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This paper introduces a novel approach for modelling time-varying connectivity in neuroimaging data, focusing on the slow fluctuations in synaptic efficacy that mediate neuronal dynamics. Building on the framework of Dynamic Causal Modelling (DCM), we propose a method that incorporates temporal basis functions into neural models, allowing for the explicit representation of slow parameter changes. This approach balances expressivity and computational efficiency by modelling these fluctuations as a Gaussian process, offering a middle ground between existing methods that either strongly constrain or excessively relax parameter fluctuations. We validate the ensuing model through simulations and real data from an auditory roving oddball paradigm, demonstrating its potential to explain key aspects of brain dynamics. This work aims to equip researchers with a robust tool for investigating time-varying connectivity, particularly in the context of synaptic modulation and its role in both healthy and pathological brain function., Comment: 17 pages, 3 figures

Macroscopic studies of cortical tissue reveal a prevalence of oscillatory activity, that reflect a fine tuning of neural interactions. This research extends neural field theories by incorporating generalized oscillatory dynamics into previous work on conservative or semi-conservative neural field dynamics. Prior studies have largely assumed isotropic connections among neural units; however, this study demonstrates that a broad range of anisotropic and fluctuating connections can still sustain oscillations. Using Lagrangian field methods, we examine different types of connectivity, their dynamics, and potential interactions with neural fields. From this theoretical foundation, we derive a framework that incorporates Hebbian and non-Hebbian learning, i.e., plasticity, into the study of neural fields via the concept of a connectivity field., Comment: 31 pages, 4 figures

The multifaceted nature of experience poses a challenge to the study of consciousness. Traditional neuroscientific approaches often concentrate on isolated facets, such as perceptual awareness or the global state of consciousness and construct a theory around the relevant empirical paradigms and findings. Theories of consciousness are, therefore, often difficult to compare; indeed, there might be little overlap in the phenomena such theories aim to explain. Here, we take a different approach: starting with active inference, a first principles framework for modelling behaviour as (approximate) Bayesian inference, and building up to a minimal theory of consciousness, which emerges from the shared features of computational models derived under active inference. We review a body of work applying active inference models to the study of consciousness and argue that there is implicit in all these models a small set of theoretical commitments that point to a minimal (and testable) theory of consciousness.

We discuss an approach to mathematically modelling systems made of objects that are coupled together, using generative models of the dependence relationships between states (or trajectories) of the things comprising such systems. This broad class includes open or non-equilibrium systems and is especially relevant to self-organising systems. The ensuing variational free energy principle (FEP) has certain advantages over using random dynamical systems explicitly, notably, by being more tractable and offering a parsimonious explanation of why the joint system evolves in the way that it does, based on the properties of the coupling between system components. Using the FEP allows us to model the dynamics of an object as if it were a process of variational inference, because variational free energy (or surprisal) is a Lyapunov function for its dynamics. In short, we argue that using generative models to represent and track relations among subsystems leads us to a particular statistical theory of interacting systems. Conversely, this theory enables us to construct nested models that respect the known relations among subsystems. We point out that the fact that a physical object conforms to the FEP does not necessarily imply that this object performs inference in the literal sense; rather, it is a useful explanatory fiction which replaces the 'explicit' dynamics of the object with an 'implicit' flow on free energy gradients - a fiction that may or may not be entertained by the object itself., Comment: 25+2 pages

Recent advances in theoretical biology suggest that basal cognition and sentient behaviour are emergent properties of in vitro cell cultures and neuronal networks, respectively. Such neuronal networks spontaneously learn structured behaviours in the absence of reward or reinforcement. In this paper, we characterise this kind of self-organisation through the lens of the free energy principle, i.e., as self-evidencing. We do this by first discussing the definitions of reactive and sentient behaviour in the setting of active inference, which describes the behaviour of agents that model the consequences of their actions. We then introduce a formal account of intentional behaviour, that describes agents as driven by a preferred endpoint or goal in latent state-spaces. We then investigate these forms of (reactive, sentient, and intentional) behaviour using simulations. First, we simulate the aforementioned in vitro experiments, in which neuronal cultures spontaneously learn to play Pong, by implementing nested, free energy minimising processes. The simulations are then used to deconstruct the ensuing predictive behaviour, leading to the distinction between merely reactive, sentient, and intentional behaviour, with the latter formalised in terms of inductive planning. This distinction is further studied using simple machine learning benchmarks (navigation in a grid world and the Tower of Hanoi problem), that show how quickly and efficiently adaptive behaviour emerges under an inductive form of active inference., Comment: 33 pages, 9 figures

This paper concerns structure learning or discovery of discrete generative models. It focuses on Bayesian model selection and the assimilation of training data or content, with a special emphasis on the order in which data are ingested. A key move - in the ensuing schemes - is to place priors on the selection of models, based upon expected free energy. In this setting, expected free energy reduces to a constrained mutual information, where the constraints inherit from priors over outcomes (i.e., preferred outcomes). The resulting scheme is first used to perform image classification on the MNIST dataset to illustrate the basic idea, and then tested on a more challenging problem of discovering models with dynamics, using a simple sprite-based visual disentanglement paradigm and the Tower of Hanoi (cf., blocks world) problem. In these examples, generative models are constructed autodidactically to recover (i.e., disentangle) the factorial structure of latent states - and their characteristic paths or dynamics.

Deep learning's immense capabilities are often constrained by the complexity of its models, leading to an increasing demand for effective sparsification techniques. Bayesian sparsification for deep learning emerges as a crucial approach, facilitating the design of models that are both computationally efficient and competitive in terms of performance across various deep learning applications. The state-of-the-art -- in Bayesian sparsification of deep neural networks -- combines structural shrinkage priors on model weights with an approximate inference scheme based on stochastic variational inference. However, model inversion of the full generative model is exceptionally computationally demanding, especially when compared to standard deep learning of point estimates. In this context, we advocate for the use of Bayesian model reduction (BMR) as a more efficient alternative for pruning of model weights. As a generalization of the Savage-Dickey ratio, BMR allows a post-hoc elimination of redundant model weights based on the posterior estimates under a straightforward (non-hierarchical) generative model. Our comparative study highlights the advantages of the BMR method relative to established approaches based on hierarchical horseshoe priors over model weights. We illustrate the potential of BMR across various deep learning architectures, from classical networks like LeNet to modern frameworks such as Vision Transformers and MLP-Mixers.

Building autonomous --- i.e., choosing goals based on one's needs -- and adaptive -- i.e., surviving in ever-changing environments -- agents has been a holy grail of artificial intelligence (AI). A living organism is a prime example of such an agent, offering important lessons about adaptive autonomy. Here, we focus on interoception, a process of monitoring one's internal environment to keep it within certain bounds, which underwrites the survival of an organism. To develop AI with interoception, we need to factorize the state variables representing internal environments from external environments and adopt life-inspired mathematical properties of internal environment states. This paper offers a new perspective on how interoception can help build autonomous and adaptive agents by integrating the legacy of cybernetics with recent advances in theories of life, reinforcement learning, and neuroscience., Comment: 28 pages, 4 figures, 3 boxes

A pervasive challenge in neuroscience is testing whether neuronal connectivity changes over time due to specific causes, such as stimuli, events, or clinical interventions. Recent hardware innovations and falling data storage costs enable longer, more naturalistic neuronal recordings. The implicit opportunity for understanding the self-organised brain calls for new analysis methods that link temporal scales: from the order of milliseconds over which neuronal dynamics evolve, to the order of minutes, days or even years over which experimental observations unfold. This review article demonstrates how hierarchical generative models and Bayesian inference help to characterise neuronal activity across different time scales. Crucially, these methods go beyond describing statistical associations among observations and enable inference about underlying mechanisms. We offer an overview of fundamental concepts in state-space modeling and suggest a taxonomy for these methods. Additionally, we introduce key mathematical principles that underscore a separation of temporal scales, such as the slaving principle, and review Bayesian methods that are being used to test hypotheses about the brain with multi-scale data. We hope that this review will serve as a useful primer for experimental and computational neuroscientists on the state of the art and current directions of travel in the complex systems modelling literature., Comment: 20 pages, 5 figures

We consider the theoretical constraints on interactions between coupled cortical columns. Each column comprises a set of neural populations, where each population is modelled as a neural mass. The existence of semi-stable states within a cortical column has been shown to be dependent on the type of interaction between the constituent neuronal subpopulations, i.e., the form of the implicit synaptic convolution kernels. Current-to-current coupling has been shown, in contrast to potential-to-current coupling, to create semi-stable states within a cortical column. In this analytic and numerical study, the interaction between semi-stable states is characterized by equations of motion for ensemble activity. We show that for small excitations, the dynamics follow the Kuramoto model. However, in contrast to previous work, we derive coupled equations between phase and amplitude dynamics. This affords the possibility of defining connectivity as a dynamic variable. The turbulent flow of phase dynamics found in networks of Kuramoto oscillators indicate turbulent changes in dynamic connectivity for coupled cortical columns. Crucially, this is something that has been recorded in epileptic seizures. We used the results we derived to estimate a seizure propagation model, which allowed for relatively straightforward inversions using variational Laplace (a.k.a., Dynamic Causal Modelling). The face validity of the ensuing seizure propagation model was established using simulated data as a prelude to future work; which will investigate dynamic connectivity from empirical data. This model also allows predictions of seizure evolution, induced by virtual lesions to synaptic connectivity: something that could be of clinical use, when applied to epilepsy surgical cases.

This white paper lays out a vision of research and development in the field of artificial intelligence for the next decade (and beyond). Its denouement is a cyber-physical ecosystem of natural and synthetic sense-making, in which humans are integral participants -- what we call ''shared intelligence''. This vision is premised on active inference, a formulation of adaptive behavior that can be read as a physics of intelligence, and which inherits from the physics of self-organization. In this context, we understand intelligence as the capacity to accumulate evidence for a generative model of one's sensed world -- also known as self-evidencing. Formally, this corresponds to maximizing (Bayesian) model evidence, via belief updating over several scales: i.e., inference, learning, and model selection. Operationally, this self-evidencing can be realized via (variational) message passing or belief propagation on a factor graph. Crucially, active inference foregrounds an existential imperative of intelligent systems; namely, curiosity or the resolution of uncertainty. This same imperative underwrites belief sharing in ensembles of agents, in which certain aspects (i.e., factors) of each agent's generative world model provide a common ground or frame of reference. Active inference plays a foundational role in this ecology of belief sharing -- leading to a formal account of collective intelligence that rests on shared narratives and goals. We also consider the kinds of communication protocols that must be developed to enable such an ecosystem of intelligences and motivate the development of a shared hyper-spatial modeling language and transaction protocol, as a first -- and key -- step towards such an ecology., Comment: 23+18 pages, one figure, one six page appendix

This paper presents a meta-theory of the usage of the free energy principle (FEP) and examines its scope in the modelling of physical systems. We consider the so-called `map-territory fallacy' and the fallacious reification of model properties. By showing that the FEP is a consistent, physics-inspired theory of inferences of inferences, we disprove the assertion that the map-territory fallacy contradicts the principled usage of the FEP. As such, we argue that deploying the map-territory fallacy to criticise the use of the FEP and Bayesian mechanics itself constitutes a fallacy: what we call the {\it map-territory fallacy fallacy}. In so doing, we emphasise a few key points: the uniqueness of the FEP as a model of particles or agents that model their environments; the restoration of convention to the FEP via its relation to the principle of constrained maximum entropy; the `Jaynes optimality' of the FEP under this relation; and finally, the way that this meta-theoretical approach to the FEP clarifies its utility and scope as a formal modelling tool. Taken together, these features make the FEP, uniquely, {\it the} ideal model of generic systems in statistical physics., Comment: 23 pages

The aim of this paper is to introduce a field of study that has emerged over the last decade called Bayesian mechanics. Bayesian mechanics is a probabilistic mechanics, comprising tools that enable us to model systems endowed with a particular partition (i.e., into particles), where the internal states (or the trajectories of internal states) of a particular system encode the parameters of beliefs about external states (or their trajectories). These tools allow us to write down mechanical theories for systems that look as if they are estimating posterior probability distributions over the causes of their sensory states. This provides a formal language for modelling the constraints, forces, potentials, and other quantities determining the dynamics of such systems, especially as they entail dynamics on a space of beliefs (i.e., on a statistical manifold). Here, we will review the state of the art in the literature on the free energy principle, distinguishing between three ways in which Bayesian mechanics has been applied to particular systems (i.e., path-tracking, mode-tracking, and mode-matching). We go on to examine a duality between the free energy principle and the constrained maximum entropy principle, both of which lie at the heart of Bayesian mechanics, and discuss its implications., Comment: 50+7 pages, six figures. Invited contribution to "Making and Breaking Symmetries in Mind and Life," a special issue of the journal Royal Society Interface. First two authors have contributed equally

How is information processed in the brain during perception? Mechanistic insight is achieved only when experiments are employed to test formal or computational models. In analogy to lesion studies, phantom perception may serve as a vehicle to understand the fundamental processing principles underlying auditory perception. With a special focus on tinnitus -- as the prime example of auditory phantom perception -- we review recent work at the intersection of artificial intelligence, psychology, and neuroscience. In particular, we discuss why everyone with tinnitus suffers from hearing loss, but not everyone with hearing loss suffers from tinnitus. We argue that the increase of sensory precision due to Bayesian inference could be caused by intrinsic neural noise and lead to a prediction error in the cerebral cortex. Hence, two fundamental processing principles - being ubiquitous in the brain - provide the most explanatory power for the emergence of tinnitus: predictive coding as a top-down, and stochastic resonance as a complementary bottom-up mechanism. We conclude that both principles play a crucial role in healthy auditory perception., Comment: arXiv admin note: substantial text overlap with arXiv:2010.01914

We apply Dynamic Causal Models to electrocorticogram recordings from two macaque monkeys performing a problem-solving task that engages working memory, and induces time-on-task effects. We thus provide a computational account of changes in effective connectivity within two regions of the fronto-parietal network, the dorsolateral prefrontal cortex and the pre-supplementary motor area. We find that forward connections between the two regions increased in strength when task demands increased, and as the experimental session progressed. Similarities in the effects of task demands and time on task allow us to interpret changes in frontal connectivity in terms of increased attentional effort allocation that compensates cognitive fatigue.

To interact seamlessly with robots, users must infer the causes of a robot's behavior and be confident about that inference. Hence, trust is a necessary condition for human-robot collaboration (HRC). Despite its crucial role, it is largely unknown how trust emerges, develops, and supports human interactions with nonhuman artefacts. Here, we review the literature on trust, human-robot interaction, human-robot collaboration, and human interaction at large. Early models of trust suggest that trust entails a trade-off between benevolence and competence, while studies of human-to-human interaction emphasize the role of shared behavior and mutual knowledge in the gradual building of trust. We then introduce a model of trust as an agent's best explanation for reliable sensory exchange with an extended motor plant or partner. This model is based on the cognitive neuroscience of active inference and suggests that, in the context of HRC, trust can be cast in terms of virtual control over an artificial agent. In this setting, interactive feedback becomes a necessary component of the trustor's perception-action cycle. The resulting model has important implications for understanding human-robot interaction and collaboration, as it allows the traditional determinants of human trust to be defined in terms of active inference, information exchange and empowerment. Furthermore, this model suggests that boredom and surprise may be used as markers for under and over-reliance on the system. Finally, we examine the role of shared behavior in the genesis of trust, especially in the context of dyadic collaboration, suggesting important consequences for the acceptability and design of human-robot collaborative systems.

The evolution of epidemiological parameters, such as instantaneous reproduction number Rt, is important for understanding the transmission dynamics of infectious diseases. Current estimates of time-varying epidemiological parameters often face problems such as lagging observations, averaging inference, and improper quantification of uncertainties. To address these problems, we propose a Bayesian data assimilation framework for time-varying parameter estimation. Specifically, this framework is applied to Rt estimation, resulting in the state-of-the-art DARt system. With DARt, time misalignment caused by lagging observations is tackled by incorporating observation delays into the joint inference of infections and Rt; the drawback of averaging is overcome by instantaneously updating upon new observations and developing a model selection mechanism that captures abrupt changes; the uncertainty is quantified and reduced by employing Bayesian smoothing. We validate the performance of DARt and demonstrate its power in revealing the transmission dynamics of COVID-19. The proposed approach provides a promising solution for accurate and timely estimating transmission dynamics from reported data., Comment: Xian Yang, Shuo Wang and Yuting Xing contribute equally

This technical report describes the rationale and technical details for the dynamic causal modelling of mitigated epidemiological outcomes based upon a variety of timeseries data. It details the structure of the underlying convolution or generative model (at the time of writing on 6-Nov-20). This report is intended for use as a reference that accompanies the predictions in following dashboard: https://www.fil.ion.ucl.ac.uk/spm/covid-19/dashboard

The principle of stationary action is a cornerstone of modern physics, providing a powerful framework for investigating dynamical systems found in classical mechanics through to quantum field theory. However, computational neuroscience, despite its heavy reliance on concepts in physics, is anomalous in this regard as its main equations of motion are not compatible with a Lagrangian formulation and hence with the principle of stationary action. Taking the Dynamic Causal Modelling neuronal state equation, Hodgkin-Huxley model, and the Leaky Integrate-and-Fire model as examples, we show that it is possible to write complex oscillatory forms of these equations in terms of a single Lagrangian. We therefore bring mathematical descriptions in computational neuroscience under the remit of the principle of stationary action and use this reformulation to express symmetries and associated conservation laws arising in neural systems.

An interesting inference drawn by some Covid-19 epidemiological models is that there exists a proportion of the population who are not susceptible to infection -- even at the start of the current pandemic. This paper introduces a model of the immune response to a virus. This is based upon the same sort of mean-field dynamics as used in epidemiology. However, in place of the location, clinical status, and other attributes of people in an epidemiological model, we consider the state of a virus, B and T-lymphocytes, and the antibodies they generate. Our aim is to formalise some key hypotheses as to the mechanism of resistance. We present a series of simple simulations illustrating changes to the dynamics of the immune response under these hypotheses. These include attenuated viral cell entry, pre-existing cross-reactive humoral (antibody-mediated) immunity, and enhanced T-cell dependent immunity. Finally, we illustrate the potential application of this sort of model by illustrating variational inversion (using simulated data) of this model to illustrate its use in testing hypotheses. In principle, this furnishes a fast and efficient immunological assay--based on sequential serology--that provides a (i) quantitative measure of latent immunological responses and (ii) a Bayes optimal classification of the different kinds of immunological response (c.f., glucose tolerance tests used to test for insulin resistance). This may be especially useful in assessing SARS-CoV-2 vaccines.

This technical note introduces parametric dynamic causal modelling, a method for inferring slow changes in biophysical parameters that control fluctuations of fast neuronal states. The application domain we have in mind is inferring slow changes in variables (e.g., extracellular ion concentrations or synaptic efficacy) that underlie phase transitions in brain activity (e.g., paroxysmal seizure activity). The scheme is efficient and yet retains a biophysical interpretation, in virtue of being based on established neural mass models that are equipped with a slow dynamic on the parameters (such as synaptic rate constants or effective connectivity). In brief, we use an adiabatic approximation to summarise fast fluctuations in hidden neuronal states (and their expression in sensors) in terms of their second order statistics; namely, their complex cross spectra. This allows one to specify and compare models of slowly changing parameters (using Bayesian model reduction) that generate a sequence of empirical cross spectra of electrophysiological recordings. Crucially, we use the slow fluctuations in the spectral power of neuronal activity as empirical priors on changes in synaptic parameters. This introduces a circular causality, in which synaptic parameters underwrite fast neuronal activity that, in turn, induces activity-dependent plasticity in synaptic parameters. In this foundational paper, we describe the underlying model, establish its face validity using simulations and provide an illustrative application to a chemoconvulsant animal model of seizure activity.

At the inception of human brain mapping, two principles of functional anatomy underwrote most conceptions - and analyses - of distributed brain responses: namely functional segregation and integration. There are currently two main approaches to characterising functional integration. The first is a mechanistic modelling of connectomics in terms of directed effective connectivity that mediates neuronal message passing and dynamics on neuronal circuits. The second phenomenological approach usually characterises undirected functional connectivity (i.e., measurable correlations), in terms of intrinsic brain networks, self-organised criticality, dynamical instability, etc. This paper describes a treatment of effective connectivity that speaks to the emergence of intrinsic brain networks and critical dynamics. It is predicated on the notion of Markov blankets that play a fundamental role in the self-organisation of far from equilibrium systems. Using the apparatus of the renormalisation group, we show that much of the phenomenology found in network neuroscience is an emergent property of a particular partition of neuronal states, over progressively larger scales. As such, it offers a way of linking dynamics on directed graphs to the phenomenology of intrinsic brain networks.

This technical report addresses a pressing issue in the trajectory of the coronavirus outbreak; namely, the rate at which effective immunity is lost following the first wave of the pandemic. This is a crucial epidemiological parameter that speaks to both the consequences of relaxing lockdown and the propensity for a second wave of infections. Using a dynamic causal model of reported cases and deaths from multiple countries, we evaluated the evidence models of progressively longer periods of immunity. The results speak to an effective population immunity of about three months that, under the model, defers any second wave for approximately six months in most countries. This may have implications for the window of opportunity for tracking and tracing, as well as for developing vaccination programmes, and other therapeutic interventions., Comment: 20 pages, 8 figures, 3 tables (technical report)

By equipping a previously reported dynamic causal model of COVID-19 with an isolation state, we modelled the effects of self-isolation consequent on tracking and tracing. Specifically, we included a quarantine or isolation state occupied by people who believe they might be infected but are asymptomatic, and only leave if they test negative. We recovered maximum posteriori estimates of the model parameters using time series of new cases, daily deaths, and tests for the UK. These parameters were used to simulate the trajectory of the outbreak in the UK over an 18-month period. Several clear-cut conclusions emerged from these simulations. For example, under plausible (graded) relaxations of social distancing, a rebound of infections within weeks is unlikely. The emergence of a later second wave depends almost exclusively on the rate at which we lose immunity, inherited from the first wave. There exists no testing strategy that can attenuate mortality rates, other than by deferring or delaying a second wave. A sufficiently powerful tracking and tracing policy--implemented at the time of writing (10th May 2020)--will defer any second wave beyond a time horizon of 18 months. Crucially, this deferment is within current testing capabilities (requiring an efficacy of tracing and tracking of about 20% of asymptomatic infected cases, with less than 50,000 tests per day). These conclusions are based upon a dynamic causal model for which we provide some construct and face validation, using a comparative analysis of the United Kingdom and Germany, supplemented with recent serological studies., Comment: 31 pages, 12 figures, one table. Technical report

Seizure activity is a ubiquitous and pernicious pathophysiology that, in principle, should yield to mathematical treatments of (neuronal) ensemble dynamics - and therefore interventions on stochastic chaos. A seizure can be characterised as a deviation of neural activity from a stable dynamical regime, i.e. one in which signals fluctuate only within a limited range. In silico treatments of neural activity are an important tool for understanding how the brain can achieve stability, as well as how pathology can lead to seizures and potential strategies for mitigating instabilities, e.g. via external stimulation. Here, we demonstrate that the (neuronal) state equation used in Dynamic Causal Modelling generalises to a Fokker-Planck formalism when propagation of neuronal activity along structural connections is considered. Using the Jacobian of this generalised state equation, we show that an initially unstable system can be rendered stable via a reduction in diffusivity (i.e., connectivity that disperses neuronal fluctuations). We show, for neural systems prone to epileptic seizures, that such a reduction can be achieved via external stimulation. Specifically, we show that this stimulation should be applied in such a way as to temporarily mirror epileptic activity in the areas adjoining an affected brain region - thus 'fighting seizures with seizures'. We offer proof of principle using simulations based on functional neuroimaging data collected from patients with idiopathic generalised epilepsy, in which we successfully suppress pathological activity in a distinct sub-network. Our hope is that this technique can form the basis for real-time monitoring and intervention devices that are capable of suppressing or even preventing seizures in a non-invasive manner.

We recently described a dynamic causal model of a COVID-19 outbreak within a single region. Here, we combine several of these (epidemic) models to create a (pandemic) model of viral spread among regions. Our focus is on a second wave of new cases that may result from loss of immunity--and the exchange of people between regions--and how mortality rates can be ameliorated under different strategic responses. In particular, we consider hard or soft social distancing strategies predicated on national (Federal) or regional (State) estimates of the prevalence of infection in the population. The modelling is demonstrated using timeseries of new cases and deaths from the United States to estimate the parameters of a factorial (compartmental) epidemiological model of each State and, crucially, coupling between States. Using Bayesian model reduction, we identify the effective connectivity between States that best explains the initial phases of the outbreak in the United States. Using the ensuing posterior parameter estimates, we then evaluate the likely outcomes of different policies in terms of mortality, working days lost due to lockdown and demands upon critical care. The provisional results of this modelling suggest that social distancing and loss of immunity are the two key factors that underwrite a return to endemic equilibrium., Comment: Technical report: 35 pages, 14 figures, 1 table

This technical report describes a dynamic causal model of the spread of coronavirus through a population. The model is based upon ensemble or population dynamics that generate outcomes, like new cases and deaths over time. The purpose of this model is to quantify the uncertainty that attends predictions of relevant outcomes. By assuming suitable conditional dependencies, one can model the effects of interventions (e.g., social distancing) and differences among populations (e.g., herd immunity) to predict what might happen in different circumstances. Technically, this model leverages state-of-the-art variational (Bayesian) model inversion and comparison procedures, originally developed to characterise the responses of neuronal ensembles to perturbations. Here, this modelling is applied to epidemiological populations to illustrate the kind of inferences that are supported and how the model per se can be optimised given timeseries data. Although the purpose of this paper is to describe a modelling protocol, the results illustrate some interesting perspectives on the current pandemic; for example, the nonlinear effects of herd immunity that speak to a self-organised mitigation process., Comment: Technical report: 40 pages, 13 figures and 2 tables

We used Bayesian model inversion to estimate epidemic parameters from the reported case and death rates from seven countries using data from late January 2020 to April 5th 2020. Two distinct generative model types were employed: first a continuous time dynamical-systems implementation of a Susceptible-Exposed-Infectious-Recovered (SEIR) model and second: a partially observable Markov Decision Process (MDP) or hidden Markov model (HMM) implementation of an SEIR model. Both models parameterise the size of the initial susceptible population (S0), as well as epidemic parameters. Parameter estimation (data fitting) was performed using a standard Bayesian scheme (variational Laplace) designed to allow for latent unobservable states and uncertainty in model parameters. Both models recapitulated the dynamics of transmissions and disease as given by case and death rates. The peaks of the current waves were predicted to be in the past for four countries (Italy, Spain, Germany and Switzerland) and to emerge in 0.5-2 weeks in Ireland and 1-3 weeks in the UK. For France one model estimated the peak within the past week and the other in the future in two weeks. Crucially, Maximum a posteriori (MAP) estimates of S0 for each country indicated effective population sizes of below 20% (of total population size), under both the continuous time and HMM models. With a Bayesian weighted average across all seven countries and both models, we estimated that 6.4% of the total population would be immune. From the two models the maximum percentage of the effective population was estimated at 19.6% of the total population for the UK, 16.7% for Ireland, 11.4% for Italy, 12.8% for Spain, 18.8% for France, 4.7% for Germany and 12.9% for Switzerland. Our results indicate that after the current wave, a large proportion of the total population will remain without immunity.

In contrast to the symmetries of translation in space, rotation in space, and translation in time, the known laws of physics are not universally invariant under transformation of scale. However, the action can be invariant under change of scale in the special case of a scale free dynamical system that can be described in terms of a Lagrangian, that itself scales inversely with time. Crucially, this means symmetries under change of scale can exist in dynamical systems under certain constraints. Our contribution lies in the derivation of a generalised scale invariant Lagrangian - in the form of a power series expansion - that satisfies these constraints. This generalised Lagrangian furnishes a normal form for dynamic causal models (i.e., state space models based upon differential equations) that can be used to distinguish scale invariance (scale symmetry) from scale freeness in empirical data. We establish face validity with an analysis of simulated data and then show how scale invariance can be identified - and how the associated conserved quantities can be estimated - in neuronal timeseries.

Active inference is a first principle account of how autonomous agents operate in dynamic, non-stationary environments. This problem is also considered in reinforcement learning (RL), but limited work exists on comparing the two approaches on the same discrete-state environments. In this paper, we provide: 1) an accessible overview of the discrete-state formulation of active inference, highlighting natural behaviors in active inference that are generally engineered in RL; 2) an explicit discrete-state comparison between active inference and RL on an OpenAI gym baseline. We begin by providing a condensed overview of the active inference literature, in particular viewing the various natural behaviors of active inference agents through the lens of RL. We show that by operating in a pure belief-based setting, active inference agents can carry out epistemic exploration, and account for uncertainty about their environment in a Bayes-optimal fashion. Furthermore, we show that the reliance on an explicit reward signal in RL is removed in active inference, where reward can simply be treated as another observation; even in the total absence of rewards, agent behaviors are learned through preference learning. We make these properties explicit by showing two scenarios in which active inference agents can infer behaviors in reward-free environments compared to both Q-learning and Bayesian model-based RL agents; by placing zero prior preferences over rewards and by learning the prior preferences over the observations corresponding to reward. We conclude by noting that this formalism can be applied to more complex settings if appropriate generative models can be formulated. In short, we aim to demystify the behavior of active inference agents by presenting an accessible discrete state-space and time formulation, and demonstrate these behaviors in a OpenAI gym environment, alongside RL agents.

Models of coupled oscillators are used to describe a wide variety of phenomena in neuroimaging. These models typically rest on the premise that oscillator dynamics do not evolve beyond their respective limit cycles, and hence that interactions can be described purely in terms of phase differences. Whilst mathematically convenient, the restrictive nature of phase-only models can limit their explanatory power. We therefore propose a generalisation of dynamic causal modelling that incorporates both phase and amplitude. This allows for the separate quantifications of phase and amplitude contributions to the connectivity between neural regions. We establish, using model-generated data and simulations of coupled pendula, that phase-only models perform well only under weak coupling conditions. We also show that, despite their higher complexity, phase-amplitude models can describe strongly coupled systems more effectively than their phase-only counterparts. We relate our findings to four metrics commonly used in neuroimaging: the Kuramoto order parameter, cross-correlation, phase-lag index, and spectral entropy. We find that, with the exception of spectral entropy, the phase-amplitude model is able to capture all metrics more effectively than the phase-only model. We then demonstrate, using local field potential recordings in rodents and functional magnetic resonance imaging in macaque monkeys, that amplitudes in oscillator models play an important role in describing neural dynamics in anaesthetised brain states., Comment: arXiv admin note: text overlap with arXiv:1812.06315

Scale free dynamics are observed in a variety of physical and biological systems. These include neural activity in which evidence for scale freeness has been reported using a range of imaging modalities. Here, we derive the ways in which connections within a network must transform - relative to system size - in order to maintain scale freeness and test these theoretical transformations via simulations. First, we explore the known invariance of planetary motion for orbits varying in size. Using parametric empirical Bayesian modelling and a generic dynamical systems model, we show that we recover Kepler's third law from orbital timeseries, using our proposed transformations; thereby providing construct validation. We then demonstrate that the dynamical critical exponent is inversely proportional to the time rescaling exponent, in the context of coarse graining operations. Using murine calcium imaging data, we then show that the dynamical critical exponent can be estimated in an empirical biological setting. Specifically, we compare dynamical critical exponents - associated with spontaneous and task states in two regions of imaged cortex - that are classified as task-relevant and task-irrelevant. We find, consistently across animals, that the task-irrelevant region exhibits higher dynamical critical exponents during spontaneous activity than during task performance. Conversely, the task-relevant region is associated with higher dynamical critical exponents in task vs. spontaneous states. These data support the idea that higher dynamical critical exponents, within relevant cortical structures, underwrite neuronal processing due to the implicit increase in cross-scale information transmission.

This paper asks whether integrating multimodal EEG and fMRI data offers a better characterisation of functional brain architectures than either modality alone. This evaluation rests upon a dynamic causal model that generates both EEG and fMRI data from the same neuronal dynamics. We introduce the use of Bayesian fusion to provide informative (empirical) neuronal priors - derived from dynamic causal modelling (DCM) of EEG data - for subsequent DCM of fMRI data. To illustrate this procedure, we generated synthetic EEG and fMRI timeseries for a mismatch negativity (or auditory oddball) paradigm, using biologically plausible model parameters (i.e., posterior expectations from a DCM of empirical, open access, EEG data). Using model inversion, we found that Bayesian fusion provided a substantial improvement in marginal likelihood or model evidence, indicating a more efficient estimation of model parameters, in relation to inverting fMRI data alone. We quantified the benefits of multimodal fusion with the information gain pertaining to neuronal and haemodynamic parameters - as measured by the Kullback-Leibler divergence between their prior and posterior densities. Remarkably, this analysis suggested that EEG data can improve estimates of haemodynamic parameters; thereby furnishing proof-of-principle that Bayesian fusion of EEG and fMRI is necessary to resolve conditional dependencies between neuronal and haemodynamic estimators. These results suggest that Bayesian fusion may offer a useful approach that exploits the complementary temporal (EEG) and spatial (fMRI) precision of different data modalities. We envisage the procedure could be applied to any multimodal dataset that can be explained by a DCM with a common neuronal parameterisation.

This technical note presents a framework for investigating the underlying mechanisms of neurovascular coupling in the human brain using multi-modal magnetoencephalography (MEG) and functional magnetic resonance (fMRI) neuroimaging data. This amounts to estimating the evidence for several biologically informed models of neurovascular coupling using variational Bayesian methods and selecting the most plausible explanation using Bayesian model comparison. First, fMRI data is used to localise active neuronal sources. The coordinates of neuronal sources are then used as priors in the specification of a DCM for MEG, in order to estimate the underlying generators of the electrophysiological responses. The ensuing estimates of neuronal parameters are used to generate neuronal drive functions, which model the pre or post synaptic responses to each experimental condition in the fMRI paradigm. These functions form the input to a model of neurovascular coupling, the parameters of which are estimated from the fMRI data. This establishes a Bayesian fusion technique that characterises the BOLD response - asking, for example, whether instantaneous or delayed pre or post synaptic signals mediate haemodynamic responses. Bayesian model comparison is used to identify the most plausible hypotheses about the causes of the multimodal data. We illustrate this procedure by comparing a set of models of a single-subject auditory fMRI and MEG dataset. Our exemplar analysis suggests that the origin of the BOLD signal is mediated instantaneously by intrinsic neuronal dynamics and that neurovascular coupling mechanisms are region-specific. The code and example dataset associated with this technical note are available through the statistical parametric mapping (SPM) software package.

Dynamic Causal Modelling (DCM) is the predominant method for inferring effective connectivity from neuroimaging data. In the 15 years since its introduction, the neural models and statistical routines in DCM have developed in parallel, driven by the needs of researchers in cognitive and clinical neuroscience. In this tutorial, we step through an exemplar fMRI analysis in detail, reviewing the current implementation of DCM and demonstrating recent developments in group-level connectivity analysis. In the first part of the tutorial (current paper), we focus on issues specific to DCM for fMRI, unpacking the relevant theory and highlighting practical considerations. In particular, we clarify the assumptions (i.e., priors) used in DCM for fMRI and how to interpret the model parameters. This tutorial is accompanied by all the necessary data and instructions to reproduce the analyses using the SPM software. In the second part (in a companion paper), we move from subject-level to group-level modelling using the Parametric Empirical Bayes framework, and illustrate how to test for commonalities and differences in effective connectivity across subjects, based on imaging data from any modality.

This tutorial provides a worked example of using Dynamic Causal Modelling (DCM) and Parametric Empirical Bayes (PEB) to characterise inter-subject variability in neural circuitry (effective connectivity). This involves specifying a hierarchical model with two or more levels. At the first level, state space models (DCMs) are used to infer the effective connectivity that best explains a subject's neuroimaging timeseries (e.g. fMRI, MEG, EEG). Subject-specific connectivity parameters are then taken to the group level, where they are modelled using a General Linear Model (GLM) that partitions between-subject variability into designed effects and additive random effects. The ensuing (Bayesian) hierarchical model conveys both the estimated connection strengths and their uncertainty (i.e., posterior covariance) from the subject to the group level; enabling hypotheses to be tested about the commonalities and differences across subjects. This approach can also finesse parameter estimation at the subject level, by using the group-level parameters as empirical priors. We walk through this approach in detail, using data from a published fMRI experiment that characterised individual differences in hemispheric lateralization in a semantic processing task. The preliminary subject specific DCM analysis is covered in detail in a companion paper. This tutorial is accompanied by the example dataset and step-by-step instructions to reproduce the analyses.

Models of coupled oscillators are useful in describing a wide variety of phenomena in physics, biology and economics. These models typically rest on the premise that the oscillators are weakly coupled, meaning that amplitudes can be assumed to be constant and dynamics can therefore be described purely in terms of phase differences. Whilst mathematically convenient, the restrictive nature of the weak coupling assumption can limit the explanatory power of these phase-coupled oscillator models. We therefore propose an extension to the weakly-coupled oscillator model that incorporates both amplitude and phase as dependent variables. We use the bilinear neuronal state equations of dynamic causal modelling as a foundation in deriving coupled differential equations that describe the activity of both weakly and strongly coupled oscillators. We show that weakly-coupled oscillator models are inadequate in describing the processes underlying the temporally variable signals observed in a variety of systems. We demonstrate that phase-coupled models perform well on simulations of weakly coupled systems but fail when connectivity is no longer weak. On the other hand, using Bayesian model selection, we show that our phase-amplitude coupling model can describe non-weakly coupled systems more effectively despite the added complexity associated with using amplitude as an extra dependent variable. We demonstrate the advantage of our phase-amplitude model in the context of model-generated data, as well as of a simulation of inter-connected pendula, neural local field potential recordings in rodents under anaesthesia and international economic gross domestic product data., Comment: 17 pages, 5 figures

In this technical note, we address an unresolved challenge in neuroimaging statistics: how to determine which of several datasets is the best for inferring neuronal responses. Comparisons of this kind are important for experimenters when choosing an imaging protocol - and for developers of new acquisition methods. However, the hypothesis that one dataset is better than another cannot be tested using conventional statistics (based on likelihood ratios), as these require the data to be the same under each hypothesis. Here we present Bayesian data comparison, a principled framework for evaluating the quality of functional imaging data, in terms of the precision with which neuronal connectivity parameters can be estimated and competing models can be disambiguated. For each of several candidate datasets, neuronal responses are inferred using Dynamic Casual Modelling (DCM) - a commonly used Bayesian procedure for modelling neuroimaging data. Next, the parameters from subject-specific models are summarised at the group level using a Bayesian General Linear Model (GLM). A series of measures, which we introduce here, are then used to evaluate each dataset in terms of the precision of (group-level) parameter estimates and the ability of the data to distinguish similar models. To exemplify the approach, we compared four datasets that were acquired in a study evaluating multiband fMRI acquisition schemes. To enable people to reproduce these analyses using their own data and experimental paradigms, we provide general-purpose Matlab code via the SPM software., Comment: Equal contribution by Peter Zeidman and Samira M Kazan

Convolutional and Recurrent, deep neural networks have been successful in machine learning systems for computer vision, reinforcement learning, and other allied fields. However, the robustness of such neural networks is seldom apprised, especially after high classification accuracy has been attained. In this paper, we evaluate the robustness of three recurrent neural networks to tiny perturbations, on three widely used datasets, to argue that high accuracy does not always mean a stable and a robust (to bounded perturbations, adversarial attacks, etc.) system. Especially, normalizing the spectrum of the discrete recurrent network to bound the spectrum (using power method, Rayleigh quotient, etc.) on a unit disk produces stable, albeit highly non-robust neural networks. Furthermore, using the $\epsilon$-pseudo-spectrum, we show that training of recurrent networks, say using gradient-based methods, often result in non-normal matrices that may or may not be diagonalizable. Therefore, the open problem lies in constructing methods that optimize not only for accuracy but also for the stability and the robustness of the underlying neural network, a criterion that is distinct from the other.

Hierarchies feature prominently in anatomical accounts of cortical organisation. An open question is which computational (algorithmic) processes are implemented by these hierarchies. One renowned hypothesis is that cortical hierarchies implement a model of the world's causal structure and serve to infer environmental states from sensory inputs. This view, which casts perception as hierarchical Bayesian inference, has become a highly influential concept in both basic and clinical neuroscience. So far, however, a direct correspondence between the predicted order of hierarchical Bayesian computations and the sequence of evoked neuronal activity has not been demonstrated. Here, we present evidence for this correspondence from neuroimaging and electrophysiological data in healthy volunteers. Trial-wise sequences of hierarchical computations were inferred from participants' behaviour during a social learning task that required multi-level inference about intentions. We found that the temporal sequence of neuronal activity matched the order of computations as predicted by the theory. These findings provide strong evidence for the operation of hierarchical Bayesian inference in human cortex. Furthermore, our approach offers a novel strategy for the combined computational-physiological phenotyping of patients with disorders of perception, such as schizophrenia or autism., Comment: 34 pages, 5 figures, 3 tables

This technical report revisits the analysis of family-wise error rates in statistical parametric mapping - using random field theory - reported in (Eklund et al., 2015). Contrary to the understandable spin that these sorts of analyses attract, a review of their results suggests that they endorse the use of parametric assumptions - and random field theory - in the analysis of functional neuroimaging data. We briefly rehearse the advantages parametric analyses offer over nonparametric alternatives and then unpack the implications of (Eklund et al., 2015) for parametric procedures.

Neuroimaging produces data that are continuous in one or more dimensions. This calls for an inference framework that can handle data that approximate functions of space, for example, anatomical images, time--frequency maps and distributed source reconstructions of electromagnetic recordings over time. Statistical parametric mapping (SPM) is the standard framework for whole-brain inference in neuroimaging: SPM uses random field theory to furnish $p$-values that are adjusted to control family-wise error or false discovery rates, when making topological inferences over large volumes of space. Random field theory regards data as realizations of a continuous process in one or more dimensions. This contrasts with classical approaches like the Bonferroni correction, which consider images as collections of discrete samples with no continuity properties (i.e., the probabilistic behavior at one point in the image does not depend on other points). Here, we illustrate how random field theory can be applied to data that vary as a function of time, space or frequency. We emphasize how topological inference of this sort is invariant to the geometry of the manifolds on which data are sampled. This is particularly useful in electromagnetic studies that often deal with very smooth data on scalp or cortical meshes. This application illustrates the versatility and simplicity of random field theory and the seminal contributions of Keith Worsley (1951--2009), a key architect of topological inference., Comment: Published in at http://dx.doi.org/10.1214/10-AOAS337 the Annals of Applied Statistics (http://www.imstat.org/aoas/) by the Institute of Mathematical Statistics (http://www.imstat.org)