picture of Prof. Giulio Tononi


Giulio Tononi, MD, PhD

Professor of Psychiatry
David P. White Chair in Sleep Medicine
Distinguished Prof. of Consciousness Science
Director of the Wisconsin Institute for Sleep and Consciousness

University of Wisconsin-Madison
School of Medicine
Department of Psychiatry
6001 Research Park Blvd
Madison, WI 53719


Giulio Tononi received his medical degree from the University of Pisa, Italy, where he specialized in Psychiatry. After serving as a medical officer in the Army, he obtained a Ph.D. in neuroscience as a fellow of the Scuola Superiore for his work on sleep regulation. From 1990 to 2000, he was a member of The Neurosciences Institute, first in New York and then in San Diego. He is currently Professor of Psychiatry, Distinguished Professor in Consciousness Science, the David P. White Chair in Sleep Medicine at the University of Wisconsin–Madison, and the Director of the Wisconsin Institute for Sleep and Consciousness.

Dr. Tononi’s laboratory studies consciousness and its disorders as well as the mechanisms and functions of sleep. For his work on sleep, he has received the NIH Director’s Pioneer Award (2005) and the Harvard Medical School’s Farrell Prize in Sleep Medicine (2017) for his outstanding lifetime contributions to the field. For his work on consciousness, he has received the Max Planck Institute’s Klaus Joachim Zülch Prize (2017) and the Leibniz Institute for Neurobiology’s Leibniz Chair (2018). Dr. Tononi is also the recipient of the Academy of Sleep and Consciousness’s Bernese Sleep Award (2018) and the Humboldt Foundation’s distinguished Humboldt Research Prize (2018).

Dr. Tononi’s main contribution in the study of sleep has been the development of a comprehensive hypothesis about the function of sleep, the synaptic homeostasis hypothesis. According to the hypothesis, sleep serves to renormalize synaptic strength, counterbalancing a net increase of synaptic strength due to plasticity during wakefulness. On the basis of the synaptic homeostasis hypothesis, Dr. Tononi and his long-term collaborator Dr. Chiara Cirelli have shown (using molecular and electrophysiological markers in both invertebrates, rodents, and humans) that synaptic strength does in fact increase during wake and decrease during sleep in much of the brain. Moreover, using a combination of genetic, molecular, and electrophysiological approaches, they have found that sleep need is increased by the amount of plastic changes during wake and that sleep can be induced on a local basis by learning and plasticity. Studies in humans also show that sleep slow waves are important for brain restoration and performance enhancements, with implications for the understanding of sleep disorders (e.g., parasomnias), the neurobiology of mood disorders, and the beneficial effects of sleep deprivation on depression. Dr. Tononi’s laboratory has also demonstrated that patients with schizophrenia show a characteristic reduction of sleep spindles—a hallmark of the sleep EEG—pointing to an impairment of a specific region of the thalamus that could account for many of the symptoms.

Dr. Tononi’s main contribution in the study of consciousness has been the development of the integrated information theory. This is a comprehensive theory of what consciousness is, what determines its quantity and quality, and how it emerges from causal structures such as neural networks. The theory provides a parsimonious account of many neuropsychological observations, among them why certain parts of the brain give rise to experience and others do not, why consciousness vanishes during slow wave sleep and seizures despite continuing neural activity, and how unconscious processes interact with conscious ones. The theory has implications for the unfolding of consciousness across development and phylogeny, and predicts which ingredients are necessary and sufficient to construct sentient machines.

On the basis of the integrated information theory, Dr. Tononi and collaborators have:

1. developed theoretical approaches aimed at defining and measuring the quantity and quality of information integration;
2. constructed large-scale computer models based on the anatomy and physiology of the thalamocortical system to study the mechanisms of information integration;
3. addressed the problem of how the activities of functionally specialized areas of the brain can be integrated to give rise to a unified conscious experience; and
4. pioneered experimental approaches aimed at characterizing the neural substrate of conscious experience by using neuroimaging and transcranial magnetic stimulation.

In several recent experiments, Dr. Tononi and collaborators have shown that the loss of consciousness during slow waves sleep, general anesthesia, and vegetative states is associated with a breakdown of information integration, as predicted by the theory. These approaches may provide an objective marker to evaluate the presence of consciousness and guide rehabilitation and treatment in non-communicating patients.


Massimini M, Tononi G (2018). Sizing up Consciousness: Towards an objective measure of the capacity for experience. Oxford University Press.

Tononi G (2012). Phi: A Voyage from the Brain to the Soul. Random House LLC.

Laureys S, Tononi G (2011). The neurology of consciousness: cognitive neuroscience and neuropathology. Academic Press.

Tononi G (2003). Galileo e il fotodiodo. Bari: Laterza.

Edelman GM, Tononi G (2000). A universe of consciousness: How matter becomes imagination. Basic books.



Marshall, W, Albantakis, L, Tononi, G. “Black-boxing and cause-effect power”. PLoS Comput Biol 2018; 14(4):e1006114.

Nir, Y, Andrillon, T, Marmelshtein, A, Suthana, N, Cirelli, C, Tononi, G, Fried, I. “Selective neuronal lapses precede human cognitive lapses following sleep deprivation”. Nat Med 2017; 23:1474-1480.

Funk, C, Peelman, K, Bellesi, M, Marshall, W, Cirelli, C, Tononi, G. “Role of Somatostatin-Positive Cortical Interneurons in the Generation of Sleep Slow Waves”. The Journal of Neuroscience 2017; 37(38):9132–9148.

Siclari, F, Tononi, G. “Local aspects of sleep and wakefulness”. Current Opinion in Neurobiology 2017; 44:222–227.

de Vivo, L, Bellesi, M, Marshall, W, Bushong, E, Ellisman, M, Tononi, G, Cirelli, C. “Ultrastructural evidence for synaptic scaling across the wake/sleep cycle”. Science 2017; 355(6324):507–510.

Tononi, G, Boly, M, Massimini, M, Koch, C. “Integrated information theory: from consciousness to its physical substrate”. Nature Reviews Neuroscience 2016; 17:450–461.

Koch, C, Massimini, M, Boly, M, Tononi, G. “Neural correlates of consciousness: progress and problems”. Nature Reviews Neuroscience 2016; 17:307–321.

Bellesi, M, de Vivo, L, Tononi, G, Cirelli, C. “Effects of sleep and wake on astrocytes: clues from molecular and ultrastructural studies”. BMC biology 2015; 13(1):66.

Ferrarelli, F, Riedner, B, Peterson, M, Tononi, G. “Altered prefrontal activity and connectivity predict different cognitive deficits in schizophrenia”. Human brain mapping 2015.

Andrillon, T, Nir, Y, Cirelli, C, Tononi, G, Fried, I. “Single-neuron activity and eye movements during human REM sleep and awake vision”. Nature communications 2015; 6.

Bellesi, M, Tononi, G, Cirelli, C, Serra, P. “Region-specific dissociation between cortical noradrenaline levels and the sleep/wake cycle”. Sleep 2015; 39(1):143–154.

Cirelli, C, Tononi, G. “Cortical Development, Electroencephalogram Rhythms, and the Sleep/Wake Cycle”. Biological psychiatry 2015; 77(12):1071–1078.

Tononi, G, Koch, C. “Consciousness: here, there and everywhere?”. Philosophical Transactions of the Royal Society of London B: Biological Sciences 2015; 370(1668):20140167.

Nir, Y, Vyazovskiy, V, Cirelli, C, Banks, M, Tononi, G. “Auditory responses and stimulus-specific adaptation in rat auditory cortex are preserved across NREM and REM sleep”. Cerebral cortex 2015; 25(5):1362–1378.

Bushey, D, Tononi, G, Cirelli, C. “Sleep-and wake-dependent changes in neuronal activity and reactivity demonstrated in fly neurons using in vivo calcium imaging”. Proceedings of the National Academy of Sciences 2015; 112(15):4785–4790.

Bernardi, G, Siclari, F, Yu, X, Zennig, C, Bellesi, M, Ricciardi, E, Cirelli, C, Ghilardi, M, Pietrini, P, Tononi, G. “Neural and behavioral correlates of extended training during sleep deprivation in humans: evidence for local, task-specific effects”. Journal of Neuroscience 2015; 35(11):4487–4500.

Boly, M, Sasai, S, Gosseries, O, Oizumi, M, Casali, A, Massimini, M, Tononi, G. “Stimulus set meaningfulness and neurophysiological differentiation: a functional magnetic resonance imaging study”. PLoS One 2015; 10(5).

Albantakis, L, Hintze, A, Koch, C, Adami, C, Tononi, G. “Evolution of Integrated Causal Structures in Animats Exposed to Environments of Increasing Complexity”. PLoS Computational Biology 2014; 10(12):e1003966.

Dentico, D, Cheung, B, Chang, JY, Guokas, J, Boly, M, Tononi, G, Van Veen, B. “Reversal of cortical information flow during visual imagery as compared to visual perception”. NeuroImage 2014; 100:237–243.

Vyazovskiy, V, Cui, N, Rodriguez, A, Funk, C, Cirelli, C, Tononi, G. “The dynamics of cortical neuronal activity in the first minutes after spontaneous awakening in rats and mice”. Sleep 2014; 37(8):1337.

*Oizumi, M, *Albantakis, L, Tononi, G. “From the Phenomenology to the Mechanisms of Consciousness: Integrated Information Theory 3.0”. PLoS Computational Biology 2014; 10(5):e1003588.

de Vivo, L, Faraguna, U, Nelson, A, Pfister-Genskow, M, Klapperich, M, Tononi, G, Cirelli, C. “Developmental patterns of sleep slow wave activity and synaptic density in adolescent mice”. Sleep 2014; 37(4):689.

Tononi, G, Cirelli, C. “Sleep and the price of plasticity: from synaptic and cellular homeostasis to memory consolidation and integration”. Neuron 2014; 81(1):12–34.

Bellesi, M, Riedner, B, Garcia-Molina, G, Cirelli, C, Tononi, G. “Enhancement of sleep slow waves: underlying mechanisms and practical consequences”. Frontiers in systems neuroscience 2014; 8.

Hoel, E, Albantakis, L, Tononi, G. “Quantifying causal emergence shows that macro can beat micro”. PNAS 2013; 110(49):19790–19795.

Bellesi, M, Pfister-Genskow, M, Maret, S, Keles, S, Tononi, G, Cirelli, C. “Effects of sleep and wake on oligodendrocytes and their precursors”. The Journal of Neuroscience 2013; 33(36):14288–14300.

Tononi, G, Cirelli, C. “Perchance to prune”. Scientific American 2013; 309(2):34–39.

Vyazovskiy, V, Olcese, U, Cirelli, C, Tononi, G. “Prolonged wakefulness alters neuronal responsiveness to local electrical stimulation of the neocortex in awake rats”. Journal of sleep research 2013; 22(3):239–250.

Nelson, A, Faraguna, U, Zoltan, J, Tononi, G, Cirelli, C. “Sleep patterns and homeostatic mechanisms in adolescent mice”. Brain sciences 2013; 3(1):318–343.

Siclari, F, Bernardi, G, Riedner, B, LaRocque, J, Benca, R, Tononi, G. “Two distinct synchronization processes in the transition to sleep: a high-density electroencephalographic study”. Sleep 2013; 37(10).

Balduzzi, D, Tononi, G. “What can neurons do for their brain? Communicate selectivity with bursts”. Theory in Biosciences 2013; 132(1):27–39.

Hashmi, A, Nere, A, Tononi, G. “Sleep-dependent synaptic down-selection (II): single-neuron level benefits for matching, selectivity, and specificity”. Frontiers in neurology 2013; 4.

Nere, A, Hashmi, A, Cirelli, C, Tononi, G. “Sleep-dependent synaptic down-selection (I): modeling the benefits of sleep on memory consolidation and integration”. Frontiers in neurology 2013; 4.

Siclari, F, LaRocque, J, Postle, B, Tononi, G. “Assessing sleep consciousness within subjects using a serial awakening paradigm”. Frontiers in psychology 2013; 4.

Dash, M, Bellesi, M, Tononi, G, Cirelli, C. “Sleep/wake dependent changes in cortical glucose concentrations”. Journal of neurochemistry 2013; 124(1):79–89.

Hung, CS, Sarasso, S, Ferrarelli, F, Riedner, B, Ghilardi, M, Cirelli, C, Tononi, G. “Local experience-dependent changes in the wake EEG after prolonged wakefulness”. Sleep 2013; 36(1):59–72.

Tononi, G. “Integrated information theory of consciousness: an updated account”. Arch Ital Biol 2012; 150(2-3):56–90.

Ferrarelli, F, Sarasso, S, Guller, Y, Riedner, B, Peterson, M, Bellesi, M, Massimini, M, Postle, B, Tononi, G. “Reduced natural oscillatory frequency of frontal thalamocortical circuits in schizophrenia”. Archives of general psychiatry 2012; 69(8):766–774.

Dash, M, Tononi, G, Cirelli, C. “Extracellular levels of lactate, but not oxygen, reflect sleep homeostasis in the rat cerebral cortex”. Sleep 2012; 35(7):909.

Massimini, M, Ferrarelli, F, Sarasso, S, Tononi, G, others. “Cortical mechanisms of loss of consciousness: insight from TMS/EEG studies”. Arch Ital Biol 2012; 150(2-3):44–55.

Tononi, G, Cirelli, C. “Time to be SHY? Some comments on sleep and synaptic homeostasis”. Neural plasticity 2012; 2012.

Nere, A, Olcese, U, Balduzzi, D, Tononi, G. “A neuromorphic architecture for object recognition and motion anticipation using burst-STDP”. PLoS One 2012; 7(5):e36958.

Andrillon, T, Nir, Y, Staba, R, Ferrarelli, F, Cirelli, C, Tononi, G, Fried, I. “Sleep spindles in humans: insights from intracranial EEG and unit recordings”. The Journal of Neuroscience 2011; 31(49):17821–17834.

Maret, S, Faraguna, U, Nelson, A, Cirelli, C, Tononi, G. “Sleep and waking modulate spine turnover in the adolescent mouse cortex”. Nature neuroscience 2011; 14(11):1418–1420.

Bushey, D, Tononi, G, Cirelli, C. “Sleep and synaptic homeostasis: structural evidence in Drosophila”. Science 2011; 332(6037):1576–1581.

Vyazovskiy, V, Olcese, U, Hanlon, E, Nir, Y, Cirelli, C, Tononi, G. “Local sleep in awake rats”. Nature 2011; 472(7344):443–447.

Nir, Y, Staba, R, Andrillon, T, Vyazovskiy, V, Cirelli, C, Fried, I, Tononi, G. “Regional slow waves and spindles in human sleep”. Neuron 2011; 70(1):153–169.

Olcese, U, Esser, S, Tononi, G. “Sleep and synaptic renormalization: a computational study”. Journal of neurophysiology 2010; 104(6):3476–3493.

Ferrarelli, F, Peterson, M, Sarasso, S, Riedner, B, Murphy, M, Benca, R, Bria, P, Kalin, N, Tononi, G. “Thalamic dysfunction in schizophrenia suggested by whole-night deficits in slow and fast spindles”. American Journal of Psychiatry 2010; 167(11):1339–1348.

Ferrarelli, F, Massimini, M, Sarasso, S, Casali, A, Riedner, B, Angelini, G, Tononi, G, Pearce, R. “Breakdown in cortical effective connectivity during midazolam-induced loss of consciousness”. Proceedings of the National Academy of Sciences 2010; 107(6):2681–2686.

Nir, Y, Tononi, G. “Dreaming and the brain: from phenomenology to neurophysiology”. Trends in cognitive sciences 2010; 14(2):88–100.

Massimini, M, Boly, M, Casali, A, Rosanova, M, Tononi, G. “A perturbational approach for evaluating the brain’s capacity for consciousness”. Progress in brain research 2009; 177:201–214.

Vyazovskiy, V, Olcese, U, Lazimy, Y, Faraguna, U, Esser, S, Williams, J, Cirelli, C, Tononi, G. “Cortical firing and sleep homeostasis”. Neuron 2009; 63(6):865–878.

Balduzzi, D, Tononi, G. “Qualia: the geometry of integrated information”. PLoS Computational Biology 2009; 5(8):e1000462.

Landsness, E, Crupi, D, Hulse, B, Peterson, M, Huber, R, Ansari, H, Coen, M, Cirelli, C, Benca, R, Ghilardi, M, others. “Sleep-dependent improvement in visuomotor learning: a causal role for slow waves”. Sleep 2009; 32(10):1273.

Tononi, G. “Consciousness as integrated information: a provisional manifesto”. The Biological Bulletin 2008; 215(3):216–242.

Alkire, M, Hudetz, A, Tononi, G. “Consciousness and anesthesia”. Science 2008; 322(5903):876–880.

Vyazovskiy, V, Cirelli, C, Pfister-Genskow, M, Faraguna, U, Tononi, G. “Molecular and electrophysiological evidence for net synaptic potentiation in wake and depression in sleep”. Nature neuroscience 2008; 11(2):200–208.

Ferrarelli, F, Huber, R, Peterson, M, Massimini, M, Murphy, M, Riedner, B, Watson, A, Bria, P, Tononi, G. “Reduced sleep spindle activity in schizophrenia patients”. American Journal of Psychiatry 2007; 164(3):483–492.

Massimini, M, Ferrarelli, F, Esser, S, Riedner, B, Huber, R, Murphy, M, Peterson, M, Tononi, G. “Triggering sleep slow waves by transcranial magnetic stimulation”. Proceedings of the National Academy of Sciences 2007; 104(20):8496–8501.

Massimini, M, Ferrarelli, F, Huber, R, Esser, S, Singh, H, Tononi, G. “Breakdown of cortical effective connectivity during sleep”. Science 2005; 309(5744):2228–2232.

Cirelli, C, Bushey, D, Hill, S, Huber, R, Kreber, R, Ganetzky, B, Tononi, G. “Reduced sleep in Drosophila Shaker mutants”. Nature 2005; 434(7037):1087–1092.

Tononi, G. “An information integration theory of consciousness”. BMC neuroscience 2004; 5(1):42.

Huber, R, Ghilardi, M, Massimini, M, Tononi, G. “Local sleep and learning”. Nature 2004; 430(6995):78–81.

Tononi, G, Edelman, G. “Consciousness and complexity”. science 1998; 282(5395):1846–1851.