Afference Copy As A Quantitative Neurophysiological Model For Consciousness

Journal of Integrative Neuroscience, Vol. 13, No.
2 (2014) 363402
c Imperial College Press
DOI: 10.1142/S0219635214400020
Aerence copy as a quantitative neurophysiological

model for consciousness
J. Integr. Neurosci. 2014.13:363-402. Downloaded from www.worldscientific.com

by Dr. Allan Coop on 07/12/14. For personal use only.
Hugo Cornelis and Allan D. Coop*

Three Way Street, P. O. Box 5160, Braddon
Australian Capital Territory 2612, Australia
*allan.coop@gmail.com
[Received 27 March 2014; Accepted 7 April 2014; Published 20 June 2014]
Consciousness is a topic of considerable human curiosity with a long history of philosophical
analysis and debate. We consider there is nothing particularly complicated about consciousness when viewed as a necessary process of the vertebrate nervous system. Here, we propose a
physiological \explanatory gap" is created during each present moment by the temporal
requirements of neuronal activity. The gap extends from the time exteroceptive and proprioceptive stimuli activate the nervous system until they emerge into consciousness. During this
\moment", it is impossible for an organism to have any conscious knowledge of the ongoing
evolution of its environment. In our schematic model, a mechanism of \aerence copy" is
employed to bridge the explanatory gap with consciously experienced percepts. These percepts
are fabricated from the conjunction of the cumulative memory of previous relevant experience
and the given stimuli. They are structured to provide the best possible prediction of the
expected content of subjective conscious experience likely to occur during the period of the
gap. The model is based on the proposition that the neural circuitry necessary to support
consciousness is a product of sub/preconscious reexive learning and recall processes. Based on
a review of various psychological and neurophysiological ndings, we develop a framework
which contextualizes the model and briey discuss further implications.
Keywords: Consciousness; explanatory gap; aerence copy; reex; inhibition; learning; memory; prediction; qualia; review.
1. Introduction and Philosophical Context

Although much is currently known about the vertebrate central nervous system
(CNS), little is known about the emergence of consciousness. Guided by philosophical
considerations, we review known psychoneurophysiological data to provide a plausible framework for conscious experience. This framework is developed from demonstrated neural properties that are widely considered fundamental to CNS
operations, including: reex activity (Sechenov, 1863), its organization (Sherrington,
1906) and timing (Maniadakis & Trahanias, 2014); and the role of inhibitory processes (Lee & Maguire, 2014; Schel et al., 2014), learning (Pavlov, 1927; Thorndike,
1911) and memory (Brown et al., 2007; Hebb, 1949). In doing this, we nd a possible
explanation for the origin of qualia within the CNS.
363

364
H. CORNELIS & A. D. COOP
Following satiation of biological imperatives such as survival and reproduction,

probably no other issues have come to stimulate human curiosity more, in one way or
another, than the descriptive, functional and explanatory mysteries surrounding the
idea of consciousness. It is an idea with which all thinking persons can engage as it is
central to the problem of how we understand the world

including ourselves

and our knowledge as part of the world, see Popper (1959). The challenge is to make
sucient room in our models of cosmology and the general nature of reality to permit
a consistent account of consciousness, and vice versa

our notions of consciousness
must accommodate an understanding of what it means for its content to be \reality
as a whole" (Bohm, 1980). However, over 2000 years of curiosity (Koch, 2009) seems
to have added little of substance to either discussion or knowledge.
Three considerations provide a context for the framework we develop. The rst
concerns Sherrington's (1940) contemplation of the planet, \furnace of molten rocks
and metals, now yielding thoughts and values. Magic furnace. Beside its alchemy and
transmutations the most impassioned dreams of Hermes Trismegistus and all his
fellowship dwindle to paltry nothing". The second proposes that, \problems are
solved not by giving new information, but by arranging what we have known since
long". (Wittgenstein, 1953). The third states that, \At stake are central key concepts
that directly involve fundamental convictions regarding the nature of man's inner
being, physical reality, the meaning of existence and related matters of ultimate
concern. . . . perspectives in this area profoundly shape human value systems and
societal decision-making and hence human destiny". (Sperry, 1980).
In short, (1) A remarkable structural evolution of matter has occurred, (2) we
already know much of what is needed to solve the problem of consciousness and (3)
we interpret and structure the world based on our understanding of how the brain
works. Importantly, we consider many problems of consciousness would evaporate if
it is accepted that construction of internal representations or replicas of the \outside"
world is unnecessary (Velmans, 2009). For as O'Regan (1992) and many others have
proposed it is continuously available \out there".
Depending upon the gure of speech chosen, consciousness has been identied with:
a state of being, a substance, a process, a place, an epiphenomenon, an emergent aspect
of matter, or the only true reality (Miller, 1962); and described as a fascinating but
elusive phenomenon for which it is impossible to specify what it is, what it does, or why
it evolved; with little worth reading written about it (Sutherland, 1989).
In the cognitive realm, nowhere has a dominant metaphor been more central and
susceptible to \sporadic reformulation" in terms of the technology of the day than
when theorizing about the brain (Daugman, 1993). For example, the following
metaphors, ranging from cosmological to mathematical, have all been employed:
embodied spirits and helmsmen (Arbib, 1972); hydraulics and mechanics (Vartanian,
1973, 1953; Resniko, 1988); electricity (von Helmholtz, 1850a,b; Hebb, 1949;
Hodgkin & Huxley, 1952) and optics (Pribram, 1969); networks of simple automata
and societies of mind (Dennett, 1978; Minsky, 1988); determinism of the unconscious
and automatic (Freud, 1904/1914); logical calculus of neurons (McCulloch & Pitts,

AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL
365
1943); the stochastic Boltzmann machine, spin-glass latices, Ising spins (Hopeld,
1982; Hinton & Sejnowski, 1986; Gutfreund et al., 1988) and chaotic dynamics and
attractors (Skarda & Freeman, 1987). Given such historical antecedents, it is
probably shortsighted to regard the current metaphors of the brain as information
processor and computer (Pylyshyn, 1986) or container of representations (Nadel &
Piattelli-Palmarini, 2003) as entirely dierent kinds of breakthrough in the history
of ideas.
Ever since publication of the classic paper by Nagel (1974), \What is it like to be a
bat?", the \intractable problem of consciousness" Churchland (1996), has been
identied as, how is it that consciousness of subjective experience is possible? This is
known as the \Hard Problem" (Chalmers, 1996), and concerns, \how and why
consciousness arises from physical processes in the brain" (Chalmers, 1997).
The \explanatory gap" is a philosophical term related to the \Hard Problem". It
refers to the lack of an explanation of how physical properties give rise to the qualia of
experience (Levine, 1983). We take a more pragmatic view of this problem than
those who consider it represents the limits of current knowledge (Churchland, 1996),
or of cognitive abilities (McGinn, 1989), or necessarily entails a metaphysical gap
(Chalmers, 1996).
Nevertheless, consciousness continues to be a prominent mystery for philosophy
and science (Chalmers, 1996). Initially, the preserve of philosophy, more recently
quantitive empirical studies have become both more accepted, and possible; with
rapid advances over the last two decades (Price & Barrell, 2012), as ongoing development of experimental techniques and new observations drive theory and research
(Blackmore, 2001).
We are, of course, unhappy with much proposed for the \problem" of consciousness, whether hard or otherwise. The fact that philosophers are forced to deny its
existence (Dennett, 1991), solve it by attribution of as yet unidentied irreducible
properties (Chalmers, 1996), or claim it is beyond the limits of human knowledge
(McGinn, 2004), seems somewhat unsatisfactory.
Our starting position is therefore similar to that of Churchland (1996); to partition
the mind-brain problem as Chalmers (1996) has done, \poses the danger of inventing
an explanatory chasm where (all that really exists is) a broad eld of ignorance". The
only conclusion from the fact that consciousness is mysterious, is that its mechanisms
are not understood. From the vantage point of ignorance, it is dicult to tell which
problems actually are harder and which will be solved rst. As Churchland (1996)
suggests, \learn the science, do the science, (then) see what happens".
The essence of the model presented here is a transguration of the philosophical
explanatory gap into a neurophysiological context. This allows the introduction
of a plausible \aerence copy" mechanism which bridges an otherwise inescapable
physiological explanatory gap. In doing so, we address the confounding observations
reported by Libet (2004, 1985), and more recently by Soon et al. (2008) and Bode
et al. (2011). The issues involved concern the timecourse of sensory stimulation and
voluntary motor acts and their emergence into consciousness.
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In proceeding, we aim to contribute to improved clarity of understanding by

rening concepts which may not previously have been well articulated. To this end,
we aim to explain, \How it is that subjectively we seem to exist in a seamlessly
perpetual and detailed present moment".

2. Considerations Developed About Consciousness in Dierent

Temporal and Phylogenetic Domains
In this section, we introduce biological considerations which assist in framing the
model presented here. We begin by noting that Penrose & Hamero (2011) report the
recognition of three general possibilities for the origin and place of consciousness in
the universe: (1) It is a quality that has always been in the universe, (2) precursors
have always been in the universe and biology evolved a mechanism to convert conscious precursors to actual consciousness or (3) it is not an independent quality but
has arisen as a natural evolutionary consequence of the biological adaptation of
brains and nervous systems,
Alternatively, the three major theories of consciousness taken most seriously by
neuroscientists include the view that (Block, 2009): Consciousness (1) must be
described in terms of higher order states, (2) is a property of a global workspace or
(3) is a biological state of the brain.
Solms (1997) has proposed the totality of human consciousness consists of three
domains: (1) Primary external perceptions experienced as material reality; (2) primary internal perceptions (aect) and (3) perceptions of activated traces of previous
experiences (memory and cognition), the latter two being experienced as introspective awareness (or psychic reality).
Alternatively, Tulving (2002) has proposed three forms of consciousness:
(1) Anoetic or \unthinking", which may be aectively intense, (2) noetic or cognitive
activity linked to the physical instantiation of the objects of exteroceptive perception
and (3) autonoetic or higher neocortical functions including, abstract perceptions
and cognition, conscious (self) awareness and reection on episodic memory, predictions of the future and fantasy or hallucination.
Solms & Panksepp (2012) mapped the preceding scheme to phylogenetic evolution
of: (1) Upper brainstem and septal areas (anoetic phenomenal experience), (2) lower
subcortical ganglia and upper limbic structures of the cortical midline (noetic consciousness and learning) and (3) association cortex, providing the critical substrates
for \reexive experiential blends that yield the stream of everyday awareness"
(autonoetic consciousness).
2.1. Consciousness

a reduction to fragments
If self-awareness is taken as evidence for consciousness, then in evolutionary terms
consciousness may have rst appeared about 5 million years ago and be at least as old
as the placental divide in mammals (Wildman et al., 2007), with the foundations
of consciousness and self-consciousness possibly formed as early as the amniote

367
radiation (Warren et al., 2008). The core of human consciousness is thought to have
originated in phylogenetically ancient structures activated by primitive emotions
mediating arousal, in conjunction with limited connectivity with frontalparietal
areas (Mashour & Alkire, 2013).
It has been well demonstrated that the operations of the CNS add color, sound and
a multiplicity of sensations to continual sampling of reality. This sampling is recorded
with remarkable veracity from atomic and quantum domains (Kaupp, 2010; Smith,
2002; Hecht et al., 1942). Such detail is likely captured at a resolution sucient for
the existence of physical phenomena to be demonstrated, quantied, modeled and
ultimately, understood.
The cerebral cortex is currently considered to be the primary site containing the
neural correlates of awareness (Mashour & Alkire, 2013). From a top-down perspective, evidence for this includes the desynchronized nature of the electroencephalogram,
activity in the thalamocortical system and widespread brain activity (Seth et al., 2005).
The coupling of an internally based need system with an externally directed situational awareness system is considered to provide a basis for the emergence of
consciousness and has been shown to be closely related to the mental machinery seen
in humans for generating arousal and awareness (Mashour & Alkire, 2013). The most
basic emotions and arousal states are associated with internal feedback networks that
guide an organism's behavior to the best possible outcomes. This functionality is
considered to underly essentially all behavioral choices in the vertebrate brain
(Mashour & Alkire, 2013).
One feature of experience putatively identied by philosophers is an insubstantial
component of consciousness called qualia. These have been dened as the internal
and subjective components of sense perceptions, arising from stimulation of the
senses by phenomena; they are recognizable qualitative characters of the given, which
may be repeated in dierent experiences, and are thus a sort of universal (Jackson,
1982). Qualia involve the subjective experience of the redness of red, the painfulness
of pain and so on.
At a psychological-level, Freud (1915) considered mental processes are in themselves unconscious and that their perception by consciousness is similar to the perception of the external world by the sense-organs. Unconscious mental activity is
therefore similar to all other natural processes (Solms, 1997), with most of mental life
unconscious most of the time, only becoming conscious as sensory percepts, such as in
words and images (Gray, 2002). Jackendor (1987) considers consciousness is not a
particularly high-level process, as everyone has always wanted it to be, and that \it is
not what makes us human".
Besides subjectivity, in humans at least, conscious life has two other dominant
features: unity and intentionality. The unitary nature of consciousness refers to the
appearance of subjective brain experiences as unied, integrated and a constructed
whole (Kandel, 2000b). Intentionality refers to the typical focus of consciousness
being about objects or events (Edelman, 2003), with emotional (LeDoux, 2000) and
semantic aspects (Chalmers, 1996), being central.

368
Ingvar (1985) considers conscious experience is based on three components:

memory of the past, experience of \actual events in the now-situation" and concepts
of the future or events which have not yet occurred. For him, the important question
is, \to what extent do events in the brain, related to the past, the present and
the future, take place at a conscious-(attentive) level or at a subconscious (pre- or
subattentive-level) or at both levels simultaneously".
In answering this question, Jackendor (1987) proposes that the character of the
phenomenological mind is comprised of unconscious and conscious elements but
that processing is always unconscious; in short, processes are never available to
consciousness in perception, thought, learning or action.
Many treatments of consciousness in the literature recognize, at least incipiently
(Jackendor, 1987), that consciousness is projected from mental information structures, where the stream of consciousness, made up of visual and verbal imagery, is
usually sucient to be believed to constitute thought. But Jackendor (1987) considers this may not actually be the case. Rather, visual and verbal images emerge
from intermediate structures, which are in turn generated from thought by mandatory fast processing modules. Hence, the stream of consciousness is essentially
nothing but evidence that thought is occurring in a subconscious domain, with both
the process of thought and its contents inaccessible to awareness.
Importantly, the fact that human cognitive processes are largely nonconscious has
been periodically reported, although often ignored. von Helmholtz (1910) observed
that we are not aware of the elements used to form a judgement

we make
\unconscious interferences" based on prior experience. Lashley (1956) further emphasized, \No activity of mind is ever conscious . . .Experience clearly gives no clue as
to the means by which it is organized". We are not conscious of the details that makeup a percept. Such details are inhibited from our conscious awareness. Instead, what
is seen depends largely on what is already known (Snyder, 1998; Snyder & Barlow,
1986; Gregory, 1970). Basically, every image is forced to t into a known percept
(Snyder et al., 2004).
This perspective on human perception is also strongly supported by reports of
detail blindness (No, 2007); e.g., change (Simons & Rensink, 2005; O'Regan, 2003),
inattentional (Mack & Rock, 1998) and repetition (Kanwisher, 1987) blindness; and
the attentional blink (Chun & Potter, 1995; Raymond et al., 1992). Space precludes
further review of this extensive literature, which we consider provides compelling
support for our conjectures.
Following a comprehensive review, Velmans (1991) concludes no human information processing is conscious in the sense that consciousness enters into or causally
inuences the process. What enters awareness follows the processing to which that
awareness relates.
Similarly, Berlin (2011) concludes that complex cognition can proceed in the
absence of consciousness, that the unconscious brain is active, purposeful and independent and can selectively access and activate implicit goals and motives. However,

369
he notes that exactly how unconscious emotions and evaluations help shape the
dynamics giving rise to conscious perception is still unknown.
Gray (2004) has postulated three aspects of consciousness: (1) It contains a model
of the relatively enduring features of the external world which is experienced as
though it is the external world, (2) features that are particularly relevant to ongoing
motor programs, or which depart from expectation, are monitored and emphasized,
(3) The control variables and set-points of the brain's nonconscious servomechanisms
are juxtaposed, combined and modied; in this way, error can be corrected.
At the core of his model is a \comparator", which compares actual stimuli with
expected stimuli

a function performed by a behavioral inhibition system (Corr,
2008). When there is no discrepancy, behavioral routines run uninterrupted and
stimuli are not extracted for detailed processing by higher-level cognitive processes.
When mismatch is detected between actual and expected states of the world, the
alien features of the error-triggering environment are subjected to controlled, attentional, analysis and (often) emerge into conscious awareness.
Perlovsky (2013) notes, brain imaging experiments have demonstrated that vague
mental states and the entire dynamic logic process (taking approximately 500 ms) are
unconscious. Only nal near-logical sensory percepts become available to consciousness. Most brain operations (more than 99%) are inaccessible to subjective
consciousness, while the mind operates by \jumps" among \islands" of consciouslogical states in an ocean of unconsciousness. In short, we are subjectively convinced
of our consciousness. Further, in a recent review, Aru & Bachmann (2014) argue that
attention and consciousness are independent from each other and phenomenal consciousness can emerge without attention.
2.2. Timing of the phenomenology of consciousness
Following Donders (1868), three fundamental response latencies or reaction times
have been conrmed, \simple", \recognition" and \choice" (Baayen & Milin, 2010).
For simple reexes in humans, the shortest reaction times are those of the unconscious blink reex with a mean latency of 10.8 ms (Shahani, 1970); whereas, reex
latencies for muscle in response to electrical stimulation range from 3151 ms (hand) to
5581 ms (foot) (Tarkka, 1986). Similarly, response times of 810 ms (auditory)
(Kemp, 1973), 2040 ms (visual) (Marshall et al., 1943) and 155 ms (touch) (Robinson,
1934), have been reported.
Although, subjectively, recognition of familiar objects and scenes appears virtually
instantaneous, this actually is not the case. For example, event related potentials in a
visual recognition task show the visual processing required to identify an animal in a
visual scene viewed for 20 ms requires up to 150 ms, with reaction times of 382567 ms
(median 445 ms, Thorpe et al., 1996).
We consider that on purely empirical grounds, these ndings argue directly
against any immediacy whatsoever for the phenomenology of consciousness. In
support of this position, in the following section (as one amongst several possible
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examples), we further briey explore vision in more detail. In particular, its temporal
properties and implications.

2.3. The problem of the rapid establishment of visual consciousness

Principled estimates of the timing of visual activity can be made. For example, any
activity requiring detail or color (e.g., driving or reading) must be located within the
foveal area by a saccade. The fovea occupies less than 1% of retinal area but over 50%
of the visual cortex (Krantz, 2012). It covers about two degrees of the visual eld or a
disc about 4 cm in diameter at arm's length. In vertebrates, the fovea may be compared to the location where axons converge to form the head of the optic nerve. The
resulting hole in the photoreceptor mosaic is up to six times the foveal area, which at
3 5 forms a surprisingly large \blind spot", equivalent to a 610 cm diameter disc
at arm's length (O'Regan, 1992).
Dierent saccades (Hopp & Fuchs, 2004), exhibit remarkably stereotyped
amplitude, duration and peak velocity (Becker, 1989). Their dynamics are determined by the point of origin within the visual eld and the trajectory of movement
employed to locate a target. Typically, they last about 25190 ms for movements of
5 60 , respectively (van Beers, 2008).
Reactive targeting saccades exhibit latencies of about 180 ms (Smit et al., 1987),
but may be as fast as 100135 ms (Fischer & Ramsperger, 1984; Fischer et al., 1993),
whereas, memory-guided saccades may take well over 200 ms (Hopp & Fuchs, 2004).
The duration of the minimum ocular pause time in the absence of stimulus processing is about 200 ms, with processing requiring an additional minimum 50100 ms
(Salthouse & Ellis, 1980). Although, for example, when reading, information may be
extracted from foveal input within about 50 ms (Rayner et al., 1981), with ocular
pauses of 100200 ms duration (van Diepen et al., 1995; Harris et al., 1988). Importantly, saccade optimization probably occurs during development rather than
being innate (van Beers, 2008), as the dynamics of equivalent saccades vary across
subjects (Boghen et al., 1974; Schmidt et al., 1979).
The number of foveal xations (Nf ) required to cover the area of the binocular
visual eld can be estimated from the number of square degrees (Guthrie, 1947), in
the visual eld (Av ) and fovea (Af ):
Nf
Av
Af
2
200 135 17; 189

6; 750:
2
2:5465
22
If the optimal duration of a two-degree saccade is assumed to be 30 ms (van Beers,

2008) and the minimal ocular pause 100 ms (well below the 250 ms required for visual
processing), the time required to scan the entire visual eld at foveal resolution with
7.5 xations s 1 is about 15 min. A more physiologically realistic rate of three xations s 1 would require up to 37.5 min.
Given these gures, it is not surprising that an infant takes several months to
develop, or more preciselylearn, accurate hand-eye coordination. However, what is

371
surprising is adults subjectively experience a very rapidly established high resolution global visual acuity in familiar environments. The question is, given that for
any one viewpoint this should take almost 40 min to establish, how is it so rapidly
generated?
A partial answer is provided by Hebb (1949) who suggests, \during the continuous, intensive and prolonged visual training of infancy and childhood, we learn to
recognize the direction of line and the distance between points, separately for each
grossly separate part of the visual eld". Further, \[I]t takes months for the rst
direct apprehension of a gure such as a plain, well-marked triangle to be established.
The normal human infant, apparently, reaches this stage quite early in life; but his
further training continues every moment that his eyes are open, and must extend his
capacity for prompt recognition of patterns falling outside the macula. . . . The signicant fact is that characteristic normal generalization only shows up after a prolonged and arduous training process."
There is every reason to expect that similar extended training occurs with other
senses. In summary, it would seem rapid establishment of consciously experienced
perception may actually be enabled by prior learning; following sensory activation,
percepts are evoked as required from the memory system.
2.4. The subjective antedating of perception
Two main types of organization of specic aerent systems have been reported for
vertebrates (Voronin et al., 1968): (1) Individual sensory modalities in sh partition
to dierent brain structures, whereas, (2) in mammals, all sensory modalities are
collected in the forebrain. Those of amphibians and reptiles lie between these two
bounds. Regardless of how basic human sensory modalities are classied, the dierent
perceptual qualities they generate are the constituent elements of the envelope of
consciousness and nothing else exists (Solms, 1997).
A basic problem related to the conscious perception of stimuli and initiation of
voluntary activity has been identied. Many studies conrm that, close to sensory
threshold, a delay of up to about 500 ms occurs before cerebral activities initiated by
stimulus detection systems achieve the duration and intensity of stimulation or
\neuronal adequacy" necessary to successfully elicit conscious awareness of the
stimulus (Libet et al., 1964). (Although, stronger stimuli may reduce this delay to as
little as 100 ms.) The problem is that following neuronal adequacy, the subjective
timing of the experience appears to occur without the actual delay required for
neuronal adequacy to elicit conscious experience of the stimulus.
One inference from such a result is that the experience of a skin-induced sensation
is elicited at a cerebral-level with a much shorter delay than for cortically-induced
sensation, an explanation discounted by Libet (1981).
As a solution to this problem, Libet (2004) has proposed that following neuronal
adequacy, subjective timing of the experience is automatically referred backwards to
the moment when primary sensory cortex received the stimulus (2025 ms after
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peripheral stimulation). The experience is thus \antedated" and subjectively appears

to occur without the substantial delay required for conscious experience of the
stimulus.
Below, we provide a less paradoxical explanation. It forms the basis of the
\aerence copy" model.

2.5. Emotion
For completeness, we now make a few comments about the role of emotion in neural
function. (For more details, see for example, LeDoux (1996)). In a review, Vuilleumier & Huang (2009) report that interactions between brain systems involved in
emotion and attention may contribute to regulating behavior and awareness by
enhancing relevant sensory information. Emotional stimuli may also evoke unconscious reexive and involuntary processing under many conditions. Further, involuntary monitoring of emotional stimuli and reexive boosting of perceptual processes
may reect some \default" settings or intrinsic preparedness within neural pathways.
They can also be adaptively shaped by various regulatory mechanisms that themselves operate potentially with or without conscious control. Finally, we note Solms
(1997) proposes that aect must be considered a sensory and perceptual modality.
3. Biological Framework for a Model of Consciousness
Based on the foregoing, and as outlined in more detail below; we present a model
where unconscious precursors of the conscious experience of each given moment are
molded within the memory system by integration with aerent stimuli (including
emotional aects experienced as qualia).
Our claim is that through the physiological mechanisms supporting this model,
the repetitive nature of a purely reexive existence is transcended, content created
and subjective experience expanded. Through these underlying mechanisms, the
cumulative procession of known prior events and activities inform current experience
and, via a surprisingly counter-intuitive mechanism, lay the basis for future behavior.
Each successive moment is formed from, and experienced through, the cumulative
record or memory of all relevant previous moments. In the deepest sense, each
present moment is the last remaining moment; experienced not in its encapsulation of
the past, but as an experience bounded prediction of the future.
From this perspective, we consider there is sucient empirical data to formulate
several broad conjectures from which a framework for consciousness might be
developed:
(1) With the exception of innate reexes (Zafeiriou, 2004), non-conscious activity in
the CNS is generated, organized and controlled through elaboration of reex
circuits assembled by associative learning.
(2) Inhibitory processes are the preeminent mechanism whereby the CNS controls
neural activity.
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(3) All CNS functions, from meaningful integration of sensory stimuli to consciousness and generation of motor plans, arise from ongoing (prenatally initiated) renement of learning.
(4) Memory is a repository for learning and throughout life it is continually formed,
and modulated, by accumulated moment-to-moment sampling of all available
sensory modalities.
(5) The physics of CNS function requires, and evolutionary pressures drive increased
capacity for anticipatory prediction.

We now briey explore evidence supporting these conjectures.

3.1. From spinal cord reexes to reexive behavior
The reexive nature of nervous systems is an evolutionary requirement imposed by
the need for control of reliable response in ever-changing and dynamic environments.
Thus, Sechenov (1863) was convinced that, \as far as the presence in man of three
separate mechanisms directing the phenomena of conscious and unconscious life is
concerned (viz. the mechanisms of the pure reex, and those of reex inhibition and
augmentation). Let anyone who thinks this hypothesis is doubtful, even poorly
demonstrated, or simply unacceptable, controvert it . . .If anyone nds a better
explanation, I shall be the rst to welcome it".
Since the early 19th century, it has been recognized that \just as the spinal reexes
[bear] a `genetic' anity to the irritability of the simplest animals and of plants, so
the operation of the highest nervous centers in humans [retain] the fundamental
reex character of the lower" (Clarke & Jacyna, 1987). For example, Laycock (1845)
concluded that, \the ganglia within the cranium being a continuation of the spinal
cord, must necessarily be regulated as to their reaction on external agencies by laws
identical with those governing the functions of the spinal ganglia and their analogues
in the lower animals".
Sechenov (1863) was the rst to propose that all aspects of cognition in humans
are based on behavioral reexes or more generally that reex circuits provide the
basis of all nervous function.
During the 19th century, the reex concept became basic for attempts to explain
the physiological functioning of the nervous system. By the end of the century the
tendency was to include increasing parts of animal behavior under reexive movements; that is, movements elicited through stimulation originating from an organism's external or internal milieu.
Early in the 20th century the reex arc was established as the functional unit in
CNS integration (Sherrington, 1906). Magnus (1924) demonstrated the reex nature
of all the elementary motor activities, and it was believed stereotyped movements,
such as those involved in walking, swimming and other locomotion, could be analyzed and described in terms of reexes. Sequential patterns of eector activity were
explained as arising from coordinated patterns of exteroreceptive and proprioceptive

374
excitation of the CNS. Richet (1925) introduced \the conception of the psychic reex,
in which the response following on a given stimulus is supposed to be determined by
the association of this stimulus with the traces left in the hemispheres by past
stimuli". In summary, the reex concept was used to explain the integrative functions
of the brain previously considered to result from \psychic activities" (Jrgensen,
1974).
Subsequently, motor-related activity in the CNS was shown to not necessarily
reect corresponding patterns of peripheral sensory activity. Thus, it was assumed
animal locomotion and other types of movements are determined by central pattern
generators, where the role of peripheral stimulation and sensory feedback is to
activate and modulate these generators (Bullock, 1961).
More recent interpretation suggests pattern generators may be intrinsic spinal
processors comprised of hybrid feedforward/feedback systems which optimally
compensate for both disturbances and sensor noise and can adapt central input to
this optimized peripheral input (Kuo, 2002). This is not inconsistent with pattern
generators being viewed as examples of sophisticated reex circuits.
The denition of a reex has been quite variable, ranging from the restrictive,
\stimulus-evoked response that usually involves a single muscle or a limited group of
muscles" (LeDoux et al., 2009), to the more general, \all neural processes (or neural
responses) and following eector responses evoked by any currently acting stimulus",
where a stimulus is a physical event (or a change in physical energy) which elicits the

activity of sensory receptors or higher-levels of the aerent nervous system ( Zernicki,
1968). In short, a reex is an automatic or involuntary and near instantaneous
response to a stimulus. Importantly, it is possible to conclude a reex circuit may be
activated by any sucient stimulus.
As McCulloch (1947) has clearly stated, modes of functional organization of the
cerebral cortex are reexive: \To that great extent to which its aerents inform it of
the peripheral consequences of the action of its own eerent, any system is part of the
path of a reex".
Complex or chain reexes located in the CNS have been demonstrated to consist of

a number of unitary processes, for which feedback control is essential ( Zernicki,
1968). Further, Slonim (1968), notes that complex, obligate stereotypic behavior
generated on the base of natural conditioned reexes

in contrast to articial or
facultative trainingare components of specic adaptive behaviors required under
dierent ecological conditions of existence.
Both simpler innate and more complex behavior comprising chains of individual
motor acts can generate and repeat themselves stereotypically and independently of
the conditions of postnatal development. Thus, between the \instinct" and the
\unconditional reex" there is a chain of stages dependent on either reex or the
automatic nature of nervous system activation (Slonim, 1968).
It is well-known in humans that, introspectively, some stimuli produce psychic
responses which may be assumed present in higher animals (Doty, 1967; Thorpe,
1966; Beritov, 1965). It has further been assumed that cerebral responses are

375
manifestations of denite unitary central processes or neuropsychic processes, of

which there are several kinds: eectively indierent or gnostic processes, or unpleasant or pleasant processes that are negative or positive emotions, respectively.
Psychic responses may additionally be divided into perceptions and images.
Signicantly, Konorski (1967) hypothesized that both perception and image are
manifestations of excitation of the same neurons, with the dierent psychic responses
(perception vs. image) due to dierent ways of exciting these neurons; from the
periphery and by association, respectively. There is a good reason to believe neuropsychic processes are located in the cerebrum, gnostic processes mainly in associative cortex, and negative and positive emotional processes in dierent places of the

limbic system and hypothalamus ( Zernicki,
1968; Olds, 1956). Importantly, this
seems to map well with the schemes of Tulving (2002) and Solms & Panksepp (2012)
mentioned above.
More recently, LeDoux et al. (2009) proposed that many behaviors fall into one of
four categories: reex, reaction, action and habit; where habit formation (the most
complex) and much of learning are dependent on conditioned reexes. For example,
the brain of a student typist must coordinate sensory impulses from both eye and
muscle to direct their ngers to particular keys. After sucient repetitions, the
ngers automatically nd and strike the proper keys even when the eyes are shut.
The student has \learned" to type; that is, typing has become a constellation of
conditioned reexes (Lagasse, 2000).
The simplest neuronal circuits are the monosynaptic reex arcs composed of one
sensory neuron and one motoneuron, for example, the circuits subserving the patellar
or achilles reex. Sherrington (1906) considered the whole nervous system to be based
on this type of circuit, \even those of the cortex of the cerebrum itself". However, all
but the simplest reex circuits are polysynaptic, where one or more interneurons are
interposed between the aerent (sensory) and eerent (motor) neurons of the
monosynaptic circuit.
Once again, Sechenov (1863) has asked, \how is it possible, (in the case of the
sensation of red), to speak of the paths of reexes?" Only to answer, \It is possible,
because though we do not know what is going on in the excited nerves, muscles or
brain centers, we cannot but see the laws of pure reex and cannot but consider them
true . . .we can say that the nature of the given conscious act of man is the same as
that of the reexes".
An explicit example of the complex behavior generated by a relatively simple
reex circuit is that found in the enteric nervous system. Associated with the gastrointestinal tract, it is comprised of a similar number of neurons as the spinal cord
(Furness & Costa, 1980). Within it, several types of neuromuscular activity are
autonomously controlled by intrinsic sensory neurons (Bulbring et al., 1958). They
include, stationary contractions associated with segmentation (Cannon, 1902), and
two types of reex (Cannon, 1911); one a slowly advancing contraction, the other a
\peristaltic rush" which propagates rapidly.

376
Numerical simulations have shown the circuit underlying this reex is sucient to
replicate both stationary and peristaltic activity, with the neuromuscular response
determined by the state of intestinal contents (gas, liquid, solid), i.e., the details of
the stimulus (Coop & Redman, 1992). These results suggest that more complex reex
circuits within other neural systems can reliably generate a range of seemingly plastic
behaviors in response to the specic details of circuit activation. Here, it is only
necessary to mention the \absent minded" week end drive resulting in unexpected
arrival at the workplace to recognize the ability of subconsciously active and entirely
learned reex pathways to formulate, control and sustain extended patterns of
complex behavior.
On the basis of the long tradition briey reviewed here, we consider that a majority of fundamental CNS operations are essentially reexive. Further, as reviewed
above, it is their reexive nature which renders their activity unconscious. In this
domain, equivalent stimuli reliably generate equivalent responses, modulated by
circumstance, where subtle dierences in circuit activation may evoke either nely
graded or signicantly dierent output.
Phylogenetic elaboration of the CNS, particularly within associative cortex
(Buckner & Krienen, 2013), results in both increased temporal separation between
reex pathway activation and response, and increased complexity of learned reexive
behavior; with (as we propose above) all such reexive activity constrained to the
subconscious. Ultimately, as Velmans (2009) has put it, \an entity in the world is
reexively experienced to be an entity in the world".
In summary, Sherrington (1906) originally described the vertebrate nervous system
as an orchestrated constellation of increasingly elaborated reex pathways culminating in a head ganglion or brain. From this perspective, the known record of
vertebrate evolutionary history (see e.g., Butler (2009)), and development of the
associative circuits of the hominid nervous system (Buckner & Krienen, 2013), it is
likely appropriate inhibitory processes and modulation of intrinsic and learned reexes
is fundamental to CNS function, and thus, ultimately, behavior and consciousness.
3.2. Inhibition and disinhibition of reexive behavior
Typically, a person might be considered capable of expressing a near innite repertoire
of behaviors. However, it is clear that in the normal course of events, they are never all
expressed simultaneously. Rather, sequences of behavioral patterns, each appropriate
to the given circumstances, are expressed as ongoing behavioral ows generated in
response to perceived environmental concerns. The emergence of a particular behavior
from a repertoire of possible behaviors is controlled by the (psychological) repression
or (physiological) central inhibition of alternative behaviors (Beritov, 1968). This is
the basic factor providing for the integrity of behavioral reactions. Further, Beritov
(1968), reports central inhibition caused by stimulation of sensory nerves embraces
the entire neuroaxis and follows adequate stimulation of all sensory modalities. In
humans, general central inhibition also occurs during deliberate cognitive activities,

377
such as arithmetic and suppression of conditioned and unconditioned reex activity.

We take this to mean that tonic inhibition at the physiological-level is responsible for
repression in the cognitive and behavioral domains.
As Smith (1992) has concluded, \Inhibition . . . became the controlling mechanism
that made possible those very functions, thought and deliberative action, that gave
humanity dignity and independence from nature. The possibility of inhibition was
the possibility that past associations rather than immediate sensations would
sometimes initiate movements. It was the possibility that sensation might combine
and recombine as intelligence and thought. Without inhibition, sensation merely
exhausted itself in movement".
Gray (2002) considers that the inhibitory function of consciousness solves a major
evolutionary problem: How to ensure that automatic responses are appropriately
activated; and how controlled processes are invoked only at critical junctures when a
denite choice must be made between cautious risk-assessment and prepotent
responses. At these moments, after ne-grained analysis aorded by control processing, appropriate adjustments can be made to the automatic system, such that
when the same (or similar) stimuli are encountered in the future, automatic-reexive
behavior will be more appropriate. In this way, controlled/conscious eects come to
determine automatic/nonconscious eects, albeit with a time lag.
We have previously concluded that, in the electrophysiological domain, ring of
hippocampal pyramidal and cerebellar Purkinje neurons (Coop & Reeke, 2004; Coop
& Redman, 1998), may more immediately be controlled through disinhibition and
eective connectivity, than alterations in aerent stimulation frequency.
It has recently been reported frontal cortex mediates unconsciously triggered inhibitory control (van Gaal et al., 2008), and a role for inhibition in the control of
cortical activity is further supported by disinhibition being a signicant control
mechanism in the spinal cord (Sechenov, 1863), striatum (Chevalier & Deniau, 1990)
and somatosensory cortex (Xing & Gerstein, 1996a,b,c), including olfactory bulb
(Cazako et al., 2014). More generally, there is persuasive evidence the cortex may be
under widespread and strong inhibition (Calford & Tweedale, 1991; Dykes et al.,
1984), while pathological alterations to inhibition may lead to the excitatory dysfunctions of epilepsy (Avoli, 2004) and schizophrenia (Lewis et al., 2005). A recent
review of the functional signicance of intrinsic oscillatory brain properties concluded, \it is by selection, via inhibition that the most elaborate neuronal patterns
are generated in the CNS" (Llin
as, 2014).
3.3. Learning and habituation
In a recent review, Schacter et al. (2011) consider human learning to be acquisition of
knowledge or skills through experience, study, or by being taught. However, learning
is not usually immediate, but rather typically is an incremental process building
upon, and shaped by, what is already known; to produce relatively permanent
changes in the behavior of an organism.

378
Two types of learnings have been distinguished (Kandel et al., 2000a). The simplest is non-associative. Two forms are common, habituation (Sokolov, 1963) and
sensitization (Shettleworth, 2010). Two forms of associative learning have also been
identied, classical (Pavlov, 1927) and operant (Thorndike, 1911) conditioning.
As suggested by Sechenov (1863), learning may contribute to understanding,
knowledge and as we have seen, even qualia (where qualia have been proposed to
originate in local cortical network activity (Orpwood, 2013)), \the successive reexes
acquired by learning lead to a perfect notion of the object, to knowledge in its elementary form. Indeed, the scientic knowledge of external objects is simply an innitely broad notion of each of them, i.e., the sum of all possible sensations evoked in
us by these objects under all conceivable conditions . . .I shall merely point out that
sensations from all spheres of the senses can be diversely combined, but always by
means of consecutive reexes. These combinations give rise to countless notions that
arise in childhood, notions providing, so to speak, material for the entire subsequent
psychical life".
In other words, when objects in the environment occur frequently enough, with
repeated regularities, they can be incrementally added to the memory of similar
previous encounters. Slight variations over time elaborate the memory system far
beyond that of any single sensory experience, and as Sechenov suggests, this continually enriches records of sensory experience and thus conscious phenomena.
It is notable that at the electrophysiological level, very young animals may only
receive and integrate relatively small amounts of information per unit time in circuits
which function more slowly than those of adults (Scherrer, 1968). Associated learning
may occur consciously or unconsciously. For example, the eects of internal stimuli
have been shown to be active as early as the embryonic stage, particularly in studies
of conditions under which the onset and patterning of electrical activity takes place in
the higher parts of the embryonic brain.
Even at this early stage, central inhibition displays its controlling inuence on
coordinating reex activity (Biryukov, 1968), and there is evidence for prenatal
habituation in humans as early as the 32nd week of gestation (Sandman et al., 1997),
indicating the nervous system is developed for learning and memory formation prior
to birth.
3.4. The unity of fragments of memory
The recollection of past experience is considered to be a reconstructive process. It can
be broken down into a number of constituent processes, with memories recreated
from their component parts, although little is known about the neural correlates
(Hassabis & Maguire, 2009). As such, human memory has been considered comprised
of a variety of testable components forming a \memory system"a containing past
a We recognize the nomenclature generally accepted in memory research, where \memory system" refers to the
multistore memory model (LaRocque et al., 2014). However, for convenience we use this phrase to refer to the more
state-based or unitary memory model described by Brown et al. (2007).

379
experience (Atkinson & Shiffrin, 1968). This multi-store modal memory model proposes sensory memory containing short-term records of sensory stimulation (250
500 ms duration). Those stimuli which are attended enter short-term memory (up to
18 s duration), and if suciently rehearsed are transferred to long-term memory (in
principle unlimited).
Each of these memory domains has subsequently been further decomposed. For
example, short-term memory is associated with a working memory (Baddeley &
Hitch, 1974), while long-term memory may be divided into either explicit memory
which is partitioned into episodic (Tulving, 1972), procedural (Milner, 1962) and
semantic (Tulving, 1972) components; or declarative memory, comprising implicit or
procedural memory (Schacter, 1987) or emotional memory (LeDoux, 2000).
One signicant problem with this multi-store approach is that dierent principles
are assumed to apply over dierent time scales. This runs counter to the generally
held expectation that scientic principles should hold over a wide range of temporal,
spatial, or physical scales (Barenblatt, 1996). As a consequence, Brown et al. (2007)
have proposed memory is unitary over all time scales, from milliseconds to years.
Here, the basic idea is that items are more distinctive, and hence both more
memorable and easier to identify, to the extent they are located in sparsely-populated
regions of psychological space. This implies forgetting is a consequence of reduced
local distinctiveness, not trace decay. Importantly, the same mechanisms are used for
retrieval over all time scales. There is considerable empirical support for this model as
it resolves many observations not easily accounted for by the more widely accepted
multi-store memory model.
This so-called temporal ratio model is concordant with the synaptic trace theory of
memory rst proposed by Hebb (1949). He posits the brain retains information
through learning-induced changes in the synaptic connections between neurons.
Once consolidated, a memory is then embedded and embodied through its xed
trace. Thus, learning occurs through general mechanisms of experience-dependent
synaptic plasticity, e.g. Hubel et al. (1977), which ultimately lead through a cascade
of intracellular molecular mechanisms to the formation of long-term memory. Subsequently, it has been shown memories can temporarily be rendered labile and sensitive to modication. Once an existing memory has been destabilized, it is possible to
enhance and even incorporate new content (for review, see Flavell et al. (2014)).
For over a century, psychologists have focused their studies on memory of the past.
However, a signicant function of memory is its role in allowing individuals to imagine
possible future events (Schacter et al., 2007), to guide behavior in the future based on
analogies, through memory resulting from both real and imagined experience (Bar,
2009b). Schacter & Addis (2009) consider that predicting the future and remembering
the past may be more closely related than everyday experience suggests. It has recently been shown that similarities in cognitive processes underlying past and future
events are complemented by analogous similarities in brain activity. For example, the
same \core network" of brain regions is recruited when people remember the past and
imagine the future (Buckner et al., 2008; Schacter et al., 2008, 2007).
380

In this view, past and future events draw on similar information stored in memory
and rely on similar underlying processes: in an active creative extrapolation, memory
supports the construction of future events by extracting and recombining stored
information into a simulation of a novel event. Schacter et al. (2008) further suggest
that simulation of future events requires a system that can exibly recombine details
from past events. Thus, memory can be thought of as a tool used by the prospective
brain to generate simulations of possible future events. Schacter et al. (2007) consider
such a hypothesis requires a shift of conceptual emphasis with regard to the role of
memory in cerebral activity.
3.5. Prediction as reconstruction of the future
Prediction is pervasive throughout much of brain function and is almost continually
operative at conscious and reex-levels, as success in a goal-oriented, moving system
is enhanced by such innate functionality (Llinas, 2001). For the nervous system to
predict, it must at least perform a rapid comparison of the sensory-referred properties
of the external world with a separate internal sensorimotor representation of those
properties (Llinas & Roy, 2009).
If it is to synchronize with the external events of each given moment the brain
must leave itself enough time to implement movement decisions. It cannot be stuck
doing something when required to perform another task. The consequent mode of
operation derives from a \look-ahead function", proposed to be an inherent property
of neural circuits (Llin
as, 2001).
Pellionisz & Llin
as (1979) provide a model of such a function based on the Taylor
expansion. It yields predictions of the frequency-time function of cortical input and is
an emergent property of inherently parallel distributed neural circuits. It relies on
relationships between neuronal ring rates and events in the external world. If some
neurons respond quickly, some do so at intermediate velocities, and some measure
events in real time, the output of such a circuit will be a reconstruction or prediction
of an event ahead of its time of completion. It is widely believed, in the absence of any
other possible sources, such functionality is memory-based (Bar, 2009a).
The signicance of such a mechanism is found mainly by its incorporation into
larger, cognitive states or entities. In other words, the extent to which sensory cues
aect brain function is determined by their impact upon pre-existing functional
dispositions of the brain (Llin
as, 1987, 1974). Llinas (2001) considers this indicates a
far deeper issue than might initially appear. In fact, the predictive abilities of the
brain may be profoundly more fundamental than suggested by a purely look-ahead
model.
We extend this approach to provide a signicantly more comprehensive predictive
functionality which lies at the heart of the aerence copy model. It is distinguished by
originating entirely within the memory system, an absence of immediately direct
comparison between sensory-referred properties of the external world and internal
sensorimotor correlates, and provision of a basis for conscious temporal detachment.
381
In fact, we claim that through infant learning, physiological processes of aerence

copy are evoked as sets of embedded reex functions. They lead to the creation of
temporal experience and consciousness through their ability to fabricate the phenomena of the present moment. The correlate is development of an awareness which
enables distinction of the past from the future and thereby create the subjective
temporal experience and phenomenology of each given moment.

3.6. Estimates of memory capacity and rate of formation

Unquantied beliefs concerning the characteristics of the human brain have long
been considered to make it an outlier in evolutionary terms and are frequently
employed to justify its remarkable cognitive abilities. However, the human brain has
been found to contain just the number of neurons and non-neuronal cells expected for
a primate brain of its size; with the same distribution of neurons between its cerebral
cortex and cerebellum as in other species, along with appropriately scaled energy
costs (Herculano-Houzel, 2011, 2012).
In concordance with our identication of a putative aerence copy system, in this
section we take the opportunity to provide some estimates of what might be referred
to as the content acquisition and content storage domains of the human CNS. These
may be distinguished from the content manipulation domain.
With regard to content manipulation, there is a considerable literature reporting
estimates of the transmission rate of information by neurons within the CNS. These
estimates are typically based on an assumption of either a rate or temporal code.
Theoretically, for a rate code with strictly periodic neuronal ring, the information
per spike is calculated from log2 N =N , where N gives the number of spikes per
second. This gives values of 3:3 10 1 and 6:6 10 2 bits spike 1 for ring rates of
10 and 100 spikes s 1 , respectively. Alternatively, for a temporal code, where the
variability of spike timing is taken into account, the calculation of information
transmission becomes considerably more complicated (e.g., see Stein (1967)). In
short, for example, information per spike increases to 34 bits spike1 (Softky,
1995), with a theoretical maximum of about 9 bits spike 1 (MacKay & McCulloch,
1952).
However, here we are initially interested in the content storage domain of the
memory system, in particular, an estimate of its \storage capacity". Such an estimate
then allows a \mean acquisition rate" to be determined for the content acquisition
domain. In doing this we are estimating a possible rate at which experience may be
recorded prior to transformation to states of the nervous system within the content
manipulation and storage domains.
One way to quantify these domains is in computational terms. For example, a
synapse can be represented as a single binary bit. Based on this assumption we can
employ the following data to make an \order of magnitude" estimate of storage
capacity. The aim is to give some sense of the putative capacity of human memory
in computational terms. We note, better estimates will be considerably complicated

382
by, for example, neuronal loss due to ageing in the frontal lobes (up to 20%
(Masliah et al., 1993; Gibson, 1983)), and likely continual addition of neurons in,
at least, the hippocampus (Altman, 1963), olfactory bulb (Altman, 1969) and
striatum (Ernst et al., 2014).
The human brain has been reported to contain 86:1 10 9 8:1 10 9 neurons,
of which 19%, or approximately 16:4 10 9 are cortically located (Azevedo et al.,
2009). The number of synapses in the neocortex has been reported at 1:5 10 14
(Pakkenberg et al., 2003). Assuming non-cortical neurons exhibit a similar average
number of synapses as cortical neurons, and all synapses may be modied by experience; the total estimated number of synapses in the brain may be as large as
7:89 10 14 .
At one bit per synapse, this number of synapses corresponds to a storage capacity
of about 90 Tb. Assuming a human lifespan of 70 years, with the knowledge there are
approximately 3:2 10 10 ms in a year, it can be estimated there are on average about
358 \naive" synapses available each millisecond to record the state of the nervous
system. From a computational perspective, this corresponds to storage of
44 bytes ms 1 or 43 Kb s 1 , 2.5 Mb minute 1 , 150 Mb hour 1 or 3.5 Gb day 1 . This
daily value is comparable with high denition movies which consume approximately
2 Gb hour 1 . Interestingly, by this analysis, a 90 min movie has a similar storage
requirement to our estimate of the average daily human storage capacity.
These gures may signicantly increase if it is accepted that new memories cannot
be formed during sleep, which is considered to be reserved for the stabilization,
enhancement and integration of memories (Walker et al., 2005). If it is assumed on
average a person spends approximately 8 h sleeping each day, then memory storage
capacity may increase up to about 470 synapses per millisecond, corresponding to
storage throughout waking life of 57.3 Kb s 1 or approximately 200 Mb hour 1 .
From a computational perspective, such calculations provide an indication of the
general volumes of \data storage" or average sustainable daily rate of lifetime
memory formation and provide a starting point for the development of more practical
hypotheses. Although the putative per millisecond synaptic sampling rate seems
trivial, as our calculations show, the cumulative capacity is substantial. As Sechenov
(1863) has proposed, \By means of absolutely involuntary learning . . . in all spheres
of the senses the child acquires a multitude of more or less complete ideas of objects,
i.e., elementary concrete knowledge".
4. The Explanatory Gap
The \explanatory gap" is a philosophical term referring to the lack of an explanation
of how physical properties give rise to the qualia of experience (Levine, 1983). We
appreciate such a \literary" denition is a necessary step in problem identication,
but consider it a misplaced expectation that any deep resolution is possible while
explication languishes in such purely metaphorical or narrative domains. Any solution is likely only available following location of the \gap" within an appropriate

383
neuropsychophysiological context. More specically, we consider the problem of the

explanatory gap becomes tractable when translated from a philosophical to an empirical context.
Quantication is fundamental to the scientic method and by making this conceptual transformation, the philosophical idea of the explanatory gap can be placed
rmly into the core of the scientic tradition. However, we recognize that with
quantication comes reduction, itself a characteristic of the scientic method. In doing
so we acknowledge the metaphorical nature of much written about consciousness. As
suggested in the Introduction, because of the highly multidimensional, intangible and
generally elusive nature of consciousness many of its various aspects are only available
through metaphorical language and description. It is these aspects which are greatly
reduced by transference from the philosophical to the physiological domain.
To make this transfer we propose, at its most general, that the physiological
explanatory gap is dened as the gap between the occurrence of a stimulus at externally or internally directed sensory receptors and the initiation of a response,
motor or otherwise. It is this gap which must be \explained" by the nervous system
and to which, in its own way, aspects of the philosophical formulation actually refer.
Most importantly, during this period it is not possible for a nervous system to
know anything about changes in the external environment. For this reason, a signicant evolutionary advantage likely accrues to an organism capable of correctly
predicting evolution of the environment during this period. More specically, in the
case of placental mammals, the foregoing denition might be recast as, \the temporal
duration from the time of interaction of a sucient sensory stimulus with a receptor
until the time at which a percept of that event is subjectively recognized within
conscious experience".
This transformation recasts the problem of the explanatory gap by removing it
from a philosophical domain of narrative contemplation to the domain of empirical
quantication. There it can be rigorously studied as an incontrovertible feature of the
vertebrate CNS.
As we show below, this is an illuminating example of how narrative language used
to speculate about the brain can be recast in empirical terms to more decisively
explore CNS activity and, in this case, \bridge" the explanatory gap.
5. Eerence Copy as Motor Predictor Model
Modern eerence copy models (Fig. 1(a)) derive from a proposal which originally
posited direct sensation of the motor command: \The impulse to move, which we
initiate through the innervation of our motor nerves, is immediately perceptible".
(von Helmholtz, 1878/1971). This idea was later integrated into motor physiology
through \corollary discharge" (von Holst, 1954; Sperry, 1950).
As reviewed by Mulliken & Andersen (2009), because sensory information is
substantially delayed, it has been proposed the brain makes use of an internal forward model which relies on eerence or eerent copy. This is an internal copy of an
384

(a)
(b)
Fig. 1. A. Eerence Copy: Model proposes that upon motor preparation and intention, copies of
eerent motor information are fed back and used centrally in an emulation algorithm, which calculates
the anticipated somatosensory changes expected as a consequence of the planned motor execution.
Subsequently, peripheral changes at the level of the muscles, the joints and the skin generate actual
proprioceptive feedback, which will eectively balance the predicted sensory feedback in somatosensory
cortex (at the level of a so-called \comparator"). B. Aerence copy: A model complementary to eerence
copy proposes that upon sensory stimulation, receptor stimuli are fed forward and maintained subconsciously in the memory system while they are integrated with the relevant memories they evoke.
Once integration is complete, a prediction of the percept expected on the basis of the received sensory
stimulus emerges as an \aerence copy" into the explanatory gap created by the time taken for integration to occur. If the aerence copy eectively matches the environment, predicted experience and
actual experience match suciently that no further action is required. Otherwise, surprise is experienced
and attention may be redirected. Panel A adapted from (Bazan, 2012).
outowing, movement-producing signal generated by the motor system. It is proposed to allow integration of sensory and motor feedback signals to estimate current
and upcoming positions and motions of a limb during movement (Blakemore et al.,
2000). Upon eective execution, the actual proprioceptive feedback of a motor action
eectively balances the predicted sensory feedback. Thereby, the eerence copy is an
early-warning signal sent by motor production areas to the corresponding somatosensory areas specialized in the proprioception of motor execution. It also allows
perceptual structures to distinguish between self and externally mediated signals.
6. Aerence Copy as Sensory Predictor Model
An explanatory gap exists for the time it takes a sensory stimulus to evoke fabrication of a current percept and for the percept to emerge into consciousness. Our
claim is, this physiological explanatory gap is bridged by conscious experience, which
originates through aerence copy as a construct evoked from within the memory
system.
The special character of this consciously experienced percept is that it is not
actually just a record of sensory activation entering the nervous system. Rather, it is
a memory-based prediction (activated by stimulus presence) of the state of the
internal and external environment of the nervous system expected at the time

385
integration of the given sensory input is complete. It is in this sense that aerence
copy is a predictor of sensory experience. This process is illustrated and explained in
more detail in Fig. 2.
Aerence copy can be conceptualized as complementary to the eerence copy
system (Fig. 1(b)), but applied to inowing sensory activation rather than the outowing, movement-producing activity of the motor system. It is the physiological
mechanism which enables and completes the cyclic ow of experience and behavior
through the cognitive domain of the nervous system and the physical world within
which it is embedded.
Following, for example, Dehaene & Naccache (2001) and Gray (2004), it is clear
the neural activity driving aerence copy is widely distributed, although the details
of precise location and mechanism are as yet unclear. Nevertheless, in the normal
course of events, the construct fabricated by aerence copy processes would be based
on, and exist in, the \past" due to the temporal delay introduced by the neural
activity enabling its conscious appearance. This situation is only compounded by the
fact that each successive moment is also a past moment prior to being experienced as
the present.
The aerence copy model is proposed as a solution as to how it is that subjective
experience appears to exist in the present moment when the time required for sensations to consciously appear necessarily require sensory events evoking a conscious
percept to occur in the past.
The model resolves this apparent conict in the following way. At any one moment, entry of sensory input into consciousness is delayed while its eects are integrated into ongoing neural activity to fabricate withinb the memory system a
comprehensive percept of the contents expected or predicted for a given moment.
Importantly, we further claim that the ability to do this is learned, as is the
content of the behavioral repertoire it ultimately subserves, with the complexity of
the processes involved being a major reason for the extended developmental period
seen in humans. As they are learned, the processes subserving aerence copy, similarly to other learning, become a \reexive" mechanism residing in the subconscious.
It is in this way the explanatory gap is bridged and conscious experience generated. The content of consciousness is this memory-based prediction of the composition of the next actual future moment, subjectively experienced as the present
moment. It is a reexive mechanism learned during development to accommodate
ongoing function in the ubiquitous presence of the explanatory gap.
Ultimately, ongoing neural activity is modulated as required by sensory content
and the associations it evokes from within the memory system. What is commonly
referred to and experienced as subjective phenomenal consciousness is this
b It does not make sense to say \from" memory as this implies memory is consulted by some lookup procedure. This
creates a signicant problem as retrieved content must be collected and displayed somewhere. Attempts to resolve
this additional complexity has spurred previous solutions resulting in homunculi and the mechanics of the Cartesian
Theatre. Importantly, following Nikoli (2014), we consider \monitor-and-act" is a more complete descriptor than
the notion of representation as the former incorporates both \memory storage" and \processing".

386
Fig. 2. Framework for consciousness: The schema is given for a single external stimulus. Model features are given at the top (A. Explanatory gap, C. Predicted percept) and subjective time (Past moment, Present moment, Future moment) indicated at the bottom of each panel for three consecutive
550 ms moments (separated by dark vertical bars). Upper horizontal dotted line distinguishes two
domains within each panel. An upper physical domain contains a stimulus originating externally to the
nervous system. A lower cognitive domain (within the nervous system) is further subdivided into two
\internal" domains; a sub/pre-conscious domain (top) containing the memory system (comprising
sensory, short and long-term memory); and a conscious domain (bottom). Stimulus evoked sensory
arousal activates the memory system () to evoke relevant memories of previous stimulus experience
which together are integrated ( ! ) to generate an appropriately fabricated percept then released to
the conscious domain () as the best prediction of immediate future experience. Panel 1: An Explanatory gap 1A exists in the past moment (1B) for the time it takes ( ! ) a stimulus to evoke a
comprehensive prediction within the memory system of future stimulus-modulated experience. Once
fabricated, the Predicted percept (1C) emerges into the conscious domain (indicated by vertical stripes
projecting from the sub/preconscious domain into the conscious domain and perception) as a subjective
conscious experience in the Present moment (1D). The large upper arrow head indicates appropriate
ongoing subconscious elaboration of memory by incorporation of correctly predicted perceptual components. The large lower arrow indicates the ongoing evolution of predicted perceptual experience
within the conscious domain. Panel 2: Illustrates temporal evolution of the relationships given in Panel 1
into the next moments. During the Explanatory gap (2A), integration of a new sensory stimulus now
occurs within memory system while the predicted percept generated during the explanatory gap of the
(just passed) moment (1B) has now emerged as an appropriate stimulus modulated percept into the
conscious domain of the present moment (illustrated in 1D, but for clarity not shown in 2B). This
process continues as the Predicted percept (2C) being generated during the current Explanatory gap
(2A) will be consciously experienced in the subsequent Future moment (2D). It is in this way ongoing
perceptual experience is continuously generated through aerence copy to bridge the sliding window of
the explanatory gap. Consciously experienced subjective phenomenology is a product of a previously
constructed prediction of the experiential contents expected to occur during the present moment.

387
physiological process of creating a prediction of the experience of the present moment

based on prior experience of the given situation or environment.
When this functionality is paired with the ability of the CNS and its memory system
to discriminate and cumulatively \record" in the domain of single photons (Gregory,
1966) and atoms (Green, 1976); the elements available for fabrication create eectively
\seamless" and entirely \resolvable" percepts. Naturally, environmental needs and
requirements create circumstances considerably more complex than the basic process
of aerence copy described here, but in principle, it can be expanded to any degree of
complexity. From this perspective, consciousness can be seen to act as an interface
between the aerence and eerence copy systems, thereby integrating and embedding
a nervous system into the spatiotemporal and physical ows of its experience.
7. Discussion
We begin this discussion by describing an expanded role for mirror neurons within
the framework we have developed here. This is followed by comments on evolutionary implications of the temporal extent of the explanatory gap. We then employ
the model to explore how the low rate and high variability reported for cortical
neuron ring may originate. Based on these considerations, we nally and briey
discuss a putative origin of qualia in the vertebrate CNS.
7.1. Mirror neurons as recruiters of predictive percepts
In humans, the subjective elements of a reproduced image are in general identical
with those of the perception, with the image of a perceived object usually produced
immediately during rst perception. The processes of perception and image reproduction are identical in their essence and their structural and physiological bases
absolutely equal (Beritashvili, 1968).
Evidence for a signicant claim of the aerence copy model may be found in the
behavior of mirror neurons. Their key characteristic is activity modulated both by
the execution and observation of action (di Pellegrino et al., 1992). Amongst other
properties, they have been reported to respond to the sound, relative distance, reward
value, viewpoint (whether visible or hidden), overall goal, action performed in different contexts and various combinations of single or dual hand and mouth actions
(Kilner & Lemon, 2013).
Historically, mirror neurons have been proposed to play a signicant role in social
cognition (Heyes, 2010). Here, we propose their properties provide further compelling
evidence for the aerence copy model. In short, if the explanatory gap is bridged by a
predictive construct fabricated by integration of sensory stimulation into the memory
system, which subsequently emerges as conscious experience; then, when viewing a
given activity, neurons in the memory system associated with previous views are the
repository of the memories from which appropriate predictive percepts may subsequently be recruited. When an observed activity is internalized by learning, and
occurrence is matched to performance, activity performance may be mapped to the
388
neurons that recognize occurrence as to the neurons involved, they are one and the
same event. An ability to distinguish between occurrence and performance conceivibly occurs as the nervous system learns to distinguish itself from everything else
(or the \other" (see Marchetti & Koster, 2014)). Similar to eerence copy, in this way
aerence copy can connect observed with performed activity to conrm a given
behavior has actually been implemented. Thus, the behavioral loop established by
eerence copy and its conrmation of successful motor plan execution, is completed.

7.2. Temporal duration of the conscious moment

Evolutionary centralization of the vertebrate nervous system creates a problem not
faced by the decentralized, ganglionated plexuses of invertebrates. With increased
evolutionary elaboration of the CNS, (although controllable in part by bre diameter, myelination, and spatial reconguration) the time between stimulus and response must increase due to increased system size and complexity (Biryukov, 1968).
Concomitant with competitive evolutionary pressures, the tension between rapidity
of reex response and selection of an optimal behavior from an ever increasing repertoire; CNS elaboration allows for increased behavioral choice. Here, there is one
question, is there a temporal limit for separation of stimulus and response?
One answer comes from recent experiments with virtual reality simulators and
computerized game playing. Here, for example, driving and ight control start to be
seriously impacted by delays of 50 ms (Kemeny, 2000) and 150 ms (Blana, 1996),
respectively. Although, with practice, operators can adapt to a constant delay of up
to 230 ms with no signicant eect on performance (Cunningham et al., 2001a,b).
This suggests the current hominid nervous system is capable of supporting an explanatory gap of at least 770 ms (550 ms (Libet, 2004) 230 ms (Cunningham et al.,
2001a,b)), with actual duration likely determined by the requirements of task complexity, familiarity and response. It is interesting to speculate on the future
computational and digital roles which might support expansion in this domain, and
the eect adolescent game playing in virtual environments may already be having.
Finally, in this section, we note two provocative implications related to duration of
the explanatory gap. In brief, as the vertebrate CNS has evolved, the lengthening
explanatory gap has necessarily required an extended aerence copy mechanism. It is
clear that this development has at least two highly signicant evolutionary ramications: (1) The longer the duration of the explanatory gap successfully bridged by
aerence copy, the larger the amount of subconscious processing that can occur. The
inevitable consequence of this is predicted to be a concomitant increase in consciousness. (2) Alternatively, the greater the stability of an environment, the more
likely fabricated predictions of aerence copy will match what occurs, thus better
supporting an increased explanatory gap with its attendant consequences, i.e.,
increased consciousness.
It has not escaped our notice that consideration of the aerence copy model we
have postulated here suggests a multiplicity of novel features of considerable
389
theoretical and practical interest which may also exhibit signicant ethical and, thus,
political repercussions.

7.3. Aerence copy and neural ring variability

In the intervening period of the explanatory gap, irrespective of its duration, it is
likely valuable to minimize generation of spurious neural activity or \noise". One
mechanism suited for this purpose is the comprehensive central inhibition exerted by,
for example, the \behavioral inhibition system" (Gray, 1972, 1981). The role of such
inhibition can be signicant, particularly when, in conjunction with disinhibition, it
assists either in the rapid extraction of certain types of subconsciously \processed"
automatic reex activity subsequently subject to higher-level cognitive analysis, or
provision of a basis for mismatch error signals between expected and actual states of
the world (Corr, 2011).
The low number and high variability of impulses reported for ongoing cortical
activity is widely considered the result of \balanced" excitation and inhibition
(Shadlen & Newsome, 1998). As an alternative explanation, and in keeping with the
requirements of selection from a behavioral repertoire, we propose that during the
explanatory gap only necessary disinhibition occurs. This allows minimal impulse
generation within widely distributed circuits; which may both explain these data and
provide direct evidence for the neural activity that might be predicted during the
process of aerence copy.
If aerent sensory stimulation is reexively, thus subconsciously, integrated into
the memory system prior to entering consciousness within a fabricated prediction of
future perceptual experience, it is likely that considerable neural activity may be
necessary, particularly as the required memory content must simultaneously be
recruited and integrated to form a dynamic and unied conscious experience.
In such an environment, moment to moment sensory changes may recruit the
same cells but congured, for example, to form dierent functional circuits as different aspects of requisite memory components are integrated to fabricate the constantly changing detail of relevant percepts. In these circumstances, the same cortical
neurons may contribute to the tracking and prediction of subtle modulations of
experience with signicant variation in ring time. The actual ring time of such
neurons may be deterministically controlled by their precise contribution (Reinagel
& Reid, 2002) and eective connectivity (Friston, 2011).
7.4. Qualia as emotional states re-evoked by aerence copy
Along with activation of neural circuits, the physical events of sensory receptor
activation elicit short-lived cognitive, physiological and somatic reactions, otherwise
known as emotions (Kensinger, 2009). Emotions are the product of complicated
collections of pattern forming chemical and neural responses, with all emotions
having some kind of regulatory role for neural activity within the nervous system
(Damasio, 1999). It is through the involvement of hormones and neurotransmitters

390
such as dopamine, noradrenaline, serotonin, oxytocin, cortisol and GABA, with

feelings, moods and emotions that humans probably come closest to direct contact
with the internal molecular reality driving this most ancient and fundamental modality of CNS function.
Similarly, the interaction of light with photoreceptors in the retina results in the
generation of patterns of neural impulses reecting the combinations of wavelengths
to which they respond. In the world, the waveforms and quanta of light are comprised
entirely of their own particular physical structures and relations. Any further
properties they acquire are due to interactions outside of their own structural features. For example, color is a property of the brain (Zeki, 1993), determined when the
visual system integrates responses across populations of photoreceptors (specically,
diering absorption spectra recognized by three types of cone cell).
Following the moment of occurrence, the consequences of sensory stimulation are
embedded within a complex ongoing ow of internally generated emotional states. It
should be no surprise that moment to moment records of sensory experience consequent upon activation of all sensory modalities, including emotional aect (Solms,
1997), are cumulatively recorded and updated, maintained and reinterpreted, by the
memory system (Beritashvili, 1968). Thus, at some later moment, as relevant
memories are evoked through aerence copy, it is only to be expected that according
to this model, associated emotional aects emerge as part of each fabricated percept
or experiential prediction.
In brief, qualia are those \remembered" emotional states associated with previous
experience, re-evoked from memory during the process of aerence copy. The immediate \redness" of a rose, taste of an apple, smell of a fresh tomato, the frying of
bacon and eggs or the glimpse of a childhood sweetheart, are never as comprehensively complete or emotionally intense as the enriched multiply re-experienced history of such sensations as they accumulate in memory. A simple relationship may be
predicted: the nature and quality of a quale is directly proportional to the extent its
referent is re-exposed to the nervous system.
This may provide an explanation for why little is ever as \good" or \pleasing" as
the positive memories of the past. Vice versa, this mechanism may also explain the
severe ongoing eects of trauma. It is the mechanism underlying the powerful nostalgic attractions of reminiscence as explored, for example, by Proust (1913/2000). It
is why olives, blue cheese, wine or true love must rst be experienced, before the
intensity of their appreciation (whether innate or acquired) can be enriched through
re-exposure. If the rst pu of a cigarette is the only indication of future enjoyment,
how could one ever proceed?
7.5. Conclusion
We have presented here a framework for consciousness. It is delimited in the neurophysiological domain by an \explanatory gap" which is a transguration of the
widely explicated philosophical explanatory gap. Here, a physiological mechanism is
proposed whereby this gap may be bridged. The surprising result is that what is

391
experienced as subjective consciousness is a learned predictive construct fabricated

from the quantal contents of a cumulative memory system, to emerge as the subjective conscious experience of the \present" moment. Consequently, the aerence
copy proposed to support this fabrication comprises a model that may provide
considerable insight into the neurophysiological underpinnings and mechanisms of
consciousness. It is a process likely applicable to, and property of, every centralized
vertebrate CNS.
The implications are complicated and diverse. In principle, it allows biological
mechanisms to account for many dierent aspects of brain structure and function
while lending a potential to unify dierent perspectives as to how subjective phenomenological states arise and could be experienced as consciousness. It is likely
driven by evolutionary requirements and the ontogenetic need to develop, organize
and thus more eectively employ, the physical constituents we are individually
composed of, embedded within and directly linked to.
In its essence aerence copy, and thus its correlate

consciousness, allows
temporal decoupling by the neocortex of innate and learned reex activity (as required) from immediate behavior. This occurs through the subconscious inhibitory
modulation of otherwise near immediate reex responses. It putatively allows behavioral expression independent of its learned origin. As for the mechanisms generating and framing consciousness; as we have shown here, a sucient fragment, for
example the result of Libet (1985), is equal to the task of solution.
It takes remarkably little eort to determine that much of what is believed about
ourselves and the world is merely convenient ction. As a renowned investigator once
said, \when you have eliminated all which is impossible, then whatever remains,
however improbable, must be the truth" (Doyle, 1926). This leaves us with only one
certainty. The shoulders from which further details of the generation and subjective
experience of conscious phenomenal content might be viewed are, as yet, entirely
unknown.
Acknowledgments
The authors would like to acknowledge Jan Braeken for his valuable discussions
about this paper.
REFERENCES
Altman, J. (1969) Autoradiographic and histological studies of postnatal neurogenesis. IV.
Cell proliferation and migration in the anterior forebrain, with special reference to persisting neurogenesis in the olfactory bulb. J. Comp. Neurol., 137, 433457.
Altman, J. (1963) Autoradiographic investigation of cell proliferation in the brains of rats and
cats. Anat. Rec., 145, 573591.
Arbib, M.A. (1972) The Metaphorical Brain. Wiley Inter-Science.
Aru, J. & Bachmann, T. (2014) Phenomenal awareness can emerge without attention. Front.
Hum. Neurosci., 7, e891.

392
Atkinson, R.C. & Shirin, R.M. (1968) Human memory: A proposed system and its control
processes. In: KW Spence and JT Spence, eds. The Psychology of Learning and Motivation,
Vol. 2. Academic Press, pp. 89195.
Avoli, M. (2004) History of neuroscience: Inhibition and epilepsy, IBRO Hist Neurosci,
http://www.ibro.info/Pub/Pub Main Display.asp?LC Docs ID=3535.
Azevedo, F.A.C., Carvalho, L.R.B., Grinberg, L.T., Farfel, J.M., Ferretti, R.E.L., Leite, R.E.
P., Filho, W.J., Lent, R. & Herculano-Houzel, S. (2009) Equal numbers of neuronal and
nonneuronal cells make the human brain an isometrically scaled-up primate brain.
J. Comp. Neurol., 513, 532541.
Baayen, R.H. & Milin, P. (2010) Analyzing reaction time. Int. J. Psychol. Res. (Medellin),
3, 1228.
Baddeley, A.D. & Hitch, G. (1974) Working memory. In: G.H. Bower, ed. The Psychology of
Learning and Motivation: Advances in Research and Theory, Vol. 8. Academic Press,
pp. 4789.
Bar, M. (2009a) Predictions: A universal principle in the operation of the human brain.
Philos. Trans. R Soc. Lond B Biol. Sci., 364, 11811182.
Bar, M. (2009b) The proactive brain: Memory for predictions. Philos. Trans. R Soc. Lond B
Biol. Sci., 364, 12351243.
Bazan, A. (2012) From sensorimotor inhibition to Freudian repression: Insights from psychosis applied to neurosis. Front Psychol., 3, e432.
Barenblatt, G.I. (1996) Scaling, Self-Similarity, and Intermediate Asymptotics. Cambridge
University Press.
Becker, W. (1989) Metrics. In: B Wurtz and M Goldberg, eds. The Neurobiology of Saccadic
Eye Movements (Elsevier), pp. 1367.
Beritashvili, I.S. (1968) A modern interpretation of the mechanism of I. M. Sechenov's
psychical reex medium member. Prog. Brain Res., 22, 252264.
Beritov, I.S. (1965) Neural Mechanisms of Higher Vertebrate Behavior. Little Brown Co.
Beritov, I.S. (1968) Central inhibition according to I. M. Sechenov's experiments and concepts, and its modern interpretation. Prog. Brain Res., 22, 2131.
Berlin, H.A. (2011) The neural basis of the dynamic unconscious. Neuropsychoanalysis,
13, 531.
Biryukov, D.A. (1968) I. M. Sechevov and some problems on evolution of nervous activity.
Prog. Brain Res., 22, 403413.
Blackmore, S. (2001) State of the artThe psychology of consciousness. The Psychologist,
14, 522525.
Blakemore, S.-J., Wolpert, D.M. & Frith, C.D. (2000) Why cant you tickle yourself ?
Neuroreport, 11, 1115.
Blana, E. (1996) A survey of driving research simulators around the world. Institute of
Transport Studies, University of Leeds, Working Paper 481. Available at: http://eprints.
whiterose.ac.uk/2110/1/ITS170 WP481 uploadable.pdf.
Block, N. (2009) Comparing the major theories of consciousness, In: M.S. Gazzaniga, ed. The
Cognitive Neurosciences, 4th edn. The MIT Press, pp. 11111122.
Bode, S., He, A.H., Soon, C.S., Trampei, R.R., Turner, R. & Haynes, J.-D. (2011) Tracking
the unconscious generation of free decisions using ultra-high eld fMRI. PLoS One,
6, e21612.

393
Boghen, D., Troost, B.T., Daro, R.B., DellOsso, L.F. & Birkett, J.E. (1974) Velocity
characteristics of normal human saccades. Invest Ophthalmol., 13, 619623.
Bohm, D. (1980) Wholeness and the Implicate Order. Routledge & Kegan Paul.
Brown, G.D.A., Neath, I. & Chater, N. (2007). A ratio model of scale-invariant memory and
identication. Psychol. Rev., 114, 539576.
Buckner, R.L. and Krienen, F.M. (2013) The evolution of distributed association networks in
the human brain. Trends Cogn Sci., 17, 648665.
Buckner, R.L., Andrews, J.R. & Schacter, D.L. (2008) The brain's default network: Anatomy, function, and relevance to disease. Ann NY Acad Sci., 1124, 138.
Bulbring, E., Lin, R.C.Y. & Schoeld, G.C. (1958) An investigation of the peristaltic reexin
relation to anatomical observations. Q J. Exp. Physiol., 43, 2637.
Bullock, H.T. (1961). Origins of patterned nervous discharge. Behaviour, 17, 4859.
Butler, A.B. (2009) Evolution of vertebrate brains. In: L.R. Squire, ed. Encyclopedia of
Neuroscience, Vol. 4. Academic Press, pp. 5766.
Calford, M.B. & Tweedale, R. (1991) Acute changes in cutaneous receptive elds in primary
somatosensory cortex after digit denervation in adult ying fox. J. Neurophysiol., 65,
178187.
Cannon, W.B. (1911) The Mechanical Factors of Digestion. Longmans.
Cannon, W.B. (1902) The movements of the intestines studied by means of Rontgen rays.
Am J. Physiol., 6, 251277.
Cazako, B.N., Lau, B.Y.B, Crump, K.L., Demmer, H.S. & Shea, S.D. (2014) Broadly tuned
and respiration-independent inhibition in the olfactory bulb of awake mice. Nat. Neurosci.,
17, 569576, http://dx.doi.org/10.1038/nn.3669.
Chalmers, D.J. (1996) The Conscious Mind: In Search of a Fundamental Theory. Oxford
University Press.
Chalmers, D.J. (1997) Moving forward on the problem of consciousness. J. Conscious. Stud.,
4, 346.
Chevalier, G. & Deniau, J.M. (1990) Disinhibition as a basic process in the expression of
striatal functions. Trends Neurosci., 13, 277280.
Chun & Potter, M.C. (1995) A two-stage model for multiple target detection in rapid serial
visual presentation. J. Exp. Psychol. Hum. Percept Perform., 21, 109127.
Churchland, P.S. (1996) The hornswoggle problem. J. Conscious. Stud., 3, 402408.
Clarke, E. & Jacyna, L.S. (1987) Nineteenth-Century Origins of Neuroscientic Concepts.
University of California Press.
Coop, A.D. & Reeke, G.N. (2004) Control of neuronal discharge timing by aerent ber
number and the temporal pattern of aerent impulses. J. Integr. Neurosci., 3, 319342.
Coop, A.D. & Redman, S.J. (1998) The inhibitory control of pyramidal cell discharge in a
neural network simulation of a local circuit in hippocampus area CA1. In: J.M. Bower, ed.
Computational Neuroscience: Trends in Research. Plenum, pp. 343348.
Coop, A.D. & Redman, S.J. (1992) A model of the peristaltic reex. In: F.H. Eeckman and
J.M. Bower, eds. Computation and Neural Systems. Kluwer, pp. 313320.
Corr, P.J. (2011) Anxiety: Splitting the phenomenological atom. Pers. Individ. Dif., 50,
889897.
Cunningham, D.W. Billock, V.A. & Tsou, B.H. (2001a) Sensorimotor adaptation to violations of temporal contiguity. Psychol. Sci., 12, 532535.

394
Cunningham, D.W. Chatziastros, A. von der Heyde, M. & Bltho, H.H. (2001b) Driving in
the future: Temporal visuomotor adaptation and generalization. J. Vis., 1, 8898.
Damasio, A. (1999) The Feeling of What Happens: Body and Emotion in the Making of
Consciousness. Houghton Milin Harcourt.
Daugman, J.G. (1993) Brain metaphor and brain theory. In: E.L. Schwartz, ed. Computational Neuroscience. The MIT Press, pp. 918.
Dehaene, S. & Naccache, L. (2001) Towards a cognitive neuroscience of consciousness: Basic
evidence and a workplace framework. Cognition, 79, 137.
Dennett, D.C. (1978) Articial intelligence as philosophy and as psychology. In: M. Ringle,
ed. Philosophical Perspectives on Articial Intelligence. Humanities Press.
Dennett, D.C. (1991) Consciousness Explained. Little, Brown.
di Pellegrino, G. Fadiga, L. Fogassi, L. Gallese, V. & Rizzolatti, G. (1992) Understanding
motor events, a neurophysiological study. Exp. Brain Res., 91, 176180.
Donders, F. (1868/1969) On the speed of mental processes. W.G. Koster (translation). Acta
Psychol (Amst), 30, 412431.
Doty, R.W. (1967) On butteries in the brain (Russian, English summary). In: Sovremennye
problemy electroziologii centralnoi nervnoi systemy. Nauka, pp. 96103.
Doyle, A.C. (1926) The adventure of the blanched soldier. In: The Case-Book of Sherlock
Holmes. John Murray.
Dykes, R.W., Landry, P., Metherate, R. & Hicks, T.P. (1984) Functional role of GABA in cat
primary somatosensory cortex: Shaping receptive elds of cortical neurons. J. Neurophysiol., 52, 10661093.
Edelman, G.M. (2003) Naturalizing consciousness: A theoretical framework. Proc. Natl.
Acad. Sci. USA, 100, 5520.
Ernst, A., Alkass, K., Bernard, S., Salehpour, M., Perl, S., Tisdale, J., Possnert, G., Druid, H.,
Frisen, J. (2014) Neurogenesis in the striatum of the adult human brain. Cell, http://dx.
doi.org/10.1016/j.cell.2014.01.044.
Fischer, B. & Ramsperger, E. (1984) Human express saccades: Extremely short reaction times
of goal directed eye movements. Exp. Brain Res., 57, 191195.
Fischer, B., Weber, H., Biscaldi, M., Aiple, F., Otto, P. & Stuhr, V. (1993) Separate populations of visually guided saccades in humans: Reaction times and amplitudes. Exp. Brain
Res., 92, 528541.
Flavell, C.R., Lambert, E.A., Winters, B.D. & Bredy, T.W. (2014) Mechanisms governing
the reactivation-dependent destabilization of memories and their role in extinction. Front
Behav. Neurosci., 7, e214.
Freud, S. (1904/1914) The Psychopathology of Everyday Life. A.A. Brill (translation).
T. Fisher. Unwin Limited.
Freud, S. (1915/1964) The unconscious. In: J. Strachey, ed. Sigmund Freud SE 14. Macmillan, pp. 166215.
Friston, K. (2011) Functional and eective connectivity: A review. Brain Connect, 1, 1336.
Furness, J.B. & Costa, M. (1980) Types of nerves in the enteric nervous system. Neuroscience, 5, 120.
Gibson, P.H. (1983) EM study of the number of cortical synapses in the brains of ageing
people and people with Alzheimer-type dementias. Acta Neuropathol., 62, 127133.
Gray, J. (2002) The sound of one hand clapping. Psyche, 8, e11.

395
Gray, J.A. (1972) The psychophysiological basis of introversion-extraversion: A modication

of Eysenck's theory. In: V.D. Nebylitsyn & J.A. Gray, eds. The Biological Bases of Individual Behaviour. Academic Press, pp. 182205.
Gray, J.A. (1981) A critique of Eysenck's theory of personality. In: H.J. Eysenck, ed. A Model
for Personality. Springer-Verlag, pp. 246276.
Gray, J.A. (2004) Consciousness: Creeping up on the Hard Problem. Oxford University
Press.
Green, D.M. (1976) An Introduction to Hearing. Erlbaum.
Gregory, R.L. (1966) Eye and Brain: The Psychology of Seeing. Weidenfeld & Nicolson.
Gregory, R.L. (1970) The Intelligent Eye. Weidenfeld.
Gutfreund, H., Reger, J.D. & Young, A.P. (1988) The nature of attractors in an asymmetric
spin glass with deterministic dynamics. J. Phys. A Math Gen., 2, 27752797.
Guthrie, D.V. (1947) The square degree as a unit of celestial area. Pop Astron, 55, 200203.
Harris, C.M., Hainline, L., Abramov, I., Lemerise, E. & Camenzuli, C. (1988) The distribution of xation durations in infants and naive adults. Vision Res., 28, 419432.
Hassabis, D. & Maguire, E.A. (2009) The construction system of the brain. Philos Trans R
Soc. Lond B Biol. Sci., 364, 12631271.
Hebb, D.O. (1949) The Organization of Behavior. Wiley.
Hecht, S., Schlaer, S. & Pirenne, M.H. (1942) Energy, quanta and vision. J. Opt. Soc. Am.,
38, 196208.
Herculano-Houzel, S. (2011) Not all brains are made the same: New views on brain scaling in
evolution. Brain Behav. Evol., 78, 2236.
Herculano-Houzel, S. (2012) The remarkable, yet not extraordinary, human brain as a scaledup primate brain and its associated cost. Proc. Natl. Acad. Sci. USA, 109, 1066110668.
Heyes, C. (2010) Mesmerising mirror neurons. Neuroimage, 51, 789791.
Hinton, G.E. & Sejnowski, T.J. (1986). Learning and relearning in boltzmann machines.
In: D.E. Rumelhart, J.L. McClelland and the PDP Research Group. Parallel Distributed
Processing: Explorations in the Microstructure of Cognition, Vol. 1. Foundations.
MIT Press, pp. 282317.
Hodgkin, A.L. & Huxley, A.F. (1952) A quantative description of membrane current and its
application to conduction and excitation in nerve. J. Physiol., 117, 500544.
Hopeld, J.J. (1982) Neural networks and physical systems with emergent collective
computational abilities. Proc. Natl. Acad. Sci. USA, 79, 22542258.
Hopp, J.J. & Fuchs, A.F. (2004) The characteristics and neuronal substrate of saccadic eye
movement plasticity. Prog. Neurobiol., 72, 2753.
Hubel, D.H., Wiesel, T.N. & LeVay, S. (1977) Plasticity of occuler dominance columns in
monkey striate cortex. Philos. Trans. R Soc. Lond B Biol. Sci., 278, 377409.
Ingvar, D.H. (1985) \Memory of the future": An essay on the temporal organization of
conscious awareness. Hum. Neurobiol., 4, 127136.
Jackendor, R. (1987) Consciousness and the Computational Mind. The MIT Press.
Jackson, F. (1982) Epiphenomenal qualia. Philos. Q, 32, 127136.
Jrgensen, C.B. (1974) Integrative functions of the brain. In: B. Lofts, ed. Physiology of the
Amphibia. Academic Press, pp. 151.
Kandel, E.R., Kupferman, I. & Iversen, S. (2000a) Learning and memory. In: E.R. Kandel, J.
H. Schwartz and T.M. Jessel, eds. Principles of Neural Science. 4th edn. McGraw-Hill
Medical Publishing Division, pp. 12271246.

396
Kandel, E.R. (2000b) From nerve cells to cognition: The internal cellular representation
required for perception and action. In: E.R. Kandel, J.H. Schwartz & T.M. Jessel, eds.
Principles of Neural Science. 4th edn. McGraw-Hill Medical Publishing Division,
pp. 381403.
Kanwisher, N.G. (1987) Repetition blindness: Type recognition without token individuation.
Cognition, 27, 117143.
Kaupp, U.B. (2010) Olfactory signalling in vertebrates and insects: Dierences and commonalities. Nat. Rev. Neurosci., 11, 188200.
Kemeny, A. (2000) Simulation and perception of movement, In: Proc. Driving Simulation
Conf. 2000. Paris, pp. 1322.
Kemp, B.J. (1973) Reaction time of young and elderly subjects in relation to perceptual
deprivation and signal-on versus signal-o condition. Dev. Psychol., 8, 268272.
Kensinger, E.A. (2009) Phases of inuence: How emotion modulates the formation and
retrieval of declarative memories. In: M.S. Gazzaniga, ed. The Cognitive Neurosciences.
4th edn. The MIT Press, pp. 725737.
Kilner, J.M. & Lemon, R.N. (2013) What we currently know about mirror neurons. Curr
Biol, 23, R1057R1062.
Koch, C. (2009) Introduction. In: M.S. Gazzaniga, ed. The Cognitive Neurosciences. 4th edn.
The MIT Press, pp. 11071109.
Konorski, J. (1967) Integrative Activity of the Brain: An Attempt at an Interdisciplinary
Synthesis. University of Chicago Press.
Krantz, J. (2012) Experiencing Sensation and Perception. Pearson Education.
Kuo, A.R. (2002) The relative roles of feedforward and feedback in the control of rhythmic
movements. Motor Control, 6, 129145.
Lagasse, P. (2000) Nervous system: The nervous system and reexes. In: P. Lagasse, ed.
Columbia Encyclopedia. 6th edn. Columbia University Press.
LaRocque, J.J., Lewis-Peacock, J.A. & Postle, B.R. (2014) Multiple neural states of representation in short-term memory? Its a matter of attention. Front Hum Neurosci., 8, e5.
Lashley, K. (1956) Cerebral organization and behavior. In: H. Solomon, S. Cobb and
W. Peneld, eds. The Brain and Human Behavior. Williams & Wilkins, pp. 118.
Laycock, T. (1845) Letter to George Coombe, 27 February 1845, in Coombe Manuscripts,
National Library of Scotland. 7276, pp. 34.
LeDoux, J.E., Schiller, D. & Cain, C. (2009) Emotional reaction and action: From threat
processing to goal-directed behavior, In: M.S. Gazzaniga, ed. The Cognitive Neurosciences. 4th edn. The MIT Press, pp. 905924.
LeDoux, J.E. (1996) The Emotional Brain: The Mysterious Underpinnings of Emotional Life.
Simon & Schuster.
LeDoux, J.E. (2000) Emotion circuits in the brain. Annu. Rev. Neurosci., 23, 155184.
Lee, V. & Maguire, J. (2004) The impact of tonic GABAA receptor-mediated inhibition on
neuronal excitability varies across brain region and cell type. Front Neural Circuits, 8, e3.
Levine, J. (1983) Materialism and qualia: The explanatory gap. Pac. Philos. Q, 64, 354361.
Lewis, D.A., Hashimoto, T. & Volk, D.W. (2005) Cortical inhibitory neurons and schizophrenia. Nat. Neurosci., 6, 312324.
Libet, B. (1981) The experimental evidence for subjective referral of a sensory experience
backwards in time: Reply to P. S. Churchland. Philos. Sci., 48, 182197.

397
Libet, B. (1985) Unconscious cerebral initiative and the role of conscious will in voluntary
action. Behav. Brain Sci., 8, 529566.
Libet, B. (2004) Mind Time: The Temporal Factor in Consciousness. Harvard University
Press.
Libet, B., Alberts, W.W., Wright, E.W. Jr., Delattre, L.D., Levin, G. & Feinstein, B. (1964)
Production of threshold levels of conscious sensation by electrical stimulation of human
somatosensory cortex. J Neurophysiol., 27, 546578.
Llin
as, R. (1974) La forme et la function des cellules nerveuses. La Recherche, 5, 232240.
Llin
as, R. (1987) Microphysiology of the cerebellum. In: V.B. Brooks ed. Handbook of Physiology, Vol. II. The Nervous System. Part II. American Physiology Society, pp. 831976.
Llin
as, R.R. (2001) I of the Vortex: From Neurons to Self. The MIT Press.
Llin
as, R.R. (2014) The olivo-cerebellar system: A key to understanding the functional signicance of intrinsic oscillatory brain properties. Front Neural Circuits, 7, e96.
Llin
as, R.R. & Roy, S. (2009) The `prediction imperative' as the basis for self-awareness.
Philos. Trans. R Soc. Lond B Biol. Sci., 364, 13011307.
Mack, A. & Rock, I. (1998) Inattentional Blindness. The MIT Press.
MacKay, D.M. & McCulloch, W.S. (1952) The limiting information capacity of a neuronal
link. Bull Math Biophys, 14, 127135.
Magnus, R. (1924) K
orperstellung: Experimental-Physiological Investigations of the Reexes
Involved in Body Posture, their Cooperation and Disturbances. Julius Springer.
Maniadakis, M. & Trahanias, P. (2014) Time models and cognitive processes: A review. Front
Neurorobot, 8, e7.
Marchetti, I. & Koster, E.H.W. (2014) Brain and intersubjectivity: A Hegelian hypothesis on
the self-other neurodynamics. Front Hum Neurosci., 8, e11.
Mashour, G.A. & Alkire, M.T. (2013) Evolution of consciousness: Phylogeny, ontogeny, and
emergence from general anesthesia. Proc. Natl. Acad. Sci. USA, 110 (Suppl. 2), 1035710364.
Masliah, E., Mallory, M., Hansen, L., De Teresa, R. & Terry, R.D. (1993) Quantitative
synaptic alterations in the human neocortex during normal ageing. Neurology 43, 192197.
Marshall, W.H., Talbot, S.A. & Ades, H.W. (1943) Cortical response of the anaesthesized cat
to gross photic and electrical aerent stimulation. J. Neurophysiol., 6, 115.
McCulloch, W.S. (1947) Modes of functional organization of the cerebral cortex. Fed Proc.,
6, 445452.
McCulloch, W.S. & Pitts, W. (1943) A logical calculus of the ideas immanent in nervous
activity. Bull Math. Biophys., 5, 115133.
McGinn, C. (1989) Can we solve the mind-body problem? Mind, 98, 349366.
McGinn, C. (2004) Consciousness and Its Objects. Oxford University Press.
Miller, G.A. (1962) Psychology. The Science of Mental Life. Harper and Row.
Milner, B. (1962) Les troubles de la memoire accompagnant des lesions hippocampiques
bilaterales, In: P Passouant, ed. Physiologie de Lhippocampe. Centre National de la
Recherche Scientique, pp. 257272.
Minsky, M. (1988) Society of Mind. Simon and Schuster.
Mulliken, G.H. & Andersen, R.A. (2009) Forward models and state estimation in posterior
parietal cortex. In: M.S. Gazzaniga, ed. The Cognitive Neurosciences. 4th edn. The MIT
Press, pp. 599611.
Nadel, L. & Piattelli-Palmarini, M. (2003). What is cognitive science? Introduction to the
Encyclopedia of Cognitive Science. Nature Publishing Group.

398
Nagel, T. (1974) What is it like to be a bat? Philos. Rev., 83, 435450.

Nikoli, D. (2014) Practopoiesis: Or how life fosters a mind, arXiv:1402.5332 [q-bio.NC].
No, A. (2007) Inattentional blindness, change blindness, and consciousness. In: M. Velmans
and S. Schneider, eds., The Blackwell Companion to Consciousness. Oxford: Blackwell
Publishing, pp. 501511.
Olds, J. (1956) A preliminary mapping of electrical reinforcing eects in the rat brain.
J. Comp. Physiol. Psychol., 49, 281285.
O'Regan, J.K. (1992) Solving the \real" mysteries of visual perception: The world as an
outside memory. Can J. Psychol., 46, 461488.
O'Regan, J.K. (2003) Change Blindness. In: L. Nadel, ed. Encyclopedia of Cognitive Science.
Nature Publishing Group.
Orpwood, R. (2013) Qualia could arise from information processing in local cortical networks.
Front Psychol, 4, e121.
Pakkenberg, B., Pelvig, D., Marner, L. & Bundgaard, M.J. (2003) Aging and the human
neocortex. Exp. Gerontol., 38, 9599.
Pavlov, I.P. (1927) Conditioned Reflexes: An Investigation of the Physiological Activity of the
Cerebral Cortex. G.V. Anrep (translation, ed.) Oxford University Press.
Pellionisz, A. & Llin
as, R. (1979) Brain modelling by tensor network theory and computer
simulation. The cerebellum: distributed processor for predictive coordination. Neuroscience, 4, 323348.
Penrose, R. & Hamero, S. (2011) Consciousness in the universe: Neuroscience, quantum
space-time geometry, and Orch OR theory. J. Cosmol., 14
Perlovsky, L. (2013) Learning in brain and machinecomplexity, Godel, Aristotle. Front
Neurorobot., 7, e23.
Popper, K. (1959) The Logic of Scientic Discovery. Hutchinson & Co., Ltd.
Pribram, K. (1969) The neurophysiology of remembering. Sci. Am., 200, 7386.
Price, D.D. & Barrell, J.J. (2012) Inner Experience and Neuroscience: Merging Both
Perspectives. The MIT Press.
Proust, M. (1913/2000) In Search of Lost Time. 6 Vols. C. Prendergast, ed. Allen Lane.
Pylyshyn, Z.W. (1986) Computation and Cognition: Towards a Foundation for Cognitive
Science. The MIT Press.
Quiroga, R.Q., Reddy, L., Kreiman, G., Koch, C. & Fried, I. (2005) Invariant visual representation by single neurons in the human brain. Nature, 435, 11021107.
Raymond, J.E., Shapiro, K.L. & Arnell, K.M. (1992) Temporary suppression of visual processing in an RSVP task: An attentional blink? J. Exp. Psychol. Hum. Percept Perform,
18, 849860.
Rayner, K., Inho, A.W., Morrison, R.E., Slowiaczek, M.L. & Bertera, J.H. (1981) Masking
of foveal and parafoveal vision during eye xations in reading. J Exp Psychol Hum Percept
Perform, 7, 167179.
Reinagel, P. & Reid, C.P. (2002) Precise ring events are conserved across neurons.
J. Neurosci., 22, 68376841.
Resniko, H.L. (1988) The Illusion of Reality: Topics in Information Science. SpringerVerlag.
Richet, C. (1925) Reexes psychiques, reexes conditionels, automatisme mental. Pavlov's
Jubilee Volume.

399
Robinson, E.S. (1934) Work of the integrated organism. In: C. Murchison, ed. Handbook of
General Experimental Psychology. Clark University Press, pp. 645650.
Salthouse, T.A. & Ellis, C.L. (1980) Determinants of eye-xation duration. Am J. Psychol.,
93, 207234.
Sandman, C.A., Wadhwa, P., Hetrick, W., Porto, M. & Peeke, H.V.S. (1997) Human fetal
heart rate dishabituation between thirty and thirty-two weeks gestation. Child Dev.,
68, 10311040.
Schacter, D.L. (1987) Implicit memory: History and current status. J. Exp. Psychol. Learn
Mem. Cogn., 13, 501518.
Schacter, D.L. & Addis, D.R. (2009) On the nature of medial temporal lobe contributions to
the constructive simulation of future events. Philos. Trans. R Soc Lond B Biol. Sci., 364,
12451253.
Schacter, D.L., Addis, D.R. & Buckner, R.L. (2007) Remembering the past to imagine the
future: The prospective brain. Nat Rev Neurosci, 8, 657661.
Schacter, D.L., Addis, D.R. & Buckner RL (2008) Episodic simulation of future events:
Concepts, data, and applications. Ann NY Acad Sci., 1124, 3961.
Schacter, D.L., Gilbert, D.T. & Wegner, D.M. (2011) Learning. In: Psychology. 2nd edn.
Worth Publishers, pp. 263306.
Schel, M.A., Khn, S., Brass, M., Haggard, P., Ridderinkhof, K.R. & Crone, E.A. (2014)
Neural correlates of intentional and stimulus-driven inhibition: A comparison. Front Hum
Neurosci., 8, e27.
Scherrer, J. (1968) Electrophysiological aspects of cortical development. Prog. Brain Res.,
22, 480489.
Schmidt, D., Abel, L.A., Dellosso, L.F. & Daro, R.B. (1979) Saccadic velocity characteristics: Intrinsic variability and fatigue. Aviat Space Environ. Med., 50, 393395.
Schr
odinger, E. & Born, M. (1935) Discussion of probability relations between separated
systems. Math. Proc. Cambridge Phil. Soc., 31, 555563.
Sechenov, I.M. (1863/1965) Reexes of the Brain. In: K. Koshtoyants, ed. S. Belsky (trans.)
The MIT Press.
Seth, A.K., Baars, B.J. & Edelman, D.B. (2005) Criteria for consciousness in humans and
other mammals. Conscious Cogn., 14, 119139.
Shadlen, M.N. & Newsome, W.T. (1998) The variable discharge of cortical neurons: Implications for connectivity, computation, and information coding. J Neurosci., 18, 3870
3896.
Sherrington, C.S. (1906) The Integrative Action of the Nervous System. Yale University
Press.
Sherrington, C.S. (1940) Man on His Nature. Cambridge University Press.
Shettleworth, S.J. (2010) Cognition, Evolution and Behavior. 2nd edn. Oxford University
Press.
Shahani, B. (1970) The human blink reex. J. Neurol. Neurosurg. Psychiatry, 33, 792800.
Simons, D.J. & Rensink, R.A. (2005) Change blindness: Past, present, and future, Trends
Cogn. Sci., 9, 1620.
Skarda, A. & Freeman, W.J. (1987) How brains make chaos in order to make sense of the
world. Behav Brain Sci., 10, 161195.
Slonim, A.D. (1968) Unconditioned reex as a specic character and problem of studying
instinct. Prog. Brain Res., 22, 506517.

400
Smit, A., van Gisbergen, A. & Cools, A. (1987) A parametric analysis of human saccades in
dierent experimental paradigms. Vision Res., 27, 17451762.
Smith, R. (1992) Inhibition: History and Meaning in the Sciences of Mind and Brain.
University of California Press.
Smith, S.W. (2002) Audio processing. In: S. Smith, ed. Digital Signal Processing: A Practical
Guide for Engineers and Scientists. Newnes, pp. 351354.
Snyder, A. (1998) Breaking mindsets. Mind Lang., 13, 110.
Snyder, A. & Barlow, H.B. (1986) Revealing the artist's touch. Nature, 331, 117118.
Snyder, A., Bossomaier, T. & Mitchell, D.J. (2004) Concept formation: `Object' attributes
dynamically inhibited from consciousness awareness. J. Integr. Neurosci., 3, 3146.
Softky, W.R. (1995) Simple codes versus ecient codes. Curr Opin Neurobiol., 5, 239247.
Sokolov, E.N. (1963) Perception and the Conditioned Reex. Pergamon Press.
Solms, M. (1997) What is consciousness? J. Am. Psychoanal. Assoc., 45, 681703.
Solms, M. & Panksepp, S. (2012) The \Id" knows more than the \Ego" admits: Neuropsychoanalytic and primal consciousness perspectives on the interface between aective
and cognitive neuroscience, Brain Sci., 2, 147175.
Soon, C.S., Brass, M., Heinze, H.-J. & Haynes, J.-D. (2008) Unconscious determinants of free
decisions in the human brain. Nat Neurosci., 11, 543545.
Sperry, R.W. (1950) Neural basis of the spontaneous optokinetic response produced by visual
inversion. J. Comp. Physiol. Psychol., 43, 482489.
Sperry, R.W. (1980) Mind-brain interaction: Mentalism, yes; dualism, no. Neuroscience,
5, 195206.
Stein, R.B. (1967) The information capacity of nerve cells using a frequency code. Biophys. J.,
7, 797826.
Sutherland, S. (1989) International Dictionary of Psychology. Continuum.
Tarkka, I.M. (1986) Short and long latency reexes in human muscles following electrical and
mechanical stimulation. Acta Physiol. Scand., 557, S1S32.
Thorndike, E.L. (1911) Animal Intelligence: Experimental Studies. The Macmillan Company.
Thorpe S, Fize, D. & Mariot, C. (1996) Speed of procession in the human visual system.
Nature, 381, 520522.
Thorpe, W.H. (1966) Ethology and consiousness. In: JC Eccles, ed. Brain and Conscious
Experience. Springer-Verlag, pp. 470505.
Tulving, E. (1972) Episodic and semantic memory. In: E. Tulving and W. Donaldson, eds.
Organization of Memory. Academic Press, pp. 381402.
Tulving, E. (2002) Episodic memory: From mind to brain. Annu. Rev. Psychol., 53, 125.
van Beers, R.J. (2008) Saccadic eye movements minimize the consequences of motor noise.
PLoS One, 3, e2070.
van Diepen, P.M.J., De Graef, P. & dYdewalle, G. (1995) Chronometry of foveal information extraction during scene perception. In: J.M. Findlay, R. Walker & R.W. Kentridge,
eds. Eye Movement Research: Mechanisms, Processes and Applications. Elsevier,
pp. 349362.
van Gaal, S., Ridderinkkhof, K.J., Farenfort, J.J., Scholte, H.S. & Lamme, V.A. (2010)
Frontal cortex mediates unconsciously triggered inhibitory control. J. Neurosci., 28,
80538062.

401
Vartanian, A. (1973) Man-machine from the Greeks to the computer. In: P.P. Weiner, ed.
Dictionary of the History of Ideas: Studies of Selected Pivotal Ideas. Charles Scribner's
Sons, pp. 131146.
Vartanian, A. (1953) Diderot and Descartes: A Study of Scientic Naturalism in the
Enlightenment. Princton University Press.
Velmans, M. (2009) Understanding Consciousness. Routledge.
Velmans, M. (1991) Is human information processing conscious? Behav. Brain Sci., 14,
651669.
von Helmholtz, H. (1910) Handbuch der Physiologischen Optic, Vol. III. Leopald Voss.
von Helmholtz, H. (1878/1971) The facts of perception. In: R. Kahl, ed. Selected Writings of
Hermann von Helmholtz. Wesleyan University Press, pp. 366408.
von Helmholtz, H. (1850a) Vorluger Bericht ber die Fortpanzungs-Geschwindigkeit der
Anatomie, Physiologie und Wissenschaftliche Medicin, 7173.
Nervenreizung. Archiv fur
von Helmholtz, H. (1850b) Messungen ber den zeitlichen Verlauf der Zuckung animalischer
Muskeln und die Fortpanzungsgeschwindigkeit der Reizung in den Nerven. Archiv fur
Anatomie, Physiologie und wissenschaftliche Medicin, 276364.
von Holst, E. (1954) Relations between the central nervous system and the peripheral organs.
Br. J. Anim. Behav., 2, 8994.
Voronin, L.G., Gusselnikova, K.G., Gusselnikov, V.I. & Supin, A.J. (1968) On the problem of
the evolution of the vertebrate aerent systems. Prog. Brain Res., 22, 541565.
Vuilleumier, P. & Huang, Y.-M. (2009) Emotional attention: Uncovering the mechanisms of
aective biases in perception. Curr. Dir. Psychol. Sci., 34, 148152.
Walker, M.P., Stickgold, R., Alsop, D., Gaab, N. & Schlaug, G. (2005) Sleep-dependent
motor memory plasticity in the human brain. Neuroscience, 133, 911917.
Warren, W.C., Hillier, L.W., Graves, J.A.M., Birney, E., Ponting, C.P., Grtzner F., Belov,
K., Miller, W., Clarke, L., Chinwalla, A.T., Yang, S.-P., Heger, A., Locke, D.P., Miethke,
P., Waters, P.D., Veyrunes, F., Fulton, L., Fulton, B., Graves, T., Wallis, J., Puente, X.S.,
L
opez-Otn, C., Ord
oez, G.R., Eichler, E.E., Chen, L., Cheng, Z., Deakin, J.E., Alsop, A.,
Thompson, K., Kirby, P., Papenfuss, A.T., Wakeeld, M.J., Olender, T., Lancet, D.,
Huttley, G.A., Smit, A.F.A., Pask, A., Temple-Smith, P., Batzer, M.A., Walker, J.A.,
Konkel, M.K., Harris, R.S., Whittington, C.M., Wong, E.S.W., Gemmell, N.J.,
Buschiazzo, E., Jentzsch, I.M.V., Merkel, A., Schmitz, J., Zemann, A., Churakov, G.,
Kriegs, J.O., Brosius, J., Murchison, E.P., Sachidanandam, R., Smith, C., Hannon, G.J.,
Tsend-Ayush, E., McMillan, D., Attenborough, R., Rens, W., Ferguson-Smith, M.,
Lefevre, C.M., Sharp, J.A., Nicholas, K.R., Ray, D.A., Kube, M., Reinhardt, R., Pringle,
T.H., Taylor, J., Jones, R.C., Nixon, B., Dacheux, J.-L., Hitoshi Niwa, H., Sekita, Y.,
Huang, X., Stark, A., Kheradpour, P., Kellis, M., Flicek, P., Chen, Y., Webber, C.,
Hardison, R., Nelson, J., Hallsworth-Pepin, K., Delehaunty, K., Markovic, C., Minx, P.,
Feng, Y., Kremitzki, C., Makedonka Mitreva, M., Glasscock, J., Wylie, T., Wohldmann,
P., Thiru, P., Nhan, M.N., Pohl, C.S., Smith, S.M., Hou, S., Renfree, M.B., Mardis, E.R. &
Wilson, R.K. (2008) Genome analysis of the platypus reveals unique signatures of evolution. Nature, 453, 175183.
Wildman, D.E., Uddin, M., Opazo, J.C., Liu, G., Lefort, V., Guindon, S., Gascuel, O.,
Grossman, L.I., Romero, R. & Goodman, M. (2007) Genomics, biogeography, and the
diversication of placental mammals. Proc Natl Acad Sci USA, 104, 1439514400.
Wittgenstein, L. (1953) Philosophical Investigations. Blackwell.

402
Xing, J.X. & Gerstein, G.L. (1996a) Networks with lateral connectivity. I. Dynamic properties mediated by the balance of intrinsic excitation and inhibition. J Neurophysiol., 75,
184199.
Xing, J.X. & Gerstein, G.L. (1996b) Networks with lateral connectivity. II. Plasticity and
reorganization of somatosensory cortex. J Neurophysiol, 75, 200216.
Xing, J.X. & Gerstein, G.L. (1996c) Networks with lateral connectivity. III. Dynamic
properties mediated by the balance of intrinsic excitation and inhibition. J Neurophysiol,
75, 217232.
Zafeiriou, D. (2004) Primitive reexes and postural reactions in the neurodevelopmental
examination. Pediatric Neurology, 31, 18.
Zeki, S. (1993) A Vision of the Brain. (Blackwell).

Zernicki,
B. (1968) Reex as the unit of neuro-behavioral activity: A theoretical multidisciplinary approach to the reex activity in higher animals. Acta Biol Exp (Warsaw), 28,
291315.

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Journal of Integrative Neuroscience, Vol. 13, No.

Aerence copy as a quantitative neurophysiological

J. Integr. Neurosci. 2014.13:363-402. Downloaded from www.worldscientific.com

Hugo Cornelis and Allan D. Coop*

1. Introduction and Philosophical Context

J. Integr. Neurosci. 2014.13:363-402. Downloaded from www.worldscientific.com

H. CORNELIS & A. D. COOP

Following satiation of biological imperatives such as survival and reproduction,

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