Publications

Planning and navigation as active inference

Abstract This paper introduces an active inference formulation of planning and navigation. It illustrates how the exploitation–exploration dilemma is dissolved by acting to minimise uncertainty (i.e., expected surprise or free energy). We use simulations of a maze problem to illustrate how agents can solve quite complicated problems using context sensitive prior preferences to form subgoals. Our focus is on how epistemic behaviour – driven by novelty and the imperative to reduce uncertainty about the world – contextualises pragmatic or goal-directed behaviour. Using simulations, we illustrate the underlying process theory with synthetic behavioural and electrophysiological responses during exploration of a maze and subsequent navigation to a target location. An interesting phenomenon that emerged from the simulations was a putative distinction between ‘place cells’ – that fire when a subgoal is reached – and ‘path cells’ – that fire until a subgoal is reached.

Anticipation in architectural experience: a computational neurophenomenology for architecture?

The perceptual experience of architecture is enacted by the sensory and motor system. When we act, we change the perceived environment according to a set of expectations that depend on our body and the built environment. The continuous process of collecting sensory information is thus based on bodily affordances. Affordances characterize the fit between the physical structure of the body and capacities for movement in the built environment. Since little has been done regarding the role of architectural design in the emergence of perceptual experience on a neuronal level, this paper offers a first step towards the role of architectural design in perceptual experience. An approach to synthesize concepts from computational neuroscience with architectural phenomenology into a computational neurophenomenology is considered. The outcome is a framework under which studies of architecture and cognitive neuroscience can be cast.

PsPM-SCRV1: Skin conductance responses to aversive/neutral pictures at different inter trial intervals.

This dataset includes skin conductance response (SCR) measurement, keypress responses and keypress response times to stimuli drawn from the International Affective Picture System for each of 24 healthy unmedicated participants (12 males and 12 females aged 27+/-4.6 years). The experiment used a 2x3 factorial design with the factors picture type (aversive, neutral), and mean ISI (3s, 9s, and 19s).

Computational psychiatry

Computational ideas pervade many areas of science and have an integrative explanatory role in neuroscience and cognitive science. However, computational depictions of cognitive function have had surprisingly little impact on the way we assess mental illness because diseases of the mind have not been systematically conceptualized in computational terms. Here, we outline goals and nascent efforts in the new field of computational psychiatry, which seeks to characterize mental dysfunction in terms of aberrant computations over multiple scales. We highlight early efforts in this area that employ reinforcement learning and game theoretic frameworks to elucidate decision-making in health and disease. Looking forwards, we emphasize a need for theory development and large-scale computational phenotyping in human subjects.

A neural mass model of spectral responses in electrophysiology

We present a neural mass model of steady-state membrane potentials measured with local field potentials or electroencephalography in the frequency domain. This model is an extended version of previous dynamic causal models for investigating event-related potentials in the time-domain. In this paper, we augment the previous formulation with parameters that mediate spike-rate adaptation and recurrent intrinsic inhibitory connections. We then use linear systems analysis to show how the model's spectral response changes with its neurophysiological parameters. We demonstrate that much of the interesting behaviour depends on the non-linearity which couples mean membrane potential to mean spiking rate. This non-linearity is analogous, at the population level, to the firing rate-input curves often used to characterize single-cell responses. This function depends on the model's gain and adaptation currents which, neurobiologically, are influenced by the activity of modulatory neurotransmitters. The key contribution of this paper is to show how neuromodulatory effects can be modelled by adding adaptation currents to a simple phenomenological model of EEG. Critically, we show that these effects are expressed in a systematic way in the spectral density of EEG recordings. Inversion of the model, given such non-invasive recordings, should allow one to quantify pharmacologically induced changes in adaptation currents. In short, this work establishes a forward or generative model of electrophysiological recordings for psychopharmacological studies.

Learning and generalization under ambiguity: An fMRI Study

Adaptive behavior often exploits generalizations from past experience by applying them judiciously in new situations. This requires a means of quantifying the relative importance of prior experience and current information, so they can be balanced optimally. In this study, we ask whether the brain generalizes in an optimal way. Specifically, we used Bayesian learning theory and fMRI to test whether neuronal responses reflect context-sensitive changes in ambiguity or uncertainty about experience-dependent beliefs. We found that the hippocampus expresses clear ambiguity-dependent responses that are associated with an augmented rate of learning. These findings suggest candidate neuronal systems that may be involved in aberrations of generalization, such as over-confidence.

In vitro neural networks minimise variational free energy

Abstract In this work, we address the neuronal encoding problem from a Bayesian perspective. Specifically, we ask whether neuronal responses in an in vitro neuronal network are consistent with ideal Bayesian observer responses under the free energy principle. In brief, we stimulated an in vitro cortical cell culture with stimulus trains that had a known statistical structure. We then asked whether recorded neuronal responses were consistent with variational message passing (i.e., belief propagation) based upon free energy minimisation (i.e., evidence maximisation). Effectively, this required us to solve two problems: first, we had to formulate the Bayes-optimal encoding of the causes or sources of sensory stimulation, and then show that these idealised responses could account for observed electrophysiological responses. We describe a simulation of an optimal neural network (i.e., the ideal Bayesian neural code) and then consider the mapping from idealised in silico responses to recorded in vitro responses. Our objective was to find evidence for functional specialisation and segregation in the in vitro neural network that reproduced in silico learning via free energy minimisation. Finally, we combined the in vitro and in silico results to characterise learning in terms of trajectories in a variational information plane of accuracy and complexity.

Brain-Inspired Computational Intelligence via Predictive Coding

Artificial intelligence (AI) is rapidly becoming one of the key technologies of this century. The majority of results in AI thus far have been achieved using deep neural networks trained with the error backpropagation learning algorithm. However, the ubiquitous adoption of this approach has highlighted some important limitations such as substantial computational cost, difficulty in quantifying uncertainty, lack of robustness, unreliability, and biological implausibility. It is possible that addressing these limitations may require schemes that are inspired and guided by neuroscience theories. One such theory, called predictive coding (PC), has shown promising performance in machine intelligence tasks, exhibiting exciting properties that make it potentially valuable for the machine learning community: PC can model information processing in different brain areas, can be used in cognitive control and robotics, and has a solid mathematical grounding in variational inference, offering a powerful inversion scheme for a specific class of continuous-state generative models. With the hope of foregrounding research in this direction, we survey the literature that has contributed to this perspective, highlighting the many ways that PC might play a role in the future of machine learning and computational intelligence at large.

Statistical parametric mapping for event-related potentials (II): a hierarchical temporal model

Evoked brain responses are generated by feedback loops

Neuronal responses to stimuli, measured electrophysiologically, unfold over several hundred milliseconds. Typically, they show characteristic waveforms with early and late components. It is thought that early or exogenous components reflect a perturbation of neuronal dynamics by sensory input bottom-up processing. Conversely, later, endogenous components have been ascribed to recurrent dynamics among hierarchically disposed cortical processing levels, top-down effects. Here, we show that evoked brain responses are generated by recurrent dynamics in cortical networks, and late components of event-related responses are mediated by backward connections. This evidence is furnished by dynamic causal modeling of mismatch responses, elicited in an oddball paradigm. We used the evidence for models with and without backward connections to assess their likelihood as a function of peristimulus time and show that backward connections are necessary to explain late components. Furthermore, we were able to quantify the contribution of backward connections to evoked responses and to source activity, again as a function of peristimulus time. These results link a generic feature of brain responses to changes in the sensorium and a key architectural component of functional anatomy; namely, backward connections are necessary for recurrent interactions among levels of cortical hierarchies. This is the theoretical cornerstone of most modern theories of perceptual inference and learning.

A physiological correlate of the meaning of words

Second waves, social distancing, and the spread of COVID-19 across America

<ns4:p>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.</ns4:p>

Testing for anatomically specified regional effects

We present a simple method that allows statistical inferences to be made about the significance of regional effects in statistical parametric maps (SPMs) when the approximate location of the effect is specified in advance. The test can be thought of as analogous to assessing activations with uncorrected P values based on the height of SPMs but, in this instance, using the spatial extent or volume of the nearest activated region. The advantage of the current test is that it eschews a correction for multiple comparisons even though the exact location of the expected activation may not be known. Hum. Brain Mapping 5:133–136, 1997. © 1997 Wiley-Liss, Inc.

Osteopathic Care as (En)active Inference: A Theoretical Framework for Developing an Integrative Hypothesis in Osteopathy

Osteopathy is a person-centred healthcare discipline that emphasizes the body’s structure-function interrelationship—and its self-regulatory mechanisms—to inform a whole-person approach to health and wellbeing. This paper aims to provide a theoretical framework for developing an integrative hypothesis in osteopathy, which is based on the enactivist and active inference accounts. We propose that osteopathic care can be reconceptualised under (En)active inference as a unifying framework. Active inference suggests that action-perception cycles operate to minimize uncertainty and optimize an individual’s internal model of the lived world and, crucially, the consequences of their behaviour. We argue that (En)active inference offers an integrative framework for osteopathy, which can evince the mechanisms underlying dyadic and triadic (e.g., in paediatric care) exchanges and osteopathic care outcomes. We propose that this theoretical framework can underpin osteopathic care across the lifespan, from preterm infants to the elderly and those with persistent pain and other physical symptoms. In situations of chronicity, as an ecological niche, the patient-practitioner dyad provides the osteopath and the patient with a set of affordances, i.e., possibilities for action provided by the environment, that through shared intentionally, can promote adaptations and restoration of productive agency. Through a dyadic therapeutic relationship, as they engage with their ecological niche’s affordances—a structured set of affordances shared by agents—osteopath and patient actively construct a shared sense-making narrative and realise a shared generative model of their relation to the niche. In general, touch plays a critical role in developing a robust therapeutic alliance, mental state alignment, and biobehavioural synchrony between patient and practitioner. However, its role is particularly crucial in the fields of neonatology and paediatrics, where it becomes central in regulating allostasis and restoring homeostasis. We argue that from an active inference standpoint, the dyadic shared ecological niche underwrites a robust therapeutic alliance, which is crucial to the effectiveness of osteopathic care. Considerations and implications of this model—to clinical practice and research, both within- and outside osteopathy—are critically discussed.

The computational neurology of movement under active inference

We propose a computational neurology of movement based on the convergence of theoretical neurobiology and clinical neurology. A significant development in the former is the idea that we can frame brain function as a process of (active) inference, in which the nervous system makes predictions about its sensory data. These predictions depend upon an implicit predictive (generative) model used by the brain. This means neural dynamics can be framed as generating actions to ensure sensations are consistent with these predictions-and adjusting predictions when they are not. We illustrate the significance of this formulation for clinical neurology by simulating a clinical examination of the motor system using an upper limb coordination task. Specifically, we show how tendon reflexes emerge naturally under the right kind of generative model. Through simulated perturbations, pertaining to prior probabilities of this model's variables, we illustrate the emergence of hyperreflexia and pendular reflexes, reminiscent of neurological lesions in the corticospinal tract and cerebellum. We then turn to the computational lesions causing hypokinesia and deficits of coordination. This in silico lesion-deficit analysis provides an opportunity to revisit classic neurological dichotomies (e.g. pyramidal versus extrapyramidal systems) from the perspective of modern approaches to theoretical neurobiology-and our understanding of the neurocomputational architecture of movement control based on first principles.

Hierarchical disruption in the Bayesian brain: Focal epilepsy and brain networks

In this opinion paper, we describe a combined view of functional and effective brain connectivity along with the free-energy principle for investigating persistent disruptions in brain networks of patients with focal epilepsy. These changes are likely reflected in effective connectivity along the cortical hierarchy and construct the basis of increased local functional connectivity in focal epilepsy. We propose a testable framework based on dynamic causal modelling and functional connectivity analysis with the capacity of explaining commonly observed connectivity changes during interictal periods. We then hypothesise their possible relation with disrupted free-energy minimisation in the Bayesian brain. This may offer a new approach for neuroimaging to specifically develop and address hypotheses regarding the network pathomechanisms underlying epileptic phenotypes.

Modeling Ketamine Effects on Synaptic Plasticity During the Mismatch Negativity

This paper presents a model-based investigation of mechanisms underlying the reduction of mismatch negativity (MMN) amplitudes under the NMDA-receptor antagonist ketamine. We applied dynamic causal modeling and Bayesian model selection to data from a recent ketamine study of the roving MMN paradigm, using a cross-over, double-blind, placebo-controlled design. Our modeling was guided by a predictive coding framework that unifies contemporary "adaptation" and "model adjustment" MMN theories. Comparing a series of dynamic causal models that allowed for different expressions of neuronal adaptation and synaptic plasticity, we obtained 3 major results: 1) We replicated previous results that both adaptation and short-term plasticity are necessary to explain MMN generation per se; 2) we found significant ketamine effects on synaptic plasticity, but not adaptation, and a selective ketamine effect on the forward connection from left primary auditory cortex to superior temporal gyrus; 3) this model-based estimate of ketamine effects on synaptic plasticity correlated significantly with ratings of ketamine-induced impairments in cognition and control. Our modeling approach thus suggests a concrete mechanism for ketamine effects on MMN that correlates with drug-induced psychopathology. More generally, this demonstrates the potential of modeling for inferring on synaptic physiology, and its pharmacological modulation, from electroencephalography data.

Microstructural plasticity in nociceptive pathways after spinal cord injury

Objective To track the interplay between (micro-) structural changes along the trajectories of nociceptive pathways and its relation to the presence and intensity of neuropathic pain (NP) after spinal cord injury (SCI). Methods A quantitative neuroimaging approach employing a multiparametric mapping protocol was used, providing indirect measures of myelination (via contrasts such as magnetisation transfer (MT) saturation, longitudinal relaxation (R1)) and iron content (via effective transverse relaxation rate (R2*)) was used to track microstructural changes within nociceptive pathways. In order to characterise concurrent changes along the entire neuroaxis, a combined brain and spinal cord template embedded in the statistical parametric mapping framework was used. Multivariate source-based morphometry was performed to identify naturally grouped patterns of structural variation between individuals with and without NP after SCI. Results In individuals with NP, lower R1 and MT values are evident in the primary motor cortex and dorsolateral prefrontal cortex, while increases in R2* are evident in the cervical cord, periaqueductal grey (PAG), thalamus and anterior cingulate cortex when compared with pain-free individuals. Lower R1 values in the PAG and greater R2* values in the cervical cord are associated with NP intensity. Conclusions The degree of microstructural changes across ascending and descending nociceptive pathways is critically implicated in the maintenance of NP. Tracking maladaptive plasticity unravels the intimate relationships between neurodegenerative and compensatory processes in NP states and may facilitate patient monitoring during therapeutic trials related to pain and neuroregeneration.