Theoretical
Modeling of Conscious Access
The
above experiments provide a convergent database of observations. In
the present section, we examine which theoretical principles may
account for these findings. We briefly survey the major theories of
conscious processing, with the goal to try to isolate a core set of
principles that are common to most theories and begin to make sense
of existing observations. We then describe in more detail a specific
theory, the Global Neuronal Workspace (GNW), whose simulations
coarsely capture the contrasting physiological states underlying
nonconscious versus conscious processing.
Convergence
toward a Set of Core Concepts for Conscious Access
Although
consciousness research includes wildly speculative proposals (Eccles,
1994; Jaynes, 1976; Penrose, 1990), research of the past
decades has led to an increasing degree of convergence toward a set
of concepts considered essential in most theories (for review, see
Seth, 2007). Four such concepts can be
isolated.
A
supervision system.
In
the words of William James, ‘‘consciousness’’ appears as ‘‘an
organ added for the sake of
steering
a nervous system grown too complex to regulate itself’’ (James,
1890, chapter 5). Posner (Posner
and Rothbart, 1998; Posner and Snyder, 1975)
and Shallice (Shallice, 1972, 1988;
Norman and Shallice, 1980) first
proposed that information is conscious when it is represented in an
‘‘executive attention’’
or ‘‘supervisory attentional’’
system that controls the activities of lower-level sensory-motor
routines and is associated with prefrontal cortex (Figure
6). In other words, a chain of sensory, semantic,
and motor processors can unfold without our awareness, as
reviewed in the previous section, but conscious perception seems
needed for the flexible control of their execution, such as their
onset, termination, inhibition, repetition, or serial chaining.
A
serial processing system.
Descartes
(1648) first observed that ‘‘ideas impede each other.’’
Broadbent (1958) theorized conscious
perception as involving access to a limited-capacity channel where
processing is serial, one object at a time. The attentional blink and
psychological refractory period effects indeed confirm that conscious
processing of a first stimulus renders us temporarily unable to
consciously perceive other stimuli presently shortly thereafter.
Several psychological models now incorporate the idea that initial
perceptual processing is parallel and nonconscious and that conscious
access is serial and occurs at the level of a later central
bottleneck (Pashler, 1994) or second
processing stage of working memory consolidation
(Chun and Potter, 1995).
A
coherent assembly formed by re-entrant or top-down loops.
In
the context of the maintenance of invariant representations of the
body/world through reafference (von Holst and
Mittelstaedt, 1950), Edelman (1987)
proposed re-entry as an essential component
of
the creation of a unified percept: the bidirectional exchange of
signals across parallel cortical maps coding for different aspects of
the same object. More recently, the dynamic core hypothesis (Tononi
and Edelman, 1998) proposes that information encoded by a
group of neurons is conscious only if it achieves
not only differentiation (i.e.,
the isolation of one specific content out of a vast repertoire of
potential internal representations) but also integration
(i.e., the formation of a single,
coherent, and unified representation, where the whole carries more
information than each part alone). A notable feature of the dynamic
core hypothesis is the proposal of a quantitative mathematical
measure of information integration
called F,
high values of which are achieved only through a hierarchical
recurrent connectivity and would be necessary and sufficient to
sustain conscious experience: ‘‘consciousness is integrated
information’’ (Tononi, 2008).
This measure has been shown to be operative for some
conscious/nonconscious distinctions such as anesthesia (e.g., Lee
et al., 2009b; Schrouff et al., 2011),
but it is computationally complicated and, as a result, has not yet
been broadly applied to most of the minimal empirical contrasts
reviewed above.
In
related proposals, Crick and Koch (1995, 2003,
2005) suggested that conscious access involves forming a
stable global neural coalition. They initially introduced
reverberating gamma band oscillations around 40 Hz as a crucial
component, then proposed an essential role of connections to
prefrontal cortex. Lamme and colleagues (Lamme
and Roelfsema, 2000; Super et al., 2001) produced data
strongly suggesting that feed forward or bottom-up processing alone
is not sufficient for conscious access
and that top-down or
feedback signals
forming recurrent loops are
essential to conscious visual perception. Llinas and colleagues
(Llina´s et al., 1998; Llina´s and
Pare, 1991) have also argued that
consciousness is fundamentally a thalamocortical closed-loop property
in which the ability of cells to be intrinsically active plays a
central role.
A
global workspace for information sharing.
The
theater metaphor (Taine,
1870) compares consciousness to a narrow
scene that allows a single actor to diffuse his message. This view
has been criticized because, at face value, it implies a conscious
homunculus
watching the scene, thus leading to infinite regress (Dennett,
1991). However, capitalizing on the
earlier concept of a blackboard system
in artificial intelligence (a common
data structure shared and updated by many specialized modules), Baars
(1989) proposed a homunculus-free
psychological model where the current conscious content is
represented within a distinct mental space called global
workspace, with the capacity to
broadcast this
information to a set of other processors (Figure
6). Anatomically, Baars speculated that
the neural bases of his global workspace might comprise the
‘‘ascending reticular formation of the brain stem and midbrain,
the outer shell of the thalamus and the set of neurons projecting
upward diffusely from the thalamus to the cerebral cortex.’’
We
introduced the Global Neuronal Workspace (GNW) model as an
alternative cortical mechanism capable of integrating the
supervision, limited-capacity, and re-entry properties (Changeux
and Dehaene, 2008; Dehaene and Changeux, 2005; Dehaene et al., 1998a,
2003b, 2006; Dehaene and Naccache, 2001). Our proposal is that
a subset of cortical pyramidal cells with long-range excitatory
axons, particularly dense in prefrontal, cingulate, and parietal
regions, together with the relevant thalamocortical loops, form a
horizontal ‘‘neuronal workspace’’ interconnecting the
multiple specialized, automatic, and nonconscious processors (Figure
6). A conscious content is assumed to
be encoded by the sustained activity of a fraction of GNW neurons,
the rest being inhibited. Through their numerous reciprocal
connections, GNW neurons amplify and maintain a specific neural
representation. The long-distance axons of GNW neurons then broadcast
it to many other processors brain-wide. Global broadcasting allows
information to be more efficiently processed (because it is no longer
confined to a subset of nonconscious circuits but can be flexibly
shared by many cortical processors) and to be verbally reported
(because these processors include those involved in formulating
verbal messages). Nonconscious stimuli can be quickly and efficiently
processed along automatized or
preinstructed processing routes before quickly decaying within a few
seconds. By contrast, conscious stimuli would be distinguished by
their lack of ‘‘encapsulation’’ in specialized processes and
their flexible circulation to various processes of verbal report,
evaluation, memory, planning, and intentional action, many seconds
after their disappearance (Baars, 1989; Dehaene
and Naccache, 2001). Dehaene and
Naccache (2001) postulate that ‘‘this global availability
of information (.) is what we
subjectively experience as a conscious state.’’
“More
generally, these simulations provide a partial neural implementation
of the psychophysical framework according to which conscious access
corresponds to a ‘‘decision’’ based on the accumulation of
stimulus-based evidence, prior knowledge, and biases (Dehaene,
2008; for specific implementations, see
Lau, 2008,
and the mathematical appendix in Del Cul
et al., 2009).
Modeling
Spontaneous Activity and Serial Goal-Driven Processing
An
original feature of the GNW model, absent from many other formal
neural network models, is the occurence of highly structured
spontaneous activity (Dehaene and
Changeux, 2005). Even in the absence of
external inputs, the simulated GNW neurons are assumed to fire
spontaneously, in a top-down manner, starting from the highest
hierarchical levels of the simulation and propagating downward to
form globally synchronized ignited states. (209-212)
Conclusion
and Future Research Directions
The
present review was deliberately limited to conscious access. Several
authors argue, however, for additional, higher-order concepts of
consciousness. For Damasio and Meyer (2009),
core consciousness of incoming sensory information requires
integrating it with a sense of self (the specific subjective point of
view of the perceiving organism) to form a representation of how the
organism is modified by the information; extended consciousness
occurs when this representation is additionally related to the memorized
past and anticipated future (see also Edelman,
1989). For Rosenthal
(2004), a higher-order
thought, coding for the very fact that
the organism is currently representing a piece of information, is
needed for that information to be conscious. Indeed, metacognition,
or the ability to reflect upon thoughts and draw judgements upon
them, is often proposed as a crucial ingredient of consciousness
(Cleeremans et al., 2007; Lau, 2008)
(although see Kanai et al., 2010, for
evidence that metacognitive judgements can occur without conscious
perception). In humans, as opposed to other animals, consciousness
may also involve the construction of a verbal narrative of the
reasons for our behavior (Gazzaniga
et al., 1977). Although this narrative
can be fictitious (Wegner, 2003),
it would be indispensable to interindividual communication (Bahrami
et al., 2010; Frith, 2007).
Metacognition
and self-representation have only recently begun to be studied
behaviorally with paradigms simple enough to extend to nonhuman
species (Kiani and Shadlen, 2009; Terrace and
Son, 2009) and to be related to specific
brain measurements, notably anterior prefrontal cortex (Fleming
et al., 2010). Thus, our view is that
these concepts, although essential, have not yet received a
sufficient empirical and neurophysiological definition to figure in
this review. Following Crick and Koch
(1990), we focused solely here on the
simpler and well-studied question of what neurophysiological
mechanisms differentiate conscious access to some information from
nonconscious processing of the same information. Additional work will
be needed to explore, in the future,
these important aspects of higher-order consciousness. (218-19).
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