Medical Hypotheses (2005) 65, 766784

http://intl.elsevierhealth.com/journals/mehy

A scientific paradigm for consciousness:

A theory of premotor relations q
C. Vakalopoulos *

171 McKean Street, North Fitzroy, 3068 Melbourne, Australia

Received 5 February 2005; accepted 11 April 2005

Summary Consciousness has become a holy grail for scientific research only in the last few decades. In spite of the
extensive recent research in the field agreement on a correct approach to a theory has been elusive. We all have an
intuitive idea of what we mean by the term and at the very least relate awareness to a descriptive phenomenology. I
present a theory of consciousness based on motor capacity. An organism may exert control through movement and any
action it performs on its surroundings results in a reaction. This type of reafference generated by movement provides
the organism with a unique opportunity to compare perceptual information, for example, a retinal image of a ball with
what the body is telling the agent about the object. In other words, limb movements will describe a certain physical
distance to the object and the shape will be conveyed by the arc described around it by any part of the body that comes
into contact with it. The reafference, also called motor efference copy, can modulate neocortical networks strictly
associated with concurrent perceptual information via input to thalamocortical projections. Concurrent perceptual
and motor reafferent input provide the critical organizing principles bestowing on a neural assembly the capacity of
conscious awareness. The cortical neural coding represents an interactive history and is what I call a premotor relation.
It avoids the trap of behaviorism by emphasizing a developmental causality such that, once the networks are
established no further motoric component is required. Supportive evidence for this position comes from the classic
psychological studies of perceptual adaptation to distorting lenses facilitated by movement and elegant animal
experiments dissociating sensorimotor development from visual exposure. One of the most innovative and useful
conceptions advanced in the paper is a component theory of motor efference copy analyzing the two key parameters of
any action namely force and proprioceptive change. There is massive neocortical input to the basal ganglia and
cerebellum and recent research has implicated cognitive roles for these structures. I propose they serve the respective
component parameters of motor efference copy. Finally, a theory of premotor relations provides a model for
understanding how dysfunction in the basal ganglia and cerebellum relates to cognitive features of schizophrenia and
autism, respectively. Motor dysfunction figures prominently in the early literature on these syndromes and I make the
case for a fundamental causal connection between motor and cognitive symptoms.
c 2005 Elsevier Ltd. All rights reserved.

q
Introduction
This work was presented by invitation at a Cognitive
Neuroscience Meeting, hosted by Jason B. Mattingley at the
Department of Psychology, University of Melbourne.
Living organisms generally manifest input and out-
* Tel.: +61408810220. put systems as a fundamental organization of their
E-mail address: hinemoa@bigpond.net.au. behaviour. Lower life forms, including those with a


0306-9877/$ - see front matter c 2005 Elsevier Ltd. All rights reserved.
doi:10.1016/j.mehy.2005.04.016
A scientific paradigm for consciousness
simple nervous system, will respond to a stimulus
or percept in a fixed or very limited fashion with
simple modulation consisting of desensitization or
onoff mechanisms. These responses can be gener-
ally classed as reflexive with only a very limited
capacity for adaptation. A classic example is the
habituation of the gill withdrawal mechanism to
repetitive stimulation of the mantle or siphon of
the mollusc Aplysia [1]. As one ascends the phylo-
genetic tree the same perceptual stimulus is asso-
ciated with increasing behavioral complexity. One
is more likely to ascribe consciousness to these
organisms, which often possess well-developed
nervous systems. Throughout this paper, the use
of the term consciousness will refer to phenomenal
states. Language will not be considered as a neces-
sary antecedent to consciousness nor will I distin-
guish between primary and secondary or higher
order consciousness.
A requirement of the model is that motor output
be directly implicated in the organization of an
intermediate stage of perceptual processing. This
can be achieved by a feedback limb called motor
efference copy. The reentrant property of a system
will be used to define a cognitive structure by
allowing an emergent complexity not normally
available to a reflex arc. The reentrant property
of motor actions is assumed to provide feedback di-
rectly modulating and organizing neocortical struc-
tures associated with each primary sensory
modality. The terminology of premotor relations
derives from a description of the basic mecha-
nisms. The property of conscious awareness of a
percept emerges from the reentrant modulation
of primary sensory networks by behavioural inter-
action with the environment, a process I will call
categorization. Once established, subsequent reac-
tivation of a neural network will possess phenome-
nal properties without any explicit movement.
However, at some stage in the developmental his-
tory of the organism an interaction with the envi-
ronment is essential for the emergence of
conscious networks. Thus neocortical activation al-
ready reflects previous behaviour coded in early
sensory networks. This coding represents the simul-
taneous association of action and percept with con-
comitant changes in the distribution of networks
and synapses. (A percept here is not synonymous
with phenomenal experience, but simply means
the information registered by a receptor such as
the retina.)
The underlying hypothesis is that neural net-
works, from the moment of conception, that associ-
ate perceptual information only with a stereotyped
output will, by definition, be unconscious. An
unconscious process will be differentiated from a
767
reflex action to a stimulus by its learned and com-
plex, albeit stereotypical nature. Conscious net-
works may process a percept in parallel to reflex
and unconscious circuits in the same organism,
but will exhibit diversification of behavioral associ-
ations in addition to complexity for that network.
The advantage of conceptualizing consciousness
as causally linked to a prior motor experience is
that it makes evolutionary sense by allowing an
organism behavioural choices and most importantly
embodying the whole nature of consciousness
bridging the explanatory gap. Consciousness of an
organism according to this view is not an epiphe-
nomenon but an inextricable product of its motor
history. Once it is established it can function largely
independently of overt action and indeed in turn al-
lows motor or behavioural diversification in the
context of any fixed situation. However, the pro-
cess has an ongoing dynamic, as we shall see, in
experimental situations such as wearing distorting
prisms or pathological conditions such as schizo-
phrenia where several clinical features of cognitive
dysfunction can be explained by aberrant motor
reafferent function.
A neuron or the neural substrate it is part of
does not have an intrinsic conscious nature. For
example, infants that are born without sight mani-
fest an extraordinary plasticity to their cortical cir-
cuitry. The occipital cortex, typically associated
with vision, is still active during inputs from other
sensory modalities. An MRI study on patients suffer-
ing from congenital blindness of peripheral (ocular)
origin showed no atrophy of the occipital cortex
[2]. Furthermore, the functional deficit associated
with the surgical removal of part of the brain in
early infancy can be almost completely compen-
sated by spared cortical tissue. What then, confers
consciousness to the processing of a neural circuit?
Logically, an intact sensory apparatus such as the
eye or ear is a necessary, but not sufficient condi-
tion for this dynamic process to occur. Further-
more, not all aspects of perception are made
consciously available. Priming by unconscious stim-
uli, as shown by altered performance of a subject
on certain tasks, is a well-documented phenome-
non. Brain-imaging techniques during the presenta-
tion of masked numerical primes revealed that
perceptual, semantic and motor processes do oc-
cur unconsciously [3].
It will be proposed here that the motor history of
an organism is a critical element to an adequate
theory of consciousness. The initial and ongoing
interactions with the environment will organize
neocortical networks associated with a specific
sensory input by a feedback mechanism, known as
motor efference copy (MEC). This process will be
768
described as categorization and states that motor
reentrant organization of perceptual information
may under certain circumstances, to be specified,
endow a neural network with the property called
a conscious act. During the neonatal period the link
between sensory information and motor behaviour
will be either reflexive or stochastic (random) in
nature. The emergence of a percept as conscious
will be directly related to the concurrent develop-
ment of input-output associations that prove fruit-
ful and is a measure of the diversity of the
behavioral capacity to respond to a stimulus.
Experiments in support of this hypothesis were
conducted on kittens raised from birth by restrict-
ing their exposure to the visual environment. Work
by Held and Hein [4] carefully dissociated activity
as the controlling variable. Kittens reared in the
same visual environment were either constrained
in a gondola or allowed to initiate movement.
The kittens were then subjected to the visual cliff
trial. Unlike the proactive kitten which always
chose the shallow drop, the one riding the gondola
failed to acknowledge the danger of the deeper
side and did not display visually guided paw place-
ment. On the other hand, because of periods of
free movement confined to total darkness, their
motor development was comparable to normals.
The authors conclusion was that variation in visual
stimulation can be effective in the development of
normal visually guided behaviours only when it is
concurrent with and systematically dependent on
self-produced movements. A further qualification
not made explicit in this classic study is that inter-
action may be a prerequisite for the development
of visual awareness. The kitten did not lack coordi-
nation but, as will be argued, failed to perceive
depth. In an experiment with similar implications
Riesen and Aarons [5] showed that kittens re-
strained from moving, but allowed a fully pat-
terned vision with both stationary and moving
objects, failed to learn movement discrimination
tasks. The authors concluded that a lack of mean-
ingful contact with the visual environment, may be
the sufficient causal antecedent. Motor efference
copy provides a solution to the problem of how an
effective motor history may be essential to the
emergence of awareness suggested by these
studies.
Behavioral diversification
Every percept that we describe as conscious has a
unique subset of possible motor relations. Each ob-
ject of perception will imply a unique history of
interaction for the organism in spatiotemporal
Vakalopoulos
terms. Conversely, the theory contends that an
organism will remain unaware of an uncategorized
perceptual system, that is, one not based on previ-
ous sensorimotor interactions. Given that a concur-
rent sensory stimulus is not likely to be identical to
a previous one, then a conscious relation to the
percept is predictive of what a previous motor his-
tory would likely have been. This confers a capac-
ity for generalization to established networks and
also a flexibility for dealing with novelty. The pro-
posed motor history is not the relationship of a sin-
gle act to a concurrent percept. It necessarily
relates to all possible exploratory movements
which form motor repertoires associated with that
stimulus. It is this history of multiplicity which is
intrinsic to the conscious act. A conscious act need
not translate to the execution of any specific motor
act, but only implies an historical association to its
concurrent motor relation.
According to this theory of premotor relations,
there is an a priori capacity of the conscious act
to modulate behaviour. As an example of restricted
behavioral repertoire one can look at a lower phy-
logenetic order. Simple organisms have small ner-
vous systems and neurons are collected into
discrete groups called ganglia. The brain of the
lower invertebrate worm, Ascaris, consists of sev-
eral ganglia containing 162 neurons. This number
never varies from animal to animal and each occu-
pies a characteristic position [1]. The large marine
snail Aplysia has a similar, but less complete invari-
ance. Even in this higher invertebrate the easily
identifiable cells of its ganglia make precise con-
nections with one another such that each cell ex-
erts control over a specific behaviour. Most
neurons connect invariably with the same follower
cells and it is possible to trace an individual com-
mand cell that controls an entire behavioral se-
quence. The finding that behaviour is always
mediated by the same cells that interconnect in
invariant ways suggests that the circuitry of inver-
tebrates does not evolve the required complexity
specifying cognitive structures.
What I am claiming is that given the underlying
potential of neuroplasticity in the brains of higher
vertebrates, the divergence of sensory neural net-
works is not a simple function of the richness of
perceptual input, but critically, depends on how
the organism responds to the stimuli generating
those percepts. A thought experiment would be
useful here. If an organism behaved repetitively
in an identical manner to all objects on every occa-
sion during its development (leaving aside whether
this is in fact likely) then, according to the theory
no retinal image can generate a state of conscious
awareness. A single neural network would suffice
A scientific paradigm for consciousness
to deal with visuomotor contingencies, as it would
represent the totality of prior interactive experi-
ence. There would be massive synaptic pruning
and atrophy of the occipital visual cortex.
Defining cognitive structures
The following discussion is a formal theory of cog-
nitive structures and how they relate to network
behaviour. As such it is a hypothesis and although
I give some empirical support under the heading
perceptual adaptation further experiments will be
required to determine the validity of my proposal.
A reflex act is not considered equivalent to an
unconscious behaviour. A homology may, nonethe-
less, be drawn between networks that serve ste-
reotypical responses to a percept and the lack of
awareness of that stimulus. The sense of the
unconscious will be restricted here to complex cog-
nitive processes that are acquired by a similar
mechanism of motor reentrant organization of neo-
cortical networks, but differ fundamentally from
those that support conscious categories by a defin-
ing association of the former with a compulsory
stereotypical output. This is not meant to indicate
that the subject is compelled to act but that an
unconscious process implies a compulsory associa-
tion between perception and a particular behaviour
for a specified network. Of course, there may be
many such networks that serve different stereotyp-
ical behaviours to associated sensory stimuli.
Fig. 1 is a simplified schematic of proposed cog-
nitive structures. The green lines on the left and
right represent conscious and unconscious neural
networks, respectively. Each neural network may
consist of many thousands of neurons and certain
elements may be distributed in parallel, for exam-
ple, the dorsal and ventral streams of the visual
hierarchy. The orange lines are neural networks
Coordinate
transformation
Neural Networks
Percept
Conscious Motor
Figure 1 Schematic of conscious and unconscious cognitive structures.
769
that encode for various behaviours 1, 2, 3, . . ., n.
These include primary and secondary motor areas.
An envisaged behaviour may be composed of a sim-
ple movement or a more complex coordinated ac-
tion. Experimental evidence in favour of this
concept of a specified neural network associated
with a behavioral repertoire comes from the
increasing complexity of movements that can be
elicited with stimulation of primary, supplemen-
tary and premotor cortex.
The dashed vertical lines are points of coordi-
nate to somatotopic transformation where percep-
tual information is distributed to a body-centred
topography. This linking of a percept to a motor
network facilitates, for example, visually guided
movements. Transformation is non-obligatory and
has a motivational basis for selection. The gaps be-
tween neural networks represented in the colours
green and orange, on the one hand and the ensuing
behaviour, on the other, are meant to further
emphasize this motivational requirement for a
serial link between percept and behaviour. Fur-
thermore, there is a motive not only for the trans-
formation of percept to behaviour, but also its
selection from a number of possible outcomes. Ar-
rows have been omitted because information may
travel in either direction in a neural network that
has reciprocal connections between cortical areas.
The diagram does not convey the feedback limb of
motor efference copy, but is reflected in its orga-
nization. In other words, the neural networks are
already products of categorization. The execution
of a distinct behaviour by each motor neural net-
work (orange) will possess its own unique motor
efference copy. The inference from the general
scheme presented in Fig. 1 is that a conscious neu-
ral network will be associated with multiple motor
efference copies, while the organization of individ-
ual unconscious networks will be only associated
with the motor efference copy of their affiliated
Coordinate
transformation
Behaviour Neural Networks
1
2 Percept
3
n
Motor Unconscious
770
motor neural networks. As a general rule, any
feedback limb of a motor output will categorize
its respective feedforward neural networks
(green). The intended scheme clearly illustrates
that a single neural network supporting a conscious
percept may be associated directly with various
behavioral repertoires. Conversely, each reper-
toire may only be associated with a distinct neural
network supporting the unconscious processing of
that percept. The oblique dashed line on the left
rep- resents a poorly defined or as yet unpracticed
behaviour and the corresponding dashed horizontal
networks on the right represent a potential uncon-
scious process that requires training for its matura-
tion. This model has obvious implications for skill
acquisition.
The proposed scheme for cognitive structures
does not ascribe unconscious behaviour to organ-
isms we infer to be non-conscious. This is not an
arbitrary distinction in the definition of cognition.
Accordingly, the mere possession of a nervous
system is not a sufficient condition for either con-
scious or unconscious processes. Learned complex
behaviours, beyond simple modulation, require a
feedback limb of motor output as an historical
factor in the general organization of neurobiologi-
cal correlates of conscious and unconscious pro-
cesses. This assumes the property of plasticity of
the neural circuitry at a critical stage of its
evolution.
A typical example of an unconscious cognitive
process is the performance of a task, such as driv-
ing, while simultaneously thinking. The evocation
of conscious imagery that is unrelated to the com-
pletion of the task illustrates a circumstantial dis-
sociation between top-down conscious and
bottom-up unconscious processes. Most people will
recall an instance of not being aware of much of
the drive home while being intensely preoccupied
with that days events. Driving a car utilizes multi-
modal perceptual information which, in this
instance, is unconsciously, yet effectively pro-
cessed. This case reveals many properties of cogni-
tive structures.
The stereotypical motor requirement of an
unconscious act does not restrict the subject from
a parallel dynamic cognitive intervention such as
switching to a state of awareness during the task.
In fact, the paradigm predicts that making signif-
icant changes to a behavioral engagement due to
unexpected outcomes does entail conscious activ-
ity. Examples include a pedestrian stepping out in
the middle of the road or the sudden braking of
the car in front. What Fig. 1 suggests is that a
conscious process may conceive of an atypical
behavioral response to any given situation. A neu-
Vakalopoulos
ral network supporting awareness of a percept im-
plies concurrent access to all possible motor
relations. Perhaps a simple analogy with a pertur-
bation in steady state between the input and
output modes may clarify this point. The interpo-
sition of a conscious mediation can achieve a new
equilibrium which may be subsumed by uncon-
scious processes either as an ultimate conse-
quence of training or by existing automated
networks.
The act of driving involves changing gears,
accelerating at green signals, stopping at red sig-
nals or at a destination. This may be achieved as
a fluent transition between unconscious reper-
toires to changing perceptual cues without the nec-
essary invocation of consciousness. The putative
precondition for all unconscious modifications to
behaviour is the consistent association of over-
trained responses with a stream of perceptual
information. Of course, conscious and unconscious
processes need not be exclusively dissociated, but
are often coactive during a single task. Thus the
perceived threat of an accident may elicit uncon-
scious and conscious reactions. Kunde et al. [6] re-
cently demonstrated conscious modulation of
unconscious processes.
Finally, what distinguishes a conscious percept
from an unconscious one is not that the former
may not be associated with a stereotypical re-
sponse. A conscious network can empirically sup-
port many such behaviours. However, even while
executing a specific behavioral repertoire the
possibility of a rich behavioral response remains
accessible to that network and in fact defines con-
sciousness by previous association. By contrast,
unconscious neural networks associated with dif-
ferential behaviours relative to a fixed percept will
be segregated by definition. Thus, the habitual acts
of cycling, driving or walking home along a well-
traversed route will process the same unconscious
visual cues in segregated circuits (green lines on
the right of Fig. 1), whilst there is the theoretical
capacity of a single conscious visual network for
all the specified actions (green line on the left of
Fig. 1).
The execution of all possible motor outcomes
with respect to a percept remains implicit to its
awareness, although by no means need it be expli-
cit once these networks are established in the neo-
cortex. In fact, conscious percepts are often not
acted upon in any explicit manner. A qualification
of the paradigm is the necessary corollary of under-
lying motives for the translation of conscious activ-
ity to a specific behaviour, but for the purpose of
these preliminary arguments a further requirement
to the current model need only be noted.
A scientific paradigm for consciousness
Premotor theory
Perceptual adaptation
The experimental use of optical devices that dis-
place, reverse or invert retinal images form a prom-
inent part of the early psychological literature.
Subjects who donned such devices found that after
initial discrepant sensations of tactual and visual
stimuli, which induced misreaching and errors in
the localization of objects, they gradually adapted
to the distortions and learned to behave normally.
Kohler [7] and Taylor [8] assume that adaptation re-
sults from changes in visual perception. This inter-
pretation is in apparent contradiction to studies
by Harris [9] using prisms that caused a lateral dis-
placement of visual targets either to the left or
right. Subjects whose heads were held fixed pro-
duced identical adaptive shifts when pointing at vi-
sual targets, auditory targets or when asked to
point straight ahead. However, no such adaptation
resulted when pointing with the unexposed hand,
as one would expect if the adaptation was in visual
perception. Nonetheless, intermanual transfer was
observed when the head was free to move [10]. Sim-
ply walking around while wearing prisms permits
adaptation to displaced vision. When the prisms
are removed the subject mispoints with both arms
even when neither arm was seen through the prisms
during adaptation.
Stratton [11] described whole scenes appearing
upright on occasions after wearing inverting lenses
for several days. During adaptation the new
appearance. . . was able to drive the old from the
field, because the new localization by sight showed
a perfect and constant relation to the reports by
muscular and tactual perception. Stratton [12]
concludes: the whole system of visual objects
can never by itself be either inverted or upright.
It could be inverted or upright only with respect
to certain non-visual experiences with which I might
compare my visual system-in other words, my tac-
tual or motor perceptions. Adaptation to revers-
ing spectacles occurs in a piecemeal fashion [7].
Some parts of the visual field are seen in the correct
location, such as buildings or advertisements, while
other parts remain reversed, such as inscriptions
seen in mirror writing. Eventually, letters and num-
bers achieve the correct reorientation. Taylor [8]
provides a lucid account of perceptual adaptation,
which becomes more veridical as the subject ac-
quires a more extensive behavioral repertoire
including walking and reaching.
771
Harris [9] believes that adaptation to lateral dis-
placement of a visual target when the head is fixed
is primarily the result of a proprioceptive and not a
visual perceptual change based on the absence of
intermanual transfer. Premotor theory can resolve
this apparent contradiction by appealing to the
hypothesis that distinct visuomotor networks are
organized by the motor efference copy of stereo-
typical movements and will be defined here in
terms of simple premotor relations. Thus, premo-
tor theory predicts that the visual perceptual
change in this instance results from unconscious
learning and is dissociable from conscious visual
localization. However, if the subject is free to walk
around or the head remains unrestrained then
acquisition of a larger number of responses will
transform the complex premotor relations and cor-
responding cortical neural traces so that eventu-
ally, a real change in conscious visual localization
results. Even in the latter cases subjects endure a
period of dissociation between successful behav-
iour and the distorting effects of reversing specta-
cles. Kohler [7] observed that with practice
cognitive strategies were built up unconsciously
irrespective of the initial verbal strategies to deal
with incongruent sensations. Walking on the street,
opening doors and bicycling all become possible
prior to seeing correctly. The daily fencing
experiment illustrates this point. After about four-
teen days errors disappeared, even though the sub-
ject saw the rapier point approach in a reverse
direction. Kohler distinguishes between the respec-
tive roles of vision in the act of parrying and when
the subject describes the direction he experiences.
The aspects of behaviour or movement that are
relevant for perceptual adaptation and by infer-
ence, development of consciousness of the infant,
remain unclear. What is the specific role of touch
(if any) in this process? Premotor theory does not
espouse the primacy of the sensation of touch
per se, but touch does imply a physical limitation
to movement and thus, the parameter of proprio-
ceptive change.
Because conscious awareness of the spatial loca-
tion of an object predicts the proprioceptive
change and force a limb will require to reach it,
distorting prisms will create an incongruent effect
between prediction and actual proprioceptive sig-
nals associated with successfully attaining the tar-
get. In time, through the mechanism of motor
efference copy the visual networks will be trained
to the correct spatial arrangement of the environ-
ment through the lenses.
Proprioception, as will be discussed in detail be-
low, is only one of two physical components of any
772
movement. Several investigators have observed
adaptation with passive exposure or movement
[13,14]. Thus, many of the perceptual changes
inferred from adaptation can be attributed to the
kinematics of movement. Note that this differs from
the theory of changes in kinesthetic sense [9], which
Kohler [7] believes are only transient and ultimately
conform to visual localization. However, some of
the distortions involved aberrations in shape or fig-
ure, straight lines appearing curved, all objects
appearing thin or thick depending on the position
of the eyes, familiar forms dissolving and reinte-
grating in ways never before seen and a rectan- tion system is not explicable merely in terms of
gle becoming a rhomboid [7]. The world appears
rubbery and people jelly like. The fact that ob- posed by physical interaction, a principle of
jects acquire a rigid and proper aspect with adapta-
tion suggests the acquisition of knowledge of the
visual world. I mean this in the sense of extracting
invariant properties that enable one to form cate-
gories or classes of visual objects, or recognition of
individual members, irrespective of actual spatial
localization. In fact, active movement greatly facil-
itates adaptation that might not be accounted for
by simple changes in position sense [15]. No compa-
rable adaptation was shown for passive movement
of body parts even when the variation in sensory
stimulation is matched to that received with free
movement. A motor theory of consciousness will
thus, need to invoke the additional torque or force
component of an action to account for the non-
kinematic dimension to perception.
The premotor act
The premotor relation is introduced as a concep-
tual term to approach a theory of consciousness
and does not correspond to a modular view nor a
specific cortical area. It is predominantly con-
cerned with the visual system, but has general
applicability to all sensory modalities. A major
principle of premotor relations states that con-
scious activity can attend to only that which it
has a motor relation to at any moment, the intri-
cacy of the response implicit. No motor act need
actually be executed, but is strictly assumed by
the historical interaction of the organism. This is
not at all obvious since the portrayal of cognition
is one of a closed neural phenomenon of no imme-
diate behavioural consequence. The conception of
consciousness as a premotor activity borne of a
sensorimotor history sets motor limitations to sub-
sequent attentional capacity. A spatial relation
that is diffusely attended attests to a generalized
motor response, such as navigating through a clut-
tered room. A focal motor relation narrows the
Vakalopoulos
range of perceptual awareness. What premotor
theory provides is a fundamental link between
our capacity for selective attention for apparently
single or largely restricted events and the capacity
to act upon perceptual information. In a sense it in-
verts the causal relationship between action and
perception by stating that awareness reflects a
behavioural history and in turn makes a prediction
about further action and not vice versa. In other
words we are not first aware of an object and sub-
sequently learn to interact with it.
Accordingly, the threshold capacity of an atten-
neural load. A neural basis for restriction is im-
embodiment. The unity of phenomenal experience
is composed of a unique set or subsets of motor his-
tory and emerges from a correlation of global neu-
ral excitement: sensory and motor associations
that proved coherent. A phenomenal instant is a
presupposed activity in the sense that an organism
cannot be aware of or attend to that which it has
no motor relation to or preacts upon. This is not
a description of the value of the object of attention
mediated by hedonic centers. The capacity of
attention and the conscious act are high level
descriptions of the same reductive process. In
other words, the capacity of attention, one that
excludes value, and the properties of awareness
are considered equivalent in premotor theory.
A second principle of premotor relations states
that the quality of awareness, whether visual,
auditory or otherwise, is determined by the capac-
ity of the premotor application. In other words, it
is not the physical nature of the medium itself,
which dictates the quality of consciousness, but
the manner in which sensory information can be
utilized by the organism that determines the class
of awareness. This becomes more obvious when
we compare the hearing in primates and sonar in
bats [16]. Although both animals detect changes
in air pressure it has been asserted that a bats
auditory capacity has the resolution of visual acuity
[17]. That which constitutes the conscious aspect
of a percept in its equivalent premotor act is the
full potential of the motor relation implied by that
percept. The premotor act is a term meant to con-
vey the historical significance of motor reentry in
the emergence of consciousness. It emphasizes
that the categorization of a percept as a conscious
act is predictive in a closed neural system of a his-
tory of effective motor feedback. The idea that
motor efference copy underlies consciousness is
not unique in itself. However, as discussed in the
next section current theories largely attempt to
rationalize an active motor representation with
A scientific paradigm for consciousness
its inherent problems of behaviourism and the
untenable view that concurrent activity in motor
areas directly subserve consciousness. The present
theory resolves these issues by looking at an histor-
ical association of motor activity to the current
awareness of the organism.
When I talk of embodiment I mean that an expla-
nation of consciousness has a somatic origin, it is
not a property magically invested on a naive brain.
Embodiment then refers to the feedback the neu-
rons receive from the entire organism when it gen-
erates movement in its surroundings. This includes
feedback from joint afferents and the efferent
copy of the tone one exerts in the various muscle
groups one uses. I must stress here that interaction
has a very specific form which is lacking in current
motor theories. It is not enough to say that senso-
rimotor contigencies must be obeyed, one must
specify the exact nature of these contingencies.
This is more than mere arbitrary movement in
ones environment, but one needs to bump into
things if you like. Some sort of tactile exploration
is mandatory to register a tangible association be-
tween visual percept and motion. By tactile I mean
a physical interface to movement and not the phe-
nomenal sense of touch. The nature of this infor-
mation depends not only on self-generation but
the limits the environment imposes on these move-
ments. Eye movements and the induced retinal
changes in input are inadequate for the organism
to acquire knowledge of its surroundings. Thus, a
certain object and its place in space will dictate
reafferent information that is consistent with every
interaction the person has with it. Objects must re-
act to ones skillful manoeuvring. Every action
must generate an opposite reaction for an object
to acquire phenomenological form. Thus, aware-
ness emerges from the consistency in the associa-
tion of the act on and the percept of the object.
Once a concrete knowledge is felt, so to speak,
then can one abstract this self-referential knowl-
edge at a distance. This predicts that if there is
no interaction with the world then perception will
remain at a reflexive level one cannot be aware
of. Thus, a percept is a scene or an object one acts
upon and its awareness is preaction if you like. This
term originates from the idea that awareness is the
prediction of an interaction based on previous
experience or an extrapolation of previous
experience.
This brings us to the next point about how one
gains knowledge if there is no possible contact with
1
I am grateful here for discussions with William Banks, which
has helped clarify my position.
773
certain objects such as the moon or the picture on
a currency note.1 The beauty of the current theory
is that it is a prediction based on the historical fact
of interaction and as such extrapolation is possible
to novel elements of the environment. One does
not need to have previous experience of the moon
in a physical sense to have knowledge of the moon.
Premotor theory simply predicts what that physical
interaction is likely to be like if one ever has the
opportunity to visit the moon. This is based on past
experience of other objects, which will have pre-
sumably similar textures shapes luminance, etc.
Ofcourse the prediction may be wrong. Perhaps
our past experience was proven incorrect with re-
spect to the moon. Perhaps the atmosphere is a
distorting prism and what we thought we knew
was incorrect. If we could reach out with a ex-
tended arm device and touch the moon and find
its position is not where we thought it was, perhaps
15 to the right of where we tried to grab it. In fact
its shape was rectangular on palpation. Once we
have gained considerable experience with it and
retract our artificial arm we now see the moon as
a rectangular object 15 to the right of everyone
else.
Every novel object is described in terms of pre-
vious experience, i.e., it has contours, texture,
colour, size, shape that in that instant may be
new in its arrangement but its elements have been
experienced before. Thus, when one holds a small
ball the afferent information is very specific in
the way this solid object deflects ones movements.
In fact it dictates all possible movements in rela-
tion to it, i.e., one cannot simply put ones finger
through the ball or walk through a wall without
altering the relationship. This given set of laws be-
tween person and object is crucial to the establish-
ment of awareness of the object. A larger but
similarly textured ball means that running ones
hand over the surface describes a greater arc and
has its corresponding afferent information. It is
not simply the retinal image of the larger ball
which tells us that our grip must be larger, but
the past reafferent experience that interprets the
retinal image and tells us the ball occupies a larger
dimension of space. Previous afferent information
described for us the arc of space and its locus
and now perception of it predicts this. The predic-
tion depends on a stable association between per-
cept and action. Lets take the thought
experiment where no such stability is present.
Say that the usual laws of physics do not apply
and the object changes shape in a physical sense
at random but it still subtends the same arc of ret-
inal image. Our premotor relation would predict
the same laws but we either completely miss it in
774
attempting to grasp it or we break it because it has
transformed to glass rather than being this rubbery
ball our senses were predicting. If the new form re-
mains stable enough then with repeated interac-
tion we will establish another set of laws in
relation to this object and we will now be aware
of not the same ball but a crystal figurine in a dif-
ferent spatial locus. This is exactly what experi-
ments with glass prisms have taught us. For
example, if a prism were to make a big ball small
and a small ball big, the incongruence of our phys-
ical experience would end up altering our percep-
tion of the balls. A new equilibrium is established
that is congruent between action and perception,
irrespective of the size of the image on the retina.
Most importantly this occurs independently of eye
or head movements unless these are an interface
for a reaction from the object in question (not
likely for eye movements but infants use their
mouth to explore). It is the physical contact that
informs the brain about what it is seeing. Logically,
the changing retinal image brought about by move-
ment cannot be what motivates visual adaptation
as the image is the same and does not inform us
of the falseness of perception, only action upon
the balls does. Some other medium of information
must be at work to inform us of changes in the real
world with respect to the image and this is medi-
ated by motor reafference. The body cannot lie
to you when it bumps into a wall.
Another way of looking at it is the world and its
objects deform the range of movements available
to an organism and this is the only mechanism by
which it (organism) can be informed of its (world)
true visual or for that matter auditory nature.
Thus, movement through space etches out a land-
scape. This is correlated with the types of percep-
tual input one is receiving and imposes a quality on
the modal experience that will depend on the way
the body in its movements can use the information,
i.e., the resolution of auditory or visual input into
specific actions. By use, I mean motor sequelae
based on what previous movements might have re-
sulted in those sounds or the visual percept of an
object associated with instances of tactile explora-
tion and not utility. An object has a certain form
and egocentric spatial relationship that only bodily
movement can resolve. A set of forces and propri-
oceptive changes corresponding to all sets of possi-
ble movements associated with that object are
unique in defining that relationship. To put it sim-
ply, a square object at one locus will require a dif-
ferent grasping posture and trajectory of the limb
than a spherical object at another locus. These
movements tell the brain what the eyes are see-
ing and what the ears are hearing. As argued fur-
Vakalopoulos
ther below, the physical laws laid down by a
motor theory of consciousness will comprise of
two orthogonal dimensions of an action, force
and proprioceptive change.
Obviously, a sound wave does not have the
resolving capacity of visual input in terms of motor
response. So the way the movement of the body
through space etches out its phenomenal aware-
ness of a sensory medium depends on a certain con-
cordance between action and perception. What I
mean by this when considering sound for example,
it reflects a subset of the entire knowledge gained
by highgrained movement and it can only inform
the brain of the previous bodily experience in a
specific and limited way. That is every reaction of
an object on a movement will have a certain visual
input associated with it and a certain auditory
character both of which when played back so to
speak can only convey a specific portion of the bod-
ily experience with vision having a far better reso-
lution. The ability for the body (bat) to resolve its
environment through sonar allows it to see. Audi-
tory waves emanating from a particular source
present a different landscape of the same objects
as judged by reactions to movements and thus
one can hear ones experience of movement but
not see it. Different sensory modalities alter the
reality of the substrates the body has to work with
even though it is describing the same objective
world. The landscape the body etches out is differ-
ent for different sensory faculties.
Preaction has the property of generalization.
Once the networks have been established one in-
fers but does not need to interact physically again.
This also deals with the other potential flaw of
motor theories of consciousness in that paralysis
can occur after categorization of perceptual infor-
mation without affecting what is perceived. For
want of a better term, by categorization of a per-
cept I simply use this term to describe the estab-
lishment of networks within the brain that serve
information processing whether implicit or explicit
by the mechanism of reafference. They can be
simple visual features or a general class of object,
which is the more usual meaning of the term.
Once established a neural network that serves
the conscious perception of any object has
evolved a life of its own that is no longer depen-
dent on further interaction but is nonetheless
inherently assumed to be part of the representa-
tion of any percept. It is in this context that I refer
to virtual action (no single term can convey the
full significance of the concept, of course). The
term preaction or premotor theory came to me
for a number of reasons. It alludes to the impor-
tance of motor activity in the evolution of neural
A scientific paradigm for consciousness
awareness, that it predicts action but does not re-
quire its execution.
How does colour fit into this picture. Every ob-
ject has a border and this border describes the ex-
tent of an object and the extent of interaction of
the organism. Ones experience with one object
of one colour may be identical with the same ob-
ject of another colour except for a distinct spatio-
temporal relation. Wavelength then adds another
perceptual feature that allows discrimination but
it is the separate spatiotemporal dimension that al-
lows the distinct wavelengths to be perceived as
different colours. When an object subsequently is
perceived and is composed of two colours say blue
and yellow these will form a border even under the
same intensity of illumination. This border repre-
sents a theoretical border that one can act on again
based on the experience of separately coloured ob-
jects. This border however is fictitious in the sense
that no reafference can differentiate it on a
smooth continuous surface and it is inferred from
the previous separation of colour in distinct ob-
jects. Conversely, this predicts that if a pair of col-
ours were always present on a single continuous
surface without the option of a tangible physical
experience of distinct borders, such as between
separate objects with those colours, then this bor-
der would not exist in perception, i.e., it cannot be
categorized and the surface would appear as a sin-
gle colour or colourless.
This is supported by the work of Kohler [7] with
split-coloured lenses (blue or yellow) for each eye-
piece, where there was no option of differentiating
the objects according to colour. In other words,
the laws between movement and colouring of the
world would become arbitrary. The lenses do not
provide any useful information in a discriminative
sense. There are no longer any fixed laws about
the colour of objects that can be discerned from
different perspectives. The colours actually faded
over time while the subject was wearing the
lenses. Adaptation results in a loss of information
and not normalization of colour perception: Sec-
ond day: . . .when the eyes remain fixed, even
for a short time all colour contrasts rapidly disap-
pear; even the dividing line vanishes after a
while. And Thirty-first day: Even though a thing there is to know about a visual object can be
great variety of yellow and blue stimuli are trans-
mitted by my spectacles and keep impinging on
my fovea, I no longer experience the corresponding
colour sensations. This is extraordinary support
for the current position.
A third principle relates to a component theory
of the premotor act which I believe is the most inno-
vative aspect of the theory and provides a truly use-
ful and entirely original application of a high level
775
paradigm to cortical structure and hierarchy. Mus-
cular contraction due to a specified motorneuron
discharge may have several effective outcomes that
are not identical. In the organisms interaction with
the environment movement may be limited by the
objective milieu with theoretically, no change in
motor intention. That is, an isotonic force may or
may not, result in actual movement. This added
informational state is conveyed by proprioception,
its function a component of the totality of the mo-
tor ensemble that specifies a categorical mecha-
nism of reentry.
What truths are discovered by movement that
can be set to laws of a component theory of action.
I do not think proprioceptive changes associated
with body movements to and from objects presents
any real conceptual problem for understanding the
acquisition of knowledge of a spatial locus coordi-
nate system and thus spatial awareness. Any set
of movements to each point in space are associated
with a specific subset of reafferent inputs of joint
position changes. The ability, of course, to per-
ceive a specific spatial locus will depend on the
capacity for modal resolution. Differences in sound
waves generated from closely contiguous spatial
loci will be difficult to detect and thus, only knowl-
edge of a more diffuse spatial localization can be
gained by movement. But what determines the
phenomenology of form independently of spatial
locus, i.e., a sphere, for example, retains its intrin-
sic shape irrespective of its spatial relationship to
the body. If movement has an equal and opposite
reaction from the objects it acts upon, then this
can be represented by the specific forces muscle
groups exert on these objects and that are unique
to them. For example, the exploration of a rectan-
gular object requires distinct grip forces of intrinsic
hand muscles to that of a sphere. The sounds gen-
erated by walking on autumn leaves, biting into an
apple or knocking on a wooden door offer a unique
pattern of resistance or decelerations that can be
described and quantified in terms of sets of specific
forces exerted by a range of movements. We now
have powerful tools for a physical description or
coding of the phenomenological world in terms
of torque (force) and proprioception. Thus, every-
mapped onto a physical system of registered body
movements.2 This is certainly true for form (size,
shape, texture) and spatial locus. It represents a
2
I want to make it absolutely clear I am not talking about the
emergence of tactile sensation (although it is applicable to all
modalities), but what the reactions of the world to bodily
movement mean for the visual (or auditory) system to gain
knowledge.
776
definitive bridge between mind and body, even if
one considers the limiting condition of a colourless
world the most satisfying under this system. It will
not escape the notice of the reader that the pro-
posed laws of motor behaviour reduce the subjec-
tive nature of phenomenology (even in this
limited sense) to the absolute principles of physics.
The special case of colour
Now consider the situation of two identical objects
(balls) in terms of shape and size, but differing in
colour, red and green. Colour in this situation does
not suggest any discriminating properties that can
be discovered through manipulation except for a
spatiotemporal difference. Is this the limit of reso-
lution of the current theory or, are there situations
where an organism can gain direct knowledge of
colour through the proposed form-resolving com-
ponent of action, force? Instead of balls lets take
a fruit like a tomato or banana and identical exem-
plars of each except for the colour, green and red
for the two tomatoes and green and yellow for
the two bananas. The colour differences signify
ripeness and tactile exploration, both manual or
by biting into them, can reveal inherent differ-
ences in compressibility or density that are re-
flected in the types of forces one uses to assess
the fruit. Let us call one of these properties
squishiness. Even though there are many arbitrary
relations between colour and object, the paradigm
cases still allow us to gain adequate knowledge.
Even arbitrary cases can be paradigmatic (see be-
low). What this means is that shape, size or texture
may not reveal the physical relation in this context
and only colour maps onto the force differential
discovered through manipulating these objects.
The fact there are colourobject relations, which
do not mimic or predict defining laws such as the
example of the balls, does not negate the discovery
of the paradigm cases. We reconcile our sense of
being fooled as long as other laws of form and loca-
tion are not being breached, unless one introduces
absolute arbitrariness such as wearing divided Koh-
ler lenses, where colour fades.
In fact, it is likely a child learns to appreciate
colour through arbitrary associations. Blue may sig-
nify a hard round ball, yellow a soft toy, red a rub-
ber object. Each colour within the context of the
childs world predicts a specific mechanical rela-
tionship even if not absolute in the real world. This
can be applied generally to the theory. Our discov-
ery of laws allows us to infer them from other
external representations such as books, photos,
pictures. The contents of the latter are symbolic
Vakalopoulos
of a real world even if they do not themselves obey
laws of action.
Ecological approach to perception
I would like to place premotor theory in its proper
historical context by discussing what it can add to
the ongoing debate about whether perception is di-
rect [18] or mediated by an internal representa-
tion, i.e., indirect [19]. Gibsons approach is a
functional analysis of perception constrained by
ecological imperatives. The organism is insepara-
ble from its environment. Thus, knowledge is
gained through self-movement, which causes
changes in the patterns of stimulus parameters
from which the organism extracts invariants. Gib-
son has a major influence on current thinking on
perception. In support of Gibson, Clancey [20] pur-
sues the idea of perception as action arguing that
categories are learned by interacting with the envi-
ronment. A problem with Gibsons theory is the
nature of the invariants picked up from the flow
of an optical array and its relation to self-move-
ment appears to me unsatisfactory. It is rather sim-
plistic to say that knowledge of the world can be
gained by changes in the flow of retinal patterns
brought on by movements of the eyes or head or
general bodily motion. One revisits the same limi-
tations in contemporary sensorimotor contingency
theories [21]. The change in perspective to the
world or an object as one scans or moves around
still refers to a pictorial image on the retina, it is
not an explanation of perceptual experience.
Clearly, premotor theory differs in that it does
not refer to feedback as perspectival change. More
sophisticated interpretations are discussed by
Clancey [20] and Shaw and Todd [22] who empha-
size the reciprocal relation between action and
perception and who regard the perception of things
as a function of actions they afford. Gibson does
mention that certain facts about objects can only
be revealed by reaching and picking up, but there
is no explicit account of an organizing principle
for the visual system.
Turvey and Shaw [23] attempt to address this
shortcoming by investigating physical laws that de-
scribe perception in terms of behaviour. This is the
main agenda of a theory of premotor relations. A
theory of motor reafference collapses percept
(stimulus) and motor outcome because motor reen-
trant function upon the cortical networks serving
perception implies neural properties (including
the genesis of phenomenology) are a direct result
of the confluence of both stimulus and action.
A scientific paradigm for consciousness
One may conceive of the elementary components
of motor efference copy, torque and propriocep-
tion, as an algorithm for perception. The algorithm
refers directly to the world because it is discovered
through previous physical interaction. The algo-
rithm codes for the physical constraints the envi-
ronment imposes on the active seeking body in its
entirety, head, mouth. tongue, arms, legs etc. If
perception of an object is an algorithm as premotor
theory claims, specified in numerical vectors for
torque and proprioceptive change, then one can
use this algorithm to judge say the movement re-
quired to pick up an apple from a bench top (visu-
omotor coordination). This movement is inherent
in the perception of the apple regardless of
whether the person decides to perform this activ-
ity. To remain consistent with my initial descrip-
tion of conscious versus unconscious processes,
being consciously aware of the apple allows one
to use a number of different algorithms. One can
pick it up with the left hand or the right hand or
with ones feet if one so desires or even with ones
teeth as in a game. Thus, within the conscious con-
struct of the apple is the implicit influence of the
different specified algorithms on the design of the
neural networks from the activity of which con-
sciousness emerges, i.e., these algorithms endow
the neural circuit with the property of phenome-
nology. At least, this is my theory. By contrast, if
one picks up the apple almost automatically, i.e.,
without really consciously acknowledging it, then
a single algorithm specifies this parallel perceptual
network. This satisfies the stereotypical presuppo-
sition of unconscious networks. The stimulus de-
coded by the algorithm and perceived (either
consciously or unconsciously) as occurring now
is a prediction it will be compatible with the knowl-
edge it has discovered of perceptaction relations.
On this last point premotor theory reconciles the
polarized views on cognition because action di-
rectly determines the categorization of perceptual
neural structures that form a representation of the
world. Premotor theory is also a prediction based on
the experiential history of the organism and as
such, is a belief system satisfying the inferential
nature of cognition. Gregory [24] is also an advo-
cate of perception as prediction. This allows one
to recover or insert information absent from stim-
uli (forbidden by Clancey) because of the temporal
disconnection of perceptual categorization from its
prediction about the meaning of the current stimu-
lus. The basis for completion of an illusion (e.g.,
Kanisza traingles) and for what we expect to see
changing what we see can be modeled this way as
can overt hallucination and is compatible with the
modern cognitive view of top-down and bottom-
777
up interactions. Of course, prediction is generally
proved correct. According to the premotor model
reentrant categorization occurs at all levels of rep-
resentation. In fact the very structure of the visual
hierarchy is a product of reentrant function. Per-
ception reflects the symbolic nature of action.
Thus, all internal states are predicated or embodied
in action. It is the embodiment of perception
through action that proposes an objective nature
to subjective experience dissolving the dualistic
hold mental experience has on the philosophical
discourse about consciousness.
A solution premotor theory offers to the prob-
lem of perception without action (one is aware of
ones surroundings without any compulsion to per-
form an act concurrent with that perception) is a
solution argued by Shaw and Todd [22] against
internal representation. In classical cognitive the-
ory perception is regarded as a function of internal
state Q(t) and stimulus S(t) at time t. Shaw and
Todd replace the term Q(t) with H(t) which refers
to the entire history of transactions of the animal
with its environment and thus, do away with the
need for internal states mediating perception.
The logical conclusion, which they are reluctant
to adopt, is on the equivalence of the terms such
that Q(t) = H(t). Premotor theory, justifies the iso-
morphism of internal states and history of transac-
tions by way of an elementary physical mechanism
that ascribes values or an algorithm to the neural
architecture and which is generally lacking in the
ecological approach.
Contemporary action theories of
consciousness
A number of recent articles have examined the
aetiology of motor behaviour in consciousness.
Although promising novel insights, they have gener-
ally been awkward in the face of criticisms of
behaviourism or functionalism. ORegan and Noe
[21] propose a sensorimotor contingency model of
visual experience, but do not offer a satisfactory
explanation of its relation to cortical structure. I
believe that only a theory of motor efference copy
(MEC) can bridge this gap. Secondly, the authors
acknowledge that sensorimotor contingencies are
not sufficient for consciousness, unless they are
linked to thoughts and planning, but the model fails
to clearly demonstrate how the differential nature
of such contingencies relates to conscious and
unconscious processes. Subliminal priming experi-
ments have demonstrated how unconscious mecha-
nisms can modify cognitively mediated behaviours.
778
Most sophisticated motor theories of conscious-
ness attempt to circumvent the criticism inherent
in any requirement of overt behaviour by postulating
some representation or knowledge of a motor pro-
cess. Thus, ORegan and Noe are forced to reassess
their invocation of a correlation of awareness with
eye movement or a dynamic probing process. The
current paradigm is a significant departure from pre-
vious theories, by proposing a developmental model
where awareness emerges from a historical associa-
tion between percept and motor activity. In fact,
proponents of Gibsons ecological approach to per-
ception do emphasize that perceptual output is a
product of the stimulus and the entire history of
transactions of the organism with its environment
[22]. Neither a concurrent act, nor an active repre-
sentation of motor contingencies is necessary. Pre-
motor theory avoids the premotor cortex and thus
the pitfalls of behaviourism. In other words, it does
not require that sensorimotor contingencies are
currently being obeyed [21]. Premotor theory is
simply a prediction that these laws will be obeyed
and thus, may be dissociably configured from the
present, for example, under circumstances of
dreaming or hallucinations. Of course, these laws
will determine adaptation when demanded by a
change in the history of sensorimotor relations, for
example, by wearing inverting goggles.
Of far greater concern is the perspectival senso-
rimotor contingencies envisaged by ORegan and
Noe are inadequate to specify bodily knowledge
of perceptual features. The statement that visual
qualities are determined by the character of the
sensorimotor contingencies set up by the visual
apparatus is difficult to support. Although ext-
raretinal signals are important in specifying an ac-
tual spatial locus, I will now argue that saccadic
activity is not critical to consciousness because
the objects that are so explored, do not provide a
limiting condition to ocular movements. The
authors do mention that other body movements
may change the perspective of an object, sensori-
motor contingencies are rules governing the sen-
sory changes produced by various motor actions,
but this appears to refer to the changes induced
on the retina and not the physical limits imposed
by the object on any motor interaction. In other
words, visualness cannot be explained alone pathway that could drive motor reentry. Cortical
by the character of the sensorimotor contingen-
cies produced by exploration mediated by the vi-
sual apparatus. The ability to follow contours of
an object with our gaze must already be inferen-
tial. Premotor theory states that such inferences
are derived from the previous limits imposed by
the environment on bodily movements. Patterns
of retinal stimulation must be simultaneously
Vakalopoulos
associated, at a cortical level, with the MECs that
such limits generate. Movements (saccadic or bod-
ily) that initiate changes in the retinal signal
without any limiting contact have a certain arbit-
rariness with respect to the developmental history
of neural networks that might engender perceptual
experience. The MEC of ocular movement has no
limiting conditions imposed by the visual environ-
ment (in terms of physical barriers to movement)
and thus, cannot extract invariants associated with
retinal input. Purposeful scanning of an object al-
ready presupposes knowledge!
Categorization
The hierarchy of receptive field properties of corti-
cal neurons is the postulated result of motor reen-
trant categorization. It is generated and refined
by the motor reafference of an interacting organ-
ism.There is a massive cortical projection to the
striatum [25] and by extension, the pallidum which
is a major source of efferents to the motor thala-
mus. It appears to play a role in motor behaviour
as judged from the extrapyramidal signs exhibited
by Parkinsonian patients. The basal ganglia also
have projections to the thalamic reticular nucleus
(TRN) which modulates the activity of the entire
dorsal thalamus. The physiological role of this path-
way is even more obscure, but it does provide a suit-
able candidate pathway for a reentrant mechanism
as proposed by the model of premotor relations.
The categorical role of motor reentry may feasi-
bly be mediated by reciprocal thalamocortical pro-
jections. The dorsolateral geniculate nucleus
(dLGN) and the pulvinar (Pul), parts of the latter
also displaying retinotopy, are the putative nodes
of motor reentry for the visual cortex. Release from
the inhibitory effects of the TRN may differentially
enhance cortical networks. The reticular collater-
als of the corticothalamic and thalamocortical pro-
jections serve to increase the signal-to-noise ratio
[26]. The external segment of the globus pallidus
(GPe) projects to the reticular nucleus [27]. Its neu-
rotransmitter is GABA. The primary motor topogra-
phy of the dorsolateral subthalamic nucleus and its
glutaminergic projection to GPe [28] is an ideal
subthalamic afferents originate principally, from
layer 5 including collaterals of pyramidal tract fi-
bres. These would qualify as carrying efference
copy. Reciprocal thalamocortical circuits function
as a limb of a broader loop to the basal ganglia (cor-
ticostriatal) that engages the motor complex in a
feedback mechanism. The ventromedial subthala-
mus receives an input from the premotor cortex
A scientific paradigm for consciousness
(not synonymous with the premotor act) and, in
turn, projects to GPi and SNr. There is some evi-
dence that the latter, including the ventral palli-
dum, also project to the TRN [29]. The association
of subthalamic networks with the basal ganglia
may represent a general system of motor reentry
with parallel functional subdivisions.
Component theory of motor efference
copy
Torque and change in joint position are the two
parameters that specify all motor activity including
velocity and acceleration. Motor reentry will be
based on a component theory of motor efference
copy serving distinct cognitive parameters in the
organization of neocortical networks. It is not intu-
itively obvious parameters of a motor action spec-
ify the properties of posterior cortical regions one
tends to associate with object or face recognition,
for example. It is precisely this conceptual under-
taking I am asking the reader to take. Conditions
of torque in a neural network may conceivably be
specified by the basal ganglia via TRN modulation
of thalamocortical projections. Whether the role
of the basal ganglia can be entirely defined in this
manner remains to be demonstrated. However,
several extrapyramidal features of patients with
Parkinsons disease support this claim. Signs of bra-
dykinesia are generally considered a deficiency of
dopamine regulation within the basal ganglia. An
elegant study by Morris et al. [30] showed a capac-
ity for these patients to vary their gait velocity
mostly by an increase in cadence (steps/min) al-
beit, with a reduced range compared to controls.
The authors concluded that stride length is the fun-
damental problem in their gait disturbance and
speculated an indirect mechanism affecting the
generation of force.
Proprioceptive feedback is the second critical
element of premotor category function. Joint posi-
tion will be served by the cerebellum. The organiza-
tion of cerebellar global networks parallels that of
the basal ganglia [31]. The afferent limb is a mas-
sive corticopontine and mossy fibre system. The
efferent reentrant limb is a projection of the deep
cerebellar nuclei to the motor thalamus. This loop
covaries with peripheral cerebellar input. The cor-
ticopontine network has a restricted origin, with a
distinct spatial character, in contrast to the wide-
spread input to the striatum [32]. In the visual hier-
archy, it is largely confined to the dorsal stream and
parietal cortex and fulfils the criteria of a specific
spatiotemporal relation to the organism and which
can only be specified by changes in joint position
779
during an interactive history. Conversely, the cate-
gorization of a percept in the inferotemporal cortex
is not spatial, but feature-dependent. Recognition
is generally, orientation insensitive, eschewing a
proprioceptive role in the organization of circuits
that serve it. Receptive field properties of the ven-
tral stream are tolerant of various spatial loci and
orientations for an object. This cannot be the case
for accurate visuomotor control that is the pre-
sumed function of the dorsal stream.
Empirical evidence
There is a burgeoning body of literature implicating
the basal ganglia and cerebellar loops in cognitive
circuits [33]. Localized surgical lesions of the inter-
nal globus pallidus, which alleviate some of the
motor abnormalities pathognomonic of Parkinsons
disease such as motor initiation, rigidity and brady-
kinesia, produce further cognitive and learning def-
icits compared to the preoperative state [34].
Maguire and Ogden [35] analyzed the MRI scans of
patients with persisting unilateral neglect. Lesions
most commonly involved the basal ganglia suggest-
ing a fundamental role for this structure in percep-
tual or attentional processes. The acute syndrome
is often characterized by a lack of awareness of
contralateral space generally results from damage
to the inferior parietal lobule and resolution is typ-
ical. Persisting neglect which is rare and is associ-
ated with extensive lesions of cortical and
subcortical regions, implicates the basal ganglia
in cortical adaptation to ischaemic insults.
Patients with Parkinsons disease are impaired
at perceiving affect [36] and on a recognition mem-
ory test for both familiar and unfamiliar faces [37].
The study by Jacobs et al. [36] showed that the
deficits extended to imagining and also expression
of facial emotion. The authors concluded that a
correlation exists between motor and imagery gen-
eration that is not accounted for by current theo-
ries of perception. Monkeys with lesions of the
tail of the caudate nucleus produce deficits of pat-
tern discrimination comparable to lesions of the
temporal lobe [38]. Dewick et al. [37] showed in
an elegant series of experiments that parkinsonian
subjects suffered from two distinct face processing
impairments. In addition to a perceptual deficit,
when patients and controls were matched in per-
formance on a structural face test, the Parkinsons
group showed a relative disproportional deficit in
memory for faces. These findings support a role
for the basal ganglia in the ongoing categorization
of ventral stream networks believed to serve rec-
780
ognition. Recently, the subthalamic nucleus has
also been implicated in cognitive function.
Weiner et al. [39] showed that patients with cer-
ebellar dysfunction, but not Parkinsons disease,
were impaired in perceptual adaptation to lateral
displacement of an image by prisms. They
demonstrated no negative after-effect after re-
moval of the prisms. Negative after-effect is a sign
of persisting adaptation and refers to misreaching
in the opposite direction. In a further analysis of
the right hemisphere damaged patients, only the
occipital subgroup demonstrated both an impaired
error reduction and diminished after-effect, simi-
lar to the cerebellar group, consistent with a pri-
mary perceptual dysfunction. A parsimonious
explanation of such a correlation is that cerebellar
damage results in a secondary impairment in the
reorganization of visual cortical networks associ-
ated with spatial perception. There is much specu-
lation regarding the cognitive role of the
phylogenetically newest part of the cerebellum.
Akshoomoff and Courchesne [40] showed impaired
rapid shifts in covert attention between the audi-
tory and visual modalities in patients with damage
to the neocerebellum.
A model for psychiatry
Up until now we have only considered how external
distortions of perceptual input (e.g., prisms) lead to
incongruent visuomotor actions and how motor dis-
covery of a new reality determines a new set of laws
are translated into a new phenomenal awareness.
However, what if the reafferent information con-
veys an inaccurate or false copy of motor behav-
iour. In this situation, the nature of forces
generated by reacting objects is a degraded repre-
sentation or the range of reafferent proprioceptive
input is unavailable to the exploring organism and
brain circuitry. Such dysfunction of proprioception
and force have been quite clearly observed in aut-
ism and schizophrenia, respectively. Premotor the-
ory predicts a causal relationship between observed
motor dysfunction and the perceptual anomalies
observed clinically. Although, not a novel idea, a
convincing causal link between motor and cognitive
dysfunction in psychiatry has never previously been
made. Not only do motor symptoms figure promi-
nently in schizophrenia and autism, but the patho-
physiology of either implicates the basal ganglia
and cerebellum, respectively. Below I propose a
model of cognitive dysfunction based on a compo-
nent theory of motor efference copy, providing a
heuristic framework for understanding some of
the most perplexing clinical features.
Vakalopoulos
Evidence from schizophrenia research
The corticostriatal and thalamocortical cir-
cuits have been implicated in the pathogenesis of
the positive symptoms of schizophrenia including
hallucinations. Hallucinations are most commonly
auditory with verbal content, but also include vi-
sual and tactile elements [41,42]. Reports of per-
ceptual distortions of familiar objects are also
prevalent, including brightness, colour, size, depth
and movement [43,44]. The dopamine hypothesis
of psychosis has endured since the discovery that
the efficacy of neuroleptic treatment is related
specifically to affinity for D2 receptor antagonism
[45]. The predominant postsynaptic localization
of the D2 receptor is in the basal ganglia and spe-
cifically on striatal GABAergic projection neurons
to GPe, the so-called indirect pathway (see review
by Gerfen and Wilson [46]). Indeed, Carlsson et al.
[47] explicitly implicates dopaminergic disinhibi-
tion of GPe output to the thalamic reticular nu-
cleus in psychosis by releasing thalamocortical
projections from inhibitory control. This may be
exacerbated by the glutamatergic projection to
the external globus pallidus by the subthalamus.
The nucleus accumbens or ventral striatum paral-
lels the organization of the dorsal striatum [48].
Stimulation of the ventral pallidum increases activ-
ity of the dorsal thalamus via its projections to TRN
[29] and may well be a primary target of neurolep-
tic treatment.
Premotor theory predicts that an abnormal cor-
relation between motor efference copy and sensory
input will result in cortical disorganization and
aberrations in perception. In fact, it predicts the
findings of Delevoye-Turrell et al. [49] which dem-
onstrate a deficit in scaling of grip force to a varia-
tion of load task in a group of schizophrenic
patients. The delusional nature of positive symp-
toms can partly be attributed to primary disorders
of perceptual awareness [43]. Abnormal efference
copy due to hyperdopaminergic release of thalamo-
cortical networks may be exaggerated by
movement, as described above. If the torque
para-meters of motor efference copy fluctuate
arbitrarily with respect to a set of motor acts in a
normal individual, then object inconstancy results
under the current model. In other words, an under-
lying dysfunction of scaling of torque impairs differ-
ential categorization of objects and results in
unstable perceptual conditions. Thus, Chapman
[43] notes that immobility attenuates cognitive dis-
ruption: Everything is all right if I stop. If I move
everything I see keeps changing, everything Im
looking at gets broken up and I stop to put it to-
A scientific paradigm for consciousness
gether again. If abnormal MEC mediates move-
ment related impairment of perceptual constancy
then this may induce the delusion of empowerment
to alter the shape and size of people and objects.
An analogy can be drawn with the descriptions
of paroxysmal upright perception facilitated by
activity while wearing inverting lenses [12]. In the
case of schizophrenic subjects the dysfunctional
adaptation is the result of abnormally generated
motor contingencies and their efferent copies.
Thus, the reality of belief systems and loss of insight
may reflect fundamental changes in perceptual net-
works. A unique behavioural marker was proposed
by Rosen et al. [50], with the discovery of a grip ten-
sion deficit that was unrelated to maximal strength
or motivational factors, but analogous to the
impediment in scaling grip force described by
Delevoye-Turrell et al. [49]. This motor anomaly
was not shared by another subgroup of psychiatric
patients with affective disorder. The deficit was
also present in first degree relatives of schizo-
phrenic probands implying a marker of vulnerability
for the disease. Rosen et al. concluded that the task
tapped asymptotic performance as it remained sta-
ble both within and across test sessions.
Blocking is characterized by transient periods of
immobility, blank expression and fixed gaze occur-
ring in the vast majority of schizophrenic patients.
At a cognitive level this involves a period of inatten-
tion and sudden arrest of the stream of thought. In
the build up to such an episode Chapman [43] ob-
served patient difficulties in coordinating simple
motor sequences which are referred to as ideoki-
netic dyspraxias, that is, the patient knows what
to do, but is unable to do it. The blocking phenom-
enon commonly results in catatonic stupor and a
disruption of perception itself: You realize that
you are not actually seeing anything or hearing any-
thing. This state is often accompanied by persis-
tence of awkward postures and varying degrees of
tonic muscular contracture. There is a fundamental
link between selective attention, awareness and
motility which is explained under the premotor par-
adigm, and which can be interpreted in terms of a
marked reduction in the capacity to modulate tor-
que. The global cognitive deficit cannot be simply
attributed to a deficit in brainstem arousal mecha-
nisms and only bears superficial resemblance to
some types of epilepsy. Chapman [43] reported that
ten patients, most affected by blocking phenom-
ena, had a normal electroencephalogram (EEG).
781
Autism
Autism is a pervasive developmental disorder that
severely affects cognition and social behaviour.
The most consistent neuropathological abnormality
revealed by autopsy studies is a marked reduction in
Purkinje cells of the cerebellar vermis and hemi-
spheres [51]. Townsend et al. [52] demonstrated
spatial attention deficits both in patients with
acquired cerebellar lesions and autism. Their per-
formance on cued spatial detection and spatial dis-
crimination tasks was marked by an increase in
reaction times and reduced accuracy, respectively.
The attention orienting deficit in the spatial discrim-
ination task correlated inversely with the size of ver-
mal lobules VIVII. The authors conclude damage
to the cerebellum disrupts both the spatial encoding
of a location for an attentional shift and the subse-
quent gaze shift. Cerebellar lesions produce meta-
bolic abnormalities (crossed diaschisis) in frontal
and parietal cortex [53]. The empirical evidence de-
rived from autistic and cerebellar patients implies a
dynamic developmental role for the cerebellum in
the evolution of spatial awareness within cortical
networks, a key proposal of premotor theory.
Sensory processing impairments can be rather
severe in a subgroup of autistic subjects. Grandin
[54] describes the experience of Donna Williams
who can only use one sensory channel at a time.
When she listens to a friends conversation, she is
unable to perceive a cat jumping on her lap. An-
other autistic person describes that there were
no clear boundaries to anything. Personal ac-
counts of sensory overload or mixing of modalities
are well-documented [55]. Sounds which cause a
loss of orientation can be unpleasant or even
frightening. Jim says being touched on the face
causes some confusion about the precise location
and nature of the stimulus [55]. Sometimes he
knows that something is coming in somewhere,
but I cant tell right away what sense its coming
through. A component theory of perception
would predict poor spatial localization and thus,
the jumbling [54] of multiple sensory input.
All 17 autistic children recently studied by
Teitelbaum et al. [56] showed disturbances of move-
ment. Abnormal behavioural milestones were de-
tected in infancy from videotapes, prior to a
diagnosis of autism. A critical study of the specific
kinematic nature of the motor impairment of autis-
tic subjects was demonstrated by Schmitz et al.
[57]. An efficient electromyographic (EMG) pattern
underlying forearm stabilization of an external force
was comparable to a control group. However, during
voluntary unloading, unlike the control group who
782
showed anticipatory or feedforward postural adjust-
ment of flexor inhibition (mean latency 15 ms), the
autistic group showed a feedback mode of control
(mean latency +51 ms). This extraordinary finding
of an increase in the duration of unloading differs
qualitatively from the kinetic disturbance shown
by schizophrenic patients [49]. A component premo-
tor theory of perception is an attempt to address the
explanatory gap between the respective cerebellar
and basal ganglia dysfunction, motor disorders and
the higher cognitive impairments that are a corner-
stone of psychiatry. Attentional and language defi-
cits are major aspects of these disorders and Harris
et al. [58] propose a heuristic model of slowed atten-
tional orienting that may underlie social and behav-
ioural development in autism. For example, slowed
deployment of attention may interfere with learning
and the interpretation of rapid communicative
signals.
A testable hypothesis
Probably the most promising line of research in-
volves prism distortions where studies could be de-
vised to further explore the role of movement in
facilitating perceptual adaptation. In particular,
where movement is dissociated from visual input
akin to the kitten experiments. The animal experi-
ments could be extended to subhuman primates
where a more sophisticated analysis and the pro-
gress of visual development could be undertaken
using modern intracellular recording techniques
and scanning devices. This could be controlled for
in terms of an active or passive interaction with
the environment. A prediction of the hypothesis is
to look and the auditory cortex of the bat and
researchers should be able to draw a basic isomor-
phism with the visual cortex, assuming the bat
sees using sonar. I discuss What it is like to be
a bat elsewhere [16]. Such a demonstration would
vindicate the proposal that the structure of a sen-
sory hierarchy and its associated phenomenology
(in this case visual and sonar are proposed to be,
in many respects, equivalent) are directly deter-
mined by an organisms motor relation to the
environment. The importance of resolving the
mindbody dualism in a substantive biological the-
ory could lead to a greater understanding of mental
disorder with the prospect of better management
and the establishment of a research paradigm for
further effective treatment.
Conclusion
Vakalopoulos
Acts upon certain objects within a concurrent vi-
sual scene will have aspects of this visual informa-
tion that remain constant or, correlate with the
limiting features to the acts, and irrelevant infor-
mation with no bearing on particular acts. The lat-
ter will or, can change in the percept without
altering the motor parameters. The visual associa-
tions that limit or alter a set of motor parameters
are those that are ultimately categorized. Con-
versely, non-complete overlap of motor activity
defines categories of information. What the para-
digm resolves is that a percept associated with
any modality is coincident with boundaries to act
upon. These may be graduating, but there is a spa-
tiotemporal dimension relative to the organism
where there is an active change or removal of per-
ceptual context such that there remains a differen-
tial motor relation. Percepts present a textured
milieu that, according to the theory, achieves con-
sciousness only through a history of active engage-
ment. The premotor act defines its borders. It
categorizes quality by acting on changes in quality.
Its capacity to deal with a medium will ultimately
determine the sensory quality. However, categori-
zation is a historical process and not dependent on
any active motor contingency.
Acknowledgements
Thanks to David Darby (Mental Health Research
Institute) and Paul Marouf (Latrobe University) for
insightful discussions. I would especially thank
Jason B. Mattingley (University of Melbourne) for
his enduring support of the work and expert
commentary.
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