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Jaan Aru

34 papers in the library · 918 citations · publishing 2009-2026

Papers

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Separating the conscious wheat from the unconscious chaff: Towards a biologically grounded computationalism.

Behavioral and Brain Sciences September 17, 2026 Matthew E Larkum, Jaan Aru, Christopher J. Whyte et al.

We agree that consciousness, as we know it, is inextricably biological - but not every biological detail is essential. Identifying which features are necessary, across scales, is vital for assessing the limits of computationalism. This process will reveal a richer, more biologically informed computationalism - one that future machines could, in principle, instantiate, but only after much deeper...

Cellular mechanisms of serotonergic psychedelics - apical hypercontextualisation.

Neuroscience & Biobehavioral Reviews July 1, 2026 Karl Kristjan Kaup, Javier Hidalgo Jiménez, Jaan Aru

Classical serotonergic psychedelics primarily exert their profound effects through agonism at the serotonin 2A (5-HT2A) receptor, which is abundantly expressed in many cortical regions, particularly in layer V pyramidal neurons of associative and visual areas. At the cellular level, these receptors are predominantly localized postsynaptically on the soma and apical dendrites. This review...

Electrophysiological mechanisms of psychedelic drugs: A systematic review.

Neuroscience & Biobehavioral Reviews June 1, 2026 Javier Hidalgo Jiménez, Karl Kristjan Kaup, Jaan Aru 1 citation

Serotonergic psychedelics are known for their profound effects on consciousness and are gaining renewed interest as potential psychiatric treatments. These advances underscore the need to clarify the mechanisms of action of these compounds. This systematic review compiles and critically evaluates 23 in vitro and 26 in vivo electrophysiological studies on psychedelic compounds, with an emphasis...

Loops All the Way Up: Recurrency as an Implementation-first Primitive for Consciousness

PsyArXiv May 29, 2026 Zefan Zheng, Robert Chis-Ciure, Peter Thestrup Waade et al. 1 citation preprint

Consciousness science is fragmented, and empirical investigations are confined within each theory of consciousness’ (TOC) framework. Proliferating ToCs accumulate anomalies without progress, operating orthogonally. The principle of recurrency – functional and architectural – offers a unifying, mechanistic scaffold for consciousness to arise. We show that all theories tacitly invoke feedback...

Effects of LSD, DMT and psilocybin on cognitive and psychological functions: A systematic review of the literature

Journal of Psychopharmacology February 16, 2026 Marten Kase, Karl Kristjan Kaup, Jaan Aru 1 citation

We carried out a systematic review of modern-era (1990-2025) placebo-controlled studies assessing the acute and post-acute effects of lysergic acid diethylamide, dimethyltryptamine and psilocybin on cognitive and psychological functions. From February 28 to March 19, 2025, PubMed and APA PsychINFO were systematically searched for placebo-controlled studies examining the influence of...

On biological and artificial consciousness: A case for biological computationalism.

Neuroscience & Biobehavioral Reviews February 1, 2026 Borjan Milinkovic, Jaan Aru 7 citations

The rapid advances in the capabilities of Large Language Models (LLMs) have galvanised public and scientific debates over whether artificial systems might one day be conscious. Prevailing optimism is often grounded in computational functionalism: the assumption that consciousness is determined solely by the right pattern of information processing, independent of the physical substrate. Opposing...

DOI: 10.1016/j.neubiorev.2025.106524 (opens in new tab) Full text (opens in new tab) Artificial consciousness Artificial intelligence Artificial neural networks Biological computation +8

Realization experiences: a convergent account of insight and mystical experiences

Philosophical Psychology 2026 Juensung J. Kim, Kadi Tulver, Jaan Aru et al. 1 citation

We argue that the powerful transformative effects of mystical-type experiences can be understood using the same machinery that underlies insight problem solving, and that both mystical experiences and insight are species of a genus of “realization experiences”: experiences wherein some phenomenon or piece of information becomes suddenly experienced as real or true. Specifically, we argue that...

Consciousness in Artificial Intelligence? A Framework for Classifying Objections and Constraints

arXiv (Cornell University) November 20, 2025 Andres Campero, Derek Shiller, Jaan Aru et al. preprint

We develop a taxonomical framework for classifying challenges to the possibility of consciousness in digital artificial intelligence systems. This framework allows us to identify the level of granularity at which a given challenge is intended (the levels we propose correspond to Marr's levels) and to disambiguate its degree of force: is it a challenge to computational functionalism that leaves...

The assumptions that restrain us from understanding consciousness

arXiv Preprint Archive June 26, 2025 Jaan Aru preprint

The science of consciousness has been successful over the last decades. Yet, it seems that some of the key questions remain unanswered. Perhaps, as a science of consciousness, we cannot move forward using the same theoretical commitments that brought us here. It might be necessary to revise some assumptions we have made along the way. In this piece, I offer no answers, but I will question some...

Full text (opens in new tab) Consciousness studies Q-bio.nc Consciousness research Science of consciousness +15

Does neural computation feel like something?

Frontiers in Neuroscience 2025 Albert Gidon, Jaan Aru, Matthew E Larkum 1 citation

Artificial neural networks are becoming more advanced and human-like in detail and behavior. The notion that machines mimicking human brain computations might be conscious has recently caused growing unease. Here, we explored a common computational functionalist view, which holds that consciousness emerges when the right computations occur-whether in a machine or a biological brain. To test...

DOI: 10.3389/fnins.2025.1511972 (opens in new tab) Full text (opens in new tab) Computational functionalism Computer simulation Consciousness simulation Counterfactual eraser +12

An integrative, multiscale view on neural theories of consciousness.

Neuron May 15, 2024 Johan F. Storm, P Christiaan Klink, Jaan Aru et al. 108 citations

How is conscious experience related to material brain processes? A variety of theories aiming to answer this age-old question have emerged from the recent surge in consciousness research, and some are now hotly debated. Although most researchers have so far focused on the development and validation of their preferred theory in relative isolation, this article, written by a group of scientists...

The road to Aha: a recipe for mental breakthroughs

February 12, 2024 Kadi Tulver, Kristjan Kaup, Jaan Aru preprint

We present a novel framework for understanding the diverse spectrum of mental breakthrough events, ranging from problem-solving insights to profound personal transformations. We propose that these events, while varied in expression and impact, share common underlying mechanisms of representational change. We also hypothesise that the differences in phenomenological intensity can be...

DOI: 10.31234/osf.io/gq5p6 (opens in new tab) Transformative learning Salience neuroscience Perception Scope computer science +7

The feasibility of artificial consciousness through the lens of neuroscience.

Trends in Neurosciences December 1, 2023 Jaan Aru, Matthew E Larkum, James M Shine

Interactions with large language models (LLMs) have led to the suggestion that these models may soon be conscious. From the perspective of neuroscience, this position is difficult to defend. For one, the inputs to LLMs lack the embodied, embedded information content characteristic of our sensory contact with the world around us. Secondly, the architectures of present-day artificial intelligence...

DOI: 10.1016/j.tins.2023.09.009 (opens in new tab) Full text (opens in new tab) Agency Artificial intelligence Generative ai Large language models +2

A thalamocortical substrate for integrated information via critical synchronous bursting.

Proceedings of the National Academy of Sciences November 14, 2023 Brandon R Munn, Eli J Müller, Jaan Aru et al.

Understanding the neurobiological mechanisms underlying consciousness remains a significant challenge. Recent evidence suggests that the coupling between distal-apical and basal-somatic dendrites in thick-tufted layer 5 pyramidal neurons (L5PN), regulated by the nonspecific-projecting thalamus, is crucial for consciousness. Yet, it is uncertain whether this thalamocortical mechanism can support...

DOI: 10.1073/pnas.2308670120 (opens in new tab) Full text (opens in new tab) Biophysical network Dendrites Integrated information Nonspecific thalamus +1

Facilitating real-life creative insight through psychedelic virtual reality

May 28, 2023 Timo Siimon, Kadi Tulver, Kristjan Kaup et al. 2 citations preprint

The goal of the present work was to study the potential of facilitating creative insight in a real-life setting. To that end, participants were recruited who were struggling with a creative problem and had reached an impasse. Based on previous research, we hypothesized that solutions might be facilitated through virtual reality that replicates visual content experienced under the influence of...

DOI: 10.31234/osf.io/zth7f (opens in new tab) Full text (opens in new tab) Virtual reality Creativity Thematic analysis Human–computer interaction +4

What happens in the brain when we die? Deciphering the neurophysiology of the final moments in life

Frontiers in Aging Neuroscience May 9, 2023 Nathan A. Shlobin, Jaan Aru, Raul Vicente et al. 11 citations

When do we die and what happens in the brain when we die? The mystery around these questions has engaged mankind for centuries. Despite the challenges to obtain recordings of the dying brain, recent studies have contributed to better understand the processes occurring during the last moments of life. In this review, we summarize the literature on neurophysiological changes around the time of...

Psychedelic replications in virtual reality and their potential as a therapeutic instrument: an open-label feasibility study.

Frontiers in Psychiatry March 1, 2023 Karl Kristjan Kaup, Madis Vasser, Kadi Tulver et al. 22 citations

Background: Recent research has shown promising results for the therapeutic benefits of psychedelics. One popular view claims that these benefits are mediated by the subjective experiences induced by these substances. Based on this, we designed a virtual reality experience, Psyrreal, that mimics the phenomenological components of psychedelic experiences. Aims: We aimed to investigate the...

DOI: 10.3389/fpsyt.2023.1088896 (opens in new tab) Full text (opens in new tab) Virtual-reality Psychedelics Mental-health Consciousness-research +1

Conscious interpretation: A distinct aspect for the neural markers of the contents of consciousness.

Consciousness and Cognition February 1, 2023 Talis Bachmann, Jaan Aru

Progress in the science of consciousness depends on the experimental paradigms and varieties of contrastive analysis available to researchers. Here we highlight paradigms where the object is represented in consciousness as a set of its features but the interpretation of this set alternates in consciousness. We group experimental paradigms with this property under the label "conscious...

Corrigendum: Psychedelic replications in virtual reality and their potential as a therapeutic instrument: an open-label feasibility study.

Frontiers in Psychiatry 2023 Karl Kristjan Kaup, Madis Vasser, Kadi Tulver et al. 3 citations correction

Karl Kristjan Kaup1*, Madis Vasser1, Kadi Tulver1, Mari Munk2, Juhan Pikamäe1,3, Jaan Aru1*correcting author names or adding authors. Insert the correct version of the author list)1 Institute of Computer Science, University of Tartu, Estonia2 Psychiatry Clinic of North Estonia Medical Centre3 Institute of Molecular and Cell Biology, University of Tartu, Estonia* Correspondence:...

Understanding the effects of serotonin in the brain through its role in the gastrointestinal tract.

Brain September 14, 2022 James M Shine, Claire O'Callaghan, Ishan C Walpola et al. 63 citations

The neuromodulatory arousal system imbues the nervous system with the flexibility and robustness required to facilitate adaptive behaviour. While there are well understood mechanisms linking dopamine, noradrenaline and acetylcholine to distinct behavioural states, similar conclusions have not been as readily available for serotonin. Fascinatingly, despite clear links between serotonergic...

Does brain activity cause consciousness? A thought experiment

PLoS Biology June 10, 2022 Albert Gidon, Jaan Aru, Matthew E Larkum 20 citations

Rapid advances in neuroscience have provided remarkable breakthroughs in understanding the brain on many fronts. Although promising, the role of these advancements in solving the problem of consciousness is still unclear. Based on technologies conceivably within the grasp of modern neuroscience, we discuss a thought experiment in which neural activity, in the form of action potentials, is...

Restructuring insight: An integrative review of insight in problem-solving, meditation, psychotherapy, delusions and psychedelics

November 26, 2021 Kadi Tulver, Karl Kristjan Kaup, Ruben Laukkonen et al. 16 citations preprint

Occasionally, a solution or idea arrives as a sudden understanding - an insight. Insight has been considered an “extra” ingredient of creative thinking and problem-solving. Here we propose that insight is central in seemingly distinct areas of research. Drawing on literature from a variety of fields, we show that besides being commonly studied in problem-solving literature, insight is also a...

DOI: 10.31234/osf.io/8fbt9 (opens in new tab) Full text (opens in new tab) Meditation Relevance law Psychotherapist Variety cybernetics +7

Cellular Mechanisms of Conscious Processing.

Trends in Cognitive Sciences October 2020 Jaan Aru, Mototaka Suzuki, Matthew E Larkum

Recent breakthroughs in neurobiology indicate that the time is ripe to understand how cellular-level mechanisms are related to conscious experience. Here, we highlight the biophysical properties of pyramidal cells, which allow them to act as gates that control the evolution of global activation patterns. In conscious states, this cellular mechanism enables complex sustained dynamics within the...

DOI: 10.1016/j.tics.2020.07.006 (opens in new tab) Full text (opens in new tab) Anesthesia Dendrites Dendritic integration theory Pyramidal cells

Coupling the State and Contents of Consciousness

Frontiers in Systems Neuroscience August 30, 2019 Jaan Aru, Mototaka Suzuki, Renate Rutiku et al.

One fundamental feature of consciousness is that the contents of consciousness depend on the state of consciousness. Here, we propose an answer to why this is so: both the state and the contents of consciousness depend on the activity of cortical layer 5 pyramidal (L5p) neurons. These neurons affect both cortical and thalamic processing, hence coupling the cortico-cortical and thalamo-cortical...