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BCI -UNIT 2 Notesfinal - Notes

Computer Science (Government College Of Engineering, Tirunelveli)

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CBM342 BRAIN COMPUTER INTERFACE AND APPLICATIONS

UNIT II ELECTROPHYSIOLOGICAL SOURCES 9

Sensorimotor activity – Mu rhythm, Movement Related Potentials – Slow Cortical Potentials-P300 - Visual
Evoked Potential - Activity of Neural Cells - Multiple Neuromechanisms.

1. Introduction
 A challenge of BCI research is to be able to control complex devices using noninvasive recordings of
brain signals at high spatial and temporal resolution.
 The cerebral cortex is the area of greatest interest in BCI as
1. it is most accessible to electrode probes and scalp recording
2. it is highly involved in the executive function of motor and communication behaviours.
 The non-invasive recording of brain signals captures the changes in blood flow or fluctuations in
electric/magnetic fields caused by the activity of large populations of neurons
 Since EEG signals reflect the combined input to large populations of neurons, methods for building BCIs
from EEG signals rely on modulating the response of large neural populations either through subject
training over a period of time or through external stimuli that can activate large populations of
neurons.
1.1 Two methods for building BCIs from EEG signals
1. Self-paced (or asynchronous) BCI
 BCIs based on the subject can voluntarily initiate control at any time without being tied to a
stimulus.
 Self-paced BCIs typically utilize some form of imagery (motor or cognitive) that can generate a
robust and reliable EEG response after a period of training.
2. Stimulus-based BCIs (also called synchronous BCIs)
 Stimulus-based BCIs (also called synchronous BCIs) rely on detecting a stereotypical brain
response generated after the subject is presented with a stimulus (such as a flash) that is linked
to a BCI command or choice.
Thus the Control is not initiated by the subject but is tied to the presentation of stimuli by the
BCI.

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 Stimulus-based BCIs however are easier to use because they do not require training on the part
of the subject, and relatively high accuracies can be obtained for naïve subjects, compared to
imagery-based BCIs.

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Figure 1 Basic principle of a typical EEG BCI. (a) and (b) show the setup of a BCI.(c–f) Examples of EEG
signals used for BCI

2. ALL ELECTROPHYSIOLOGICAL SOURCES FOR BUILDING BCIS FROM EEG SIGNALS

1. Self-paced (or asynchronous) BCI

Self paced BCIs voluntarily initiate control at any time without being tied to a stimulus. They typically
utilize some form of imagery (motor or cognitive) that can generate a robust and reliable EEG response
after a period of training.
a. Oscillatory Potentials and ERD event-related desynchronization motor imagery-based BCIs, where
users modulate their brain activity to produce changes in oscillatory power.
b. Slow Cortical Potentials SCPs are type of electrical brain activity characterized by slow changes in
cortical voltage over time, typically lasting from several hundred milliseconds to several seconds. These

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potentials reflect changes in the excitability of the cortical neurons and are thought to be involved in
various cognitive processes.
c. Movement-Related Potentials Movement-related potentials (MRPs) are a type of event-related
potential (ERP) that occurs in the brain before, during, or after the execution of voluntary movements.
These potentials reflect the neural activity associated with motor planning, execution, and feedback
processes.

2. Stimulus-based BCIs (also called synchronous BCIs) A major class of EEG signals used in non-
invasive BCIs are evoked potentials (EPs), which are stereotypical EEG responses generated by the
brain when the sub ject is presented with a particular type of stimulus.
a. The P300 Potential: when a rare but task- relevant auditory, visual or somatosensory stimulus is
interspersed with fre quent and routine stimuli, the rare stimulus evokes a potential with a positive peak
at about 300 ms after the stimulus is presented. This potential is called the P300 (or P3) potential.
b. Visually Evoked Potentials (VEPs): VEPs are generated by visual stimuli such as flashing lights.
c. Auditory Evoked Potentials (AEPs): AEPs are generated by auditory stimuli such as clicks and tones.
d. Steady-State Visual Evoked Potential (SSVEP): SSVEP BCIs use visual stimuli flickering at different
frequencies. The user focuses on a specific flickering stimulus, eliciting a frequency-specific brain
response that can be detected and used for control.
e. Somatosensory Evoked Potentials (SSEPs): These BCIs use tactile or proprioceptive stimuli to evoke
brain responses. For example, vibrotactile stimuli applied to the skin can elicit event-related
desynchronization (ERD) or event-related synchronization (ERS) patterns that can be used for control.
3. Hybrid BCIs: These BCIs combine multiple types of stimuli or brain signals to improve performance
and usability. For example, a hybrid BCI may combine SSVEP and motor imagery signals for more
robust control.

3. Sensorimotor rhythms (SMRs)

 In relaxed awake people, the EEG recorded over primary sensorimotor cortical areas often displays 8–
12 Hz (mu-rhythm) and 18–26 Hz (beta-rhythm) activity. These sensorimotor rhythms (SMRs) are
thought to be produced by thalamocortical circuits.

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 The execution or imagination of limb movement induces changes in rhythmic activity recorded over
sensorimotor cortex.
 These changes in SMRs can be detected on the scalp by electroencephalography (EEG) or
magnetoencephalography (MEG) or on the surface of the brain by electrocorticography (ECoG)
 SMRs are oscillations in the electric or magnetic fields recorded over sensorimotor cortices. (ie)
Sensorimotor rhythms (SMR) refer to oscillations in brain activity recorded from somatosensory and
motor areas.
 SMR activity comprises a variety of different rhythms that are distinguished from each other by location,
frequency, and/or relationship to concurrent sensory input or motor output. Some beta rhythms are
harmonics of mu rhythms, while others are separable from mu rhythms by topography and/or timing.

Figure 2 Major brain regions

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Figure 3 Major areas and functional specialization of the neocortex

 Movement or preparation for movement is usually accompanied by SMR decrease, especially

contralateral to the movement. This decrease, known as event-related desynchronization (ERD). It
consists of a reduction in rhythmic activity related to an internally or externally paced event such as a
voluntary movement. SMRs decrease during motor behaviors. ERD is also referred to as blocking of a
rhythm. SMR ERDs can be seen as correlates of activated cortical networks.
 SMRs can also be increased in association with sensorimotor events (e.g., immediately after movement).
This is called event-related synchronization (ERS) Correspondingly, the increase of oscillatory activity
in a specific frequency band is called event-related synchronization (ERS). ERS can be seen as correlate
of a deactivated or inhibited cortical network.
 Furthermore, and most pertinent to BCI use, ERD and ERS do not require actual movement, they occur
also with motor imagery (i.e., imagined movement).
 SMRs typically fall into three major frequency bands: mu (8–12 Hz), beta (18–30 Hz), and gamma (30–
200+ Hz).
 EEG recording is largely limited to mu, beta and lower-frequency gamma activity, but ECoG and MEG
can detect higher-frequency gamma activity.
 Alpha activity recorded from sensorimotor areas is also called mu activity.
 Several laboratories have shown that people can learn to control mu or beta rhythm amplitudes in the
absence of movement or sensation.
 People with or without motor disabilities learn to control mu- and/or beta-rhythm amplitudes.

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Figure 5 SMR topographies and spectra from a BCI user controlling three-dimensional cursor movement

• Users initially employ motor imagery to control the cursor, but, as training proceeds, imagery usually
becomes less important, and users learn to move the cursor automatically.

• SMR-based BCI has been used to answer simple yes/no questions with accuracies >95%, to perform
basic word processing.

• The frequency bands that are most important for motor imagery are mu and beta in EEG signals.

• Activity invoked by right hand movement imagery is most prominent over electrode location C3. Left
hand movement imagery produces activity most prominent over [Link] is, activity invoked by hand
movement imagery is located on the contralateral (opposite) side.

• Foot movement imagery invokes activity over Cz.

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• A distinction between left and right foot movement is not possible in EEG because the corresponding
cortical areas are too close. Similarly, ERD/ERS patterns of individual fingers cannot be discriminated
in EEG.

• To produce patterns that can be detected, the cortical areas involved have to be large enough so that the
resulting activity is sufficiently prominent compared to the remaining EEG (background EEG).

• Hand areas, foot areas, and the tongue area are comparatively large and topographically different.
Therefore, BCIs have been controlled by imagining moving the left hand, right hand, feet, and tongue.

• When there is no movement, the spectrum at C4, which corresponds to the left hand, is displayed in the
following image.

Figure 6 Spectrum at C4 when there is no movement

 An imagined movement in left hand causes the SMR rhythm to decrease which is depicted below.

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Figure 7 Imagining movement of left limb causes local blocking of corresponding SMR

3.1 Mu rhythm

The mu rhythm and the beta rhythm are originated from the sensorimotor cortex, i.e., the area which is
primarily responsible for the control of hand and foot movements.
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Alpha waves (or the alpha rhythm) are electrical fluctuations in the range 8–13 Hz and can be measured in
EEG from the occipital region in awake persons when they are relaxed or their eyes are closed.

 A particular kind of alpha wave popular in BCI applications is known as the mu rhythm (8–12 Hz).
 Mu rhythm is found over sensorimotor areas in the absence of movement and is decreased or abolished
when the subject performs a movement or imagines performing a movement.

 Mu rhythms display two distinct ERD patterns. Lower frequency (8–10 Hz) mu-rhythm ERD occurs
during almost any kind of motor behaviour, is widespread over the entire sensorimotor cortex, and
probably reflects general motor preparation and attentional processes.
 In contrast, higher-frequency (10–13 Hz) mu-rhythm ERD is topographically restricted and is related
to task-specific aspects of performance. In sum, lower-frequency mu ERD appears to be nonspecific,
whereas higher-frequency mu ERD is topographically and functionally specific.
4. Slow Cortical Potentials(SCPs)
 The slowest features of the scalp-recorded EEG yet used in BCI systems are slow voltage changes
generated in cortex. These potential shifts occur over 0.5–10.0 s and are called slow cortical potentials
(SCPs).
 SMRs are EEG features that are analysed in the frequency domain (e.g., frequency on the x-axis) and
thus are often referred to as frequency-domain activity. Additional EEG features associated with motor
function are measured in the time domain and thus are referred to as time-domain activity.
 Movement or movement imagery is typically associated with relatively slow changes in the voltages
recorded over sensorimotor cortex in time domain. These are called slow cortical potentials (SCPs).
 SCPs are event-related potentials that are time-locked and phase-locked to specific sensorimotor events
(i.e., they occur at predictable times before, during, or after specific events).
 SCPs typically consist of negative potential shifts that precede actual or imagined movement or
cognitive tasks. They are thought to represent cortical activation in preparation for action. An SCP is
typically followed by a biphasic wave referred to as the movement-related potential.

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 In normal brain function, negative SCPs accompany mental preparation, while positive SCPs probably
accompany mental inhibition.
 Negative and positive SCPs probably reflect an increase and decrease, respectively, in excitation of
cortical neurons.
 The contingent negative variation is a slowly developing negative SCP that occurs between a warning
stimulus and a stimulus requiring a response [22]. It reflects increased activation and decreased firing
thresholds in cortical networks associated with the preparatory process preceding the response.
 Like SMRs, SCPs and related potentials over sensorimotor areas are associated with motor imagery as
well as with actual movements.
 They are thought to reflect a mechanism for local mobilization of excitation or inhibition in cortical
populations, caused by inputs from the thalamus.
 In one of the studies with 13 healthy subjects and 3 patients with total motor paralysis, EEG was
recorded from electrode locations Cz, C3, and C4 , and two channels were extracted: a Cz-linked
mastoid channel (i.e., 1/2 [(Cz-A1) + (Cz-A2)]) and a bipolar C3 minus C4 channel.
 The training task involved controlling a cursor to hit the top or bottom edge of the screen. The position
of the cursor was proportional to the difference between average baseline EEG amplitude and the
average EEG amplitude over the last 500 ms from the Cz channel.
 The baseline amplitude was calculated from an immediately preceding baseline period.
 Some subjects participated in a two-dimensional cursor task where the target could also be the left or
right edge of the screen. In this case, the horizontal position of the cursor was proportional to the
difference between average baseline EEG amplitude from the (C3−C4) channel and the average EEG
amplitude from this channel over the last 500 ms.

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Figure 10 Slow cortical potentials (SCPs) in healthy subjects. (Top 2 panels) Average SCP at Cz
and vertical electro-oculogram (vEOG), averaged during a single training session over 13
subjects. (Lower 2 panels) SCP difference between the left (C3) and right (C4) motor cortex and
horizontal EOG (hEOG), averaged across last 3 training sessions for 5 subjects.

Figure 11 SCP- based BCI in an ALS patient. (A) SCP and EOG for an ALS patient MP at the beginning of
training (left) and after training over several months (right). (B) Improvement in performance for the same

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patient over time, as revealed by an increasing percentage of hits (black dots) and decreasing percentage of
false positive (asterisks) over sessions spanning several months

 Figure 10 shows the average SCP waveforms generated by healthy subjects on cue after training. For
both channels, a clear deviation from baseline activity in the positive or negative direction can be seen:
this difference from baseline was used to proportionally move the cursor up/down or left/right. Out of
the 13 subjects, 4 were able to produce significant positive-going responses, 3 generated significant
negative-going responses, and 3 were able to generate both.
 Slow cortical potentials (SCPs) are slow, mainly negative, voltage shifts recorded over sensorimotor or
frontal cortical areas.
 They precede and coincide with imagined or actual motor actions or cognitive tasks.
 With extensive training, people can learn to control SCPs and use them to operate spelling programs and
other applications. Although SCP-based BCIs were used successfully in the past by people with severe
disabilities (e.g., Amyotrophic lateral sclerosis (ALS) patent), this BCI modality receives little attention
at present because it is inherently slow, allows only one dimension of control, requires extensive training,
and is prone to error.
 In the future, SCP-based BCI paradigms may prove useful as therapeutic neuro-feedback tools or as
adjuncts to other BCI or conventional control modalities (e.g., by allowing recognition of anticipation in
the user)

5. Movement-Related Potentials
 EEG signals show a small and slow potential drift prior to voluntary movements.
 These movement-related potentials (MRPs), sometimes also called readiness potentials (RPs) or
Bereitschaftspotentials (BPs), show variation in distribution over the scalp with respect to the body part
being moved.
 The Bereitschaftspotential (or readiness potential) is a negative SCP that usually begins 500–1000 ms
before a self-initiated movement.
 For example, the BP related to movement of left versus right arm shows a strong lateral asymmetry.
 This potentially allows one to not only estimate the intent to move, but also distinguish between left and
right movement intention.
 This makes them attractive targets for BCI applications but since they are typically much smaller than
other EEG phenomena such as alpha or beta rhythms, their detection is much harder.

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 It has been suggested that while ERD may reflect changes in the background oscillatory activity in wide
cortical sensorimotor areas, MRPs may represent increased, task-specific responses of supplementary
and primary motor cortical areas.
 Its amplitude and topography are affected by movement type and the muscles involved.
6 Stimulus-Evoked Potentials
 A major class of EEG signals used in noninvasive BCIs are evoked potentials (EPs), which are
stereotypical EEG responses generated by the brain when the subject is presented with a particular type
of stimulus.
 For example, when a rare but task- relevant auditory, visual or somatosensory stimulus is interspersed
with frequent and routine stimuli, the rare stimulus evokes a potential with a positive peak at about 300
ms after the stimulus is presented.
 This potential is called the P300 (or P3) potential Other types of responses include:
 visually evoked potentials (VEPs) generated by visual stimuli such as flashing lights,
 steady state visually evoked potentials (SSVEP) produced by a visual stimulus repeated at a rate greater
than 5 Hz,
 auditory evoked potentials (AEPs) generated by auditory stimuli such as clicks and tones, and
 Somatosensory evoked potentials (SSEPs) caused by somatosensory stimulation.

6.1 The P300 Potential

 The P300 (or P3) signal is so named because it is a positive deflection in the EEG signal that occurs
approximately 300 ms after a stimulus. The stimulus itself must be rare and unpredictable but relevant
to the subject (e.g., sudden intensification of an attended target).
 The amplitude of the P300 depends directly on how relevant the stimulus is and varies inversely with the
probability of the stimulus.
 The P300 is generally observed most strongly over the parietal lobe, although some components also
originate in the temporal and frontal lobes.
 Although it often occurs at a latency of about 300 msec relative to the eliciting stimulus (hence the
designation of P300), its latency may vary from 250 to 750 msec.
 This variability in latency reflects the fact that the P300 is elicited by nature of the decision.
 The P300 is usually largest over central parietal scalp and attenuates gradually as distance from this area
increases.

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 The voltages that constitute the ERP are embedded within the general EEG activity recordable from the
scalp and are usually quite small relative to the ongoing EEG.
 However, because the ERPs are time-locked to events, and follow a constant time course, they can be
extracted by averaging multiple trials of eliciting events.
 The result is a series of positive and negative voltage deflections that are referred to as components.
 Longer-latency components ( > 150 msec) tend to reflect information-processing activity that is cognitive
in nature and is thus less dependent on stimulus modality and more dependent on the significance of the
eliciting event in the subject’s concurrent tasks. They are usually referred to as endogenous components.
 Current P300-based BCIs allow users to select items displayed on a computer screen.
 The specific set of circumstances for eliciting the P300 ERP is known as the Oddball Paradigm.

This paradigm has three essential attributes:

 A subject is presented with a series of events (i.e.,stimuli), each of which falls into one of two
classes.
 The events that fall into one of the classes frequent than those that fall into the other class.
 The subject performs a task that requires classifying each event into one of the two classes.
 The events that fall into the less-frequent class elicit a P300.
 As long as an experimental design adopts the three attributes of the oddball paradigm, any stimulus and
any classification task can elicit a P300.
 A P300 can be elicited by an event that consists of the absence of a stimulus, if that absence satisfies the
conditions of the oddball paradigm.
 Figure 12 illustrates a typical P300 experiment. The letters ‘O’ and ‘X’ flash on a video screen in a
random order at a rate of one per second.
 The X occurs infrequently (e.g., 20 % of the flashes) and is thus the oddball stimulus, while the O occurs
frequently. The subject is asked to count the number of times one of the stimuli (e.g., X ) occurs.
 Each time a stimulus occurs, a marker is placed in the data file to indicate the identity of the stimulus, X
or O
 Each stimulus is presented on the screen for 100 msec, and then the screen is blank for 900 msec until
the presentation of the next stimulus.
 The ERPs elicited by the oddball stimulus at midline electrode locations Fz, Cz, and Pz of the 10–20
system for 800 msec after the stimulus.

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Figure 12. Time course of rare (i.e., oddball) (X) and common (O) stimuli in a standard oddball protocol. (B)
Average oddball ERPs from a subject for electrode locations Fz, Cz, and Pz.

 The three responses show a typical P300 scalp topography: the most prominent potential is a positive
component occurring about 350 msec after the X stimulus; and it is largest at the Pz electrode and
attenuates at more anterior and posterior locations.
 It should be noted that the results would be essentially the same even if the subject had been asked to
count the frequent O stimuli rather than the rare X stimuli.
- P300 is always elicited by the rare events.
- P300 latency may vary from 250 to 750 msec
 A famous example of an early BCI based on EEG is the P300 now-classic BCI “speller” based on the
oddball paradigm, the 26 letters of the English alphabet are displayed in the form of a 6 × 6 matrix on a
computer screen.
 In order to spell a word, the subject must select each letter comprising the word by focusing attention on
that letter in the matrix.
 While the subject is focusing on the letter or command, the rows and columns of the matrix are repeatedly
flashed in random order.
 Each flash of a row or column lasts 100 ms, and the interval between flashes is fixed at either 500 ms or
125 ms.

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 Only when the row or column containing a subject’s chosen letter or command is flashed is a large P300
generated by the subject’s brain.
 The subject’s choice of letter or command can thus be inferred by keeping track of which flashed row
and column elicited the largest P300s.

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 This signal can be detected using a classifier such as linear discriminant analysis (LDA). The subject’s
choice of letter or command can thus be inferred by keeping track of which f lashed row and column
elicited the largest P300s

6.2 STEADY-STATE VISUAL EVOKED POTENTIALS (SSVEP)

 Brain processing of a sensory stimulus or another event can produce a time-locked series of positive-
negative deflections in the EEG.
 These event-related potential (ERP) components are distinguished by their scalp locations and latencies.
 Earlier components with latencies <100 ms originate largely in primary sensory cortices and are
determined mainly by the properties of the evoking stimulus.
 Later ERP components with latencies of 100–>500 msec reflect to a greater extent ongoing brain processes
and thus are more variable in form and latency.
 SSVEP stimuli do not flash successively, but flicker continuously with different frequencies in the range
of about 6–30 Hz. Paying attention to one of the flickering stimuli elicits an SSVEP in the visual cortex
that has the same frequency as the target flicker.
 That is, if the targeted stimulus flickers at 16 Hz, the resulting SSVEP will also flicker at 16 Hz. Therefore,
an SSVEP BCI can determine which stimulus occupies the user’s attention by looking for SSVEP activity
in the visual cortex at a specific frequency.
 The BCI knows the flickering frequencies of all light sources, and when an SSVEP is detected, it can
determine the corresponding light source and its associated command.
 Consider a system where the goal is to decode one of two possible choices. One can then represent the
two choices by visual stimuli (e.g., buttons on a screen or light emitting diodes – LEDs), each blinking at
a different frequency.
 The subject focuses attention on the button corresponding to his or her choice (e.g., by looking at it).
 This results in an EEG signal in the early visual areas of the brain (the occipital region) oscillating at the
stimulus frequency – this signal is called a steady state visually evoked potential(SSVEP)
 By performing a frequency decomposition of the EEG stimulus the BCI can detect the frequency of the
stimulus the user is paying attention to and therefore, the user’s choice.
 BCI allowing selection from 13 buttons on a computer screen, representing a virtual telephone keypad
with the digits 0–9, BACKSPACE, ENTER, and an ON/OFF button.
 Each of the 13 buttons was flashed on and off at a different frequency between 6Hz and 14 Hz.

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 To reduce false positives due to alpha rhythms, a screening experiment with eyes closed was first
performed, and frequencies with power more than twice the mean power between 4 Hz and 35 Hz were
excluded from the stimulation frequencies.
 Additionally, all stimulation frequencies were odd multiples of the frequency resolution to prevent one
stimulation frequency being twice another stimulation frequency.

Figure 15 An example of an SSVEP BCI

• In other experiments, the authors found that the minimum difference in flickering frequency (i.e.,
frequency resolution) between neighbouring targets that a subject can discriminate is about 0.2 Hz, and
the frequency range in which the SSVEP can be effectively observed is approximately 6–24 Hz.
• EEG signals were recorded from electrode locations O1 and O2 according to the 10–20 system with
left/right mastoids as reference electrodes.
• A fast Fourier transform was performed every 0.3s to compute the amplitude spectrum.
• For each stimulation frequency, the sum of its amplitude and that of its second harmonic was used as
the feature for classification.
• A simple threshold classifier was used, where the threshold was chosen to be twice the mean value of
the amplitude spectrum between 4 Hz and 35 [Link] output of the classifier was the frequency with the
largest intensity.

SSVEP-based BCI operation and analysis

 The user views three red boxes each of which flickers at a different frequency. By choosing to focus on
the 8-Hz box, it elicits EEG activity at 8 Hz and its harmonic frequencies (as shown in the frequency
spectrum of EEG activity at occipital location O2; blue trace in bottom panel). Also shown for
comparison is the spectrum produced when the user is not looking at one of the boxes (red trace).

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Figure 16 SSVEP-based BCI operation and analysis.

 It lacks the 8-Hz and harmonic peaks but is otherwise similar. The 8-Hz activity is focused over occipital
areas (as shown in the topographical plot [top middle] of power at 8 Hz [with blue indicating higher
power].
 The top right panel shows that this increased activity does not occur immediately. The vertical line
indicates the time at which the user decides to look at the 8-Hz box.
 Power at 8 Hz and its harmonic frequencies increase significantly (p < 0.01 indicated by blue-green)
over the next 2 sec. (Yellow indicates significant power decrease.)

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Figure 17 Three different SSVEP or related VEP stimulus paradigms

 The standard SSVEP BCI paradigm, in which each of the repetitive stimuli occurs at a specific
frequency, is called the frequency-modulated visual evoked potential (f-VEP) BCI paradigm.
 In a t-VEP paradigm, the different stimuli are mutually independent and non-overlapping.
 The BCI computes the average VEP for each stimulus and produces the output represented by the
stimulus that elicits the largest VEP.
 In a c- (or m-sequence) VEP paradigm, each stimulus occurs in a pseudorandom pattern that is nearly
orthogonal to the patterns of all the other stimuli.
 The BCI computes the correlation between the EEG and a template computed for each stimulus.
Typically, the stimulus on which the user is fixating produces the highest correlation.
 The accuracy and speed of BCIs that used f-, c-, and t-VEP paradigms are compared.
 They found that the c-VEP paradigm performed best, with the f-VEP paradigm yielding intermediate
performance, and the t-VEP BCI much worse.

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Figure 18 Overview of an early c-VEP BCI system

7. Activity of Neural Cells

 The workhorse of the brain is a type of cell known as a neuron, a complex electrochemical device that
receives information from hundreds of other neurons, processes this information, and conveys its output
to hundreds of other neurons.
 Furthermore, the connections between neurons are plastic, allowing the brain’s networks to adapt to new
inputs and changing circumstances.
Neurons
 A neuron is a type of cell that is generally regarded as the basic computational unit of the nervous system.
The neuron can be regarded as a leaky bag of charged liquid.
 The membrane of a neuron is made up of a lipid bi-layer (Figure 19) that is impermeable except for
openings called ionic channels that selectively allow the passage of particular kinds of ions.
 Neurons reside in an aqueous medium with a larger concentration of sodium (Na+),chloride (Cl), and
calcium (Ca2+) on the outside of the cell and a greater concentration of potassium (K+) and organic
anions (A-)inside the cells (Figure 19). As a result of this imbalance, there is a potential difference of
approximately −65 to −70 mV across the neuron’s membrane when the neuron is at rest.

Figure 19 The electrochemical dance of ions in a neuron

Action Potentials or Spikes
 When the neuron receives sufficiently strong inputs from other neurons a cascade of events is triggered:
there is a rapid influx of Na+ ions into the cell, causing the membrane potential to rise rapidly, until the
opening of K+ channels triggers the outflux of K+ ions, causing a drop in the membrane potential.
 This rapid rise and fall of the membrane potential is called an action potential or spike (Figure 20), and
represents the dominant mode of communication between one neuron and another. The spike is an all-
or-one stereotyped event with little or no information in the shape of the spike itself – information is
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thought to be conveyed instead by the firing rate (number of spikes per second) and/or the timing of
spikes.
 Neurons are therefore often modeled as emitting a 0 or 1 digital output. Similarly, in extracellular
recordings typically done in awake animals a spike is often represented as a short vertical bar at the time
the spike occurred. Neural cells, or neurons, are the primary functional units of the nervous system,
responsible for transmitting information through electrical and chemical signals. The activity of neural
cells can be broadly categorized into two main types:
 Resting State: Neurons are typically in a resting state when they are not actively sending signals. During
this state, the neuron maintains a negative resting membrane potential, meaning that the inside of the cell
is more negatively charged compared to the outside. This resting potential is maintained by the selective
permeability of the cell membrane to ions such as potassium (K+) and sodium (Na+), as well as the
action of ion pumps like the sodium-potassium pump.
 Action Potential: When a neuron receives a strong enough stimulus, it can depolarize and generate an
action potential. This is a rapid change in membrane potential where the inside of the cell becomes more
positively charged. This depolarization is caused by the opening of voltage-gated ion channels, allowing
sodium ions to flow into the cell. The action potential then propagates down the length of the neuron's
axon, ultimately leading to the release of neurotransmitters at the synapse.

Figure 20 Generation of spikes or action potentials

 Neurons are typically classified into three types based on their function.
 Sensory neurons respond to stimuli such as touch, sound, or light that affect the cells of
the sensory organs, and they send signals to the spinal cord or brain.
 Motor neurons receive signals from the brain and spinal cord to control everything from muscle
contractions to glandular output.
 Interneurons connect neurons to other neurons within the same region of the brain or spinal cord.
When multiple neurons are functionally connected together, they form a neural circuit.
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Figure 21 Dendrites, soma, axon, and synapse

Dendrites and Axons
 Neurons in different regions of the brain have different morphological structures, but the typical structure
includes a cell body (called the soma) connected to a tree-like structure with branches called dendrites
and a single branch called the axon that emanates from the soma and conveys the output spike to other
neurons.
 The spike is typically initiated near the junction of the soma and axon and propagates down the length
of the axon. Many axons are covered by myelin, a white sheath that significantly boosts the speed of
propagation of the spike over long distances.
 The terms white matter and gray matter correspond respectively to the myelinated axons connecting
different brain regions and the regions containing the cell bodies.
Synapses
 Neurons communicate with each other through connections known as synapses. Synapses can be
electrical but are more typically chemical. A synapse is essentially a gap or cleft between the axon of
one neuron (called the presynaptic neuron) and a dendrite(or soma) of another neuron (called the
postsynaptic neuron).
 When an action potential arrives from a presynaptic neuron, it causes the release of chemicals known as
neurotransmitters into the synaptic cleft. These chemicals in turn bind to the ionic channels (or receptors)
on the postsynaptic neuron, causing these channels to open, thereby influencing the local membrane
potential of the postsynaptic cell.

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 Synapses can be excitatory or inhibitory. As the name suggests, excitatory synapses cause a momentary
increase in the local membrane potential of the postsynaptic cell. This increase is called an excitatory
postsynaptic potential (EPSP).
 EPSPs contribute to a higher probability of firing a spike by the postsynaptic cell. Inhibitory synapses
do the opposite – they cause inhibitory postsynaptic potentials (IPSPs), which temporarily decrease the
local membrane potential of the postsynaptic cell.
 A neuron is called excitatory or inhibitory based on the kind of synapse it forms with postsynaptic
neurons. Each neuron forms only one kind of synapse, and therefore if an excitatory neuron is to inhibit
a second neuron, it must excite an inhibitory “interneuron,” which then inhibits the desired neuron.
Spike Generation
 When the neuron receives sufficiently strong inputs from its synapses for its membrane potential to cross
a neuron-specific threshold, a spike is emitted (Figure 20).
 This makes the neuron a hybrid analog-digital computing device: digital 0/1 inputs are converted to
analog changes in the local membrane potential, followed by summation of these changes at the soma,
and a spike if the summation of changes exceeds threshold.
 This simplified model of course ignores the complex and potentially important forms of signal processing
associated with dendrites, but the threshold model of a neuron has proven to be a useful abstraction in
neural modeling and artificial neural networks.
Adapting the Connections: Synaptic Plasticity
 A critical component of the brain’s adaptive capabilities is the ability of neurons to change the strength
of the connections between themselves through synaptic plasticity.
 Numerous forms of synaptic plasticity have been experimentally observed, the most studied being long-
term potentiation (LTP) and long-term depression (LTD).
 Both involve changes to a synapse that last for hours or even days. More recently, other types of plasticity
have been characterized, including spike timing dependent plasticity (STDP), where the relative timing
of input and output spikes determines the polarity of synaptic change, and short-term
facilitation/depression, where the plasticity is rapid but not long-lasting.

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Figure 22 Synaptic plasticity

LTP
 One of the most important forms of synaptic plasticity is long-term potentiation or LTP (Figure 22). In
its simplest form, LTP involves an increase in the strength of a synaptic connection between two neurons
caused by correlated firing of the two neurons.
 LTP has been found in a number of brain areas including the hippocampus and the neocortex.

LTD
 Long-term depression or LTD involves a decrease in the strength of a synaptic connection caused, for
example, by uncorrelated firing between the two neurons involved. LTD has been observed most
prominently in the cerebellum, although it also coexists with LTP in the hippocampus, neocortex, and
other brain areas.
STDP
 Traditional experimental protocols demonstrating LTP/LTD involved stimulating a presynaptic neuron
and a postsynaptic neuron simultaneously.
 These protocols manipulate the firing rate of pre-and postsynaptic neurons but not the timing between
presynaptic and postsynaptic spikes.
 The precise timing of pre-and postsynaptic spikes can determine whether the change in synaptic strength
is positive or negative. This form of synaptic plasticity has been termed spike timing dependent plasticity
(STDP).

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The types of synaptic plasticity discussed above are called long-term plasticity because the changes they
cause can last for hours, days, or even longer periods of time.

7.1 Multiple neuromechanisms

 Multiple neuro mechanisms refer to the various processes and mechanisms through which neurons
communicate and process information in the brain. These mechanisms are essential for the functioning
of the nervous system and underlie various cognitive and behavioral processes. Some of the key
neuromechanisms include:
 Synaptic Transmission: Neurons communicate with each other through specialized junctions called
synapses. At the synapse, the presynaptic neuron releases neurotransmitters, which then bind to receptors
on the postsynaptic neuron, leading to changes in the postsynaptic neuron's membrane potential.
 Neurotransmitter Systems: Neurotransmitters are chemical messengers that transmit signals between
neurons. Different neurotransmitter systems, such as the dopamine, serotonin, and acetylcholine systems,
play crucial roles in regulating various functions such as mood, behavior, and cognition.
 Neural Plasticity: Neural plasticity refers to the brain's ability to reorganize its structure and function in
response to experience. This includes synaptic plasticity, where the strength of synaptic connections
between neurons can be modified, and structural plasticity, where new connections can be formed or
existing ones can be eliminated.
 Neural Oscillations: Neural oscillations are rhythmic fluctuations in neural activity that occur in various
frequency bands. These oscillations are thought to play a role in coordinating communication between
brain regions and are associated with different cognitive processes, such as attention, memory, and
consciousness.
 Neural Coding: Neural coding refers to the way information is represented and processed by neurons.
This includes the rate coding, where the intensity of a stimulus is encoded by the firing rate of neurons,
and temporal coding, where information is encoded in the precise timing of action potentials.
 Neural Circuits: Neural circuits are networks of interconnected neurons that are responsible for
processing specific types of information. These circuits can involve multiple brain regions and are
organized in a hierarchical manner, with higher-order regions integrating and coordinating information
from lower-order regions.

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