Neuroscience of Consciousness I
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1 1 C83MAB: Mind and Brain Neuroscience of Consciousness I Tobias Bast, School of Psychology, University of Nottingham
2 2 What is consciousness?
3 3 Consciousness State of consciousness - Being awake/alert/attentive/responsive Contents of consciousness ( consciousness of... ) particular (reportable) experience at a certain level of consciousness Laureys (2005) Trends in Cogn. Sci. 9:
4 4 Does consciousness matter to you? 1)Yes, very much! Consciousness is central to my life! 2) No, I can t see what difference consciousness makes to my life!
5 5 Great public interest in the brain substrates of consciousness reflected by an abundance of popular-science books on the topic. Etc.
6 6 Problems of consciousness what is to explain? Difference between wakefulness and sleep Difference between being responsive and unresponsive Reportability of our mental states Integration of information Easy problems Ability to access own internal states Focus of attention Deliberate control of behaviour (as opposed to automatic behaviour) The hard problem: Subjective experience ( phenomenal consciousness, qualia ) Chalmers (1995) Facing up to the problem of consciousness Journal of Consciousness Studies 2:
7 7 Today States of consciousness and neural correlates Neurological disorders of (the states of) consciousness Next week Contents of consciousness and neural correlates Neurological disorders affecting contents of consciousness What s left to explain?
8 8 States of consciousness Baars et al. (2003) Trends Neurosci. 26:
9 9 Key brain substrates of consciousness Mormann & Koch (2007) Scholarpedia 2(12):1740
10 Wakefulness 10 and sleep - the cardinal states of consciousness Sleep Hobson & Pace-Schott (2002) Nature Rev Neurosci 3:
11 11 Neurochemistry of wakefulness and sleep Jones (2005) Trends Pharmacol. Sci. 26:
12 12 Cortico-thalamic mechanisms in wakefulness and sleep Falling asleep: Reticular thalamic nucleus neurons fire rhythmic bursts - reflected by EEG spindles - which inhibits thalamo-cortical neurons, thereby blocking information transfer to cortex. Awake: Cortical and thalamic neurons show tonic singlespike firing and afferent information is transferred to cortex. EEG is largely desynchronised, but fast oscillations occur. Sleep (non- REM/slow-wave): cortical and thalamic neurons show rhythmic slow changes between down and up states - reflected by slow EEG oscillations. Waking up: Cholinergic neurons inhibit neurons in reticular thalamic nucleus and activate thalamocortical neurons reflected by the replacement of slow oscillations by faster EEG oscillations, thereby facilitating information transfer to cortex. Figure from Franks (2008) Nature Rev. Neurosci. 9: ; for detailed explanation see: Steriade (2003) Front. Biosci. 8:d
13 13 Comparison of wakefulness and dreams: 40-Hz oscillations characterise conscious states Spontaneous 40-Hz oscillations during wakefulness and REM sleep, but not slow-wave sleep Re-set of 40-Hz oscillations by sensory stimulus (tone) during wakefulness Similar to the awake state, REM sleep the main dream phase of sleep is characterised by 40 Hz oscillations as measured by surface MEG or EEG. Thus, 40-Hz oscillations are a correlate of conscious processing and have been hypothesised to play a key role in such processing. Depth recordings in animal models suggest 40-Hz oscillations are generated by thalamo-cortical circuits. Only during wakefulness, but not REM sleep, 40-Hz oscillations are re-set by sensory stimulus (tone), in line with fast oscillations being critical for conscious perception of sensory information. Llinas & Ribary (1993) Proc. Natl. Acad. Sci. USA 90:
14 Thalamo-cortical circuitry hypothesised to underlie 40-Hz 14 oscillations and consciousness Widespread cortical projections of intra-laminar thalamus Sensory information Llinas & Ribary (1993) Proc. Natl. Acad. Sci. USA 90:
15 15 Anaesthesia and consciousness Loss of consciousness is the objective of anaesthesia. Loss of consciousness is not trivial to determine, as unresponsiveness and amnesia are no proof. Neuropharmacological targets of current anaesthetics Inhibitory neuro-transmission is enhanced, excitatory transmission decreased. Note: this can lead to excitation of certain brain regions due to network effects (e.g., ketamine). Brain sites associated with anaesthetic effects Thalamus: Activity is decreased by many, but not all, anaesthetics (e.g., ketamine); may be secondary to cortical deactivation. Neocortical areas: Deactivation of, especially posterior, mesial cortical areas and of a lateral temporo-parieto-occipital complex. Note: Primary sensory cortices can often show unchanged responsiveness under anaesthesia! Anaesthesia may not necessarily involve cortical deactivation, but disruption of cortical integration, i.e. of the interaction of several cortical areas. Alkire et al. (2008) Science 322:
16 Anaesthesia disrupts cortico-cortical interactions organised 16 by fast oscillations Alkire et al. (2008) Science 322: , based on Imas et al. (2005) Neurosci. Letters 387:
17 17 Similarities between sleep and anaesthesia Regional decreases in brain metabolic activity during sleep and anaesthesia Franks (2008) Nature Rev. Neurosci. 9: ;
18 18 (Abnormal) Dissociated states of consciousness There are several abnormal conditions in which wakefulness is relatively preserved whereas consciousness (of anything) appears to be largely absent. Deactivated brain areas (in comparison to conscious wakefulness) Laureys (2005) Trends in Cogn. Sci. 9:
19 19 Loss of consciousness in epilepsy Yu & Blumenfeld (2009) Ann. N.Y. Acad. Sci. 1157:48-60
20 20 States of reduced consciousness following brain injury Resting brain metabolism in patients following brain injury and in healthy control Normal consciousness! Laureys et al. (2004) Lancet Neurol. 3: Brain injury associated with vegetative state Zeman (1997) Lancet 350:
21 21 In a nut shell states of consciousness and brain substrates Neocortical areas Thalamus Ascending activating systems Global states of consciousness depend on a distributed brain network, comprising ascending activating systems, thalamus and neocortex. Consciousness seems to involve the temporal coordination of distributed activity in thalamic and neocortical areas as reflected by fast oscillations. Neuronal correlates of specific contents of consciousness (e.g., specific percepts, specific plans, specific memories)...??? Next week!
22 22 Neuroscience of consciousness I Selected Reading General overviews: Chalmers D (1995) Facing up to the problem of consciousness. Journal of Consciousness Studies 2: ( Zeman A (2001) Consciousness. Brain 124: Tononi G & Koch C (2008) The neural correlates of consciousness: and update. Ann. N. Y. Acad. Sci. 1124: Sleep & anaesthesia: Alkire MT, Hudetz AG, Tononi G (2008) Consciousness and anaesthesia. Science 322: Franks NP (2008) General anaesthesia: from molecular targets to neuronal pathways of sleep and arousal. Nature Rev. Neurosci. 9: Dissociated states of consciousness: Laureys S (2005) The neural correlates of (un)awareness. Trends Cogn. Sci. 9:
23 Neuroscience of consciousness I Some questions to guide your revision 23 What is consciousness, what are the problems of consciousness? What strategies can neuroscientists pursue to study the brain substrates of consciousness? In terms of brain substrates, what seem conscious states to have in common? What happens in your brain, so you wake up ( regain consciousness ) after a night s sleep? How do anaesthetics act in our brains to (hopefully!) result in loss of consciousness? What goes wrong in the brains of patients that suffer from disorders of consciousness?
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