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Recent years have seen a blossoming of theories about the biological and physical basis of consciousness. Good theories guide empirical research, allowing us to interpret data, develop new experimental techniques and expand our capacity to manipulate the phenomenon of interest. Indeed, it is only when couched in terms of a theory that empirical discoveries can ultimately deliver a satisfying understanding of a phenomenon. However, in the case of consciousness, it is unclear how current theories relate to each other, or whether they can be empirically distinguished. To clarify this complicated landscape, we review four prominent theoretical approaches to consciousness: higher-order theories, global workspace theories, re-entry and predictive processing theories and integrated information theory. We describe the key characteristics of each approach by identifying which aspects of consciousness they propose to explain, what their neurobiological commitments are and what empirical data are adduced in their support. We consider how some prominent empirical debates might distinguish among these theories, and we outline three ways in which theories need to be developed to deliver a mature regimen of theory-testing in the neuroscience of consciousness. There are good reasons to think that the iterative development, testing and comparison of theories of consciousness will lead to a deeper understanding of this most profound of mysteries.Various theories have been developed for the biological and physical basis of consciousness. In this Review, Anil Seth and Tim Bayne discuss four prominent theoretical approaches to consciousness, namely higher-order theories, global workspace theories, re-entry and predictive processing theories and integrated information theory.

We review a recent shift in conceptions of interoception and its relationship to hierarchical inference in the brain. The notion of interoceptive inference means that bodily states are regulated by autonomic reflexes that are enslaved by descending predictions from deep generative models of our internal and external milieu. This re-conceptualization illuminates several issues in cognitive and clinical neuroscience with implications for experiences of selfhood and emotion. We first contextualize interoception in terms of active (Bayesian) inference in the brain, highlighting its enactivist (embodied) aspects. We then consider the key role of uncertainty or precision and how this might translate into neuromodulation. We next examine the implications for understanding the functional anatomy of the emotional brain, surveying recent observations on agranular cortex. Finally, we turn to theoretical issues, namely, the role of interoception in shaping a sense of embodied self and feelings. We will draw links between physiological homoeostasis and allostasis, early cybernetic ideas of predictive control and hierarchical generative models in predictive processing. The explanatory scope of interoceptive inference ranges from explanations for autism and depression, through to consciousness. We offer a brief survey of these exciting developments. This article is part of the themed issue 'Interoception beyond homeostasis: affect, cognition and mental health'.

Converging theories suggest that organisms learn and exploit probabilistic models of their environment. However, it remains unclear how such models can be learned in practice. The open-ended complexity of natural environments means that it is generally infeasible for organisms to model their environment comprehensively. Alternatively, action-oriented models attempt to encode a parsimonious representation of adaptive agent-environment interactions. One approach to learning action-oriented models is to learn online in the presence of goal-directed behaviours. This constrains an agent to behaviourally relevant trajectories, reducing the diversity of the data a model need account for. Unfortunately, this approach can cause models to prematurely converge to sub-optimal solutions, through a process we refer to as a bad-bootstrap. Here, we exploit the normative framework of active inference to show that efficient action-oriented models can be learned by balancing goal-oriented and epistemic (information-seeking) behaviours in a principled manner. We illustrate our approach using a simple agent-based model of bacterial chemotaxis. We first demonstrate that learning via goal-directed behaviour indeed constrains models to behaviorally relevant aspects of the environment, but that this approach is prone to sub-optimal convergence. We then demonstrate that epistemic behaviours facilitate the construction of accurate and comprehensive models, but that these models are not tailored to any specific behavioural niche and are therefore less efficient in their use of data. Finally, we show that active inference agents learn models that are parsimonious, tailored to action, and which avoid bad bootstraps and sub-optimal convergence. Critically, our results indicate that models learned through active inference can support adaptive behaviour in spite of, and indeed because of, their departure from veridical representations of the environment. Our approach provides a principled method for learning adaptive models from limited interactions with an environment, highlighting a route to sample efficient learning algorithms.

Predictive coding is an influential model of cortical neural activity. It proposes that perceptual beliefs are furnished by sequentially minimising “prediction errors”—the differences between predicted and observed data. Implicit in this proposal is the idea that successful perception requires multiple cycles of neural activity. This is at odds with evidence that several aspects of visual perception—including complex forms of object recognition—arise from an initial “feedforward sweep” that occurs on fast timescales which preclude substantial recurrent activity. Here, we propose that the feedforward sweep can be understood as performing amortized inference (applying a learned function that maps directly from data to beliefs) and recurrent processing can be understood as performing iterative inference (sequentially updating neural activity in order to improve the accuracy of beliefs). We propose a hybrid predictive coding network that combines both iterative and amortized inference in a principled manner by describing both in terms of a dual optimization of a single objective function. We show that the resulting scheme can be implemented in a biologically plausible neural architecture that approximates Bayesian inference utilising local Hebbian update rules. We demonstrate that our hybrid predictive coding model combines the benefits of both amortized and iterative inference—obtaining rapid and computationally cheap perceptual inference for familiar data while maintaining the context-sensitivity, precision, and sample efficiency of iterative inference schemes. Moreover, we show how our model is inherently sensitive to its uncertainty and adaptively balances iterative and amortized inference to obtain accurate beliefs using minimum computational expense. Hybrid predictive coding offers a new perspective on the functional relevance of the feedforward and recurrent activity observed during visual perception and offers novel insights into distinct aspects of visual phenomenology.

We review the current knowledge about nuclear star clusters (NSCs), the spectacularly dense and massive assemblies of stars found at the centers of most galaxies. Recent observational and theoretical works suggest that many NSC properties, including their masses, densities, and stellar populations, vary with the properties of their host galaxies. Understanding the formation, growth, and ultimate fate of NSCs, therefore, is crucial for a complete picture of galaxy evolution. Throughout the review, we attempt to combine and distill the available evidence into a coherent picture of NSC evolution. Combined, this evidence points to a clear transition mass in galaxies of ∼109M⊙ where the characteristics of nuclear star clusters change. We argue that at lower masses, NSCs are formed primarily from globular clusters that inspiral into the center of the galaxy, while at higher masses, star formation within the nucleus forms the bulk of the NSC. We also discuss the co-existence of NSCs and central black holes, and how their growth may be linked. The extreme densities of NSCs and their interaction with massive black holes lead to a wide range of unique phenomena including tidal disruption and gravitational-wave events. Finally, we review the evidence that many NSCs end up in the halos of massive galaxies stripped of the stars that surrounded them, thus providing valuable tracers of the galaxies’ accretion histories.

Brain–computer interfaces (BCIs) enable direct communication between the brain and external computers, allowing processing of brain activity and the ability to control external devices. While often used for medical purposes, BCIs may also hold great promise for nonmedical purposes to unlock human neurocognitive potential. In this Essay, we discuss the prospects and challenges of using BCIs for cognitive enhancement, focusing specifically on invasive enhancement BCIs (eBCIs). We discuss the ethical, legal, and scientific implications of eBCIs, including issues related to privacy, autonomy, inequality, and the broader societal impact of cognitive enhancement technologies. We conclude that the development of eBCIs raises challenges far beyond practical pros and cons, prompting fundamental questions regarding the nature of conscious selfhood and about who—and what—we are, and ought, to be.

A recent measure of 'integrated information', Φ(DM), quantifies the extent to which a system generates more information than the sum of its parts as it transitions between states, possibly reflecting levels of consciousness generated by neural systems. However, Φ(DM) is defined only for discrete Markov systems, which are unusual in biology; as a result, Φ(DM) can rarely be measured in practice. Here, we describe two new measures, Φ(E) and Φ(AR), that overcome these limitations and are easy to apply to time-series data. We use simulations to demonstrate the in-practice applicability of our measures, and to explore their properties. Our results provide new opportunities for examining information integration in real and model systems and carry implications for relations between integrated information, consciousness, and other neurocognitive processes. However, our findings pose challenges for theories that ascribe physical meaning to the measured quantities.

Integrated Information Theory (IIT) is a prominent theory of consciousness that has at its centre measures that quantify the extent to which a system generates more information than the sum of its parts. While several candidate measures of integrated information (“ Φ ”) now exist, little is known about how they compare, especially in terms of their behaviour on non-trivial network models. In this article, we provide clear and intuitive descriptions of six distinct candidate measures. We then explore the properties of each of these measures in simulation on networks consisting of eight interacting nodes, animated with Gaussian linear autoregressive dynamics. We find a striking diversity in the behaviour of these measures—no two measures show consistent agreement across all analyses. A subset of the measures appears to reflect some form of dynamical complexity, in the sense of simultaneous segregation and integration between system components. Our results help guide the operationalisation of IIT and advance the development of measures of integrated information and dynamical complexity that may have more general applicability.

The experience of authorship over one’s actions and their consequences—sense of agency—is a fundamental aspect of conscious experience. In recent years, it has become common to use intentional binding as an implicit measure of the sense of agency. However, it remains contentious whether reported intentional-binding effects indicate the role of intention-related information in perception or merely represent a strong case of multisensory causal binding. Here, we used a novel virtual-reality setup to demonstrate identical magnitude-binding effects in both the presence and complete absence of intentional action, when perceptual stimuli were matched for temporal and spatial information. Our results demonstrate that intentional-binding-like effects are most simply accounted for by multisensory causal binding without necessarily being related to intention or agency. Future studies that relate binding effects to agency must provide evidence for effects beyond that expected for multisensory causal binding by itself.

The broad concept of emergence is instrumental in various of the most challenging open scientific questions—yet, few quantitative theories of what constitutes emergent phenomena have been proposed. This article introduces a formal theory of causal emergence in multivariate systems, which studies the relationship between the dynamics of parts of a system and macroscopic features of interest. Our theory provides a quantitative definition of downward causation , and introduces a complementary modality of emergent behaviour—which we refer to as causal decoupling . Moreover, the theory allows practical criteria that can be efficiently calculated in large systems, making our framework applicable in a range of scenarios of practical interest. We illustrate our findings in a number of case studies, including Conway’s Game of Life, Reynolds’ flocking model, and neural activity as measured by electrocorticography.