you imagine moving your hand. Related: Sensorimotor rhythm (mu)Sensorimotor rhythm (mu) An 8 to 13 Hz rhythm over motor cortex that weakens when you move, or imagine moving, the corresponding body part., Motor imageryMotor imagery Imagining a movement without making it, which changes rhythms over motor cortex in a way a classifier can sometimes detect.. Glossary entry, even though the raw average shows nothing.
So: average the signal to see ERPsEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. , N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. , Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. , Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. . Glossary entry; average the power to see oscillatory changes. Doing the wrong one shows nothing and leads people to conclude there is nothing.
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 spectrogramSpectrogram A spectrum computed over sliding windows and plotted against time, showing how frequency content changes. Related: Wavelet transformWavelet transform A time-frequency analysis that uses short wiggles at high frequencies and long ones at low, matching the trade-off to the frequency., SpectrumSpectrum A plot of how much of a signal lives at each frequency.. Glossary entry or the wavelet transformWavelet transform A time-frequency analysis that uses short wiggles at high frequencies and long ones at low, matching the trade-off to the frequency. Related: SpectrogramSpectrogram A spectrum computed over sliding windows and plotted against time, showing how frequency content changes.. Glossary entry, applied per trial and averaged, gives a picture with time along the bottom, frequency up the side, and colour for power relative to baseline. ERDEvent-related desynchronization (ERD) The drop in rhythmic power over a cortical area when it becomes active, such as mu weakening when you imagine moving your hand. Related: Sensorimotor rhythm (mu)Sensorimotor rhythm (mu) An 8 to 13 Hz rhythm over motor cortex that weakens when you move, or imagine moving, the corresponding body part. , Motor imageryMotor imagery Imagining a movement without making it, which changes rhythms over motor cortex in a way a classifier can sometimes detect. . Glossary entry 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 spectrumSpectrum A plot of how much of a signal lives at each frequency. Related: Fast Fourier transform (FFT)Fast Fourier transform (FFT) The algorithm that breaks a stretch of signal into the sinusoids that would add up to it, giving amplitude and phase at each frequency. , Power spectral density (PSD)Power spectral density (PSD) The spectrum expressed as power per hertz, the standard way to quantify how strong alpha is. . Glossary entry 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 ERPEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. , N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. , Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. , Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. . Glossary entry can itself be thought of as a phase reset of ongoing rhythms; the debate over whether that is what ERPsEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. , N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. , Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. , Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. . Glossary entry are has run for twenty years.
Where decoders live
P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. Related: Event-related potential (ERP)Event-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. , Oddball paradigmOddball paradigm An experiment in which a rare target is mixed in among frequent standards; the rare one produces a P300. . Glossary entry spellers decode an ERPEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. , N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. , Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. , Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. . Glossary entry: the classifierClassifier An algorithm that assigns each trial a label, such as left hand or right hand, after being trained on labelled examples. Related: Linear discriminant analysis (LDA)Linear discriminant analysis (LDA) A simple, robust classifier that draws a straight boundary between classes; the workhorse of EEG decoding for two decades. , Support vector machine (SVM)Support vector machine (SVM) A classifier that finds the boundary with the widest margin between classes; strong on small datasets. , Cross-validationCross-validation Testing a model on data it never saw during training, repeatedly, so the reported accuracy means something. . Glossary entry looks at the average-like shape of each epochEpoch A fixed-length slice of the recording locked to an event, such as one second starting 200 milliseconds before each stimulus. Related: Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. , Baseline correctionBaseline correction Subtracting the mean of the pre-stimulus interval from each epoch so trials line up. . Glossary entry. Motor imageryMotor imagery Imagining a movement without making it, which changes rhythms over motor cortex in a way a classifier can sometimes detect. Related: Sensorimotor rhythm (mu)Sensorimotor rhythm (mu) An 8 to 13 Hz rhythm over motor cortex that weakens when you move, or imagine moving, the corresponding body part. , Event-related desynchronization (ERD)Event-related desynchronization (ERD) The drop in rhythmic power over a cortical area when it becomes active, such as mu weakening when you imagine moving your hand. , BCI inefficiencyBCI inefficiency The finding that around one in five people cannot control a motor-imagery BCI even after training; once called "BCI illiteracy," a label the field is retiring. . Glossary entry decodes oscillations: the classifierClassifier An algorithm that assigns each trial a label, such as left hand or right hand, after being trained on labelled examples. Related: Linear discriminant analysis (LDA)Linear discriminant analysis (LDA) A simple, robust classifier that draws a straight boundary between classes; the workhorse of EEG decoding for two decades. , Support vector machine (SVM)Support vector machine (SVM) A classifier that finds the boundary with the widest margin between classes; strong on small datasets. , Cross-validationCross-validation Testing a model on data it never saw during training, repeatedly, so the reported accuracy means something. . Glossary entry looks at band power over channels. SSVEPSteady-state visual evoked potential (SSVEP) When you stare at something flickering at a fixed rate, visual cortex oscillates at that rate, and the frequency can be read straight out of the EEG spectrum. Related: Canonical correlation analysis (CCA)Canonical correlation analysis (CCA) Finding the combination of channels that correlates best with reference sinusoids at each candidate frequency; the standard SSVEP decoder. , Brain-computer interface (BCI)Brain-computer interface (BCI) A system that turns measured brain activity into control of something outside the body, from a cursor to a wheelchair to a speech synthesizer. . Glossary entry decodes a driven oscillation: the classifierClassifier An algorithm that assigns each trial a label, such as left hand or right hand, after being trained on labelled examples. Related: Linear discriminant analysis (LDA)Linear discriminant analysis (LDA) A simple, robust classifier that draws a straight boundary between classes; the workhorse of EEG decoding for two decades. , Support vector machine (SVM)Support vector machine (SVM) A classifier that finds the boundary with the widest margin between classes; strong on small datasets. , Cross-validationCross-validation Testing a model on data it never saw during training, repeatedly, so the reported accuracy means something. . Glossary entry 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 ERPsEvent-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. Related: P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. , N170N170 A negative deflection about 170 milliseconds after seeing a face, over the back of the head on the right. , Error-related negativity (ERN)Error-related negativity (ERN) A sharp negative wave about 50 to 100 milliseconds after you make a mistake, before you consciously notice. , Trial averagingTrial averaging Adding many time-locked epochs together so the consistent response grows and the random noise shrinks by the square root of the number of trials. . Glossary entry; averaging raw signal kills induced responses.
Deep dive Latency jitter and single-trial ERPs 3 min
If the P300P300 A positive bump about 300 milliseconds after a rare or meaningful stimulus, the basis of the classic BCI speller. Related: Event-related potential (ERP)Event-related potential (ERP) The small, stereotyped voltage waveform the brain produces in response to an event, invisible in a single trial and revealed by averaging many. , Oddball paradigmOddball paradigm An experiment in which a rare target is mixed in among frequent standards; the rare one produces a P300. . Glossary entry 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 BCIsBrain-computer interface (BCI) A system that turns measured brain activity into control of something outside the body, from a cursor to a wheelchair to a speech synthesizer. Related: DecoderDecoder The algorithm that maps neural signals to intended actions, such as a classifier or a Kalman filter. , Closed loopClosed loop A system in which the output feeds back to influence the input, such as a user adjusting their brain activity while watching the cursor it controls. . Glossary entry, jitter is why classifiersClassifier An algorithm that assigns each trial a label, such as left hand or right hand, after being trained on labelled examples. Related: Linear discriminant analysis (LDA)Linear discriminant analysis (LDA) A simple, robust classifier that draws a straight boundary between classes; the workhorse of EEG decoding for two decades. , Support vector machine (SVM)Support vector machine (SVM) A classifier that finds the boundary with the widest margin between classes; strong on small datasets. , Cross-validationCross-validation Testing a model on data it never saw during training, repeatedly, so the reported accuracy means something. . Glossary entry 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 conductionVolume conduction The spreading of currents from a source through the conductive tissues of the head, which is why one source shows up on many electrodes and why scalp EEG is blurry. Related: Current dipoleCurrent dipole The simplest model of a patch of active cortex: a tiny current source and sink close together, with a strength and a direction. , Forward problemForward problem Given a source in the head, compute the voltages on the scalp; it has a unique answer. , Surface LaplacianSurface Laplacian Subtracting the average of neighbouring electrodes from each electrode, which sharpens spatial detail and suppresses distant sources. . Glossary entry puts the same source on both electrodesElectrode The conductor that turns ion currents in tissue into electron currents in a wire; its chemistry and contact quality set the noise you live with. Related: Silver/silver-chloride (Ag/AgCl)Silver/silver-chloride (Ag/AgCl) The standard electrode material for biopotentials, because its chemistry produces a stable, low-noise, low-drift interface with salty gel. , Electrode impedanceElectrode impedance How hard it is to push current through the skin-electrode contact, typically 5 to 50 kilohms with gel and megohms dry, higher at low frequencies. , Half-cell potentialHalf-cell potential The DC voltage that appears wherever a metal meets an electrolyte; two electrodes with different half-cell potentials produce an offset that can swamp the signal. . Glossary entry 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 EEGElectroencephalography (EEG) Recording the brain's electrical activity from electrodes on the scalp, a few tens of microvolts of blurred, summed cortical activity. Related: Electrocorticography (ECoG)Electrocorticography (ECoG) Recording from electrodes placed on the surface of the brain under the skull, usually during epilepsy surgery evaluation; cleaner and sharper than EEG. , Magnetoencephalography (MEG)Magnetoencephalography (MEG) Recording the tiny magnetic fields produced by cortical currents with superconducting sensors in a shielded room; less blurred by the skull than EEG, and far more expensive. , Volume conductionVolume conduction The spreading of currents from a source through the conductive tissues of the head, which is why one source shows up on many electrodes and why scalp EEG is blurry. , Microvolt (µV)Microvolt (µV) One millionth of a volt. Scalp EEG is tens of microvolts; a AA battery is 1.5 million of them. . Glossary entry connectivity is volume conductionVolume conduction The spreading of currents from a source through the conductive tissues of the head, which is why one source shows up on many electrodes and why scalp EEG is blurry. Related: Current dipoleCurrent dipole The simplest model of a patch of active cortex: a tiny current source and sink close together, with a strength and a direction. , Forward problemForward problem Given a source in the head, compute the voltages on the scalp; it has a unique answer. , Surface LaplacianSurface Laplacian Subtracting the average of neighbouring electrodes from each electrode, which sharpens spatial detail and suppresses distant sources. . Glossary entry.
Explain what this page was about to your roommate in three sentences. No jargon they would not know.
We reveal locked waveforms by averaging the raw signal across about 50 trials, which keeps what repeats and shrinks random hiss by the square root of the trial count. We reveal rhythms by averaging power instead, because alpha with random phase cancels to near nothing in the raw mean even when its power drops by half for a second after a stimulus. We protect slow waves by keeping the lowest kept frequency near half a hertz and protect rhythm changes by never averaging raw traces when we hunt them.
Yours does not have to match. Check that the three ideas are there.