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The ERP Boot Camp. What Are ERPs and What Are They Good For?. Overview. A bit of history Review : Basic neurophysiology & electricity Neural origins of ERPs Comparison of ERPs with other techniques. CNV. The Dawn of History. 1964: Gray Walter and the CNV. No Task: Click Only.

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the erp boot camp

The ERP Boot Camp

What Are ERPs and What Are They Good For?

  • A bit of history
  • Review: Basic neurophysiology & electricity
  • Neural origins of ERPs
  • Comparison of ERPs with other techniques
the dawn of history


The Dawn of History
  • 1964: Gray Walter and the CNV

No Task: Click Only

No Task: Flashes Only

No Task: Click followed by flashes

Task: Press button when flashes start

Walter, W. G., Cooper, R., Aldridge, V. J., McCallum, W. C., & Winter, A. L. (1964). Contingent negative variation: An electric sign of sensorimotor association and expectancy in the human brain. Nature, 203, 380-384.

which way is up
Which Way is Up?

René Descartes

the birth of the p300



The Birth of the P300
  • 1965: Sutton, Braren, Zubin, & John

Sound-Elicited ERPs


Cue stimulus indicating whether click or flash was likely

Delay of 3-5 seconds: Subject guesses whether stimulus will be click or flash

Click or flash occurs

Light-Elicited ERPs

Sutton, S., Braren, M., Zubin, J., & John, E. R. (1965). Evoked potential correlates of stimulus uncertainty. Science, 150, 1187-1188.

the oddball paradigm
The Oddball Paradigm

Typical Task Alternatives:

Count oddballs

Press only for oddballs

Different buttons for standard, oddball

= Standard

= Deviant/Oddball/Target

some terminology
Some Terminology

Stimulus Onset Asynchrony



Interstimulus Interval



Intertrial Interval



the next 4 decades
The Next 4 Decades
  • 1970s
    • Development and standardization of methods
    • Characterization of P3 and application to psychopathology
    • Selective attention (Hillyard, Näätänen, Harter)
  • The 1980s
    • Kutas & Hillyard (1980) discover N400
    • ERPology transitions into cognitive neuroscience
    • Multichannel recordings (16+ electrodes) and BESA
  • The 1990s
    • More components: LRP, N2pc, ERN, N170, etc.
    • fMRI takes off, leading to predictions of the demise of ERPs
the next 4 decades1
The Next 4 Decades
  • 1970s
    • Development and standardization of methods
    • Characterization of P3 and application to psychopathology
    • Selective attention (Hillyard, Näätänen, Harter)
  • The 1980s
    • Kutas & Hillyard (1980) discover N400
    • ERPology transitions into cognitive neuroscience
    • Multichannel recordings (16+ electrodes) and BESA
  • The 1990s
    • More components: LRP, N2pc, ERN, N170, etc.
    • fMRI takes off, leading to predictions of the demise of ERPs
  • The 2000s
    • Time-frequency analyses become mainstream
    • The ERP Boot Camp!
some basics of neuroscience
Some Basics of Neuroscience
  • Resting membrane potential
    • -70 mV on inside of cell
  • Action potentials
    • Triggered when membrane potential goes sufficiently positive
    • Starts at axon hillock and travels down axon
    • Rarely contributes to scalp ERPs
  • Postsynaptic potentials (PSPs)
    • Neurotransmitter binds with receptor, opens ion channels
    • Excitatory: Positive charges move into cell
    • Inhibitory: Negative charges move into cell
    • The origin of most ERPs
some basics of electricity
Some Basics of Electricity
  • Current (I for Intensity; Amperes)
    • Movement of charges across space (coulombs per second)
    • Like rate of water coming out of a hose (liters per second)
  • Voltage (E for Electromotive Force; Volts)
    • Potential for charges to move
    • Like water pressure
  • Resistance (R; Ohms [Ω])
    • Resistance to movement of charges
    • Like having a skinny or blocked hose segment
  • Impedance (Z)
    • Resistance to the flow of alternating current (AC)
    • Combines resistance, capacitance, and inductance
some basics of electricity1
Some Basics of Electricity
  • Power (Watts) = EI (typically proportional to E2)
  • Ohm’s Law: E = IR
    • If resistance increases and current is constant, voltage increases!
    • If you keep total water flow constant but constrict part of the hose, the pressure increases
some basics of electricity2
Some Basics of Electricity
  • Electricity follows the path of least resistance

Overall R < lowest individual R

Overall R = sum of individual Rs

some basics of electricity3
Some Basics of Electricity
  • Measuring Electrode Impedances

Outside of Skin

Inside of Skin

Measuring between E1 and E8 gives you the sum of E1 and E8; which impedance is high?

Measuring between E1–E7 (in parallel) and E8 gives you the sum of E8 and less than the lowest of E1–E7

some basics of electricity4
Some Basics of Electricity
  • Induction
    • If you pass an electrical current through a conductor, a magnetic field will run around it (right-hand rule)
    • If you pass a magnetic field across a conductor, an electrical current is induced in the conductor
some basics of electricity5
Some Basics of Electricity
  • AC is “Alternating Current”
    • Changes fairly rapidly over time
    • Line current (50 or 60 Hz)
    • EEG
  • DC is “Direct Current”
    • Fairly constant “offset” in voltage
    • Batteries
    • Skin potentials
    • In discussions of amplifiers, DC can also mean “Direct Coupled” (as opposed to capacitively coupled)
      • Conveniently, this means that the amplifier can amplify DC (direct current) signals
where do erps come from
Where Do ERPs Come From?

Cortical pyramidal cell (basic input-output cell of cerebral cortex)

Excitatory transmitter released on apical dendrites causes positive charges to flow into dendrites

Net negative on outside of cell

Current flows through cell, completing the circuit

Polarity reverses with inhibitory transmitter

Polarity reverses with PSP on cell body and basal dendrites

Polarity at scalp also depends on orientation of the cortical surface and position of reference electrode

where do erps come from1
Where Do ERPs Come From?

Equivalent Current Dipole

To be recorded at a distance, large numbers of neurons must have similar voltage fields

where do erps come from2
Where Do ERPs Come From?

Open Field

Closed Field

Scalp-recorded potentials are possible only for layered structures with consistent orientations

Primarily cerebral cortex

What about:


Brainstem nuclei?

Superior colliculus?

Possible role of radial glia

Local Field Potentials

where do erps come from3
Where Do ERPs Come From?

Voltages spread through the brain by “volume conduction”

Nearly speed of light

Voltage everywhere except at positive-negative transition

Skull causes lateral spread (like spraying hose on cardboard)

magnetoencephalography meg
Magnetoencephalography (MEG)

Magnetic fields travel around electrical dipoles

The skull is transparent to magnetism -- less blurring

Deep and radial dipoles are invisible from outside the head

the superposition problem
The Superposition Problem


















Voltage at an electrode at time t is a weighted sum of all components that are active at time t

There is no foolproof way to recover the underlying components from the observed waveforms





what are erps good for
What are ERPs Good For?
  • Temporal resolution
    • Continuous measure of processing between S and R
    • Many processes occur in a given region at different times
  • Determine whether an experimental manipulation influenced Process A or Process B
    • Which ERP component was affected?
  • Identifying multiple neurocognitive processes
    • A given behavioral effect is often accompanied by multiple ERP effects (different components)
    • Easy to identify multiple processes contributing to behavior
  • Covert monitoring of processing
    • Processing can be measured without a behavioral response (or from subjects who cannot easily respond)
    • Did the brain do something that was not evident in behavior?
what are erps good for1
What are ERPs Good For?
  • Link to the brain
    • Under some conditions, neural systems can be identified
    • But people often draw unwarranted conclusions about underlying neural processes from ERPs
    • Many researchers desperately want ERPs to be like fMRI or single-unit recordings, but they are not
    • “Those English boys want to play the blues so bad—and they DO play it so bad” — Sonny Boy Williamson
    • “Those ERPers want to study the brain so bad—and they DO study it so bad” — Sonny Boy Luck

Sonny Boy Williamson

erps as biomarkers
ERPs as Biomarkers

See Luck et al. (2011, Biological Psychiatry)

  • ERPs are tightly tied to neurotransmission
    • MMN may reflect current flow through NMDA receptors
    • A change in ERPs reflects a change in PSPs (not mediated through hemodynamic response)
  • Rodent/primate models available for some components
    • Potentially useful as an assay in drug discovery
  • Easily tolerated by patients
  • Some paradigms have excellent stability, reliability
  • Relatively inexpensive, feasible for large-N studies
  • May be able to predict which patients will respond to a given treatment
  • Potential roadblocks
    • Individual differences, lack of quality assurance standards
erps are bad when
ERPs Are Bad When…
  • You desire certainty about the neuroanatomical locus of an effect
  • You are interested in activity that is slow (>~2 s) or not time-locked to a sudden, observable event
  • You cannot collect large numbers of trials in each critical condition
    • Long period of time between trials
    • Severe adaptation of response over trials
    • Huge number of different control conditions
    • Need to surprise subjects
  • Subjects make frequent head or mouth movements during the time period of interest
    • Speech is particularly bad (tongue has strong dipole)
erps and fmri
ERPs and fMRI
  • Spatial and temporal resolution
    • If you want to study the sequence of events from 0-500 ms poststimulus, use ERPs
    • If you want to differentiate V2 from V4, use fMRI
  • Cost
    • $10/hour versus $500/hour
  • Tolerability
    • Infants and children, obese people, anxious people
  • Specificity for postsynaptic potentials
    • Except in rare instances, ERPs reflect PSPs and not action potentials
    • fMRI reflects PSPs and APs, plus coupling with hemodynamic response
erps and fmri1
ERPs and fMRI
  • Sustained versus transient responses
    • Hard to look at slow, sustained activity with ERPs (no block designs with ERPs)
    • Low power for infrequent, brief effects with fMRI (hard to pick up a 50-ms effect that occurs every 6 seconds)
  • Components versus anatomical regions
    • Who really cares about N170, P300, N400, per se?
  • Flexibility
    • Scalp ERPs are generated by a small fraction of brain processes
    • fMRI can pick up anything that changes overall activity level
some terminology pet peeves
Some Terminology Pet Peeves
  • ERPs are not “evoked response potentials”
    • This is like “irregardless”
    • These are fine: Evoked Responses; Evoked Potentials; VEPs; AEPs; VERs; AERs;
  • Must put an “s” on the end of “ERP” if and only if you would say “event-related potentials”
    • “We recorded event-related potentials…”
    • “We recorded ERPs…”
    • Not “We recorded ERP…”
    • “We conducted an event-related potential experiment...”
    • “We conducted an ERP experiment…”
    • Not “We conducted an ERPs experiment…”