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Journal of Integrative Neuroscience, Vol. 13, No.

2 (2014) 363–402
c Imperial College Press
DOI: 10.1142/S0219635214400020

A®erence copy as a quantitative neurophysiological
model for consciousness

J. Integr. Neurosci. 2014.13:363-402. Downloaded from
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Hugo Cornelis and Allan D. Coop*
Three Way Street, P. O. Box 5160, Braddon
Australian Capital Territory 2612, Australia
[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 \a®erence 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 re°exive learning and recall processes. Based on
a review of various psychological and neurophysiological ¯ndings, we develop a framework
which contextualizes the model and brie°y discuss further implications.
Keywords: Consciousness; explanatory gap; a®erence copy; re°ex; 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: re°ex 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.


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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
su±cient 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 identi¯ed 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,

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1943); the stochastic Boltzmann machine, spin-glass latices, Ising spins (Hop¯eld,
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 di®erent 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
identi¯ed 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 unidenti¯ed 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 di±cult 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 trans¯guration of the philosophical
explanatory gap into a neurophysiological context. This allows the introduction
of a plausible \a®erence 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.



In proceeding, we aim to contribute to improved clarity of understanding by
re¯ning 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".

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2. Considerations Developed About Consciousness in Di®erent
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 (a®ect) 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 a®ectively 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 re°ection 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 \re°exive 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

Jackendor® (1987) considers consciousness is not a particularly high-level process. For personal use only. These have been de¯ned as the internal and subjective components of sense perceptions. 2013). 2010. The most basic emotions and arousal states are associated with internal feedback networks that guide an organism's behavior to the best possible outcomes.13:363-402. 2013). 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. One feature of experience putatively identi¯ed by philosophers is an insubstantial component of consciousness called qualia. Intentionality refers to the typical focus of consciousness being about objects or events (Edelman. evidence for this includes the desynchronized nature of the electroencephalogram. conscious life has two other dominant features: unity and intentionality. 2000b). The unitary nature of consciousness refers to the appearance of subjective brain experiences as uni¯ed. they are recognizable qualitative characters of the given. The cerebral cortex is currently considered to be the primary site containing the neural correlates of awareness (Mashour & Alkire.J. in humans at least. Such detail is likely captured at a resolution su±cient for the existence of physical phenomena to be demonstrated. Unconscious mental activity is therefore similar to all other natural processes (Solms. sound and a multiplicity of sensations to continual sampling of reality. as everyone has always wanted it to be. only becoming conscious as sensory percepts. 1982). 2002). such as in words and images (Gray. which may be repeated in di®erent experiences. 2005). At a psychological-level.. It has been well demonstrated that the operations of the CNS add color. 2003). with emotional (LeDoux. Smith. with most of mental life unconscious most of the time. 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. From a top-down perspective. This functionality is considered to underly essentially all behavioral choices in the vertebrate brain (Mashour & Alkire. 1996). in conjunction with limited connectivity with frontal–parietal areas (Mashour & Alkire. and are thus a sort of universal (Jackson. Allan Coop on 07/12/14. modeled and ultimately. 2014. This sampling is recorded with remarkable veracity from atomic and quantum domains (Kaupp. the painfulness of pain and so on. integrated and a constructed whole (Kandel. Downloaded from www. and that \it is not what makes us human". Besides subjectivity.. quanti¯ed. 2008). being central. by Dr. 2013).. .worldscientific. 1997). arising from stimulation of the senses by phenomena. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 367 radiation (Warren et al. activity in the thalamocortical system and widespread brain activity (Seth et al. Qualia involve the subjective experience of the redness of red. 1942). 2000) and semantic aspects (Chalmers. Hecht et al. 2013). 2002. Integr. The core of human consciousness is thought to have originated in phylogenetically ancient structures activated by primitive emotions mediating arousal. understood.

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. Space precludes further review of this extensive literature. Following a comprehensive by Dr. But Jackendor® (1987) considers this may not actually be the case.13:363-402. This perspective on human perception is also strongly supported by reports of detail blindness (Noë. Basically. For personal use only. change (Simons & Rensink. is usually su±cient to be believed to constitute thought. 368 H. CORNELIS & A. Hence. 1987). 1987) blindness. e. 2003). 2007). 1986. Importantly. that the unconscious brain is active. Many treatments of consciousness in the literature recognize. every image is forced to ¯t into a known percept (Snyder et al. at least incipiently (Jackendor®. . 1970). experience of \actual events in the now-situation" and concepts of the future or events which have not yet occurred. \No activity of mind is ever conscious .. learning or action. the stream of consciousness is essentially nothing but evidence that thought is occurring in a subconscious domain. the present and the future. O'Regan. For him. 1992). purposeful and independent and can selectively access and activate implicit goals and motives.J. Snyder & Barlow. Velmans (1991) concludes no human information processing is conscious in the sense that consciousness enters into or causally in°uences the process. 2005. Neurosci. thought.worldscientific. We are not conscious of the details that makeup a percept. Gregory. which are in turn generated from thought by mandatory fast processing modules. the fact that human cognitive processes are largely nonconscious has been periodically reported. Lashley (1956) further emphasized. \to what extent do events in the brain. Instead. although often ignored. processes are never available to consciousness in perception. the important question is. What enters awareness follows the processing to which that awareness relates. which we consider provides compelling support for our conjectures.Experience clearly gives no clue as to the means by which it is organized". Similarly.g. with both the process of thought and its contents inaccessible to awareness. 1998) and repetition (Kanwisher. that consciousness is projected from mental information structures. Raymond et al. where the stream of consciousness. However. take place at a conscious-(attentive) level or at a subconscious (pre. In answering this question. . Such details are inhibited from our conscious awareness. Jackendor® (1987) proposes that the character of the phenomenological mind is comprised of unconscious and conscious elements but that processing is always unconscious. COOP Ingvar (1985) considers conscious experience is based on three components: memory of the past. made up of visual and verbal imagery. 1998. Allan Coop on 07/12/14. related to the past.. visual and verbal images emerge from intermediate structures. and the attentional blink (Chun & Potter.or subattentive-level) or at both levels simultaneously". in short. Integr. . inattentional (Mack & Rock. 2014. Berlin (2011) concludes that complex cognition can proceed in the absence of consciousness.. D. Downloaded from www. 1995. Rather. 2004). what is seen depends largely on what is already known (Snyder.

Downloaded from www. 1970). AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 369 he notes that exactly how unconscious emotions and evaluations help shape the dynamics giving rise to conscious perception is still unknown. these ¯ndings argue directly against any immediacy whatsoever for the phenomenology of consciousness.. Perlovsky (2013) notes. 1934). have been reported. For simple re°exes in humans.. whereas. For personal use only. \recognition" and \choice" (Baayen & Milin. we are subjectively convinced of our consciousness. At the core of his model is a \comparator". brain imaging experiments have demonstrated that vague mental states and the entire dynamic logic process (taking approximately 500 ms) are unconscious. 2. When mismatch is detected between actual and expected states of the world. 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. 1986). (3) The control variables and set-points of the brain's nonconscious servomechanisms are juxtaposed. When there is no discrepancy. re°ex latencies for muscle in response to electrical stimulation range from 31–51 ms (hand) to 55–81 ms (foot) (Tarkka. the shortest reaction times are those of the unconscious blink re°ex with a mean latency of 10. \simple". this actually is not the case.worldscientific. combined and modi¯ed. in this way. which compares actual stimuli with expected stimuli    a function performed by a behavioral inhibition system (Corr. in a recent review. 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. 1943) and 155 ms (touch) (Robinson. attentional. Allan Coop on 07/12/14. while the mind operates by \jumps" among \islands" of consciouslogical states in an ocean of unconsciousness. by Dr. error can be corrected. 2008). 1996). 1973). (2) features that are particularly relevant to ongoing motor programs. recognition of familiar objects and scenes appears virtually instantaneous. Thorpe et al. in the following section (as one amongst several possible . are monitored and emphasized. Only ¯nal near-logical sensory percepts become available to consciousness. Similarly. with reaction times of 382–567 ms (median 445 ms. 20–40 ms (visual) (Marshall et al. For example. or which depart from expectation. the alien features of the error-triggering environment are subjected to controlled. We consider that on purely empirical grounds.J. In support of this position. response times of 8–10 ms (auditory) (Kemp. 2014. In short. behavioral routines run uninterrupted and stimuli are not extracted for detailed processing by higher-level cognitive processes. 2010).2.13:363-402. Timing of the phenomenology of consciousness Following Donders (1868). Although. Aru & Bachmann (2014) argue that attention and consciousness are independent from each other and phenomenal consciousness can emerge without attention. Most brain operations (more than 99%) are inaccessible to subjective consciousness. subjectively.8 ms (Shahani. Neurosci. three fundamental response latencies or reaction times have been con¯rmed. Further. analysis and (often) emerge into conscious awareness.

In vertebrates. any activity requiring detail or color (e. with processing requiring an additional minimum 50–100 ms (Salthouse & Ellis. Harris et al. Downloaded from www. 2008).. 1989). The duration of the minimum ocular pause time in the absence of stimulus processing is about 200 ms.3. However.5 ¯xations s 1 is about 15 min. duration and peak velocity (Becker. 1993). Although. 2014. 1974. For personal use only. for example. CORNELIS & A. in the visual ¯eld (Av ) and fovea (Af ): Nf ¼ Av ¼ Af 2   200  135 17. whereas.370 H. Reactive targeting saccades exhibit latencies of about 180 ms (Smit et al. exhibit remarkably stereotyped amplitude. the fovea may be compared to the location where axons converge to form the head of the optic nerve. saccade optimization probably occurs during development rather than being innate (van Beers. 1987). we further brie°y explore vision in more detail.. it is not surprising that an infant takes several months to develop. 2012). accurate hand-eye coordination. 2008) and the minimal ocular pause 100 ms (well below the 250 ms required for visual processing). equivalent to a 6–10 cm diameter disc at arm's length (O'Regan. the time required to scan the entire visual ¯eld at foveal resolution with 7. Integr. its temporal properties and implications. driving or reading) must be located within the foveal area by a saccade.worldscientific.g.. which at 3  –5  forms a surprisingly large \blind spot". 1981). The resulting hole in the photoreceptor mosaic is up to six times the foveal area. Typically.. 1947).. 2008). For example. D. as the dynamics of equivalent saccades vary across subjects (Boghen et al. Fischer et al. J. information may be extracted from foveal input within about 50 ms (Rayner et al. respectively (van Beers.13:363-402. 2004). Neurosci. by Dr. It covers about two degrees of the visual ¯eld or a disc about 4 cm in diameter at arm's length. A more physiologically realistic rate of three ¯xations s 1 would require up to 37. 1995. but may be as fast as 100–135 ms (Fischer & Ramsperger.. The fovea occupies less than 1% of retinal area but over 50% of the visual cortex (Krantz. 189 ¼ 6. Allan Coop on 07/12/14. 1984. Schmidt et al. 1992).5 min. 2004). 1979). In particular. 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. Given these ¯gures.. they last about 25–190 ms for movements of 5  –60  . memory-guided saccades may take well over 200 ms (Hopp & Fuchs.. with ocular pauses of 100–200 ms duration (van Diepen et al. COOP examples). what is . Their dynamics are determined by the point of origin within the visual ¯eld and the trajectory of movement employed to locate a target. 1980). Di®erent saccades (Hopp & Fuchs. 1988). or more precisely–learn. when reading. The problem of the rapid establishment of visual consciousness Principled estimates of the timing of visual activity can be made. 2. 750: ¼ 2 2:5465 22 If the optimal duration of a two-degree saccade is assumed to be 30 ms (van Beers.

percepts are evoked as required from the memory system. (2) in mammals. . given that for any one viewpoint this should take almost 40 min to establish. Many studies con¯rm that. but his further training continues every moment that his eyes are open. close to sensory threshold.. subjective timing of the experience is automatically referred backwards to the moment when primary sensory cortex received the stimulus (20–25 ms after . reaches this stage quite early in life. 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. For personal use only. separately for each grossly separate part of the visual ¯eld". Libet (2004) has proposed that following neuronal adequacy.J.. the di®erent perceptual qualities they generate are the constituent elements of the envelope of consciousness and nothing else exists (Solms. we learn to recognize the direction of line and the distance between points. In summary. the subjective timing of the experience appears to occur without the actual delay required for neuronal adequacy to elicit conscious experience of the stimulus. (Although. an explanation discounted by Libet (1981). A basic problem related to the conscious perception of stimuli and initiation of voluntary activity has been identi¯ed. Allan Coop on 07/12/14. 1964). it would seem rapid establishment of consciously experienced perception may actually be enabled by prior learning. Neurosci. The question is. Those of amphibians and reptiles lie between these two bounds. all sensory modalities are collected in the forebrain. Integr. .4. whereas. . Downloaded from www. As a solution to this problem. well-marked triangle to be established.13:363-402. The subjective antedating of perception Two main types of organization of speci¯c a®erent systems have been reported for vertebrates (Voronin et al.) The problem is that following neuronal adequacy. 2014. how is it so rapidly generated? A partial answer is provided by Hebb (1949) who suggests. and must extend his capacity for prompt recognition of patterns falling outside the macula. 1997). 2." There is every reason to expect that similar extended training occurs with other senses. by Dr. \during the continuous. stronger stimuli may reduce this delay to as little as 100 ms. \[I]t takes months for the ¯rst direct apprehension of a ¯gure such as a plain.worldscientific. 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. The normal human infant. 1968): (1) Individual sensory modalities in ¯sh partition to di®erent brain structures. following sensory activation. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 371 surprising is adults subjectively experience a very rapidly established high resolution global visual acuity in familiar environments. Regardless of how basic human sensory modalities are classi¯ed. The signi¯cant fact is that characteristic normal generalization only shows up after a prolonged and arduous training process. Further. intensive and prolonged visual training of infancy and childhood.

involuntary monitoring of emotional stimuli and re°exive boosting of perceptual processes may re°ect some \default" settings or intrinsic preparedness within neural pathways. the repetitive nature of a purely re°exive existence is transcended. 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. CORNELIS & A. Finally. 3. we consider there is su±cient empirical data to formulate several broad conjectures from which a framework for consciousness might be developed: (1) With the exception of innate re°exes (Zafeiriou. 2014. (2) Inhibitory processes are the preeminent mechanism whereby the CNS controls neural activity. D. content created and subjective experience expanded. non-conscious activity in the CNS is generated. Allan Coop on 07/12/14. the cumulative record or memory of all relevant previous moments. each present moment is the last remaining moment.372 H. From this perspective. Biological Framework for a Model of Consciousness Based on the foregoing. experienced not in its encapsulation of the past. LeDoux (1996)). 2004). In a review.worldscientific. Integr. Our claim is that through the physiological mechanisms supporting this model. Below. For personal use only. Further. J. the cumulative procession of known prior events and activities inform current experience and.5. we note Solms (1997) proposes that a®ect must be considered a sensory and perceptual modality. Emotion For completeness. Emotional stimuli may also evoke unconscious re°exive and involuntary processing under many conditions. 2. They can also be adaptively shaped by various regulatory mechanisms that themselves operate potentially with or without conscious control. The experience is thus \antedated" and subjectively appears to occur without the substantial delay required for conscious experience of the by Dr. see for example.13:363-402. but as an experience bounded prediction of the future. In the deepest sense. Each successive moment is formed from. organized and controlled through elaboration of re°ex circuits assembled by associative learning. It forms the basis of the \a®erence copy" model. via a surprisingly counter-intuitive mechanism. . we now make a few comments about the role of emotion in neural function. and as outlined in more detail below. Through these underlying mechanisms. (For more details. we provide a less paradoxical explanation. we present a model where unconscious precursors of the conscious experience of each given moment are molded within the memory system by integration with a®erent stimuli (including emotional a®ects experienced as qualia). Downloaded from www. lay the basis for future behavior. COOP peripheral stimulation). and experienced through. Neurosci.

Sechenov (1863) was the ¯rst to propose that all aspects of cognition in humans are based on behavioral re°exes or more generally that re°ex circuits provide the basis of all nervous function. 3. J. and evolutionary pressures drive increased capacity for anticipatory prediction. For personal use only. Let anyone who thinks this hypothesis is doubtful. Allan Coop on 07/12/14. the re°ex concept became basic for attempts to explain the physiological functioning of the nervous system. Early in the 20th century the re°ex arc was established as the functional unit in CNS integration (Sherrington. 1987).If anyone ¯nds a better explanation. and it was believed stereotyped movements. (4) Memory is a repository for learning and throughout life it is continually formed. 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". swimming and other locomotion. We now brie°y explore evidence supporting these conjectures. During the 19th century. arise from ongoing (prenatally initiated) re¯nement of learning. it has been recognized that \just as the spinal re°exes [bear] a `genetic' a±nity to the irritability of the simplest animals and of plants. by accumulated moment-to-moment sampling of all available sensory modalities. Neurosci.AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 373 (3) All CNS functions. and modulated. movements elicited through stimulation originating from an organism's external or internal by Dr. Integr.1. I shall be the ¯rst to welcome it". even poorly demonstrated. such as those involved in walking. By the end of the century the tendency was to include increasing parts of animal behavior under re°exive movements.13:363-402. . Thus. Magnus (1924) demonstrated the re°ex nature of all the elementary motor activities. that is. Since the early 19th century. From spinal cord re°exes to re°exive behavior The re°exive nature of nervous systems is an evolutionary requirement imposed by the need for control of reliable response in ever-changing and dynamic environments. \as far as the presence in man of three separate mechanisms directing the phenomena of conscious and unconscious life is concerned (viz. and those of re°ex inhibition and augmentation). Sequential patterns of e®ector activity were explained as arising from coordinated patterns of exteroreceptive and proprioceptive . or simply unacceptable. Downloaded from www. Laycock (1845) concluded that. Sechenov (1863) was convinced that. could be analyzed and described in terms of re°exes. \the ganglia within the cranium being a continuation of the spinal cord. controvert it . . 1906). For example. so the operation of the highest nervous centers in humans [retain] the fundamental re°ex character of the lower" (Clarke & Jacyna. 2014. (5) The physics of CNS function requires. from meaningful integration of sensory stimuli to consciousness and generation of motor plans.worldscientific. the mechanisms of the pure re°ex.

In summary. It has further been assumed that cerebral responses are . Beritov. 1967. a re°ex is an automatic or involuntary and near instantaneous response to a stimulus. 1965). 2009). The de¯nition of a re°ex has been quite variable. where the role of peripheral stimulation and sensory feedback is to activate and modulate these generators (Bullock. Thorpe. D. As McCulloch (1947) has clearly stated.. 1968). any system is part of the path of a re°ex". Downloaded from www. Subsequently. 1968). 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). Richet (1925) introduced \the conception of the psychic re°ex. for which feedback control is essential ( Zernicki. Integr. 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 a®erent nervous system ( Zernicki. \stimulus-evoked response that usually involves a single muscle or a limited group of muscles" (LeDoux et al.13:363-402. CORNELIS & A. it is possible to conclude a re°ex circuit may be activated by any su±cient by Dr. 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 short. obligate stereotypic behavior generated on the base of natural conditioned re°exes    in contrast to arti¯cial or facultative training–are components of speci¯c adaptive behaviors required under di®erent ecological conditions of existence. 1961). 1974). modes of functional organization of the cerebral cortex are re°exive: \To that great extent to which its a®erents inform it of the peripheral consequences of the action of its own e®erent. Further. 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. introspectively. Complex or chain re°exes located in the CNS have been demonstrated to consist of  a number of unitary processes. 1966. the re°ex concept was used to explain the integrative functions of the brain previously considered to result from \psychic activities" (Jørgensen.J. COOP excitation of the CNS. For personal use only. Thus. Importantly. \all neural processes (or neural responses) and following e®ector responses evoked by any currently acting stimulus". 2014. motor-related activity in the CNS was shown to not necessarily re°ect corresponding patterns of peripheral sensory activity. some stimuli produce psychic responses which may be assumed present in higher animals (Doty. to the more general. Allan Coop on 07/12/14. This is not inconsistent with pattern generators being viewed as examples of sophisticated re°ex circuits. 374 H. it was assumed animal locomotion and other types of movements are determined by central pattern generators. 1968). ranging from the restrictive.worldscientific. notes that complex. between the \instinct" and the \unconditional re°ex" there is a chain of stages dependent on either re°ex or the automatic nature of nervous system activation (Slonim. Neurosci. It is well-known in humans that. Thus. Slonim (1968).

\It is possible. Associated with the gastrointestinal tract. the brain of a student typist must coordinate sensory impulses from both eye and muscle to direct their ¯ngers to particular keys. the other a \peristaltic rush" which propagates rapidly. where one or more interneurons are interposed between the a®erent (sensory) and e®erent (motor) neurons of the monosynaptic circuit. it is comprised of a similar number of neurons as the spinal cord (Furness & Costa. with the di®erent psychic responses (perception vs. The student has \learned" to type. this seems to map well with the schemes of Tulving (2002) and Solms & Panksepp (2012) mentioned above.we can say that the nature of the given conscious act of man is the same as that of the re°exes". Downloaded from www. we cannot but see the laws of pure re°ex and cannot but consider them true . image) due to di®erent ways of exciting these neurons. However. 1911). Within it. Olds. Integr. LeDoux et al. 1956). typing has become a constellation of conditioned re°exes (Lagasse. For example. More recently. They include. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 375 manifestations of de¯nite unitary central processes or neuropsychic processes. one a slowly advancing contraction. respectively. muscles or brain centers. 2014. of which there are several kinds: e®ectively indi®erent or gnostic processes.J. \even those of the cortex of the cerebrum itself". Konorski (1967) hypothesized that both perception and image are manifestations of excitation of the same neurons. . gnostic processes mainly in associative cortex. . from the periphery and by association. because though we do not know what is going on in the excited nerves. Sherrington (1906) considered the whole nervous system to be based on this type of circuit. to speak of the paths of re°exes?" Only to answer. respectively. the circuits subserving the patellar or achilles re°ex. several types of neuromuscular activity are autonomously controlled by intrinsic sensory neurons (Bulbring et al. that is. stationary contractions associated with segmentation (Cannon. (2009) proposed that many behaviors fall into one of four categories: re°ex. Psychic responses may additionally be divided into perceptions and images. or unpleasant or pleasant processes that are negative or positive emotions. the ¯ngers automatically ¯nd and strike the proper keys even when the eyes are shut. for example.. and two types of re°ex (Cannon. 2000). and negative and positive emotional processes in di®erent places of the  limbic system and hypothalamus ( Zernicki. 1968. 1958). . Importantly. Once again.13:363-402. \how is it possible. Allan Coop on 07/12/14. 1980). reaction. For personal use only. (in the case of the sensation of red). where habit formation (the most complex) and much of learning are dependent on conditioned re°exes. Sechenov (1863) has asked. 1902). all but the simplest re°ex circuits are by Dr. The simplest neuronal circuits are the monosynaptic re°ex arcs composed of one sensory neuron and one motoneuron.worldscientific. After su±cient repetitions. Signi¯cantly. There is a good reason to believe neuropsychic processes are located in the cerebrum. An explicit example of the complex behavior generated by a relatively simple re°ex circuit is that found in the enteric nervous system. action and habit. Neurosci.

it is likely appropriate inhibitory processes and modulation of intrinsic and learned re°exes is fundamental to CNS function.e. are expressed as ongoing behavioral °ows generated in response to perceived environmental concerns. 1992). the known record of vertebrate evolutionary history (see e. In summary.. they are never all expressed simultaneously. \an entity in the world is re°exively experienced to be an entity in the world".worldscientific. Ultimately. In this domain.g. 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 re°ex pathways to formulate. as reviewed above. Phylogenetic elaboration of the CNS. Butler (2009)). we consider that a majority of fundamental CNS operations are essentially re°exive. Beritov (1968). control and sustain extended patterns of complex behavior.J. each appropriate to the given circumstances. Integr. However. Downloaded from www. results in both increased temporal separation between re°ex pathway activation and response. 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. For personal use only. 2013). COOP Numerical simulations have shown the circuit underlying this re°ex is su±cient to replicate both stationary and peristaltic activity. Allan Coop on 07/12/14. 2014. CORNELIS & A.. where subtle di®erences in circuit activation may evoke either ¯nely graded or signi¯cantly di®erent output. Rather. and thus. From this perspective. with (as we propose above) all such re°exive activity constrained to the subconscious. D. with the neuromuscular response determined by the state of intestinal contents (gas. i. Sherrington (1906) originally described the vertebrate nervous system as an orchestrated constellation of increasingly elaborated re°ex pathways culminating in a head ganglion or brain. sequences of behavioral patterns. particularly within associative cortex (Buckner & Krienen. This is the basic factor providing for the integrity of behavioral reactions. behavior and consciousness. Inhibition and disinhibition of re°exive behavior Typically. it is clear that in the normal course of events. Further. 1968). Here. . 2013). On the basis of the long tradition brie°y reviewed here. reports central inhibition caused by stimulation of sensory nerves embraces the entire neuroaxis and follows adequate stimulation of all sensory modalities. equivalent stimuli reliably generate equivalent responses. solid).13:363-402. modulated by circumstance. These results suggest that more complex re°ex circuits within other neural systems can reliably generate a range of seemingly plastic behaviors in response to the speci¯c details of circuit activation. liquid. by Dr. ultimately. and development of the associative circuits of the hominid nervous system (Buckner & Krienen. general central inhibition also occurs during deliberate cognitive activities.2. the details of the stimulus (Coop & Redman. it is their re°exive nature which renders their activity unconscious. 376 H. Further. a person might be considered capable of expressing a near in¯nite repertoire of behaviors. as Velmans (2009) has put it. In humans. and increased complexity of learned re°exive behavior. 3.

1991. Schacter et al. Integr. and how controlled processes are invoked only at critical junctures when a de¯nite choice must be made between cautious risk-assessment and prepotent responses.. may more immediately be controlled through disinhibition and e®ective connectivity.J. thought and deliberative action. 2014). A recent review of the functional signi¯cance of intrinsic oscillatory brain properties concluded. to produce relatively permanent changes in the behavior of an organism. but rather typically is an incremental process building upon. . \Inhibition . there is persuasive evidence the cortex may be under widespread and strong inhibition (Calford & Tweedale.c). study. what is already known.13:363-402. 1990) and somatosensory cortex (Xing & Gerstein. Dykes et al. and a role for inhibition in the control of cortical activity is further supported by disinhibition being a signi¯cant control mechanism in the spinal cord (Sechenov. . It has recently been reported frontal cortex mediates unconsciously triggered inhibitory control (van Gaal et al. 2004) and schizophrenia (Lewis et al. The possibility of inhibition was the possibility that past associations rather than immediate sensations would sometimes initiate movements. 2014. Allan Coop on 07/12/14. For personal use only. became the controlling mechanism that made possible those very functions. Neurosci. controlled/conscious e®ects come to determine automatic/nonconscious e®ects.. 2005).b. In this by Dr. including olfactory bulb (Cazako® et al. albeit with a time lag. Without inhibition. while pathological alterations to inhibition may lead to the excitatory dysfunctions of epilepsy (Avoli.worldscientific. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 377 such as arithmetic and suppression of conditioned and unconditioned re°ex activity. automatic-re°exive behavior will be more appropriate. . (2011) consider human learning to be acquisition of knowledge or skills through experience.3. appropriate adjustments can be made to the automatic system.. However.. We take this to mean that tonic inhibition at the physiological-level is responsible for repression in the cognitive and behavioral domains. 1863). \it is by selection. that gave humanity dignity and independence from nature. 2014). after ¯ne-grained analysis a®orded by control processing. Downloaded from www. 2008). 1984). learning is not usually immediate. 1996a. Learning and habituation In a recent review. and shaped by. striatum (Chevalier & Deniau. 3. At these moments. via inhibition that the most elaborate neuronal patterns are generated in the CNS" (Llin as. or by being taught. 2004. We have previously concluded that. It was the possibility that sensation might combine and recombine as intelligence and thought. Coop & Redman. ¯ring of hippocampal pyramidal and cerebellar Purkinje neurons (Coop & Reeke. As Smith (1992) has concluded. than alterations in a®erent stimulation frequency. in the electrophysiological domain. 1998). such that when the same (or similar) stimuli are encountered in the future. Gray (2002) considers that the inhibitory function of consciousness solves a major evolutionary problem: How to ensure that automatic responses are appropriately activated. sensation merely exhausted itself in movement". More generally.

learning may contribute to understanding.. 1927) and operant (Thorndike. The unity of fragments of memory The recollection of past experience is considered to be a reconstructive process. Even at this early stage. D. Associated learning may occur consciously or unconsciously. 1968). COOP Two types of learnings have been distinguished (Kandel et al. Allan Coop on 07/12/14. Downloaded from www.worldscientific. In other words.4. It is notable that at the electrophysiological level. the e®ects of internal stimuli have been shown to be active as early as the embryonic stage. Two forms of associative learning have also been identi¯ed. . although little is known about the neural correlates (Hassabis & Maguire. Indeed. classical (Pavlov. habituation (Sokolov. 1997). . when objects in the environment occur frequently enough. notions providing.e. 2000a). but always by means of consecutive re°exes. and as Sechenov suggests. even qualia (where qualia have been proposed to originate in local cortical network activity (Orpwood. For personal use only. 2014. \the successive re°exes acquired by learning lead to a perfect notion of the object. i. central inhibition displays its controlling in°uence on coordinating re°ex activity (Biryukov.I shall merely point out that sensations from all spheres of the senses can be diversely combined. 1911) conditioning. However.13:363-402. Integr. CORNELIS & A. the sum of all possible sensations evoked in us by these objects under all conceivable conditions .J. so to speak. 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. they can be incrementally added to the memory of similar previous encounters. As suggested by Sechenov (1863). 378 H. Two forms are common. As such. 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.. 2013)). this continually enriches records of sensory experience and thus conscious phenomena. to knowledge in its elementary form. (2007).. 2014). . for convenience we use this phrase to refer to the more state-based or unitary memory model described by Brown et al. with memories recreated from their component parts. 2009). Slight variations over time elaborate the memory system far beyond that of any single sensory experience. 3. Neurosci. It can be broken down into a number of constituent processes. 1963) and sensitization (Shettleworth. For example. where \memory system" refers to the multistore memory model (LaRocque et al. the scienti¯c knowledge of external objects is simply an in¯nitely broad notion of each of them. 1968). These combinations give rise to countless notions that arise in childhood. knowledge and as we have seen. material for the entire subsequent psychical life". with repeated by Dr. The simplest is non-associative.. and there is evidence for prenatal habituation in humans as early as the 32nd week of gestation (Sandman et al. 2010). particularly in studies of conditions under which the onset and patterning of electrical activity takes place in the higher parts of the embryonic brain. indicating the nervous system is developed for learning and memory formation prior to birth.

e.J. This implies forgetting is a consequence of reduced local distinctiveness. This multi-store modal memory model proposes sensory memory containing short-term records of sensory stimulation (250– 500 ms duration). the basic idea is that items are more distinctive. For example. through memory resulting from both real and imagined experience (Bar. from milliseconds to years.g. For example. Subsequently. learning occurs through general mechanisms of experience-dependent synaptic plasticity. which ultimately lead through a cascade of intracellular molecular mechanisms to the formation of long-term memory. a memory is then embedded and embodied through its ¯xed trace. 2007). . For over a century. short-term memory is associated with a working memory (Baddeley & Hitch. 1972) components. Those stimuli which are attended enter short-term memory (up to 18 s duration). AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 379 experience (Atkinson & Shiffrin. Downloaded from by Dr. the same \core network" of brain regions is recruited when people remember the past and imagine the future (Buckner et al.worldscientific. One signi¯cant problem with this multi-store approach is that di®erent principles are assumed to apply over di®erent time scales. (2007) have proposed memory is unitary over all time scales. while long-term memory may be divided into either explicit memory which is partitioned into episodic (Tulving. (1977). 2014. 1962) and semantic (Tulving. comprising implicit or procedural memory (Schacter. Integr. This runs counter to the generally held expectation that scienti¯c principles should hold over a wide range of temporal. Schacter et al. Here. Thus.13:363-402. it has been shown memories can temporarily be rendered labile and sensitive to modi¯cation. 2009b).. 1968). the same mechanisms are used for retrieval over all time scales. 1996). Hubel et al. and hence both more memorable and easier to identify. it is possible to enhance and even incorporate new content (for review. or physical scales (Barenblatt. However. Importantly. to guide behavior in the future based on analogies.. As a consequence. 1987) or emotional memory (LeDoux. a signi¯cant function of memory is its role in allowing individuals to imagine possible future events (Schacter et al. to the extent they are located in sparsely-populated regions of psychological space. Brown et al. not trace decay. 2000). 2007). 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. Once consolidated. Neurosci. procedural (Milner. see Flavell et al. 2008. Each of these memory domains has subsequently been further decomposed. This so-called temporal ratio model is concordant with the synaptic trace theory of memory ¯rst proposed by Hebb (1949). 1974). (2014)). spatial. It has recently been shown that similarities in cognitive processes underlying past and future events are complemented by analogous similarities in brain activity. Allan Coop on 07/12/14. 1972). psychologists have focused their studies on memory of the past. For personal use only. He posits the brain retains information through learning-induced changes in the synaptic connections between neurons. or declarative memory. 2008. Once an existing memory has been destabilized.. Schacter & Addis (2009) consider that predicting the future and remembering the past may be more closely related than everyday experience suggests. and if su±ciently rehearsed are transferred to long-term memory (in principle unlimited).

It is distinguished by originating entirely within the memory system.13:363-402. The consequent mode of operation derives from a \look-ahead function". D. It is widely believed. Allan Coop on 07/12/14. Integr. the extent to which sensory cues a®ect brain function is determined by their impact upon pre-existing functional dispositions of the brain (Llin as. memory can be thought of as a tool used by the prospective brain to generate simulations of possible future events. cognitive states or entities. For personal use only. We extend this approach to provide a signi¯cantly more comprehensive predictive functionality which lies at the heart of the a®erence copy by Dr. . Llinas (2001) considers this indicates a far deeper issue than might initially appear. and provision of a basis for conscious temporal detachment. 2009). 1987. past and future events draw on similar information stored in memory and rely on similar underlying processes: in an active creative extrapolation. (2008) further suggest that simulation of future events requires a system that can °exibly recombine details from past events. such functionality is memory-based (Bar. For the nervous system to predict. Schacter et al. It yields predictions of the frequency-time function of cortical input and is an emergent property of inherently parallel distributed neural circuits. In this view. Thus. some do so at intermediate velocities.380 H. the output of such a circuit will be a reconstruction or prediction of an event ahead of its time of completion. 1974). Schacter et al.5. In other words. The signi¯cance of such a mechanism is found mainly by its incorporation into larger. 2001). In fact. in the absence of any other possible sources. Pellionisz & Llin as (1979) provide a model of such a function based on the Taylor expansion. 2009a). Neurosci. It cannot be stuck doing something when required to perform another task. moving system is enhanced by such innate functionality (Llinas. memory supports the construction of future events by extracting and recombining stored information into a simulation of a novel event.worldscientific. 2001). (2007) consider such a hypothesis requires a shift of conceptual emphasis with regard to the role of memory in cerebral activity. and some measure events in real time. 3. proposed to be an inherent property of neural circuits (Llin as. Downloaded from www. 2014. COOP J. Prediction as reconstruction of the future Prediction is pervasive throughout much of brain function and is almost continually operative at conscious and re°ex-levels. If some neurons respond quickly. an absence of immediately direct comparison between sensory-referred properties of the external world and internal sensorimotor correlates. CORNELIS & A. 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. as success in a goal-oriented. If it is to synchronize with the external events of each given moment the brain must leave itself enough time to implement movement decisions. the predictive abilities of the brain may be profoundly more fundamental than suggested by a purely look-ahead model. It relies on relationships between neuronal ¯ring rates and events in the external world.

there is a considerable literature reporting estimates of the transmission rate of information by neurons within the CNS. physiological processes of a®erence copy are evoked as sets of embedded re°ex functions. 1995). With regard to content manipulation. 2014. with the same distribution of neurons between its cerebral cortex and cerebellum as in other species. 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. In short. One way to quantify these domains is in computational terms. where N gives the number of spikes per second. information per spike increases to 3–4 bits spike1 (Softky. an estimate of its \storage capacity". The aim is to give some sense of the putative capacity of human memory in computational terms. Integr. for a rate code with strictly periodic neuronal ¯ring. the information per spike is calculated from log2 ðN Þ=N . here we are initially interested in the content storage domain of the memory system.AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 381 In fact. in particular. the calculation of information transmission becomes considerably more complicated (e. Alternatively. with a theoretical maximum of about 9 bits spike 1 (MacKay & McCulloch.6. better estimates will be considerably complicated . These may be distinguished from the content manipulation domain. for example. 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. Theoretically. For personal use only.13:363-402. We note. 3. In concordance with our identi¯cation of a putative a®erence copy system. Based on this assumption we can employ the following data to make an \order of magnitude" estimate of storage capacity. a synapse can be represented as a single binary bit. 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. They lead to the creation of temporal experience and consciousness through their ability to fabricate the phenomena of the present moment. 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 . where the variability of spike timing is taken into account. 1952). These estimates are typically based on an assumption of either a rate or temporal code. J. Estimates of memory capacity and rate of formation Unquanti¯ed 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. for a temporal code. Downloaded from www. Neurosci. see Stein (1967)).g. 2011. Allan Coop on 07/12/14.. we claim that through infant learning. along with appropriately scaled energy costs (Herculano-Houzel. 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. However.worldscientific. Such an estimate then allows a \mean acquisition rate" to be determined for the content acquisition by Dr. For example. 2012). respectively.

it can be estimated there are on average about 358 \naive" synapses available each millisecond to record the state of the nervous system. 2005). 1993. 2. COOP by. neuronal loss due to ageing in the frontal lobes (up to 20% (Masliah et al. Allan Coop on 07/12/14.5 Mb minute 1 .e. and all synapses may be modi¯ed by experience.. elementary concrete knowledge". If it is assumed on average a person spends approximately 8 h sleeping each day. We appreciate such a \literary" de¯nition is a necessary step in problem identi¯cation. From a computational perspective.. corresponding to storage throughout waking life of 57. Gibson. 1963). this number of synapses corresponds to a storage capacity of about 90 Tb. 2003). D..5 Gb day 1 . which is considered to be reserved for the stabilization. in all spheres of the senses the child acquires a multitude of more or less complete ideas of objects. Assuming a human lifespan of 70 years. This daily value is comparable with high de¯nition movies which consume approximately 2 Gb hour 1 . CORNELIS & A.13:363-402. this corresponds to storage of 44 bytes ms 1 or 43 Kb s 1 . Downloaded from www. 2009).com by Dr. but consider it a misplaced expectation that any deep resolution is possible while explication languishes in such purely metaphorical or narrative domains.3 Kb s 1 or approximately 200 Mb hour 1 . 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. at least. for example. enhancement and integration of memories (Walker et al. 2014). the cumulative capacity is substantial. The human brain has been reported to contain 86:1  10 9  8:1  10 9 neurons. . 2014. 4. For personal use only. These ¯gures may signi¯cantly increase if it is accepted that new memories cannot be formed during sleep. the hippocampus (Altman. . a 90 min movie has a similar storage requirement to our estimate of the average daily human storage capacity. As Sechenov (1863) has proposed. Integr. the total estimated number of synapses in the brain may be as large as 7:89  10 14 . of which 19%. 150 Mb hour 1 or 3. Neurosci. 1969) and striatum (Ernst et al.. olfactory bulb (Altman. then memory storage capacity may increase up to about 470 synapses per millisecond. 1983)). Assuming non-cortical neurons exhibit a similar average number of synapses as cortical neurons. 382 H. Interestingly. The number of synapses in the neocortex has been reported at 1:5  10 14 (Pakkenberg et al.worldscientific.. i. with the knowledge there are approximately 3:2  10 10 ms in a year. 1983). From a computational perspective. by this analysis. At one bit per synapse. as our calculations show. Although the putative per millisecond synaptic sampling rate seems trivial. and likely continual addition of neurons in. or approximately 16:4  10 9 are cortically located (Azevedo et al. Any solution is likely only available following location of the \gap" within an appropriate . 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..J. \By means of absolutely involuntary learning .

For this reason. Neurosci. However. Allan Coop on 07/12/14. which we initiate through the innervation of our motor nerves. Most importantly. we consider the problem of the explanatory gap becomes tractable when translated from a philosophical to an empirical context. This is an internal copy of an . because of the highly multidimensional. itself a characteristic of the scienti¯c method. in this case. 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. As suggested in the Introduction. 5. in the case of placental mammals. To make this transfer we propose. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 383 neuropsychophysiological context. As reviewed by Mulliken & Andersen (2009). Downloaded from www. More speci¯cally.13:363-402. a signi¯cant evolutionary advantage likely accrues to an organism capable of correctly predicting evolution of the environment during this period. This transformation recasts the problem of the explanatory gap by removing it from a philosophical domain of narrative contemplation to the domain of empirical quanti¯cation. 1878/1971). 2014. 1954. Sperry. at its most general. More speci¯cally. \the temporal duration from the time of interaction of a su±cient sensory stimulus with a receptor until the time at which a percept of that event is subjectively recognized within conscious experience". E®erence Copy as Motor Predictor Model Modern e®erence copy models (Fig. As we show below. intangible and generally elusive nature of consciousness many of its various aspects are only available through metaphorical language and description. because sensory information is substantially delayed. we recognize that with quanti¯cation comes reduction. it has been proposed the brain makes use of an internal forward model which relies on e®erence or e®erent copy. Integr. during this period it is not possible for a nervous system to know anything about changes in the external environment. the philosophical idea of the explanatory gap can be placed ¯rmly into the core of the scienti¯c tradition. It is this gap which must be \explained" by the nervous system and to which.worldscientific. in its own way. is immediately perceptible". aspects of the philosophical formulation actually refer.J. This idea was later integrated into motor physiology through \corollary discharge" (von Holst. 1(a)) derive from a proposal which originally posited direct sensation of the motor command: \The impulse to move. Quanti¯cation is fundamental to the scienti¯c method and by making this conceptual transformation. \bridge" the explanatory gap. It is these aspects which are greatly reduced by transference from the philosophical to the physiological domain. 1950). In doing so we acknowledge the metaphorical nature of much written about consciousness. (von Helmholtz. There it can be rigorously studied as an incontrovertible feature of the vertebrate by Dr. For personal use only. motor or otherwise. the foregoing de¯nition might be recast as. that the physiological explanatory gap is de¯ned as the gap between the occurrence of a stimulus at externally or internally directed sensory receptors and the initiation of a response.

copies of e®erent motor information are fed back and used centrally in an emulation algorithm. 2012).worldscientific. Thereby. this physiological explanatory gap is bridged by conscious experience. COOP J. 1. A. 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.. which originates through a®erence copy as a construct evoked from within the memory system. Otherwise. surprise is experienced and attention may be redirected. Upon e®ective execution. It also allows perceptual structures to distinguish between self and externally mediated signals. Once integration is complete. Our claim is. The special character of this consciously experienced percept is that it is not actually just a record of sensory activation entering the nervous system. B. the joints and the skin generate actual proprioceptive feedback. which calculates the anticipated somatosensory changes expected as a consequence of the planned motor execution. receptor stimuli are fed forward and maintained subconsciously in the memory system while they are integrated with the relevant memories they evoke. (a) (b) Fig. 6. peripheral changes at the level of the muscles. the actual proprioceptive feedback of a motor action e®ectively balances the predicted sensory feedback. Rather. D. movement-producing signal generated by the motor by Dr. out°owing. 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 . If the a®erence copy e®ectively matches the environment. E®erence Copy: Model proposes that upon motor preparation and intention. predicted experience and actual experience match su±ciently that no further action is required. which will e®ectively balance the predicted sensory feedback in somatosensory cortex (at the level of a so-called \comparator").13:363-402. 2014.384 H. Integr. Downloaded from www. A®erence copy: A model complementary to e®erence copy proposes that upon sensory stimulation. CORNELIS & A. For personal use only. the e®erence copy is an early-warning signal sent by motor production areas to the corresponding somatosensory areas specialized in the proprioception of motor execution. Subsequently. A®erence 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. Neurosci. Panel A adapted from (Bazan. a prediction of the percept expected on the basis of the received sensory stimulus emerges as an \a®erence copy" into the explanatory gap created by the time taken for integration to occur. Allan Coop on 07/12/14. 2000).

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. It is in this way the explanatory gap is bridged and conscious experience generated. This situation is only compounded by the fact that each successive moment is also a past moment prior to being experienced as the present. Allan Coop on 07/12/14. subjectively experienced as the present moment. This creates a signi¯cant problem as retrieved content must be collected and displayed somewhere. At any one moment. Importantly. movement-producing activity of the motor system. Following. The a®erence 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. Ultimately. 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.worldscientific. This process is illustrated and explained in more detail in Fig. 1(b)). in the normal course of events. Nevertheless. we consider \monitor-and-act" is a more complete descriptor than the notion of representation as the former incorporates both \memory storage" and \processing". The model resolves this apparent con°ict in the following way. Neurosci. 2. As they are learned. as is the content of the behavioral repertoire it ultimately subserves. . the construct fabricated by a®erence copy processes would be based on. it is clear the neural activity driving a®erence copy is widely distributed.13:363-402. A®erence copy can be conceptualized as complementary to the e®erence copy system (Fig. ongoing neural activity is modulated as required by sensory content and the associations it evokes from within the memory system. for example. the \past" due to the temporal delay introduced by the neural activity enabling its conscious appearance. and exist in. the processes subserving a®erence copy. with the complexity of the processes involved being a major reason for the extended developmental period seen in humans. although the details of precise location and mechanism are as yet unclear. Integr. Downloaded from www. Importantly. entry of sensory input into consciousness is delayed while its e®ects 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. but applied to in°owing sensory activation rather than the out°owing. For personal use only. Attempts to resolve this additional complexity has spurred previous solutions resulting in homunculi and the mechanics of the Cartesian Theatre. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 385 integration of the given sensory input is complete. following Nikolić (2014). by Dr. become a \re°exive" mechanism residing in the subconscious. Dehaene & Naccache (2001) and Gray (2004).J. we further claim that the ability to do this is learned. It is a re°exive mechanism learned during development to accommodate ongoing function in the ubiquitous presence of the explanatory gap. The content of consciousness is this memory-based prediction of the composition of the next actual future moment. similarly to other learning. It is in this sense that a®erence copy is a predictor of sensory experience.

386 H. The large lower arrow indicates the ongoing evolution of predicted perceptual experience within the conscious domain. CORNELIS & A.13:363-402. Predicted percept) and subjective time (Past moment. but for clarity not shown in 2B). Neurosci. Consciously experienced subjective phenomenology is a product of a previously constructed prediction of the experiential contents expected to occur during the present moment. 2. An upper physical domain contains a stimulus originating externally to the nervous system. 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). a sub/pre-conscious domain (top) containing the memory system (comprising sensory. 2014. Downloaded from www. Present moment. A lower cognitive domain (within the nervous system) is further subdivided into two \internal" domains. Framework for consciousness: The schema is given for a single external stimulus.worldscientific. Model features are given at the top (A. Integr. D. During the Explanatory gap (2A). 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). Upper horizontal dotted line distinguishes two domains within each panel. For personal use only. COOP Fig. and a conscious domain (bottom). Panel 2: Illustrates temporal evolution of the relationships given in Panel 1 into the next moments. 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. Allan Coop on 07/12/14. short and long-term memory). . Explanatory gap. It is in this way ongoing perceptual experience is continuously generated through a®erence copy to bridge the sliding window of the explanatory gap.J. The large upper arrow head indicates appropriate ongoing subconscious elaboration of memory by incorporation of correctly predicted perceptual components. C. 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 by Dr. Future moment) indicated at the bottom of each panel for three consecutive 550 ms moments (separated by dark vertical bars). 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.

2014. overall goal. but in principle. For personal use only. consciousness can be seen to act as an interface between the a®erence and e®erence copy systems. Naturally. Their key characteristic is activity modulated both by the execution and observation of action (di Pellegrino et al. 1968). Mirror neurons as recruiters of predictive percepts In humans. relative distance. 7. 1992). and occurrence is matched to performance. it can be expanded to any degree of complexity. viewpoint (whether visible or hidden). if the explanatory gap is bridged by a predictive construct fabricated by integration of sensory stimulation into the memory system. the elements available for fabrication create e®ectively \seamless" and entirely \resolvable" percepts. Evidence for a signi¯cant claim of the a®erence copy model may be found in the behavior of mirror neurons. Historically. Allan Coop on 07/12/14. Based on these considerations. which subsequently emerges as conscious experience. Here. This is followed by comments on evolutionary implications of the temporal extent of the explanatory gap. When an observed activity is internalized by learning.1. 2013). mirror neurons have been proposed to play a signi¯cant role in social cognition (Heyes. we ¯nally and brie°y discuss a putative origin of qualia in the vertebrate CNS. thereby integrating and embedding a nervous system into the spatiotemporal and physical °ows of its by Dr. neurons in the memory system associated with previous views are the repository of the memories from which appropriate predictive percepts may subsequently be recruited. reward value. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 387 physiological process of creating a prediction of the experience of the present moment based on prior experience of the given situation or environment. action performed in different contexts and various combinations of single or dual hand and mouth actions (Kilner & Lemon. 2010). we propose their properties provide further compelling evidence for the a®erence copy model. the subjective elements of a reproduced image are in general identical with those of the perception. 7. 1966) and atoms (Green. when viewing a given activity. Integr. 1976). We then employ the model to explore how the low rate and high variability reported for cortical neuron ¯ring may originate. Neurosci. In short. environmental needs and requirements create circumstances considerably more complex than the basic process of a®erence copy described here.. activity performance may be mapped to the .worldscientific. they have been reported to respond to the sound.J. Amongst other properties. From this perspective. The processes of perception and image reproduction are identical in their essence and their structural and physiological bases absolutely equal (Beritashvili. 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. Downloaded from www. with the image of a perceived object usually produced immediately during ¯rst perception. Discussion We begin this discussion by describing an expanded role for mirror neurons within the framework we have developed here.13:363-402. then.

Here. with practice. J. the more likely fabricated predictions of a®erence copy will match what occurs. thus better supporting an increased explanatory gap with its attendant consequences. ganglionated plexuses of invertebrates. and the e®ect adolescent game playing in virtual environments may already be having. COOP neurons that recognize occurrence as to the neurons involved. 2000) and 150 ms (Blana. the tension between rapidity of re°ex response and selection of an optimal behavior from an ever increasing repertoire. 2014)). It has not escaped our notice that consideration of the a®erence copy model we have postulated here suggests a multiplicity of novel features of considerable .worldscientific. respectively. Integr. driving and °ight control start to be seriously impacted by delays of 50 ms (Kemeny. This suggests the current hominid nervous system is capable of supporting an explanatory gap of at least 770 ms (550 ms (Libet. with actual duration likely determined by the requirements of task complexity. Concomitant with competitive evolutionary pressures. myelination. Here. 7.. It is clear that this development has at least two highly signi¯cant evolutionary rami¯cations: (1) The longer the duration of the explanatory gap successfully bridged by a®erence copy. by Dr. 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. 2004) þ230 ms (Cunningham et al. 1996).b)). Allan Coop on 07/12/14. is completed.13:363-402. Temporal duration of the conscious moment Evolutionary centralization of the vertebrate nervous system creates a problem not faced by the decentralized.2.388 H. 2001a. In brief. in this way a®erence copy can connect observed with performed activity to con¯rm a given behavior has actually been implemented. Neurosci. CNS elaboration allows for increased behavioral choice. (although controllable in part by ¯bre diameter. for example. the larger the amount of subconscious processing that can occur. the lengthening explanatory gap has necessarily required an extended a®erence copy mechanism. the behavioral loop established by e®erence copy and its con¯rmation of successful motor plan execution. as the vertebrate CNS has evolved. operators can adapt to a constant delay of up to 230 ms with no signi¯cant e®ect on performance (Cunningham et al. D. (2) Alternatively. Although. and spatial recon¯guration) the time between stimulus and response must increase due to increased system size and complexity (Biryukov. increased consciousness. familiarity and response. Similar to e®erence copy. It is interesting to speculate on the future computational and digital roles which might support expansion in this domain. 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. we note two provocative implications related to duration of the explanatory gap. Finally. 2014. they are one and the same event. i. The inevitable consequence of this is predicted to be a concomitant increase in consciousness. the greater the stability of an environment. For personal use only.b). CORNELIS & A. With increased evolutionary elaboration of the CNS.. Thus. 1968). there is one question. in this section. Downloaded from www..

Integr. 7. thus subconsciously. it assists either in the rapid extraction of certain types of subconsciously \processed" automatic re°ex activity subsequently subject to higher-level cognitive analysis. J. 2009). It is through the involvement of hormones and neurotransmitters . 1981) by Dr. 2011). A®erence copy and neural ¯ring variability In the intervening period of the explanatory gap. 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). particularly when. the physical events of sensory receptor activation elicit short-lived cognitive. Allan Coop on 07/12/14.4.AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 389 theoretical and practical interest which may also exhibit signi¯cant ethical and. in conjunction with disinhibition. and in keeping with the requirements of selection from a behavioral repertoire. or provision of a basis for mismatch error signals between expected and actual states of the world (Corr. Downloaded from www. political repercussions. 7. In such an environment. moment to moment sensory changes may recruit the same cells but con¯gured. Neurosci.13:363-402. Qualia as emotional states re-evoked by a®erence copy Along with activation of neural circuits. the \behavioral inhibition system" (Gray. The actual ¯ring time of such neurons may be deterministically controlled by their precise contribution (Reinagel & Reid. for example. As an alternative explanation. One mechanism suited for this purpose is the comprehensive central inhibition exerted by. for example. irrespective of its duration. thus. 2011). For personal use only. integrated into the memory system prior to entering consciousness within a fabricated prediction of future perceptual experience. physiological and somatic reactions. with all emotions having some kind of regulatory role for neural activity within the nervous system (Damasio. 2014. 1972. In these circumstances. The role of such inhibition can be signi¯cant. Emotions are the product of complicated collections of pattern forming chemical and neural responses. the same cortical neurons may contribute to the tracking and prediction of subtle modulations of experience with signi¯cant variation in ¯ring time. to form di®erent functional circuits as different aspects of requisite memory components are integrated to fabricate the constantly changing detail of relevant percepts. particularly as the required memory content must simultaneously be recruited and integrated to form a dynamic and uni¯ed conscious experience. it is likely valuable to minimize generation of spurious neural activity or \noise". it is likely that considerable neural activity may be necessary. 1999). otherwise known as emotions (Kensinger. This allows minimal impulse generation within widely distributed circuits. If a®erent sensory stimulation is re°exively. we propose that during the explanatory gap only necessary disinhibition occurs. 2002) and e®ective connectivity (Friston.worldscientific. which may both explain these data and provide direct evidence for the neural activity that might be predicted during the process of a®erence copy.

D. determined when the visual system integrates responses across populations of photoreceptors (speci¯cally.J. color is a property of the brain (Zeki. noradrenaline. the interaction of light with photoreceptors in the retina results in the generation of patterns of neural impulses re°ecting the combinations of wavelengths to which they by Dr. blue cheese. with feelings. The surprising result is that what is . qualia are those \remembered" emotional states associated with previous experience. maintained and reinterpreted. are cumulatively recorded and updated. Downloaded from www. as relevant memories are evoked through a®erence copy. oxytocin. CORNELIS & A. for example. If the ¯rst pu® of a cigarette is the only indication of future enjoyment.13:363-402. before the intensity of their appreciation (whether innate or acquired) can be enriched through re-exposure. This may provide an explanation for why little is ever as \good" or \pleasing" as the positive memories of the past. In the world. It should be no surprise that moment to moment records of sensory experience consequent upon activation of all sensory modalities. the waveforms and quanta of light are comprised entirely of their own particular physical structures and relations. 2014. 1997). Thus. 1968). For example. by Proust (1913/2000). wine or true love must ¯rst be experienced. taste of an apple. it is only to be expected that according to this model. It is the mechanism underlying the powerful nostalgic attractions of reminiscence as explored. It is why olives. Similarly. For personal use only. Integr. Here. 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. a physiological mechanism is proposed whereby this gap may be bridged.5. cortisol and GABA.worldscientific. Any further properties they acquire are due to interactions outside of their own structural features. Following the moment of occurrence. associated emotional a®ects emerge as part of each fabricated percept or experiential prediction. 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. It is delimited in the neurophysiological domain by an \explanatory gap" which is a trans¯guration of the widely explicated philosophical explanatory gap. smell of a fresh tomato. including emotional a®ect (Solms. serotonin. are never as comprehensively complete or emotionally intense as the enriched multiply re-experienced history of such sensations as they accumulate in memory. COOP such as dopamine. re-evoked from memory during the process of a®erence copy. the consequences of sensory stimulation are embedded within a complex ongoing °ow of internally generated emotional states. Conclusion We have presented here a framework for consciousness. Vice versa. the frying of bacon and eggs or the glimpse of a childhood sweetheart. Neurosci. how could one ever proceed? 7. at some later moment. 1993). 390 H. Allan Coop on 07/12/14. by the memory system (Beritashvili. di®ering absorption spectra recognized by three types of cone cell). In brief. this mechanism may also explain the severe ongoing e®ects of trauma. The immediate \redness" of a rose.

allows temporal decoupling by the neocortex of innate and learned re°ex activity (as required) from immediate behavior.13:363-402. Cell proliferation and migration in the anterior forebrain. Consequently. organize and thus more e®ectively employ. J. then whatever remains. Comp. The implications are complicated and diverse. (1972) The Metaphorical Brain. This leaves us with only one certainty. Acknowledgments The authors would like to acknowledge Jan Braeken for his valuable discussions about this paper. to emerge as the subjective conscious experience of the \present" moment. It takes remarkably little e®ort to determine that much of what is believed about ourselves and the world is merely convenient ¯ction. Neurosci. embedded within and directly linked to.. the physical constituents we are individually composed of. AFFERENCE COPY AS A QUANTITATIVE NEUROPHYSIOLOGICAL MODEL 391 experienced as subjective consciousness is a learned predictive construct fabricated from the quantal contents of a cumulative memory system. the a®erence copy proposed to support this fabrication comprises a model that may provide considerable insight into the neurophysiological underpinnings and mechanisms of consciousness. 7. The shoulders from which further details of the generation and subjective experience of conscious phenomenal content might be viewed are. (1969) Autoradiographic and histological studies of postnatal neurogenesis. REFERENCES Altman. as by Dr. 137.worldscientific. Integr. Neurol. Anat. Allan Coop on 07/12/14. a su±cient fragment. . Arbib. 433–457. As for the mechanisms generating and framing consciousness. Altman. & Bachmann. for example the result of Libet (1985). This occurs through the subconscious inhibitory modulation of otherwise near immediate re°ex responses. (2014) Phenomenal awareness can emerge without attention. is equal to the task of solution. In principle. Downloaded from www. J. For personal use only. It putatively allows behavioral expression independent of its learned origin. and property of. 145. however improbable. \when you have eliminated all which is impossible. J.A. every centralized vertebrate CNS. with special reference to persisting neurogenesis in the olfactory bulb. It is a process likely applicable to. Aru.. 573–591. and thus its correlate    consciousness. it allows biological mechanisms to account for many di®erent aspects of brain structure and function while lending a potential to unify di®erent perspectives as to how subjective phenomenological states arise and could be experienced as consciousness. 2014.. IV. In its essence a®erence copy.J. 1926). Hum. Rec. T. must be the truth" (Doyle. Wiley Inter-Science. as we have shown here. M. Neurosci. (1963) Autoradiographic investigation of cell proliferation in the brains of rats and cats. It is likely driven by evolutionary requirements and the ontogenetic need to develop. entirely unknown. Front. As a renowned investigator once said. e891. J.

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