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Tactile sensory information from facial whiskers provides nocturnal tunnel-dwelling rodents, including mice and rats, with important spatial and textural information about their immediate surroundings. Whiskers are moved back and forth to scan the environment (whisking), and touch signals from each whisker evoke sparse patterns of neuronal activity in whisker-related primary somatosensory cortex (wS1; barrel cortex). Whisking is accompanied by desynchronized brain states and cell-type-specific changes in spontaneous and evoked neuronal activity. Tactile information, including object texture and location, appears to be computed in wS1 through integration of motor and sensory signals. wS1 also directly controls whisker movements and contributes to learned, whisker-dependent, goal-directed behaviours. The cell-type-specific neuronal circuitry in wS1 that contributes to whisker sensory perception is beginning to be defined.

Spontaneous fluctuations in hemodynamic signals in the absence of a task or overt stimulation are used to infer neural activity. We tested this coupling by simultaneously measuring neural activity and changes in cerebral blood volume (CBV) in the somatosensory cortex of awake, head-fixed mice during periods of true rest and during whisker stimulation and volitional whisking. We found that neurovascular coupling was similar across states and that large, spontaneous CBV changes in the absence of sensory input were driven by volitional whisker and body movements. Hemodynamic signals during periods of rest were weakly correlated with neural activity. Spontaneous fluctuations in CBV and vessel diameter persisted when local neural spiking and glutamatergic input were blocked, as well as during blockade of noradrenergic receptors, suggesting a non-neuronal origin for spontaneous CBV fluctuations. Spontaneous hemodynamic signals reflect a combination of behavior, local neural activity, and putatively non-neural processes. The relationship of resting-state hemodynamics signals to ongoing neural activity is poorly understood. Using optical imaging, electrophysiology, and local pharmacological infusions, Winder et al. found that resting hemodynamic signals were weakly correlated with neural activity and that these hemodynamic fluctuations persisted when neural activity was silenced.

Feedforward and feedback synaptic pathways shape how neural activity evolves across cortical areas, but they are difficult to monitor using traditional methods during behavior. The authors use pathway-specific and cellular-resolution in vivo imaging to quantify sensory and decision-related neural activity both within and propagating between two cortical areas critical for touch perception. The brain transforms physical sensory stimuli into meaningful perceptions. In animals making choices about sensory stimuli, neuronal activity in successive cortical stages reflects a progression from sensation to decision. Feedforward and feedback pathways connecting cortical areas are critical for this transformation. However, the computational functions of these pathways are poorly understood because pathway-specific activity has rarely been monitored during a perceptual task. Using cellular-resolution, pathway-specific imaging, we measured neuronal activity across primary (S1) and secondary (S2) somatosensory cortices of mice performing a tactile detection task. S1 encoded the stimulus better than S2, while S2 activity more strongly reflected perceptual choice. S1 neurons projecting to S2 fed forward activity that predicted choice. Activity encoding touch and choice propagated in an S1–S2 loop along feedforward and feedback axons. Our results suggest that sensory inputs converge into a perceptual outcome as feedforward computations are reinforced in a feedback loop.

Volitional motor control involves not only the initiation of desired movements but also the suppression of undesired movements. In this Opinion article, Ebbesen and Brecht argue that motor cortex neurons have a role in both aspects of motor control. The motor cortex is a large frontal structure in the cerebral cortex of eutherian mammals. A vast array of evidence implicates the motor cortex in the volitional control of motor output, but how does the motor cortex exert this 'control'? Historically, ideas regarding motor cortex function have been shaped by the discovery of cortical 'motor maps' — that is, ordered representations of stimulation-evoked movements in anaesthetized animals. Volitional control, however, entails the initiation of movements and the ability to suppress undesired movements. In this article, we highlight classic and recent findings that emphasize that motor cortex neurons have a role in both processes.

Sensory cortex spiking is well known to predict trial-to-trial variability in perceptual choice, but the origins of this choice-related activity are not fully understood. In the mouse somatosensory system, electrophysiology, imaging and optogenetic experiments reveal a progression of choice-related activity as touch signals flow from primary afferents to cortex. During perceptual decisions about faint or ambiguous sensory stimuli, even identical stimuli can produce different choices. Spike trains from sensory cortex neurons can predict trial-to-trial variability in choice. Choice-related spiking is widely studied as a way to link cortical activity to perception, but its origins remain unclear. Using imaging and electrophysiology, we found that mouse primary somatosensory cortex neurons showed robust choice-related activity during a tactile detection task. Spike trains from primary mechanoreceptive neurons did not predict choices about identical stimuli. Spike trains from thalamic relay neurons showed highly transient, weak choice-related activity. Intracellular recordings in cortex revealed a prolonged choice-related depolarization in most neurons that was not accounted for by feed-forward thalamic input. Top-down axons projecting from secondary to primary somatosensory cortex signaled choice. An intracellular measure of stimulus sensitivity determined which neurons converted choice-related depolarization into spiking. Our results reveal how choice-related spiking emerges across neural circuits and within single neurons.

Here the authors demonstrate a causal role for the barrel cortex in the detection of single whisker stimuli. Whisker deflection evoked an early (<50 ms) reliable sensory response that was encoded through cell-specific reversal potentials. A secondary late (50–400 ms) depolarization was enhanced in hit trials compared to misses. Optogenetic inactivation revealed a causal role for late excitation. Neocortical activity can evoke sensory percepts, but the cellular mechanisms remain poorly understood. We trained mice to detect single brief whisker stimuli and report perceived stimuli by licking to obtain a reward. Pharmacological inactivation and optogenetic stimulation demonstrated a causal role for the primary somatosensory barrel cortex. Whole-cell recordings from barrel cortex neurons revealed membrane potential correlates of sensory perception. Sensory responses depended strongly on prestimulus cortical state, but both slow-wave and desynchronized cortical states were compatible with task performance. Whisker deflection evoked an early (<50 ms) reliable sensory response that was encoded through cell-specific reversal potentials. A secondary late (50–400 ms) depolarization was enhanced on hit trials compared to misses. Optogenetic inactivation revealed a causal role for late excitation. Our data reveal dynamic processing in the sensory cortex during task performance, with an early sensory response reliably encoding the stimulus and later secondary activity contributing to driving the subjective percept.

A rich fossil record chronicles the distant origins of mammals, but the evolution of defining soft tissue characters of extant mammals, such as mammary glands and hairs is difficult to interpret because soft tissue does not readily fossilize. As many soft tissue features are derived from dermic structures, their evolution is linked to that of the nervous syutem and palaeoneurology offers opportunities to find bony correlates of these soft tissue features. Here, a CT scan study of 29 fossil skulls shows that non-mammaliaform Prozostrodontia display a retracted, fully ossified and non-ramified infraorbital canal for the infraorbital nerve, unlike more basal therapsids. The presence of a true infraorbital canal in Prozostrodontia suggests that a motile rhinarium and maxillary vibrissae were present. Also the complete ossification of the parietal fontanelle (resulting in the loss of the parietal foramen) and the development of the cerebellum in Probainognathia may be pleiotropically linked to the appearance of mammary glands and having body hair coverage since these traits are all controlled by the same homeogene, Msx2, in mice. These suggest that defining soft tissue characters of mammals were already present in their forerunners some 240 to 246 mya.

Imaging of activity in long-range axons is reported in mice performing tactile object-localization with their whiskers; the feedback projection from the motor cortex to the somatosensory cortex provides information to integrate whisker movement information and touch, which are key components of object identification. Touch perception by a whisker When rodents explore objects with their whiskers, touch signals are represented in the somatosensory cortex, and are also sent, by projections, to the motor cortex. The motor cortex also projects back into the somatosensory cortex, potentially providing whisker-movement information for integration with touch signals. This study addresses the question of what the motor cortex actually tells the somatosensory cortex. Karel Svoboda and colleagues imaged activity in the motor neuron axonal arbors of the somatosensory cortex while mice were performing an object-localization task that required integration of movement with touch and other senses. The resulting images show axons representing a diverse range of signals, including whisker movements and touch. Cortical-feedback projections to primary sensory areas terminate most heavily in layer 1 (L1) of the neocortex 1 , 2 , where they make synapses with tuft dendrites of pyramidal neurons. L1 input is thought to provide ‘contextual’ information 3 , but the signals transmitted by L1 feedback remain uncharacterized. In the rodent somatosensory system, the spatially diffuse 4 feedback projection from vibrissal motor cortex (vM1) to vibrissal somatosensory cortex (vS1, also known as the barrel cortex) may allow whisker touch to be interpreted in the context of whisker position to compute object location 5 , 6 . When mice palpate objects with their whiskers to localize object features 7 , 8 , whisker touch excites vS1 9 and later vM1 in a somatotopic manner 10 , 11 , 12 , 13 . Here we use axonal calcium imaging to track activity in vM1→vS1 afferents in L1 of the barrel cortex while mice performed whisker-dependent object localization. Spatially intermingled individual axons represent whisker movements, touch and other behavioural features. In a subpopulation of axons, activity depends on object location and persists for seconds after touch. Neurons in the barrel cortex thus have information to integrate movements and touches of multiple whiskers over time, key components of object identification and navigation by active touch.

Primary sensory cortex has long been believed to play a straightforward role in the initial processing of sensory information. Yet, the superficial layers of cortex overall are sparsely active, even during sensory stimulation; additionally, cortical activity is influenced by other modalities, task context, reward, and behavioral state. Our study demonstrates that reinforcement learning dramatically alters representations among longitudinally imaged neurons in superficial layers of mouse primary somatosensory cortex. Learning an object detection task recruits previously unresponsive neurons, enlarging the neuronal population sensitive to touch and behavioral choice. Cortical responses decrease upon repeated stimulus presentation outside of the behavioral task. Moreover, training improves population encoding of the passage of time, and unexpected deviations in trial timing elicit even stronger responses than touches do. In conclusion, the superficial layers of sensory cortex exhibit a high degree of learning-dependent plasticity and are strongly modulated by non-sensory but behaviorally-relevant features, such as timing and surprise. Activity in the superficial layers of the sensory cortex is believed to be largely driven by incoming sensory stimuli. Here the authors demonstrate how learning changes neural responses to sensations according to both behavioral relevance and timing, suggesting a high degree of non-sensory modulation.

Association areas in neocortex encode novel stimulus-outcome relationships, but the principles of their engagement during task learning remain elusive. Using chronic wide-field calcium imaging, we reveal two phases of spatiotemporal refinement of layer 2/3 cortical activity in mice learning whisker-based texture discrimination in the dark. Even before mice reach learning threshold, association cortex—including rostro-lateral (RL), posteromedial (PM), and retrosplenial dorsal (RD) areas—is generally suppressed early during trials (between auditory start cue and whisker-texture touch). As learning proceeds, a spatiotemporal activation sequence builds up, spreading from auditory areas to RL immediately before texture touch (whereas PM and RD remain suppressed) and continuing into barrel cortex, which eventually efficiently discriminates between textures. Additional correlation analysis substantiates this diverging learning-related refinement within association cortex. Our results indicate that a pre-learning phase of general suppression in association cortex precedes a learning-related phase of task-specific signal flow enhancement. Learning is a dynamic process involving many cortical areas. Here, using cortex-wide imaging, the authors show that in mice learning to discriminate between two textures a distinct task related signal flow is enhanced involving a specific association area whereas other association areas are suppressed.