WORKING MEMORY AND ELECTROENCEPHALOGRAPHIC (EEG) COHERENCE

Brežan Simon, Štukovnik Vita

Faculty of Medicine, University of Ljubljana, Ljubljana, Slovenia
Institute of Clinical Neurophysiology, University Medical Centre Ljubljana

ABSTRACT
Cognitive neuroscience searches for neuronal correlates of higher mental functions, including working memory. Electrophysiological
methods explore brain functions on the level of electrical activity of the brain with high time resolution. This allows the study of basic
mechanisms of information processing and operational system of the brain. EEG coherence, as a newer method of EEG signal analysis,
is suitable for studying integrative brain function. It measures interregional synchronized oscillatory activity of neurons as one possible
mechanism of the functional integration – binding of different communicating brain areas. The binding could serve as a mechanism of
functional coupling, present with different distributed brain processes and behavioural modes, for example working memory.

WORKING MEMORY
Memory includes the following main processes: encoding (cognitive
events during presentation of information), storage (maintenance) and
retrieval (recall) of information.
One of the best studied models of working memory is Baddeley’s model
(Baddeley and Hitch, 1974). The model consists of the attentional
control system – the central executive (CE), and three subsystems (slave
systems): the phonological loop, the visuospatial sketchpad and the
episodic buffer. CE is responsible for controlling subsystems, strategy
selection, planning and manipulation of information, providing link between
subsystems and long-term memory. Information is stored and actively
rehearsed separately in the modal specific subsystems – visuospatial
sketchpad and phonological loop. The role of episodic buffer is binding of
information from the slave systems and long-term memory and temporary
storing them in a multi-dimensional code.
NEUROANATOMICAL AND
NEUROPHYSIOLOGICAL BASIS OF
WORKING MEMORY
Various components of working memory are mediated by different
anatomically separated neuronal networks. Cell electrophysiological
studies on animal models and functional brain imaging studies on human
subjects have shown that working memory is mainly based on the activity
in prefrontal, premotor, limbic, and posterior association parietal areas of
the brain (Fletcher and Henson, 2001).
Figure 3. Illustration of EEG scalp electrodes and functional coupling between different brain areas (the colour of inter-electrode
The basic neurophysiological mechanism of working memory is supposed connections shows different coherence values).

Figure 1. Basic structure of memory with three memory stores (Atkinson and Schiffrin, 1968).
The short-term memory is also termed working memory. This emphasizes
its active role in cognitive processes – besides temporal storage of
information, the information is also available for active manipulation.
Working memory can therefore be defined as a complex of cognitive
processes for maintenance, manipulation and utilization of mental
representations.
COGNITIVE PARADIGMS FOR WORKING
MEMORY ASSESSMENT
Different cognitive paradigms are used for assessment of specific working
memory processes:
– STERNBERG PARADIGM: maintenance of information
(activity of phonological loop, visuospatial sketch pad)
– GO/NO-GO TEST: central executive component of
working memory
– STROOP COLOUR-WORD TEST: central executive
component of working memory
– TOWER OF LONDON: central executive component of
working memory, maintenance of information (activity of
phonological loop, visuospatial sketch pad).
– N-BACK TASK: maintenance of information (activity
of phonological loop, visuospatial sketchpad), central
executive component of working memory
Figure 2. Baddeley’s working memory model (Baddeley, 2000).
At the Institute of Clinical Neurophysiology our research group plans to
carry out the EEG study of working memory in the context of synchronized
brain oscillations, using the EEG coherence method. The study design will
be based on the modified Sternberg paradigm, which in comparison to
other current EEG studies allows independent assessment of both central
executive component and maintenance processes of working memory.
This modified Sternberg paradigm includes:
– maintenance of sequentially presented verbal stimuli (letters) and their
serial position (maintenance processes of working memory)
– alphabetically reordering of presented letters and maintenance of
reordered sequence of letters (executive component and maintenance
processes of working memory)
– control task (different dilemmas and problems exist what should be the
appropriate control task which would ideally exclude working memory
and other uncontrolled intrinsic mental processes)
to be based on repeated reverberations of electrical impulses in
EEG coherence and power spectra analyses are newer methods for
reverberational (feedback) loops (Štrucl, 1999). A repeated excitation of a
analyzing the EEG signal.
synapse in such an excitational loop leads to postsynaptic facilitation, and
Power spectra are a measure for a degree of the representation of a
this can be a basis for the maintenance of specific information, coded by
specific frequency band in the signal. The power spectra changes reflect
activated neural network pattern.
different levels of regional cortical activity or different levels of their
The exact mechanisms for coordination and dynamic interactions
synchronization. Synchronization usually reflects cortical inactivity, while
of anatomically separated brain regions with neural mechanisms for
desynchronization reflects activity.
information processing itself are not yet known.
EEG coherence reflects different degrees of synchronization between
The functional integration (‘binding’) of different brain areas, responsible
separate cortical regions and is independent measure of synchronized
for specific (working memory) functions, is one of the key problems in
oscillatory activity, which could mediate the functional coupling – binding
understanding the brain function – a question not yet resolved. Binding
and communication between distributed brain centers involved in specific
is perhaps mediated by a synchronized oscillatory activity of neuronal
tasks. Power spectra and coherence changes therefore reflect two
networks, which can be determined by the electroencephalographic (EEG)
coherence analysis.
different operational systems of the brain. Many studies have found that
synchronous oscillations correlate with specific behavioural contexts and
cognitive tasks (Hallet, 2000).
ELECTROENCEPHALOGRAPHY (EEG), Most important is the difference between coherence values in the test task
SIGNAL ANALYSIS: EEG COHERENCE AND compared to basal values in the control tasks (∆Cxy).
POWER SPECTRA
Electroencephalography (EEG) is a method, which measures repeated,
periodic electrical activity of cortical neurons. The summed activity
of many neurons results in field potentials, many of them constituting Cxy(ω): coherence value between signals x and y
macropotential (EEG signal). Macropotential is a result of changing pattern
Φxy(ω)- value of cross-correlation power spectrum of signals x, y
of the synchronization and desynchronization of regional brain cells; this Φxx(ω)- value of auto-correlation power spectrum of signal x
results in amplitude changes of specific frequency bands. EEG has great Φyy(ω)- value of auto-correlation power spectrum of signal y
time resolution and shows distinct patterns of activity (brain rhythms,
oscillations).
Brain rhythms can be divided in many frequency bands (delta: 0,5-4 Hz,
theta: 4-7 Hz, alpha: 8-13 Hz, beta: 13-30 Hz, gamma: over 30 Hz) with
their specific functional and behavioural correlates, activating contexts and
different spatial scales. Their possible role is switching neural networks
between different functional states with activating or inhibiting proper
neural systems.

Figure 4. Power spectrum increase (red colour) in theta
frequency band for working memory retention task compared
to control task (increases in frontal, central and parietal brain
areas); computer brain model; /personal archive material/.
Figure 5. Coherence increases (red colour) and decreases
(blue colour) in theta frequency band for working memory
retention task compared to control task (increases between
frontal, central and parietal brain areas); computer brain
model; /personal archive material/

WORKING MEMORY AND The majority of EEG studies have shown that working memory
tasks are often related to:
Spatial scales of brain oscillations involved in working memory

SYNCHRONIZED BRAIN • regional neuronal desynchronization and consequent power spectrum

OSCILLATIONS: POWER SPECTRA decrease in lower alpha band (showing a non-specific effect of
attention, mental effort);
AND EEG COHERENCE STUDIES
• regional desynchronization leading to power spectrum decrease in
upper alpha band (correlate of semantic processing);
The neurophysiological theory of
• paradoxical alpha synchronization, power spectrum increases (active
(working) memory: inhibition of disturbing neural networks not needed for the memory
task as optimization of processes; Klimesch et al., 1998, etc.);
• Brain oscillations in different frequency • increased interregional synchronization resulting in coherence
bands subserve specific (memory) increases in theta frequency band – working memory processes
(Serrien et al., 2003; Sauseng et al., 2004; Sarnthein et al., 1998;
functions and operate over different Jensen et al., 2002);
spatial scales. • increased regional synchronization with increases of power spectrum
in theta frequency band: frontal midline theta rhythm – memory
maintenance, attention, mental effort (Klimesch et al., 1997, Gevins et
• Multiple superimposed synchronized al., 1997, model Lisman, Idiart and Jensen, 1998, etc.). The source
of theta rhythm? – anterior cingulate cortex, cortico-hippocampal
(coherent) oscillations in different feedback loops;
frequency bands with different spatial • increased power spectra and coherence in gamma band (sensory,
perceptional and attentional processes, working memory processes);
patterns and functional correlates
govern specific mental functions. • changes of power spectra and coherence with different memory load
(Gevins et al., 1997; Jensen, 2000, etc.).
RESEARCH DILEMMAS: different non-consistent definitions of specific
frequency bands for different brain rhythms by authors, possible inter-
individual differences in oscillatory frequencies, usage of different
cognitive paradigms, methodological and technical problems, artefacts,
signal to noise ratio…
Mainly pre(frontal) and posterior association brain areas were found
oscillating synchronously (theta, gamma, alpha band; Sarnthein et al.,
This is in accordance with Baddeley’s model of working memory which
predicts constant interplay and functional coupling between central
1998, etc.): executive control system (probably mediated by prefrontal cortex, Collette
• prefrontal cortex itself rehearses and maintains the information
et al., 2002) and slave subsystems for storage of verbal and visuospatial
information, located in posterior brain regions (Gathercole, 1999).
transferred from posterior storage areas and manipulates with it;
• top-down control by prefrontal cortex and its central executive, gating
of irrelevant sensory signals coming from posterior regions and
memory process optimization.
Lisman/Idiart/Jensen (LIJ) model – example of
neurophysiological model of working memory
Results of studies showing the effect of different memory load on theta
oscillations confirm the LIJ model (1998). Model predicts that different
memory representations (7 items +-2) are encoded and activated
separately and sequentially in subsequent phases of theta cycle. Theta
input serves as a signal carrier and induces repeating of scanning and
working memory maintenance processes. This means that model is
based on serial information processing principles. Therefore it can
explain empirical data (Sternberg, 1964) on increase of reaction times
with respect to working memory load (number of items). The memory
code itself is mediated by synchronous gamma oscillations, induced
Figure 6. Diagrams indicating the concept of LIJ (Lisman, Idiart, Jensen, 1998) working memory model. Three memory items (A, B,
by pattern of alternating excitation and inhibition of different memory C) are loaded in memory buffer, the theta period increases by one gamma period with each item added. In retention interval (delay
period), items are maintained by activity-dependent intrinsic properties of the neurons coding these items. After probe presentation
the items can be scanned – compared with the probe as they are activated. After scanning, motor response and answer can be
initiated. RT – reaction time
codes. The source of theta oscillations could be the hippocampo-cortical
reverberating loops.

Working memory processes are thereby potentially mediated by posterior

(sensory)-hippocampal-prefrontal neural network, which is integrated
and functionally coupled by the synchronization of oscillations in theta
frequency band.

CONCLUSION WITH CLINICAL IMPLICATIONS OF EEG COHERENCE

Figure 7. LIJ model of a network functioning as a multi-item short-term memory buffer.
Theta and gamma oscillations play an important role in the concept. An afterdepolarization (ADP) is triggered after a cell fires
(sensory input) and it causes depolarizing ramp that serves to trigger the same cell to fire again after delay. These ramps are
Synchronization in gamma frequency band is believed to be correlated with sensory processing and the very content of information processing, but temporarily offset for different memories, an offset that causes different memories to fire in different gamma cycles. The key function
of this buffer is to perpetuate the firing of cells in a way that retains serial order. The repeat time is determined by theta oscillations
could also reflect increased attentiveness. The increased coherence for interactions between posterior and frontal areas in theta frequency band (also due to external input. Gamma oscillations arise from alternating global feedback inhibition and excitation (the cell with most
depolarized ramp will fire again) due to separate firing of different memory codes.

with probable involvement of hippocampus) on the other hand, possibly mediates working memory processes (storage, rehearsal and scanning).
In this way, slower rhythms could act as signal carriers constituting the context whereas fast waves refer to the entire content of the representation.
The neuronal synchronization – functional coupling plays a role in interaction of posterior association cortex, where sensory information is stored, and
(pre)frontal cortex, where relevant current information is held, rehearsed and updated. Verbal memory tasks seem to activate primarily the left brain
hemisphere, while visuospatial memory tasks activate predominantly the right brain hemisphere.
REFERENCES:

The value of EEG coherence method for brain research is greatest when combined with other brain research approaches which show different
aspects of the brain function. With choosing appropriate cognitive paradigms and neuropsychological tests, it is possible to study physiological and
pathophysiological aspects of cognitive, motor and sensory brain function.
Recently, some studies (Hogan et al., 2003; Spencer et al., 2003, etc.) also showed important differences in coherence values when comparing
working memory in neurodegenerative or psychiatric patients (Alzheimer disease, Parkinson disease, schizophrenia) to healthy controls. This opens new
future perspectives for possible search of pathophysiological mechanisms and etiological factors contributing to many different neurological diseases.
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