An

introducon
to consciousness

Camilla Derchi
Lorenzo Ga
Marisa Saggio

Topics
Today:

Introducon to consciousness and aenon

The sensorymotor conngency (OReagan and No
2001)

Introducon to neural correlates of consciousness
(Tononi & Koch 2008)
Thursday:

The integrated informaon theory

Consciousness in dreams (Nir & Tononi 2010)

Discussion

 Brain & consciousness

The words conscious and consciousness are umbrella
terms that cover a wide variety of mental phenomena

Both are used with a diversity of meanings: the
adjecve conscious is heterogeneous in its range,
being applied both to whole organisms (creature
consciousness) and to parcular mental states and
processes (state consciousness) (Rosenthal 1986,
Gennaro 1995, Carruthers 2000)

Conscious mental state:

at least six major opons
1) States one is aware of
States in which some informaon is available for
control of behaviour and for guiding verbal reports
(Chalmers 1996)

Conscious mental state:

at least six major opons
1) States one is aware of
2) Qualitave states
States that involve qualitave or experienal
properes of the sort oen referred to as qualia or
raw sensory feels

Conscious mental state:

at least six major opons
1) States one is aware of
2) Qualitave states
3) Phenomenal states

Conscious mental state:

at least six major opons
1) States one is aware of
2) Qualitave states
3) Phenomenal states
4) What-it-is-like states

Conscious mental state:

at least six major opons
1) States one is aware of
2) Qualitave states
3) Phenomenal states
4) What-it-is-like states
5) Access consciousness
A states being conscious is a maer of its availability
to interact with other states and of the access that
one has to its content (Nagel 1995)

Conscious mental state:

at least six major opons
1) States one is aware of
2) Qualitave states
3) Phenomenal states
4) What-it-is-like states
5) Access consciousness
6) Narrave consciousness

Qualia

Perceptual experiences (seeing green, hearing loud
trumpets, tasng liquorice, )

Bodily sensaons (feeling a twinge of pain, feeling an
itch, feeling hungry, having a stomachache, )

Felt reacons or passions or emoons (feeling
delight, lust, fear, love, feeling grief, jealousy, ...)

Felt moods (feeling elated, depressed, calm, bored,
tense, ...)

Explanatory gap
Qualitave, phenomenic aspect of experience
vs
Representaonal, intenonal and
funconal aspect of experience

Its not possible to explain the subjecve, felt aspects
of experience in behavioral, physical or funconal
terms

A sensorimotor account of vision

and visual consciousness
ORegan & No
2001

The explanaon of ORegan & No

Qualia are meant to be properes of experiental states

or events. But experiences are not states: they are ways
of acng and things we do

There is no introspecbly available property determining

the character of ones experienal states

Qualia are an illusion, and explanatory gap is not a real

gap at all

The sensorimotor conngency

Proposal
Seeing as a way of acng: parcular way of exploring
the environment. The experience of seeing occurs
when the organism masters the governing laws of
sensorimotor conngency

The sensorimotor conngency

Proposal
Seeing as a way of acng: parcular way of exploring
the environment. The experience of seeing occurs
when the organism masters the governing laws of
sensorimotor conngency

Advantages
Provides a natural and principled way of accounng for
visual consciousness and for dierences in perceived
quality of the experience in dierent sensory
modalies

Brain & vision

General idea
For every perceptual state there is a neural correlate
sucient to produce it. Its funcon is to produce sensory
experience by generang a representaon corresponding
to the content of the experience

Brain & vision

General idea
For every perceptual state there is a neural correlate
sucient to produce it. Its funcon is to produce sensory
experience by generang a representaon corresponding
to the content of the experience

Sensorimotor conngency theory
Seeing is a skillful acvity. The brain supports vision by
enabling mastery and exercise of knowledge of
sensorimotor conngencies

Brain & vision

General idea
For every perceptual state there is a neural correlate
sucient to produce it. Its funcon is to produce sensory
experience by generang a representaon corresponding
to the content of the experience

Sensorimotor conngency theory
Seeing is a skillful acvity. The brain supports vision by
enabling mastery and exercise of knowledge of
sensorimotor conngencies

Visual consciousness
It is not a special kind of brain state or a special quality of
informaonal states of the brain: its something we do

The neural correlates of

consciousness
An update

Tononi & Koch

2008

Neural correlates of consciousness

Using the tools of neuroscience to shed light on the
minimal neural structures and acvity paerns that
underlie consciousness

Neural correlates of consciousness

How?
1)Measuring how brain acvity changes when a
smulus is experienced or not
2)Considering condions in which consciousness is
globally diminished such as sleep or anesthesia:
what has changed in the brain?

Consciousness and
sensory input/motor output
Consciousness might be ghtly linked to the ongoing
interacon we maintain with the world and the body

However:

We are conscious of our thoughts, which do not
seem to correspond to anything out there (smulus-
independent thoughts)

During dreams, we are virtually disconnected from
the environment. Nevertheless, we are vividly
conscious (Hobson, 2009)

Consciousness and self-reecon

Consciousness might arise through the ability to reect
on our own percepons: our brain would form a scene
of what it sees but we experience it subjecvely only
when we, as subject of experience, watch that scene
from the inside.

However:
When we are immersed in the rapid ow of experience
(for example watching and engrossing movie) we are
vividly conscious, without any need for reecon or
introspecon.

Consciousness and aenon

When a subjects pay aenon to an object they
become conscious of its various aributes; when the
focus of aenon shis away, the object fades from
consciousness

Are aenon and consciousness same phenomenon or
are they disnct phenomena with disnct funcon and
neuronal mechanisms?

Recent evidence argues in favor of a dissociaon
between selecve aenon and consciousness
(Bahrami et al. 2007)

Bahrami et al. 2007

Bahrami et al. 2007

The dierent fMRI

response in V1 for the two
condions

Aenon without consciousness

Results show that availability of aenonal capacity
determines neural representaons related to
unconscious processing of connuously suppressed
smuli in V1.

Spillover of aenon to corcal representaons of
invisible smuli cannot be a sucient condion for
their awareness

Consciousness in the
absence of aenon
Using the idencal renal layout, the subject either
performs the central task, the peripheral task, or both
simultaneously (Sperling & Dosher 1986; Braun & Sagi
1990; Braun & Julesz 1998)
In a dual-task paradigms, the subjects aenon is
drawn to a demanding central task, while at the same
me a secondary smulus is ashed somewhere in the
periphery (Braun & Julesz 1998)

Consciousness & aenon: summary

The existence of aenon without consciousness and
consciousness without aenon, should not be surprising when
considering their dierent funcons

Aenon: mechanisms whereby the brain selects a subset of
incoming sensory informaon for higher level processing,
while nonaended parts are analyzed at a lower bandwidth.
It can be directed by boom-up, exogenous cues or by top-
down endogenous features and can be applied to a spaally
restricted part of the image (focal, spotlight of aenon), an
aribute (e.g., all red objects), or to an enre object


Consciousness: involved in providing a kind of execuve
summary of the current situaon that is useful for decision
making, planning, and learning

An important disncon
Changes in level of
consciousness

(the degree to which
we are conscious)

How does brain acvity
change when level of
consciousness changes?

An important disncon
Changes in level of
consciousness

(the degree to which
we are conscious)

How does brain acvity
change when level of
consciousness changes?

Changes in content of
consciousness

(the parcular experience
we are having)

How does brain acvity
change when specic
content of consciousness
changes?

Changes in level of consciousness

Many situaons where the level of consciousness can
decrease:

Sleep

Anesthesia

Coma and vegetave states

Seizures

Sleep
The brain isnt (completely) shut down: vivid conscious
experiences in REM or late NREM sleep (dreams)

Sleep
The brain isnt (completely) shut down: vivid conscious
experiences in REM or late NREM sleep (dreams)

So what are the dierences between NREM and REM?

Part of the brain inacve (frontal & parietal areas)

Slow (1 Hz) up-down oscillaons in corcal and
thalamic neurons (Steriade et al. 2001)

TMS response (Massimini et al. 2005, 2007)

Steriade et al. 2001

Why is NREM not conscious?

A rst explanaon based on these slow oscillaons:

Changes in level of consciousness might be related to
the degree of bistability of thalamocorcal networks
(Tononi 2004, Massimini et al. 2007)

Massimini et al. 2004, 2005, 2007

TMS during NREM:

in lateral corcal regions remains localized to the
smulaon site and lasts for <150ms

in cetromedian parietal regions triggers a
stereotypical, long-lasng response in all cortex
TMS during REM:

resembles more the wakefulness

Massimini et al. 2005

TMS

Wakefulness

TMS

NREM sleep

Why is NREM not conscious?

The experiment show that during NREM connecvity
among corcal regions break down, and so does
corcal integraon

This suggests that during sleep brain becomes bistable:

part of it breaks down in casually independent
modules (premotor cortex)

another part shows a global stereotypical response
(sensorimotor cortex)
It loses the ability to enter states that are both
integrated and dierenated, typical of wakefulness

Anesthesia
How do anesthecs cause (brusque) loss of
consciousness (LOC)?
They reduce thalamic methabolism and blood ow, but
also a global 30%-60% reducon.


Thalamus might be a consciousness switch (Alkire et
al. 2000)

Thalamus might be eecng the corcal circuits
(deacvaon of cortex alone can cause LOC,
acvaon of thalamus alone cannot maintain it)
(Velly et al. 2007)

Alkire et al. 2000

Intersecon of eect between

halothane and isourane anesthesia

Coma and vegetave states

Apart from lesions, LOC caused by impairment in cells
acvaon in intralaminar nuclei, which enable
interacons among corcal regions

Brain metabolism reduced by 50%-60%, especially in
posterior cingulate cortex and precuneus (Laureys et al.
2004; Schi 2006). These areas acvate most reliably if
paent recovers

Schi 2006

Acvaon paerns
in response to
passive language
presentaons

Forward
speech

Reversed
speech

Overlap

Coma and vegetave states

In paents with great part of the thalamocorcal
system intact, normal acvaon paerns can be
induced by cognive smuli (Laureys et al. 2004; Schi
2006) or even by thoughts (Owen et al. 2006)

Owen et al. 2006

Acvaon paerns during tennis

and moving around a house
imagery.

Seizures
All types of seizures with momentary LOC show:

Increased upper brainstem and medial thalamus
acvity

Decreased anterior and posterior cingulate, medial
frontal cortex and precuneus acvity

Altered acvity in the lateral and orbital front cortex,
and in the lateral parietal cortex
LOC appears when neurons are excessively and
synchronously acve (Blumenfeld & Taylor 2003)

Blumenfeld & Taylor 2003

Level of consciousness: summary

Status

Region aected

Sleep

Inacve frontal and parietal areas. Slow oscillaons in corcal and

thalamic neurons

Anesthesia Reduced methabolism in thalamic area, but also everywhere else

Coma

Reduced posterior cingulate cortex and precuneus acvity. Might show

consciousness signs if thalamocorcal system is intact

Seizures

Increased upper brainstem and medial thalamus acvity

Decreased anterior and posterior cingulate, medial frontal cortex and
precuneus acvity

Changes in content of consciousness

The ulmate goal is to idenfy the NCC (neural
correlates of consciousness)

Mainly done with visual smuli because

Easy to manipulate

Visual area is the best known in primates
Uses techniques (masking, binocular rivalry, change
blindness, ) to disrupt the smulus-percept
associaon and its resulng neural acvity

Masking

Binocular rivalry

The NCC in visual cortex

There is good evidence (Wolfe 1984; Leopold &
Logothes 1996; Logothes 1998) that the inferior
temporal cortex responds to the percept instead of the
smulus.

The NCC in visual cortex

Many experiments seem to show that V1 contains
perceptually suppressed informaon (Haynes & Rees
2005; Bahrami et al. 2007)

Probably, much of the neural acvity in V1 does not
belong to the NCC (and this could be true for all the
primary sensory corces)

Consciousness and neural dynamics

Most of the cortex is acve during early NREM,
anesthesia and generalized seizures, but there is no
consciousness

This suggests that addional dynamic features of
neural acvity are necessary to generate consciousness

Models of neural dynamics

Three proposed models:

Substained vs Phasic acvity
Neural acvity needs to last a minimum amount of
me (propagaon me?) in appropriate areas.
Or maybe its the onset and oset discharge that
correlates with consciousness, and masking smulus
overlap the on and o phases and so are suppressed

Models of neural dynamics

Three proposed models:

Substained vs Phasic acvity

Reentrant vs Feedforward acvity
Simple smulus percepon is the rst, feedforward
acvity, but consciousness is due to a reentrant
wave.

Models of neural dynamics

Three proposed models:

Substained vs Phasic acvity

Reentrant vs Feedforward acvity

Synchronizaon and oscillaons
Consciousness might require ne synchronizaon
between distributed populaon of neurons. Phase
locking could be a disambiguang factor between
mulple objects

A theorecal framework
The integrated informaon theory (Tononi 2004) tries
to estabilish


What consciousness is

A theorecal framework
The integrated informaon theory (Tononi 2004) tries
to estabilish


What consciousness is

How we can measure it

A theorecal framework
The integrated informaon theory (Tononi 2004) tries
to estabilish


What consciousness is

How we can measure it

What is required to generate it in any physical
system

What is consciousness?
When we experience a conscious state, we rule out
other alternave experience

So, consciousness is informaon, because it lowers our
level of uncertainty

Why integrated informaon theory?

The level of consciousnes of a physical system is
related to the repertoire of causal states (informaon)
avaiable to the system as a whole (integraon)


Integrated because the experience is an integrated
whole, it cannot be subdivided into indipendently
experienced components

How to measure it?

We can dene a measure of integrated informaon
that quanes the reducon of uncertainty

If a system has a posive value of its called a
complex, which is connected to other elements by
means of ports-in and ports-out

Since integrated informaon is generated within a
complex, this implies subjecvness of conscience.

What does consciousness require?

Computer simulaons (Tononi 2004, 2005) show that
high requires networks that conjoin funconal
specializaon with funconal integraon

This would explain, for example, why consciousness is
related to the thalamo-corcal systems but not for the
cerebellum (which, despite its large number of
neurons, does not contribute much to consciousness
because its made of quasi-independent modules)

Evidence for this models?

Evidence for this models?

None.

Evidence for this models?

None.

But

its validity [] rests on its ability to account
coherently for some phenomenological
observaons and for some elementary but
puzzling facts about consciousness and the
brain