On the relation between complex brain activity and consciousness

Why does it feel like something to be awake? I.e. how is consciousness generated by the body, the brain in particular? Seeking to map phenomenological properties of any first person experience to neural activity patterns, theories of consciousness suggest a correlation between a specific type of neural dynamical complexity and the level of consciousness: When awake and aware, all brain regions are to a certain extent connected and there is diversity in the interactions. In support of this, Casali et al. (2013) have used EEG and transcranial magnetic stimulation to show extensively that brain response activity to direct perturbation is the more diverse across regions and time, the higher the level of consciousness. The spatio-temporal diversity of the response signal is quantified by a single index, the perturbational complexity index (PCI), using a Lempel-Ziv compression algorithm. Motivated by this result, and given that spontaneous neural signals are easier to obtain than response signals to perturbation, this thesis proposes measures - based on Lempel-Ziv compression and entropy - to quantify spontaneous neural signal diversity across channels and observations. Our measures’ sensitivity and specificity to conscious level is demonstrated by re-analysing resting state scalp EEG during propofol-induced anaesthesia and depth electrode recordings during sleep stages, resulting in consistently higher scores for subjects that are awake than being in propofol-induced anaesthesia or non-rapid eye movement sleep. In addition we demonstrate that our measures score higher for states induced by psychedelic substances by re-analysing resting state magnetoencephalography (MEG) data. We further explore in computer simulation how our measures and PCI behave as a function of connectivity of coupled oscillators, informing models of brain mechanisms associated with the loss of consciousness. While our measures may be weaker than PCI in terms of specificity and sensitivity to conscious level, they are quick and easy to compute and applicable to readily available resting state data. This thesis provides strong evidence that cortical signal diversity is a hallmark of consciousness, as predicted by integrated information and complexity theories of consciousness. i

Global and local complexity of intracranial EEG decreases during NREM sleep

Key to understanding the neuronal basis of consciousness is the characterisation of the neural signatures of changes in level of consciousness during sleep. Here we analysed three measures of dynamical complexity on spontaneous depth electrode recordings from 10 epilepsy patients during wakeful rest and different stages of sleep: (i) Lempel-Ziv complexity, which is derived from how compressible the data are; (ii) amplitude coalition entropy, which measures the variability over time of the set of channels active above a threshold; (iii) synchrony coalition entropy, which measures the variability over time of the set of synchronous channels. When computed across sets of channels that are broadly distributed across multiple brain regions, all 3 measures decreased substantially in all participants during early-night non-rapid eye movement (NREM) sleep. This decrease was partially reversed during late-night NREM sleep, while the measures scored similar to wakeful rest during rapid eye movement (REM) sleep. This global pattern was in almost all cases mirrored at the local level by groups of channels located in a single region. In testing for differences between regions, we found elevated signal complexity in the frontal lobe. These differences could not be attributed solely to changes in spectral power between conditions. Our results provide further evidence that the level of consciousness correlates with neural dynamical complexity

Informativeness of Auditory Stimuli Does Not Affect EEG Signal Diversity

Brain signal diversity constitutes a robust neuronal marker of the global states of consciousness. It has been demonstrated that, in comparison to the resting wakefulness, signal diversity is lower during unconscious states, and higher during psychedelic states. A plausible interpretation of these findings is that the neuronal diversity corresponds to the diversity of subjective conscious experiences. Therefore, in the present study we varied an information rate processed by the subjects and hypothesized that greater information rate will be related to richer and more differentiated phenomenology and, consequently, to greater signal diversity. To test this hypothesis speech recordings (excerpts from an audio-book) were presented to subjects at five different speeds (65, 83, 100, 117, and 135% of the original speed). By increasing or decreasing speed of the recordings we were able to, respectively, increase or decrease the presented information rate. We also included a backward (unintelligible) speech presentation and a resting-state condition (no auditory stimulation). We tested 19 healthy subjects and analyzed the recorded EEG signal (64 channels) in terms of Lempel-Ziv diversity (LZs). We report the following findings. First, our main hypothesis was not confirmed, as Bayes Factor indicates evidence for no effect when comparing LZs among five presentation speeds. Second, we found that LZs during the resting-state was greater than during processing of both meaningful and unintelligible speech. Third, an additional analysis uncovered a gradual decrease of diversity over the time-course of the experiment, which might reflect a decrease in vigilance. We thus speculate that higher signal diversity during the unconstrained resting-state might be due to a greater variety of experiences, involving spontaneous attention switching and mind wandering

Complexity of multi-dimensional spontaneous EEG decreases during propofol induced general anaesthesia

Emerging neural theories of consciousness suggest a correlation between a specific type of neural dynamical complexity and the level of consciousness: When awake and aware, causal interactions between brain regions are both integrated (all regions are to a certain extent connected) and differentiated (there is inhomogeneity and variety in the interactions). In support of this, recent work by Casali et al (2013) has shown that Lempel-Ziv complexity correlates strongly with conscious level, when computed on the EEG response to transcranial magnetic stimulation. Here we investigated complexity of spontaneous high-density EEG data during propofol-induced general anaesthesia. We consider three distinct measures: (i) Lempel-Ziv complexity, which is derived from how compressible the data are; (ii) amplitude coalition entropy, which measures the variability in the constitution of the set of active channels; and (iii) the novel synchrony coalition entropy (SCE), which measures the variability in the constitution of the set of synchronous channels. After some simulations on Kuramoto oscillator models which demonstrate that these measures capture distinct ‘flavours’ of complexity, we show that there is a robustly measurable decrease in the complexity of spontaneous EEG during general anaesthesia

Mitochondrial physiology

As the knowledge base and importance of mitochondrial physiology to evolution, health and disease expands, the necessity for harmonizing the terminology concerning mitochondrial respiratory states and rates has become increasingly apparent. The chemiosmotic theory establishes the mechanism of energy transformation and coupling in oxidative phosphorylation. The unifying concept of the protonmotive force provides the framework for developing a consistent theoretical foundation of mitochondrial physiology and bioenergetics. We follow the latest SI guidelines and those of the International Union of Pure and Applied Chemistry (IUPAC) on terminology in physical chemistry, extended by considerations of open systems and thermodynamics of irreversible processes. The concept-driven constructive terminology incorporates the meaning of each quantity and aligns concepts and symbols with the nomenclature of classical bioenergetics. We endeavour to provide a balanced view of mitochondrial respiratory control and a critical discussion on reporting data of mitochondrial respiration in terms of metabolic flows and fluxes. Uniform standards for evaluation of respiratory states and rates will ultimately contribute to reproducibility between laboratories and thus support the development of data repositories of mitochondrial respiratory function in species, tissues, and cells. Clarity of concept and consistency of nomenclature facilitate effective transdisciplinary communication, education, and ultimately further discovery

Mitochondrial physiology

As the knowledge base and importance of mitochondrial physiology to evolution, health and disease expands, the necessity for harmonizing the terminology concerning mitochondrial respiratory states and rates has become increasingly apparent. The chemiosmotic theory establishes the mechanism of energy transformation and coupling in oxidative phosphorylation. The unifying concept of the protonmotive force provides the framework for developing a consistent theoretical foundation of mitochondrial physiology and bioenergetics. We follow the latest SI guidelines and those of the International Union of Pure and Applied Chemistry (IUPAC) on terminology in physical chemistry, extended by considerations of open systems and thermodynamics of irreversible processes. The concept-driven constructive terminology incorporates the meaning of each quantity and aligns concepts and symbols with the nomenclature of classical bioenergetics. We endeavour to provide a balanced view of mitochondrial respiratory control and a critical discussion on reporting data of mitochondrial respiration in terms of metabolic flows and fluxes. Uniform standards for evaluation of respiratory states and rates will ultimately contribute to reproducibility between laboratories and thus support the development of data repositories of mitochondrial respiratory function in species, tissues, and cells. Clarity of concept and consistency of nomenclature facilitate effective transdisciplinary communication, education, and ultimately further discovery

Increased spontaneous MEG signal diversity for psychoactive doses of ketamine, LSD and psilocybin

What is the level of consciousness of the psychedelic state? Empirically, measures of neural signal diversity such as entropy and Lempel-Ziv (LZ) complexity score higher for wakeful rest than for states with lower conscious level like propofol-induced anesthesia. Here we compute these measures for spontaneous magnetoencephalographic (MEG) signals from humans during altered states of consciousness induced by three psychedelic substances: psilocybin, ketamine and LSD. For all three, we find reliably higher spontaneous signal diversity, even when controlling for spectral changes. This increase is most pronounced for the single-channel LZ complexity measure, and hence for temporal, as opposed to spatial, signal diversity. We also uncover selective correlations between changes in signal diversity and phenomenological reports of the intensity of psychedelic experience. This is the first time that these measures have been applied to the psychedelic state and, crucially, that they have yielded values exceeding those of normal waking consciousness. These findings suggest that the sustained occurrence of psychedelic phenomenology constitutes an elevated level of consciousness - as measured by neural signal diversity