International Journal of Psychophysiology 60 (2006) 133 – 138

www.elsevier.com/locate/ijpsycho

The theory of the whole-brain-work

Erol Başar
Dokuz Eylül University, Brain Dynamics Multidisciplinary Research Center and Faculty of Medicine Department of Biophysics, 35340, Balçova, Iżmir, Turkey
Received 10 December 2005; received in revised form 23 December 2005; accepted 23 December 2005
Available online 24 March 2006

Abstract

The theory of the whole-brain-work basically explains the oscillatory dynamics of the human and nonhuman brain during cognitive processing.
The theory is based on principles according to which brain functions are represented by the oscillatory activity. Oscillatory activity in a given
frequency band performs multiple functions since they vary on a number of response parameters. There is selective cooperation in the stimulated
brain; this produces super-binding between neural populations and super-synergy in the whole brain. The concept of super-synergy thus includes
super-binding and, additionally, entropy and the role of EEG-oscillations as control parameters in brain's responsiveness. In super-synergy, spatial
integration occurs through the selective cooperation of brain structures. Temporal integration occurs in line with the principle of superposition of
oscillations in which the comparative polarity and phase angle are critical for forming the function-specific configurations. Extension of the theory
of whole-brain-work to cognitive processing proposes that there is a constant reciprocal activation within the subprocesses of attention, perception
learning and remembering and this leads to an APLR-alliance. In such a context, all brain functions are inseparable, for instance, from memory
function and, in turn, memory states have no exact boundaries along the time space; memory states thus evolve in the APLR-alliance. The theory
claims that the reentry and the dynamic behavior of oscillations during the reciprocal activation in APLR-alliance are among the causal factors for
brain dynamics and for cognition.
© 2006 Published by Elsevier B.V.

Keywords: Brain oscillations; Spatial integration; Temporal integration; Super-synergy; APLR-alliance; Causality; Memory; Attention; Perception

“…The whole burden of philosophy seems to consist in this,
from the phenomena of motions to investigate the forces of
nature, and from these forces to demonstrate the other
phenomena.” (Newton, 1687/1726)
1. Introduction
Science seeks for causes. With the exception of the
conventional analysis of neural activity which uses single unit
recordings, the concept of causality has somewhat been
overlooked in neural sciences and the need for rules and
theories have thus been overlooked. This has generally been the
situation for also EEG-oscillation analysis.
In all natural sciences, the general questions and problems on
the concept of causality are based on or are derived from
Newtonian dynamics. According to the first law of motion in
Newtonian system, “free motion” is uniform motion in a straight
E-mail address: erol.basar@deu.edu.tr.
0167-8760/$ - see front matter © 2006 Published by Elsevier B.V.
doi:10.1016/j.ijpsycho.2005.12.007
line. When a force is applied to a body, it causes the body to
deviate from this free motion. All observed motions could be
analyzed into two components: a free component (inertia) and a
component due to the acting force. The second law states that
the force acting on a body is always proportional to the product
of its mass and acceleration. Newton (1687/1726) never
regarded the word “force” just as a name for this mathematical
product. However, as a natural scientist, he eschewed
speculation in dealing with the nature of forces and, for
scientific purposes, he thought it sufficient to calculate and
observe the effects of these forces.
Furthermore, Isaac Newton (1687/1726) was interested not
only in descriptions pertaining to motions of planets, but he also
wanted to find the mechanism of gravitation between the
planets. Likewise, Galileo Galilei did not only observe the
oscillations of clocks, but he also wanted to learn about their
machineries. Albert Einstein was interested in describing tracks
of the molecules as in the case of Brownian motion, but he also
analyzed the causes of Brownian motion. Furthermore, Einstein
was searching for causes of gravitation, but he also wanted to
134 E. Başar / International Journal of Psychophysiology 60 (2006) 133–138
understand the causes of dissipating energy. To establish what
has happened in the galactic system, Einstein predicted the
existence of “black holes” from not only the facts about
astrophysical events, but also from a combination of accumu-
lated data on the motion of stars and laws of physics. Einstein
thus offered descriptions and explanations pertaining to the
nature of stars and the galaxy, including those that were not
visible to conventional observations, i.e. the black holes.
Due to its breadth and impact, Newtonian dynamics has
become the metaphor of all natural sciences. The relevance of
the prestimulus EEG as a causal factor in attention, perception,
learning and remembering has important parallels with New-
ton's (1687/1726) first law of motion where the state of a
moving body is a causal factor for the further evolution of its
movement. The application of this law to electrophysiology is
the following: The state of the brain as reflected in the
prestimulus EEG is the causal factor for the later brain
responses.
The trajectories (EEG signals) reflecting the activity of
neuronal populations can also be analyzed as somewhat similar
to the analysis of motion. As the expression “brain dynamics”
implies, we intend to also elucidate the causes or mechanisms
that give rise to the trajectories of the electrical signals in the
brain. Similar to trajectories of missiles, or trajectories in
Brownian motion, EEG trajectories is already providing most
useful information on neural mechanisms that give rise to
different transitions. The EEG seems to serve as a fundamental
trajectory that is causally related to memory building and
integrative brain function. The application of the Newtonian
perspective or Einstein's approach when searching for the
mechanisms behind EEG trajectories has already started.
However, recent developments on the dynamics of quantum
physics and the new approach on chaos certainly brought a
different understanding to the Newtonian causality. In his highly
popular book on chaos, Gleick (1987), an advocate of the new
science, went so far as to say: “Twentieth-century science will
be remembered for just three things: relativity, quantum
mechanics and chaos.” Chaos is the century's third great
revolution in the physical sciences. Like the first two
revolutions, chaos cuts away at the tenets of Newtonian physics.
Can these development be useful and have the great impact
that the systems of Newton, Galilei and Einstein's has had. The
new development, “chaos in brain function”, is certainly
fascinating. However, in the period of 1985–2000 during
which findings on chaotic EEG were obtained, noteworthy
progress was also occurring in the study of oscillatory
phenomena and neural network resonance at the cellular level.
Fruitful findings were also obtained upon application of the
concept of entropy to brain processes (Rosso et al., 2001, 2002).
The slogan, “EEG is not simple noise”, was formulated and this
represented the conceptual renaissance in brain electrophysiol-
ogy. Interpreting their findings, the neurophysiologists stated:
“EEG is not noise, but is a quasi-deterministic signal.” All these
empirically derived approaches were in fact favorable to chaotic
dynamics of brain function.
Meanwhile, with a few exceptions, prediction and mathe-
matical description of brain behavior has not been in the
mainstream of brain research. Influential ideas such as those of
John von Neumann and Burks (1966) may have had their share
in this neglect: “…logics and mathematics in the central nervous
system, when viewed as languages, must be structurally
essentially different from those languages to which our common
experience refers.” Therefore, the “big bang” of applying
chaotic dynamics to brain activity has struck brain scientists all
too early, when they were not yet prepared to use these
concepts. Accordingly, studies that attempt to explain the brain
through chaotic dynamics could not gain the status of a coherent
research endeavor.
The present section presents a theory on brain function that
basically follows Newton's, Galilei's and Einstein's pathway. In
this theory, the language brain uses is the brain waves. The
oscillations in the different frequency bands are like the
phonemes in languages. Superimposed oscillatory responses
are the words. The selectively distributed parallel processing
pathways are the syntax of the brain language. And the whole-
brain-work that follows the super-synergy is the sentences and
the discourse in the language of the brain.
2. The theory of the whole-brain-work: an approach to
brain function by means of EEG-oscillations
Chronological evolution of our conceptual framework
evolved in the last 20–25 years. The development in the last
5 years has especially been fast. At its present state, the theory
of the whole-brain-work represents the extension of the
previously formulated neurons-brain theory.
2.1. Basic principles
The theory assumes that functions of the brain, especially
those in cognitive processing, are based on EEG and field
potentials, shortly, the oscillatory activity. The theory rests on
four basic principles.
Principle 1. Brain functions are represented by its oscillatory
activity. It is to be noted that this activity is the paradigm change
that Mountcastle (1998) had announced for brain sciences
toward the end of the last century.
Principle 2. There is cooperation between distant structures of
the brain and these can be measured by means of coherence and
phase differences. The whole brain is activated during cognitive
processing. Thus there is a super-synergy in the brain during all
percept-and memory-related processes.
Principle 3. The cooperation between brain structures is
selective. The selectivity may be demonstrated in the selective
distribution of the coherence functions over various brain
structures with values that vary between 0 and 1. The
demonstration of the principle of selective cooperation
requires the analysis of oscillations in several neural
populations and in several frequency windows. Such analyses
and the related findings have been instrumental in the further
refinement of the concepts pertaining to “whole brain” and
“cooperation”.
 E. Başar / International Journal of Psychophysiology 60 (2006) 133–138
Karl Lashley (1929) had concluded that the brain operates as
a “whole”. Oscillations that are selectively distributed in the
whole brain may be the mechanism whereby this wholistic
operation is achieved. The new tools and concepts that are used
in the analysis of brain electrical activity during sensory-
cognitive processing may also provide new interpretations on
Lashley's (1929) and Hebb's (1949) statements on brain
functioning.
Principle 4. The machinery the brain uses for super-synergy
consists of an ensemble of sub-mechanisms. These sub-
mechanisms act in synergy when stimulated by sensory and/
or cognitive input.
2.2. The theory in extended form: mechanisms in the whole-
brain-work
The theory of the whole-brain-work proposes that integrative
brain function is based on the coexistence and cooperative
action of many interwoven and interacting sub-mechanisms. In
its extension, the theory includes mechanisms which consist of
super-synergy, superbinding and reciprocal interaction of
attention, perception, learning and remembering (APLR-
alliance). These mechanisms were grouped in the theory
under four structural and/or functional levels.
2.2.1. Level A: transition from single neurons to oscillatory
dynamics of the brain
1. The neuron is the basic signaling element of the brain.
2. Since morphologically different neurons or neural net-
works are excitable upon sensory-cognitive stimulation,
the type of the structural element does not play a major
role in the frequency tuning of oscillatory networks.
Research has shown that neural populations in the
cerebral cortex, hippocampus or cerebellar cortex are all
tuned to the very same frequency ranges (Eckhorn et al.,
1988; Llinas, 1988; Singer, 1989; Steriade et al., 1990,
1992; Başar, 1998, 1999). These findings support the
suggestion that all brain networks communicate by means
of the same set of frequency codes of EEG-oscillations.
3. Intrinsic oscillatory activity of single neurons forms the
basis of the natural frequencies of neural assemblies.
Oscillatory activity of the neural assemblies or the brain
consists of the gamma, beta, alpha, theta and delta
frequencies. These frequencies are the natural frequencies
and thus the real responses of the brain (Başar et al.,
2001a,b,c).
4. Feature detectors (Sokolov, 2001), place cells and
memory cells (Fuster, 1995) are empirically established
constructs. However, a crucial turning point occurred
with the “grandmother” experiments. These experiments
showed that large groups of neural populations were
selectively activated upon complex semantic and episodic
inputs to the brain (Edelman, 1978; Bullock, 1992; Başar,
2004). These experiments and other similar studies
replaced the neuron with the neural assemblies in
135
attempts at describing the integrative functions of the
brain (Başar et al., 2001a). The emphasis on neural
assemblies differs Başar's theory from Sherrington's
“neuron doctrine” and Barlow's “new perception doc-
trine” (Barlow, 1995).
5. Sokolov (2001) has excellently described and also
constructively criticized the role of feature detectors.
However, integrative functioning of the brain needs the
selectively distributed and selectively coherent neural
populations in concert with the feature detectors.
6. The brain has response susceptibilities. These suscept-
ibilities mostly originate form its intrinsic rhythmic
activity, i.e. its spontaneous activity (Başar, 1980,
1983a,b, 1992; Narici et al., 1990). A brain system
responds to external or internal stimuli with those rhythms
or frequency components that are among its intrinsic
(natural) rhythms. Accordingly, if a given frequency range
does not exist in its spontaneous activity, it will also be
absent in the evoked activity. Conversely, if activity in a
given frequency range does not exist in the evoked
activity, it will also be absent in the spontaneous activity.
7. There is an inverse relation between EEG and event-
related potentials. The amplitude of the EEG thus serves
as a control parameter for responsiveness of the brain
which can be obtained in the form of evoked potentials or
event-related potentials (Rahn and Başar, 1993; Başar,
1998; Barry et al., 2003; Başar et al., 2003).
8. The EEG is a quasi-deterministic or a chaotic signal and
should not be considered as simple background noise.
This characteristic and the concept of response suscep-
tibility lead to the conclusion that the oscillatory activity
that form the EEG governs the most general transfer
functions in the brain (Başar, 1990).
9. Oscillatory neural tissues that are selectively distributed
in the whole brain are activated upon sensory-cognitive
input. The oscillatory activity of neural tissues may be
described through a number of response parameters.
Different tasks and the functions that they elicit are
represented by different configuration of parameters.
Due to this characteristic, the same frequency range is
used in the brain to perform not just one but multiple
functions. The response parameters of the oscillatory
activity is as follows: enhancement (amplitude), delay
(latency), blocking or desynchronization, prolongation
(duration), degree of coherence between different
oscillations, degree of entropy (Pfurtscheller et al.,
1997; Başar et al., 1999a,b; Miltner et al., 1999;
Schürmann et al., 2000; Kocsis et al., 2001; Rosso et
al., 2001, 2002; Başar, 2004).
10. The number of oscillations and the ensemble of
parameters that are obtained under a given condition
increase as the complexity of the stimulus increases or the
recognition of the stimulus becomes difficult (Başar et al.,
1999a,b, 2001a,b,c). These were demonstrated not only
through techniques of frequency analysis but also through
recent techniques of time-frequency analysis (Rosso et
al., 2001; Özdemir et al., 2005; Tağluk et al., 2005).
136 E. Başar / International Journal of Psychophysiology 60 (2006) 133–138
2.2.2. Level B: from superbinding of neural assemblies to
super-synergy
According to the theory of whole-brain-work, super-synergy
is obtained by way of the following submechanisms:
11. In simple binding, there is temporal coherence between cells
in cortical columns. This has been demonstrated by several
authors (Eckhorn et al., 1988; Gray and Singer, 1989).
12. Each function is represented in the brain by the
superposition of the oscillations in various frequency
ranges. The values of the oscillations vary on a number of
response parameters (Principle 9). The comparative
polarity and phase angle of different oscillations are
decisive in producing function-specific configurations.
Neuron assemblies do not obey the all-or-none rule that
the single neurons obey (Karakaş et al., 2000a,b;
Klimesch et al., 2000a,b; Chen and Herrmann, 2001).
13. The superposition principle indicates synergy between the
alpha, beta, gamma, theta and delta oscillations during
performance of sensory-cognitive tasks. Thus, according
to the superposition principle, integrative brain function is
obtained through the combined action of multiple
oscillations.
14. The response susceptibility of the brain activates resonant
communications in the brain by facilitating electrical
processing between networks (Başar et al., 1997a,b;
Başar, 2004). This could be also interpreted as a general
tuning process between neural populations and feature
detectors (Sokolov, 2001).
15. Parallel processing in the brain shows selectivity. The
selectivity in parallel processing is produced by variations
in the degree of spatial coherences that occur over long-
distances between brain structures/neural assemblies
(Başar, 1980, 1983a,b; Başar et al., 1999a,b; Miltner et
al., 1999; Schürmann et al., 2000; Kocsis et al., 2001).
16. Temporal and spatial changes of entropy in the brain
demonstrate that the oscillatory activity is a controlling
factor in the functions of the brain (Graben et al., 2000;
Graben, 2001; Quiroga et al., 2001; Yordanova et al., 2002).
As can be deduced from the foregoing explanations, the
concept of superbinding collectively denotes the mechanisms of
superposition, activation of selectively distributed oscillatory
systems, and the existence of selectively distributed long
distance coherences. The concept of super-synergy includes
superbinding and, additionally, entropy and the role of EEG-
oscillations as control parameter in brain's responsiveness.
2.2.3. Level C: integration, alliance and interplay in memory
Extension of the theory of whole-brain-work to cognitive
processing is governed by the following principles:
17. All brain functions are inseparable from memory function
(Hayek, 1952; Fuster, 1995, 1997). Like in all integrative
brain functions, memory is manifested as multiple and
superimposed oscillations. A specific superposition of
oscillations, each of which is characterized with the
response parameters in Item 9, represents the configura-
tion that is specific to the given type of memory.
18. Attention, perception, learning and remembering (APLR-
alliance) are interrelated. As the grandmother-experi-
ments demonstrated (Başar et al., 2003; Başar, 2004;
Özgören et al., 2005), memory-related oscillations are
selectively distributed in the brain. They have dynamic
properties and evolve upon exogenous and endogenous
input to brain. Memory states have no exact boundaries
along the time space. There is a hierarchical order that
takes place on a continuum but the boundaries of memory
states merge into each other. Memory functions from the
simplest sensory memories to the most complex semantic
and episodic memories are manifested in distributed
multiple oscillations in the whole brain.
19. In our theoretical framework, we introduced the expres-
sion “evolving memory” or “memory building”. The
critical factor in memory building is the APLR-alliance.
This concept represents a constant reciprocal activation
within its sub-processes. Evolving memory has a con-
trolling role in integrative brain functions (Edelman,
1978; Tononi et al., 1992; Barry et al., 2003). The
hierarchy of memories is not manifested with separable
states, since the memory manifests rapid transitions.
Therefore we suggest using the term “memory states”
rather than “memory stores”, a concept in which memory
is considered to take place in successive stages. These
explanations do not apply, however, to persistent memory
which can be inborn or obtained through over-learned
engrams or habits.
3. Causality in brain responsiveness
To discover the cause of an event is to discover something
among its temporal antecedents such that, if it had not been
present, the event would not have occurred. In the introduction
of this section, causality was described as Newton, Galieli and
Einstein conceptualized it. The present section considers
causality as it pertains specifically to the responsivity of the
brain. The theory of the whole-brain-work considers the
following as the critical causal factors of brain dynamics.
3.1. Genetically fixed causal factors
There are in the brain, or in the CNS-ganglia genetically
coded networks. The phyletic memory-networks that are inborn
play essential roles in the responsiveness of neural populations.
Accordingly, (a) occipital networks in the mammalian brain
respond to light stimulation with enhanced 12 Hz oscillations
(Başar, 2004). On the other hand, temporal auditory areas that
do not react to light stimulation respond to auditory stimuli with
10 Hz enhanced oscillations. (b) The ray brain reacts with 10 Hz
oscillations to electric stimuli (electroception); the human brain,
on the other hand, does not have this ability (Başar, 2004). (c)
Like alpha networks, there are selectively distributed gamma
networks in the brain. These networks show obligatory
responses to sensory stimuli (Karakaş and Başar, 1998). (d)
 E. Başar / International Journal of Psychophysiology 60 (2006) 133–138
Reflexes are genetically coded. The so-called “prepotent
responses” (Miller, 2000) in reflexive actions also partially
represent this type of causality. (e) Results of Sokolov (1975) on
the orienting response and the genetically fixed causal factors
have to be emphasized: There are expectation cells, which fire
upon expected input; sensory-reporting cells which fire in
response to actual stimulus; and comparator cells which fire
whenever there is a discrepancy between stimuli (Başar, 2004).
3.2. Dynamic causality due to reentry and to dynamic behavior
of EEG-oscillations during reciprocal activation of the APLR-
Alliance
Possible reentries (Edelman, 1978; Başar, 2004;), reciprocal
activation of attention, perception, learning and remembering,
recurrent inputs, and changes in prestimulus EEG-oscillations
(Barry et al., 2003) change the causality of the brain response in
a dynamic manner (Başar and Stampfer, 1985).
The present behavior influences the immediately following
future behavior. The plasticity in this adaptive behavior is
demonstrated in the oscillations, showing that oscillatory
plasticity is an additional causal factor in brain responsiveness.
In auditory and visual memory task experiments, the EEG-
oscillations manifest a high degree of plasticity: This is because
neural networks have susceptibility for being activated with
superimposed frequency codes, i.e., with multiple oscillations
showing varied degrees of enhancements (Başar, 2004). The
reciprocal activation of the APLR alliance (Başar, 2004) also
affects the future responsiveness of the brain, attesting for the
presence of oscillational plasticity in the higher cognitive
processes.
The existence of a significant difference in the major
operating oscillations in occipital or frontal areas gives strong
support to the possibility that spontaneous, evoked or induced
alpha rhythms or theta rhythms have fundamentally different
functional operations. But during some functional states, major
operating rhythms can change their functional roles. The nature
of the experiment and the task conditions can influence the
weight of the functional components on brain rhythms. This
behavior of brain oscillations is another instance of the dynamic
plasticity in brain responsiveness.
Recent results of Karakaş et al. (2003) showed that the early
sensory gamma response showed individual differences and,
further, the existence or nonexistence of the gamma response
could be predicted from a battery of neuropsychological tests
that measure attention, perception, learning and memory, in
short, the APLR-alliance. Such complex cognitive processes are
most probably multi-causal, including both the impact of genetic
codes and the influence of environmental conditions. A recent
study, however, showed that the presence of the early gamma
response is not to be sought for in gender differences (in press).
4. To sum up
Why did we use the term “whole-brain-work” for our theory?
This theory is based on principles and mechanisms according to
which the integrative functions that involve EEG-oscillations
137
are processed by the whole brain. The concepts of “all
functions” and that of “whole brain” are therefore brought
together in the present theoretical framework to produce the
synthetic phrase, “all-works of the brain” or shortly, “whole-
brain-work”.
The term “whole” indicates, first, the whole brain and the
holistic functions that such a brain produces. Secondly, it
indicates the fusion in function, space and time. Accordingly,
the theory of the “whole-brain-work” is an explanation of the
integration of all functions in the whole brain and the functional
duration throughout, which the brain works. Such integration is
more forcefully represented in the phrase “whole-brain-work”
than in “integrative brain function”. The term “work”, on the
other hand, indicates the additional dynamic component, one
that is not implied in the term “function”.
Influential studies and explanatory findings on electrophys-
iology have mainly been on the spike potentials. The theory of
the whole-brain-work explains brain function in cognitive
processing on the basis of oscillatory activity. The very existence
of a theory on the oscillations shows that EEG activity, be it
spontaneous or evoked, needs special emphasis. The theory of
whole-brain-work theory that this section presents provides the
rules that are to be considered when dealing with the oscillatory
dynamics of the brain.
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