A perspective on cortical layering and layer-spanning neuronal elements

This review article addresses the function of the layers of the cerebral cortex. We develop the perspective that cortical layering needs to be understood in terms of its functional anatomy, i.e., the terminations of synaptic inputs on distinct cellular compartments and their effect on cortical activity. The cortex is a hierarchical structure in which feed forward and feedback pathways have a layer-specific termination pattern. We take the view that the influence of synaptic inputs arriving at different cortical layers can only be understood in terms of their complex interaction with cellular biophysics and the subsequent computation that occurs at the cellular level. We use high-resolution fMRI, which can resolve activity across layers, as a case study for implementing this approach by describing how cognitive events arising from the laminar distribution of inputs can be interpreted by taking into account the properties of neurons that span different layers. This perspective is based on recent advances in measuring subcellular activity in distinct feed-forward and feedback axons and in dendrites as they span across layers

The laminar integration of sensory inputs with feedback signals in human cortex

The cortex constitutes the largest area of the human brain. Yet we have only a basic understanding of how the cortex performs one vital function: the integration of sensory signals (carried by feedforward pathways) with internal representations (carried by feedback pathways). A multi-scale, multi-species approach is essential for understanding the site of integration, computational mechanism and functional role of this processing. To improve our knowledge we must rely on brain imaging with improved spatial and temporal resolution and paradigms which can measure internal processes in the human brain, and on the bridging of disciplines in order to characterize this processing at cellular and circuit levels. We highlight apical amplification as one potential mechanism for integrating feedforward and feedback inputs within pyramidal neurons in the rodent brain. We reflect on the challenges and progress in applying this model neuronal process to the study of human cognition. We conclude that cortical-layer specific measures in humans will be an essential contribution for better understanding the landscape of information in cortical feedback, helping to bridge the explanatory gap

The promise of layer-specific neuroimaging for testing predictive coding theories of psychosis

Predictive coding potentially provides an explanatory model for understanding the neurocognitive mechanisms of psychosis. It proposes that cognitive processes, such as perception and inference, are implemented by a hierarchical system, with the influence of each level being a function of the estimated precision of beliefs at that level. However, predictive coding models of psychosis are insufficiently constrained—any phenomenon can be explained in multiple ways by postulating different changes to precision at different levels of processing. One reason for the lack of constraint in these models is that the core processes are thought to be implemented by the function of specific cortical layers, and the technology to measure layer specific neural activity in humans has until recently been lacking. As a result, our ability to constrain the models with empirical data has been limited. In this review we provide a brief overview of predictive processing models of psychosis and then describe the potential for newly developed, layer specific neuroimaging techniques to test and thus constrain these models. We conclude by discussing the most promising avenues for this research as well as the technical and conceptual challenges which may limit its application

Contextual Feedback to Superficial Layers of V1

Neuronal cortical circuitry comprises feedforward, lateral, and feedback projections, each of which terminates in distinct cortical layers [1-3]. In sensory systems, feedforward processing transmits signals from the external world into the cortex, whereas feedback pathways signal the brain's inference of the world [4-11]. However, the integration of feedforward, lateral, and feedback inputs within each cortical area impedes the investigation of feedback, and to date, no technique has isolated the feedback of visual scene information in distinct layers of healthy human cortex. We masked feedforward input to a region of V1 cortex and studied the remaining internal processing. Using high-resolution functional brain imaging (0.8 mm(3)) and multivoxel pattern information techniques, we demonstrate that during normal visual stimulation scene information peaks in mid-layers. Conversely, we found that contextual feedback information peaks in outer, superficial layers. Further, we found that shifting the position of the visual scene surrounding the mask parametrically modulates feedback in superficial layers of V1. Our results reveal the layered cortical organization of external versus internal visual processing streams during perception in healthy human subjects. We provide empirical support for theoretical feedback models such as predictive coding [10, 12] and coherent infomax [13] and reveal the potential of high-resolution fMRI to access internal processing in sub-millimeter human cortex

Structure Learning in Coupled Dynamical Systems and Dynamic Causal Modelling

Identifying a coupled dynamical system out of many plausible candidates, each of which could serve as the underlying generator of some observed measurements, is a profoundly ill posed problem that commonly arises when modelling real world phenomena. In this review, we detail a set of statistical procedures for inferring the structure of nonlinear coupled dynamical systems (structure learning), which has proved useful in neuroscience research. A key focus here is the comparison of competing models of (ie, hypotheses about) network architectures and implicit coupling functions in terms of their Bayesian model evidence. These methods are collectively referred to as dynamical casual modelling (DCM). We focus on a relatively new approach that is proving remarkably useful; namely, Bayesian model reduction (BMR), which enables rapid evaluation and comparison of models that differ in their network architecture. We illustrate the usefulness of these techniques through modelling neurovascular coupling (cellular pathways linking neuronal and vascular systems), whose function is an active focus of research in neurobiology and the imaging of coupled neuronal systems

Big-Loop Recurrence within the Hippocampal System Supports Integration of Information across Episodes

Recent evidence challenges the widely held view that the hippocampus is specialized for episodic memory, by demonstrating that it also underpins the integration of information across experiences. Contemporary computational theories propose that these two contrasting functions can be accomplished by big-loop recurrence, whereby the output of the system is recirculated back into the hippocampus. We use ultra-high-resolution fMRI to provide support for this hypothesis, by showing that retrieved information is presented as a new input on the superficial entorhinal cortex—driven by functional connectivity between the deep and superficial entorhinal layers. Further, the magnitude of this laminar connectivity correlated with inferential performance, demonstrating its importance for behavior. Our findings offer a novel perspective on information processing within the hippocampus and support a unifying framework in which the hippocampus captures higher-order structure across experiences, by creating a dynamic memory space from separate episodic codes for individual experiences

Six challenges for embodiment research

20 years after Barsalou's seminal perceptual symbols paper (Barsalou, 1999), embodied cognition, the notion that cognition involves simulations of sensory, motor, or affective states, has moved in status from an outlandish proposal advanced by a fringe movement in psychology to a mainstream position adopted by large numbers of researchers in the psychological and cognitive (neuro)sciences. While it has generated highly productive work in the cognitive sciences as a whole, it had a particularly strong impact on research into language comprehension. The view of a mental lexicon based on symbolic word representations, which are arbitrarily linked to sensory aspects of their referents, for example, was generally accepted since the cognitive revolution in the 1950s. This has radically changed. Given the current status of embodiment as a main theory of cognition, it is somewhat surprising that a close look at the state of the affairs in the literature reveals that the debate about the nature of the processes involved in language comprehension is far from settled and key questions remain unanswered. We present several suggestions for a productive way forward