Barrel cortex function | EPFL Graph Search

Neocortex, the neuronal structure at the base of the remarkable cognitive skills of mammals, is a layered sheet of neuronal tissue composed of juxtaposed and interconnected columns. A cortical column is considered the basic module of cortical processing present in all cortical areas. It is believed to contain a characteristic microcircuit composed of a few thousand neurons. The high degree of cortical segmentation into vertical columns and horizontal layers is a boon for scientific investigation because it eases the systematic dissection and functional analysis of intrinsic as well as extrinsic connections of the column. In this review we will argue that in order to understand neocortical function one needs to combine a microscopic view, elucidating the workings of the local columnar microcircuits, with a macroscopic view, which keeps track of the linkage of distant cortical modules in different behavioral contexts. We will exemplify this strategy using the model system of vibrissal touch in mice and rats. On the macroscopic level vibrissal touch is an important sense for the subterranean rodents and has been honed by evolution to serve an array of distinct behaviors. Importantly, the vibrissae are moved actively to touch - requiring intricate sensorimotor interactions. Vibrissal touch, therefore, offers ample opportunities to relate different behavioral contexts to specific interactions of distant columns. On the microscopic level, the cortical modules in primary somatosensory cortex process touch inputs at highest magnification and discreteness - each whisker is represented by its own so-called barrel column. The cellular composition, intrinsic connectivity and functional aspects of the barrel column have been studied in great detail. Building on the versatility of genetic tools available in rodents, new, highly selective and flexible cellular and molecular tools to monitor and manipulate neuronal activity have been devised. Researchers have started to combine these with advanced and highly precise behavioral methods, on par with the precision known from monkey preparations. Therefore, the vibrissal touch model system is exquisitely positioned to combine the microscopic with the macroscopic view and promises to be instrumental in our understanding of neocortical function. (C) 2012 Elsevier Ltd. All rights reserved.

Functionally independent columns of rat somatosensory barrel cortex revealed with voltage-sensitive dye imaging

Carl Petersen

Whisker movement is somatotopically represented in rodent neocortex by electrical activity in clearly defined barrels, which can be visualized in living brain slices. The functional architecture of this part of the cortex can thus be mapped in vitro with respect to its physiological input and compared with its anatomical architecture. The spatial extent of excitation was measured at high temporal resolution by imaging optical signals from voltage-sensitive dye evoked by stimulation of individual barrels in layer 4. The optical signals correlated closely with subthreshold EPSPs recorded simultaneously from excitatory neurons in layer 4 and layer 2/3, respectively. Excitation was initially (3 msec) spread in a columnar manner into layer 2/3 and then subsided in both layers after approximately 50 msec. The lateral extent of the response was limited to the cortical column defined structurally by the barrel in layer 4. Two experimental interventions increased the spread of excitation. First, blocking GABA(A) receptor-mediated synaptic inhibition caused excitation to spread laterally throughout wide regions of layer 2/3 and layer 5 but not into neighboring barrels, suggesting that the local excitatory connections within layer 4 are restricted to single barrels and that inhibitory neurons control spread in supragranular and infragranular layers. Second, NMDA receptor-dependent increase of the spread of excitation was induced by pairing repetitive stimulation of a barrel column with coincident stimulation of layer 2/3 in a neighboring column. Such plasticity in the spatial extent of excitation in a barrel column could underlie changes in cortical map structure induced by alterations of sensory experience.

2001

Two-photon calcium imaging of neocortical projection neurons in whisker somatosensory cortex during goal-directed sensorimotor learning

Dimitra Angeliki Vavladeli

Abstract Excitatory projection neurons of the neocortex are thought to play important roles in per-ceptual and cognitive functions of the brain by directly connecting diverse cortical and subcortical areas. However, many aspects of the anatomical and functional organization of these inter-areal connections are unknown. The mouse primary somatosensory whisk-er barrel cortex (S1) serves as an important model for investigating the mammalian neo-cortex, and, here, I firstly investigate the structure and secondly the function of a specific subset of S1 cortico-cortical long-range projection neurons. In the first part of my thesis, I studied long-range axonal projections of excitatory layer 2/3 neurons with cell bodies located in S1. As a population, these neurons densely projected to secondary whisker somatosensory cortex (S2) and primary/secondary whisker motor cortex (M1/2), with additional axon in the dysgranular zone surrounding the barrel field, perirhinal temporal association cortex and striatum. The execution of a goal-directed behavior requires the brain to process incoming sensory information from the environment in a context-, learning- and motivation-dependent manner in order to perform specific motor actions. Cortico-cortical communica-tion in the context of goal-directed sensorimotor transformation has begun to be studied, but little is known about how signaling between interconnected cortical areas is modified by sensorimotor learning, as well as in response to changes in reward contingencies. Hence, in the second part of my thesis, I studied cortico-cortical dynamics in primary whisker somatosensory barrel cortex (S1) of mice during a combined whisker and audito-ry task. Using transgenic mice expressing GCaMP6f combined with two-photon micros-copy and retrograde labeling techniques, I chronically monitored the activity of excitatory layer 2/3 neurons in S1 projecting to M1 or S2, while mice learned the behavioral switch task. The results demonstrated that both classes of neurons responded after whisker and auditory stimulation. However, the whisker stimulus evoked response was stronger than the auditory stimulus evoked response. Neurons projecting to S2 exhibited stronger re-sponses compared to neurons projecting to M1 neurons. Those responses remained rela-tively stable across training sessions and under different reward conditions. Furthermore, both classes of neurons responded during spontaneous licking, but neurons projecting to S2 had larger licking-related responses compared to neurons projecting to M1. This work therefore furthers our knowledge of the structure and function of specific types of cortical projection neurons, which is a necessary step towards detailed under-standing of how sensory information might be signaled from primary sensory areas to downstream brain regions for further processing.

EPFL2018

Structure and dynamics of the neocortical microcircuit connectivity

Jean-Vincent Le Bé

The neocortex is the most computationally advanced portion of the brain. It is currently assumed to be composed of a large number of "cortical columns" – intricate arrangements of cortical neurons approximately 300-500 µm in diameter and 2-5 mm in height in humans – that might serve as the elementary computational unit of the neocortex. Understanding the computation performed by this microcircuit is one of the keys to our comprehension of the brain. The so-called cortical column is not a static entity, however, and it evolves throughout a lifetime and continually adapts to the information from its cortical environment. Despite the differences between cortical columns across the cortex, a number of common features have been identified: a laminar structure, the dynamics of connections between identified neurons or the mechanisms for these connections to be modified (that give its specificity to each microcircuit). This thesis presents the description of the differential connectivity and synaptic dynamics across cell populations and the long term neuronal rewiring in a particular neuronal population within the cortical column. Somatic whole cell recordings have been performed to probe the connectivity, synaptic dynamics and plasticity of the connections in the rat neocortex. Two populations of layer V pyramidal neurons were studied in particular: cortico-callosally projecting pyramidal cells (CCPs) and thick tufted pyramidal cells (TTCs). The first major results from this work revealed the degree of connectivity and the linear dynamics of the CCPs population when compared to the TTCs. CCPs have nearly 4 times fewer interconnections and subsequent post-synaptic potentials were less decreasing in amplitude along a pre-synaptic series of action potentials. Long term configuration of TTC networks was explored. These experiments show for the first time the emergence of new functional synaptic connections between TTCs within hours. Activation of the slice by glutamate greatly increases their rate of emergence and this work demonstrated that metabotropic glutamate receptor 5 (mGluR5) activation and action potential firing are required for new connections to be formed in this experimental protocol. Newly formed connections respond in a more linear fashion and have weaker post-synaptic influence than already existing connections. Pre-existing connections are also modified after stimulation, requiring mGluR5, action potentials as well as α-amino-3-hydroxy-5-methyl-4-isoxazole-propionic acid (AMPA) and N-methyl-D-aspartic acid (NMDA) receptors activation. The activation of group III metabotropic glutamate receptors (mGluRs) however results in a decrease in the strength of connections. Finally, the influence of inhibitory interneurons on the activity and connectivity between TTCs was also investigated. The results of this study show that firing of inhibitory interneurons can be triggered by the input of only one pyramidal cell. They further show that stimulation of a single TTC can result in an hyperpolarization of the post-synaptic TTC mediated by an interneurone. When the pre-synaptic neuron is also directly connected to the post-synaptic neuron with an excitatory synapse, the indirect inhibitory connection serves to curtail the excitatory response. This work has provided a new insight into the dynamic nature of the cortical microcircuitry, showing that it evolves rapidly and can adapt, reconfigure and rewire itself in remarkably short time-spans. It also describes the variety of dynamics exhibited by the different types of pyramidal cells, due to either the projecting site specificity or to the action of an intermediate interneuron.