|
|
|
|
 |
| The
Sense of Self |
 |
|
|
|
Thalamocortical interactions are
involved in generating neuronal
oscillations across various parts of the cortex.
In this sense, the thalamus acts somewhat like the
conductor of an orchestra whose musicians are distributed
throughout the cortex. The conductor doesn’t
play the music for the musicians, but instead co-ordinates
them and imposes a rhythm on them. Without the thalamus,
the cortex could probably display some isolated examples
of synchrony but would not be able to bind the various properties
of a perception into a coherent concept .
This metaphor also helps us to understand
why it is pointless to look for any one seat or centre of
consciousness in the brain. The thalamic conductor might
impose the rhythm, but that would be meaningless unless the
cortical musicians were playing their sensory scores. It
is the co-ordination of all these things that makes the mental
symphony - the object of consciousness - coherent. |
The thalamus is very
well positioned to control the inputs to the cortex. Among
the various thalamic nuclei, the reticular nucleus is
known to exert an inhibiting modulation on the other specific
sensory nuclei of the thalamus. The reticular nucleus thus
helps to select the sensory inputs that can reach the cortex,
and hence enter consciousness.
The circuits in this thalamic nucleus
can favour one particular input over several others. For
example, this is what happens when a stimulus that has a
strong meaning for someone (such as the sound of their own
name) manages to clear a path through numerous other auditory
stimuli and thus reach that person’s consciousness.
This kind of activation, “from
the bottom up”,
would be controlled by the brainstem, the amygdala, or
the systems associated with the perception of pain. In
contrast, activation “from
the top down” would be controlled by
the executive functions of the frontal cortex and, according
to certain authors, would operate through the anterior
cingulate cortex. |
Brodmann area
46, located in the frontal cortex, is activated
by a wide range of tasks and seems well situated for co-ordinating
conscious thoughts. In conjunction with all the other areas
of the brain, area 46 might help us to switch from one
thought to another by facilitating certain global activation
patterns at the expense of others.
The particular content of a thought—“what
you have on your mind”— is associated with the
content of working
memory: the temporary memory that you use for
tasks such as doing arithmetic in your head, or keeping your
train of thought while formulating long sentences or advancing
complex arguments, or for assessing possible moves when you
are playing chess.
This working memory is often described
as being composed of a central
executive (identified with frontal area
46) and two main auxiliary “slave” systems.
One of these auxiliary systems is a
visual/spatial form of memory that engages several areas
in the right hemisphere. This memory holds mental images,
the pictures that you can “see in your head” and
that are so helpful for solving spatial-configuration problems.
The second auxiliary memory system
comprises an auditory form of memory, or “phonological
loop”. This is the locus of your inner discourse,
that small voice that you constantly use to talk to yourself
and that activates the areas
in the left hemisphere that are used to decode language.
But regardless of which of these two
auxiliary systems is at work, the executive processor in
the frontal lobes is always activated.
|
To try to explain the
complex role of the frontal cortex more
clearly, some authors use the metaphor of an executive committee
composed of five members, each of whom represents a sub-committee
of more posterior or subcortical areas in the brain.
The first member is the Perceiver.
Located mainly in the ventral-lateral portion of the right
frontal hemisphere, the Perceiver is the frontal extension
of the ventral
perceptual system and is focused on objects.
The second member is the Verbalizer. Dominant
in the ventral-lateral portion of the left prefrontal cortex,
it is the frontal extension of the language circuits. The third committee member is
the Motivator. Located in the ventral-medial
region of the orbitofrontal cortex, it is the cortical extension
of subcortical pathways that include the amygdala and represent the world in terms of emotional motivations.
The fourth member, the Attender, occupies
the dorsal-medial portion of the frontal cortex, as well as
the anterior cingulate cortex. The Attender is the frontal
extension of a subcortical pathway involving the hippocampus.
The Attender represents the world and self in spatiotemporal
co-ordinates and can direct attention to internal and external events. Lastly, the fifth member
is the Co-ordinator (or central executive - see preceding sidebar). It is
located in the dorsolateral region of the frontal cortex and
is the frontal extension of the dorsal
pathway. The Co-ordinator represents the world
and self in body-centred coordinates, which enables it to
control willed movements and working
memory.
 |
|
|
| CAN STATES OF CONSCIOUSNESS BE MAPPED IN
THE BRAIN? |
|
Some neurobiological models of consciousness,
such as the global
workspace theory, assume that the contents of consciousness
are widely distributed in the brain. This assumption has been
confirmed by many brain-imaging experiments, in particular those
of Stanislas Dehaene and his collaborators. In these experiments,
when the amount of time that a word was projected onto a screen
was extended just past the threshold required for subjects to
perceive it consciously, there was a major increase in activity
in their frontal, prefrontal, anterior cingulate, and parietal
cortexes.
Thus conscious sensory inputs would
appear to produce far more extensive brain activity than comparable
unconscious stimuli, and a sudden activation of the frontal and
parietal lobes would appear to be the typical signature of a
conscious perception.
But this perceptual consciousness,
or as some would call it, primary
consciousness, is
not the only form of consciousness. When we are trying to
associate consciousness with particular structures in the brain,
we must therefore clearly define what level of consciousness
we are talking about. For example, the first condition necessary
for the brain to be able to process external sensory stimuli
consciously is that it must be in an appropriate state of alertness
(for instance, awake
rather than asleep).
Starting from this premise,
authors such as Damasio distinguish a very primitive form
of consciousness that he calls the proto-self and
that is more like a moment-to-moment perception of the body’s
internal emotional state. This state is associated with activity
of such brain structures as the reticular formation, the
hypothalamus, and the somatosensory cortex.
The reticular
formation is also associated with consciousness
in the minimal sense of wakefulness. Other structures
involved in simply maintaining wakefulness include the pons,
the raphe nuclei and the locus
coeruleus. |
|
It should be noted here that the activity
of the reticular formation, like that of the primary sensory areas,
seems to be necessary but not sufficient for a more elaborate level
of consciousness. This latter level is attained with what several
authors call primary
consciousness, meaning a waking state in which we
are in relationship with our environment “here and now”.
On the basis of the research done by Swedish neuroscientist Bjorn
Merker, it seems that the
brainstem plays a more important role in primary consciousness
than was formerly believed.
Damasio calls this type of consciousness “core
consciousness” and says that it depends chiefly on the cingulate
cortex and on the intralaminar
nuclei of the thalamus. Indeed, experiments
have shown that bilateral destruction of the centromedial portion
of the intralaminar nuclei of the thalamus also eliminates consciousness,
produces a coma,
or causes other states similar to brain death. In addition, this
region of the thalamus is one of the main sites acted upon by anaesthetics and
by antipsychotic drugs.
Models of consciousness that attribute a
role to the thalamus are no recent development. As far back as
1984, Francis
Crick offered one of the first hypotheses about consciousness,
the “thalamic searchlight hypothesis”, according to
which the thalamus controlled which region of the cortex became
the focal point for consciousness. A similar but more sophisticated
idea has recently been proposed by Rodolfo
Llinas. He hypothesizes that the oscillations of certain neurons
in the thalamus serve as a sort of basic rhythm with which the
cortical oscillations of the various sensory modalities synchronize
themselves to form a unified image of the environment—somewhat
like an orchestra conductor who provides the beat for all the musicians
to follow (see sidebar). This is an original solution to the binding
problem.

|
The thalamus is
often compared to a railroad switching yard, because the
signals from all of the senses (except smell) must pass through
it before they can reach the cortex. The cortex also sends
many connections back to the thalamus. Most of the nuclei
in the thalamus are considered “specific” because
their neurons make connections with relatively circumscribed
areas in the cortex (for example, the neurons of the lateral
geniculate nucleus project to the primary
visual cortex).
The thalamus also has many “non-specific” nuclei
that send diffuse projections to wide areas of the cortex.
The intralaminar
nuclei, located in the internal medullary lamina,
are a good example of non-specific thalamic nuclei.
To complete this overview of the thalamus, we should note
that only one of its nuclei, the reticular
nucleus, which wraps around the lateral portion
of the thalamus, does not send any projections directly to
the cortex. It does, however, play a role in the thalamocortical
feedback loops, by receiving inputs from the cortex and sending
outputs to the dorsal nucleus of the thalamus. |
These thalamocortical
loops have come to play an important role in practically all
of the neurobiological theories that attempt to explain
the higher states of consciousness, for which the lower levels
of consciousness that we have been discussing up to now are
in a sense only the prerequisites. These higher levels of human
consciousness are known as reflexive consciousness and self-consciousness.
Reflexive consciousness—this
sense that “it is I who am perceiving”—is often
presented as a necessary condition for self-consciousness:
the feeling of being oneself and not someone else. This autobiographic
dimension of consciousness implies that we can form mental representations
of conscious experiences in the past or the future, and therefore
requires the support of memory and
the higher functions that make abstract conceptualization and planning
possible.
You would therefore expect that the areas
of the brain that are known to be involved in these functions,
especially in the frontal and parietal lobes,
would be actively engaged in self-consciousness. And that indeed
has been found to be so in certain studies that addressed this
specific question.
These higher levels of consciousness also
appear to involve other brain structures whose roles were long
poorly understood, partly because some of them are located deep
in the brain, which made them hard to access. Modern brain imaging
techniques have now overcome this problem.
Three
of these structures—the angular gyrus, the precuneus,
and the anterior cingulate cortex, which
are often very active in a conscious state of rest—may
be part of a functional network that makes self-consciousness
possible.
The case of the precuneus, which is the
postero-medial portion of the parietal lobe, is especially
revealing. The conscious resting state is a state in which
the subject’s eyes are generally closed and the subject’s EEG shows
an alpha
rhythm, or in which the subject is passively looking
at a simple target such as a “+” sign. Among
all the areas of the brain that are active during this state,
the precuneus is the one that shows the highest rate of neural
activity. But in contrast, the precuneus is known to be less
active during tasks that make no reference to the self. Some
authors have therefore proposed that the activation of the
precuneus, and of the posterior cingulate cortex,
which is closely connected to it, is correlated with the
feeling of selfhood and the sense of being an “agent”. |


After Wheatley et al.,
2007.
|
This hypothesis is also consistent with studies
that have shown decreased activity in the postero-medial parietal
cortex in many states of altered consciousness, such as sleep,
anaesthesia, or a vegetative state. Other studies have also shown
decreased activity in the precuneus and the posterior cingulate
cortex when the subject is under hypnosis, another altered state
of consciousness.
Lastly, the precuneus also seems to play
a role in visual/spatial imagery. For example, some experiments
have shown that the precuneus is more active when subjects are
is engaged in motor
imagery of a finger movement than when they are actually performing
this movement. This again seems to indicate that people have a
propensity to represent their own bodies in space.
The insula (also
known as the insular cortex) is another region of the brain that
remained little understood for a long time because of its position
deep in the folds of the cortex. Also, because it was not associated
with the “higher” brain functions, it was of less interest
to scientists who were investigating consciousness.
But this indifference gave way to intense
interest after Antonio
Damasio conducted research on the insula and proposed that
most of this structure consists of somatic markers. Damasio hypothesized
that this part of the cortex maps the bodily states associated
with our emotional experiences, thus giving rise to conscious feelings.
This hypothesis falls within the school of thought known as embodied
cognition, according to which conscious rational thought cannot
be separated from emotions and their incarnation in the rest of
the body.

After Wheatley et al.,
2007. |
The insula thus appears to
provide an emotional context that is suitable for a given sensory
experience. The insula is also well positioned to integrate
information about the state of the body and to make this information
available to higher-order cognitive and emotional processes.
For example, the insula receives homeostatic sensory inputs
via the thalamus and sends outputs to several structures associated
with the limbic system, such as the amygdala,
the ventral
striatum, and the orbitofrontal cortex. |
The insula has also been convincingly shown
to be associated with pain processes, as well as with several basic
emotions such as anger, fear,
disgust, joy, and sadness. Its most anterior portion is regarded
as part of the limbic
system. The insula also appears to be deeply involved in conscious
desires, such as the active search for food or drugs.
What is common to all of these states is that they affect the entire
body profoundly—which tends to strengthen the case for the
insula’s likely role in the way we represent our bodies to
ourselves and in the subjective aspect of emotional experience.
Lastly, the insula in humans, and to a lesser
extent in the great apes, appears to incorporate two evolutionary innovations that
provide these species with a greater ability to read the state
of their own bodies than any other mammals.
First, the anterior portion of the insula,
and more particularly that of the insula in the right hemisphere,
is more developed in humans and great apes than in other animal
species. This greater development might enable more precise decoding
of bodily states—the capability that translates a bad odour,
for example, into a feeling of disgust, or the touch of a lover
into a feeling of delight.
The other major evolutionary modification
in the insula is a type of neuron that is found only in the great
apes and in humans. These large,
elongated, cigar-shaped nerve cells are known as von Economo neurons
(VENs). VENs occur only in the insula and in the anterior cingulate
cortex. These neurons connect to various parts of the brain, which
would be an essential attribute for the higher functions attributed
to these two brain structures.
| Now
it is time to say a few words about the anterior cingulate
cortex, which also acts as an important
interface between emotion and cognition, and more specifically
in the conversion of feelings into intentions and actions.
This structure is involved in higher functions such as controlling
one’s own emotions, concentrating on solving problems,
recognizing one’s own mistakes, and making adaptive responses
to changing conditions. All of these functions are closely
linked with our emotions. |

After Wheatley et al.,
2007.
|
When experimental subjects are pricked with
a needle, the activity in their cingulate cortex increases; this
response is so clear-cut that the neurons in question are often
called the “pain neurons”. A fascinating side-note:
in 1999, William Hutchison and his colleagues
at the University of Toronto showed that these same neurons in
the cingulate cortex also become active when the subject sees someone
else being pricked with a needle. Thus, for these kinds of neurons,
known as mirror
neurons, there is no boundary between the self and the other.
Primates, including humans, are highly social
creatures. Knowing other individuals’ intentions has always
been crucial for our survival. That is why we are past masters
of the art of internally
simulating other people’s minds, perhaps with the help
of such mirror neurons.
Some neuroscientists,
such as V.S. Ramachandran, even suggest that this
ability to decode other individuals’ states of mind may
even have evolved first, and subsequently been applied to the self,
to become what we call self-consciousness. And in Ramachandran’s
view, not only the mirror neurons, but
all parts of the brain that contribute to language, such as
Wernicke’s area in the temporal lobe, must inevitably play
a role in this process.
This important role is ascribed to language
in several models of higher consciousness, including that proposed
by Michael Gazzaniga, who is known for his work
with “split-brain”patients. But while Gazzaniga's model
identifies the language hemisphere as the locus of this “interpreter”
that makes us conscious of ourselves, other authors, such as Edelman, argue that consciousness
cannot be attributed to any specific structure in the brain.
|
|