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Measuring Consciousness: Relating Behavioural and Neurophysiological Approaches

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Measuring Consciousness: Relating Behavioural and Neurophysiological Approaches TICS-697; No of Pages 8 Review Measuring consciousness: relating behavioural and neurophysiological approaches Anil K. Seth1, Zolta´ n Dienes2, Axel Cleeremans3, Morten Overgaard4 and Luiz Pessoa5 1 Department of Informatics, University of Sussex, Falmer, Brighton, BN1 9QJ, UK 2 Department of Psychology, University of Sussex, Falmer, Brighton, BN1 9QJ, UK 3 Cognitive Science Research Unit, Universite´ Libre de Bruxelles CP 191, Av. F.-D. Roosevelt 50, 1050 Bruxelles, Belgium 4 Cognitive Neuroscience Research Unit, Hammel Neurorehabilitation and Research Center, Aarhus University Hospital, Voldbyvej 15, 8450 Hammel, Denmark 5 Department of Psychological and Brain Sciences, Indiana University, 1101 E10th Street, Bloomington, IN 47405, USA The resurgent science of consciousness has been accom- sizing recent contributions. We find that potential and panied by a recent emphasis on the problem of measure- actual conflicts among measures suggest new experiments ment. Having dependable measures of consciousness is (Table 1); we also assess how different measures can track essential both for mapping experimental evidence to the graded nature of conscious experience (Table 2). We theory and for designing perspicuous experiments. Here, conclude that it is only by behaving sensibly in a theoreti- we review a series of behavioural and brain-based cal context that proposed measures pick themselves up by measures, assessing their ability to track graded con- their bootstraps, both validating themselves as measures sciousness and clarifying how they relate to each other of what they say they measure and the theories involved. by showing what theories are presupposed by each. We identify possible and actual conflicts among measures Theories of consciousness that can stimulate new experiments, and we conclude Worldly discrimination theory that measures must prove themselves by iteratively Perhaps the simplest theory that still impacts the exper- building knowledge in the context of theoretical frame- imental literature is that any mental state that can works. Advances in measuring consciousness have express its content in behaviour is conscious; thus, a implications for basic cognitive neuroscience, for com- person shows they are consciously aware of a feature in parative studies of consciousness and for clinical appli- the world when they can discriminate it with choice beha- cations. viour [1,2]. This theory often makes use of signal-detection theory (SDT), a statistical framework for quantifying the The problem of measurement discriminability of a stimulus [3]. SDT itself is mute on the How can we measure whether and to what extent a subject of consciousness and can, thus, be combined with particular sensory, motor or cognitive event is consciously experienced? Such measurements provide the essential data on which the current and future science of conscious- Glossary ness depends, yet there is little consensus on how they Blindsight: the capability of some individuals with visual cortical damage to should be made. The problem of measuring consciousness perform visually guided behaviours even though they report the absence of any associated conscious content [4]. differs from the problem of identifying unconscious proces- Conscious content: the continually changing phenomenal content (e.g. ‘qualia’ sing. For instance, in subliminal perception experiments it such as redness and warmth) and intentional content (e.g. explicitly held beliefs, is desirable to know whether or not a stimulus has been conscious knowledge) that is present to varying degrees at non-zero conscious levels. consciously perceived, and in implicit learning paradigms Conscious level: applies to a whole organism and refers to a scale ranging from it is interesting to know whether the relationships between total unconsciousness (e.g. death and coma) to vivid wakefulness. A different consciously represented stimuli are uncon- ‘conscious organism’ is one that is capable of having non-zero conscious levels. An organism that is dreaming has some conscious level, although the sciously inferred. Measuring consciousness, however, conscious level is reduced in dreamless sleep. A non-zero conscious level requires saying something about conscious level (Glossary) indicates the presence of some conscious content. and conscious content beyond the zero-point of uncon- Primary consciousness: conscious content consisting of a multimodal scene composed of basic perceptual and motor events. Primary consciousness is sciousness. sometimes called ‘sensory consciousness’. By contrast, higher-order conscious- Here, we review current approaches for measuring ness refers to awareness of being in a mental state. In humans it is usually consciousness, covering both behavioural measures and associated with language and an explicit sense of selfhood. In higher-order theories it is possible to have higher-order thoughts that are not themselves measures based on neurophysiological data. We outline (higher-order) conscious, but in virtue of which other (primary) contents are a variety of broad theoretical positions before describing a conscious. Steady-state visual–evoked potential: stimulus-induced components of brain range of measures in the context of these theories, empha- signals that can be identified over extended periods of time. For example a visual image flickering at 10 Hz will evoke a response at 10 Hz in the EEG or magnetoencephalography signal, readily identifiable by a Fourier transform. Corresponding author: Seth, A.K. ([email protected]). 1364-6613/$ – see front matter ß 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.tics.2008.04.008 Available online xxxxxx 1 TICS-697; No of Pages 8 Review Trends in Cognitive Sciences Vol.xxx No.x Table 1. Conflicts between measuresa Content unconscious according to: Objective Strategic control Subjective Wagering Content Objective Unconscious knowledge by In both subliminal perception Shown in blindsight and conscious Jacoby’s process dissociation and implicit learning, subjects in the Iowa gambling task according procedure is ipso facto often pass objective tasks while by [34] to: conscious by objective claiming to have no knowledge measures (e.g. Refs [60,72]) or showing no relation between confidence and accuracy (e.g. [22,73–75]) Strategic Not possible Subjects can control which Not yet shown but control grammar to employ while entirely possible (see claiming to be guessing [32] and below) hypnotized subjects can engage in strategic control while reporting no awareness [76] Subjective Not possible Shown in Stroop effects – a As yet only shown in our person can report the word’s unpublished work – a meaning but cannot control person can report its rapid use awareness but still wager indiscriminately Wagering Not possible Not yet shown but entirely Not yet shown but entirely possible (see Box 2) possible (see Box 2) Widespread Cognitive control Likely for Stroop with clearly Shown in a ‘relative blindsight’ Likely given the results activation system, including shown words [44] paradigm [31] with verbal subjective prefrontal cortex, measures, but not yet activated by objectively tested invisible stimuli [56] Synchrony g synchrony persists As left As left, also, similar levels of g As left during non-REM sleep synchrony are observed during and under anaesthesia non-REM and during (reportable) [53] REM sleep [52] Complexity Possible in theory (but As left As left As left measures see F). Not tested in practice Content unconscious according to: Widespread activation Synchrony Complexity measures Content Objective Local neuronal activity can Unlikely given current Not yet tested but entirely possible conscious support discriminatory evidence according behaviour in many non- to: conscious organisms (e.g. nematodes and worms). In humans, at least sensory and motor cortices need to be active Strategic control Unlikely: strategic control As above Possible but not tested probably requires activation in both perceptual and frontal regions Subjective As above As above As above Wagering As above As above As above Widespread Experimentally open. Some Possible in theory. Not tested activation studies show increased long- in practice range synchrony accompanying conscious access [48] Synchrony Gamma synchrony is often As above localized [47] Complexity High neural complexity (or F, Possible in theory. Not tested measures or cd) probably requires in practice widespread activity: all else being equal, larger networks will give rise to higher complexity values [17] aRows indicate a measure finds the content conscious and columns indicate the measure finds the content unconscious. Entries reflect a scale according to which a particular conflict is (i) experimentally noted, (ii) not yet shown but entirely possible, (iii) experimentally open, (iv) possible but not tested, (v) unlikely given current evidence and/or theory, or (vi) not possible. different theories. The combination of SDT with the that they can see at all. However, two properties of worldly discrimination theory (WDT) asserts that continu- blindsight indicate intuitively that the seeing is not ousinformationavailablefordiscriminationsisnecess- conscious [4]. First, blindsight patients do not spon- arily the content of conscious mental states. This theory taneously attempt to use the information practically or captures one property of conscious knowledge, namely inferentially. Second, blindsight patients themselves that it enables choice behaviour. However, rightly or think they cannot see. wrongly, it
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