Whisking with robots from rat vibrissae to biomimetic technology for active touch

This article summarizes some of the key features of the rat vibrissal system, including the actively controlled sweeping movements of the vibrissae known as whisking, and reviews the past and ongoing research aimed at replicating some of this functionality in biomimetic robots

Simultaneous mesoscopic and two-photon imaging of neuronal activity in cortical circuits.

Spontaneous and sensory-evoked activity propagates across varying spatial scales in the mammalian cortex, but technical challenges have limited conceptual links between the function of local neuronal circuits and brain-wide network dynamics. We present a method for simultaneous cellular-resolution two-photon calcium imaging of a local microcircuit and mesoscopic widefield calcium imaging of the entire cortical mantle in awake mice. Our multi-scale approach involves a microscope with an orthogonal axis design where the mesoscopic objective is oriented above the brain and the two-photon objective is oriented horizontally, with imaging performed through a microprism. We also introduce a viral transduction method for robust and widespread gene delivery in the mouse brain. These approaches allow us to identify the behavioral state-dependent functional connectivity of pyramidal neurons and vasoactive intestinal peptide-expressing interneurons with long-range cortical networks. Our imaging system provides a powerful strategy for investigating cortical architecture across a wide range of spatial scales

The role of environmental feedback in a brain state switch from passive to active sensing

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Adaptive cancelation of self-generated sensory signals in a whisking robot

Sensory signals are often caused by one's own active movements. This raises a problem of discriminating between self-generated sensory signals and signals generated by the external world. Such discrimination is of general importance for robotic systems, where operational robustness is dependent on the correct interpretation of sensory signals. Here, we investigate this problem in the context of a whiskered robot. The whisker sensory signal comprises two components: one due to contact with an object (externally generated) and another due to active movement of the whisker (self-generated). We propose a solution to this discrimination problem based on adaptive noise cancelation, where the robot learns to predict the sensory consequences of its own movements using an adaptive filter. The filter inputs (copy of motor commands) are transformed by Laguerre functions instead of the often-used tapped-delay line, which reduces model order and, therefore, computational complexity. Results from a contact-detection task demonstrate that false positives are significantly reduced using the proposed scheme

MOTOR CORTEX REGULATION OF THALAMIC-CORTICAL ACTIVITY IN THE SOMATOSENSORY SYSTEM

A prominent feature of thalamocortical circuitry in sensory systems is the extensive and highly organized feedback projection from the cortex to thalamic neurons that provide input to it. Intriguingly, many corticothalamic (CT) neurons are weakly responsive to peripheral stimuli, or silent altogether. Here using the whisker-to-barrel system, we examine whether the responses of CT neurons and their related thalamic neurons are affected by motor cortex, a prominent source of input to deep layers of the somatosensory cortex. Pharmacological facilitation of motor cortex activity produced using focal, microiontophoresis leads to enhanced whisker-evoked firing of topographically aligned layer 6 neurons, including identified CT cells, and of cells in corresponding regions of the thalamic ventral posterior medial nucleus (VPm barreloids). Together, the findings raise the possibility that cortico-thalamo-cortical circuitry in primary sensory areas is engaged by other functionally related cortical centers, providing a means for context-dependent regulation of information processing within thalamocortical circuits.We investigated how vMCx influence activity in thalamic VPm nucleus in a freely behaving rat. We examine afferent-evoked thalamic activity in animals that are either alert but voluntarily relatively motionless or actively whisking in the air without object contact. Afferent activity is evoked in VPm by means of electrical microstimulation of a single whisker follicle. In some experiments, neural processing in brainstem trigeminal nuclei was either by-passed by means of medial lemniscus stimulation, or altered by pharmacological intervention. We found that sensory responses during voluntary whisker movements, when motor cortex is likely to be active, are reduced relative to responses that occur during periods of wakeful quiescence. Enhancement of thalamic activity during whisking can be observed, however, when signal processing in sub-thalamic centers is either by-passed or experimentally altered. Findings suggest that during voluntary movement activity within the lemniscal system is globally diminished, perhaps at early, brainstem levels at the same time that activity within specific thalamocortical neuronal populations is facilitated. Though activity levels are reduced system-wide, activity within some local circuits may be subject to less net suppression. This decrease in suppression may occur on a moment-to-moment basis in a context-dependent manner. Thus, during voluntary whisker movement, sensory transmission in thalamocortical circuits may be modulated according to specific activation patterns distributed across the motor map

Serotonin and Whisking

AbstractRhythmic whisker movements, called “whisking,” are produced by a brainstem central pattern generator (CPG) that uses serotonin to induce periodic firing in facial motorneurons. During active touch, motor cortex could regulate whisking frequency by controlling the rate of firing of the serotonergic neurons

A theory of how active behavior stabilises neural activity: neural gain modulation by closed-loop environmental feedback

During active behaviours like running, swimming, whisking or sniffing, motor actions shape sensory input and sensory percepts guide future motor commands. Ongoing cycles of sensory and motor processing constitute a closed-loop feedback system which is central to motor control and, it has been argued, for perceptual processes. This closed-loop feedback is mediated by brainwide neural circuits but how the presence of feedback signals impacts on the dynamics and function of neurons is not well understood. Here we present a simple theory suggesting that closed-loop feedback between the brain/body/environment can modulate neural gain and, consequently, change endogenous neural fluctuations and responses to sensory input. We support this theory with modeling and data analysis in two vertebrate systems. First, in a model of rodent whisking we show that negative feedback mediated by whisking vibrissa can suppress coherent neural fluctuations and neural responses to sensory input in the barrel cortex. We argue this suppression provides an appealing account of a brain state transition (a marked change in global brain activity) coincident with the onset of whisking in rodents. Moreover, this mechanism suggests a novel signal detection mechanism that selectively accentuates active, rather than passive, whisker touch signals. This mechanism is consistent with a predictive coding strategy that is sensitive to the consequences of motor actions rather than the difference between the predicted and actual sensory input. We further support the theory by re-analysing previously published two-photon data recorded in zebrafish larvae performing closed-loop optomotor behaviour in a virtual swim simulator. We show, as predicted by this theory, that the degree to which each cell contributes in linking sensory and motor signals well explains how much its neural fluctuations are suppressed by closed-loop optomotor behaviour. More generally we argue that our results demonstrate the dependence of neural fluctuations, across the brain, on closed-loop brain/body/environment interactions strongly supporting the idea that brain function cannot be fully understood through open-loop approaches alone

Whisker Movements Reveal Spatial Attention: A Unified Computational Model of Active Sensing Control in the Rat

Spatial attention is most often investigated in the visual modality through measurement of eye movements, with primates, including humans, a widely-studied model. Its study in laboratory rodents, such as mice and rats, requires different techniques, owing to the lack of a visual fovea and the particular ethological relevance of orienting movements of the snout and the whiskers in these animals. In recent years, several reliable relationships have been observed between environmental and behavioural variables and movements of the whiskers, but the function of these responses, as well as how they integrate, remains unclear. Here, we propose a unifying abstract model of whisker movement control that has as its key variable the region of space that is the animal's current focus of attention, and demonstrate, using computer-simulated behavioral experiments, that the model is consistent with a broad range of experimental observations. A core hypothesis is that the rat explicitly decodes the location in space of whisker contacts and that this representation is used to regulate whisker drive signals. This proposition stands in contrast to earlier proposals that the modulation of whisker movement during exploration is mediated primarily by reflex loops. We go on to argue that the superior colliculus is a candidate neural substrate for the siting of a head-centred map guiding whisker movement, in analogy to current models of visual attention. The proposed model has the potential to offer a more complete understanding of whisker control as well as to highlight the potential of the rodent and its whiskers as a tool for the study of mammalian attention