The modular organization of nervous systems is a widely documented principle of design for both vertebrate and invertebrate brains of which the columnar organization of the neocortex is an example. The classical cytoarchitectural areas of the neocortex are composed of smaller units, local neural circuits repeated iteratively within each area. Modules may vary in cell type and number, in internal and external connectivity, and in mode of neuronal processing between different large entities; within any single large entity they have a basic similarity of internal design and operation. Modules are most commonly grouped into entities by sets of dominating external connections. This unifying factor is most obvious for the heterotypical sensory and motor areas of the neocortex. Columnar defining factors in homotypical areas are generated, in part, within the cortex itself. The set of all modules composing such an entity may be fractionated into different modular subsets by different extrinsic connections. Linkages between them and subsets in other large entities form distributed systems. The neighborhood relations between connected subsets of modules in different entities result in nested distributed systems that serve distributed functions. A cortical area defined in classical cytoarchitectural terms may belong to more than one and sometimes to several distributed systems. Columns in cytoarchitectural areas located at some distance from one another, but with some common properties, may be linked by long-range, intracortical connections.

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The columnar organization of the neocortex.

It has been clear for almost two decades that cortical representations in adult animals are not fixed entities, but rather, are dynamic and are continuously modified by experience. The cortex can preferentially allocate area to represent the particular peripheral input sources that are proportionally most used. Alterations in cortical representations appear to underlie learning tasks dependent on the use of the behaviorally important peripheral inputs that they represent. The rules governing this cortical representational plasticity following manipulations of inputs, including learning, are increasingly well understood. In parallel with developments in the field of cortical map plasticity, studies of synaptic plasticity have characterized specific elementary forms of plasticity, including associative long-term potentiation and long-term depression of excitatory postsynaptic potentials. Investigators have made many important strides toward understanding the molecular underpinnings of these fundamental plasticity processes and toward defining the learning rules that govern their induction. The fields of cortical synaptic plasticity and cortical map plasticity have been implicitly linked by the hypothesis that synaptic plasticity underlies cortical map reorganization. Recent experimental and theoretical work has provided increasingly stronger support for this hypothesis. The goal of the current paper is to review the fields of both synaptic and cortical map plasticity with an emphasis on the work that attempts to unite both fields. A second objective is to highlight the gaps in our understanding of synaptic and cellular mechanisms underlying cortical representational plasticity.

/pdf/cortical-plasticity-from-synapses-to-maps-2j9eoi9428.pdf

CORTICAL PLASTICITY: From Synapses to Maps

http://yael-nissan.weebly.com/uploads/5/2/9/2/5292348/the_introduction_to_pain_an_integrative_review_-_millan.pdf

The induction of pain: an integrative review

Intrinsic firing patterns of diverse neocortical neurons.

https://www.shadlenlab.columbia.edu/publications/publications/mike/shadlen_newsome_Noise94.pdf

Noise, neural codes and cortical organization

Extracellular recordings of 209 neurons were obtained with carbon fiber-containing multibarrel micropipettes. The cells were isolated in the primary somatosensory cortex of cats anesthetized with barbiturate and classified according to the nature of their response to natural stimuli, the nature of the surrounding multiunit responses to the same stimuli, the response to thalamic stimulation, and their depth in the cortex. To study factors controlling the excitability of somatosensory neurons, their receptive fields were examined in the presence of iontophoretically administered gamma-aminobutyric acid (GABA), glutamate, and bicuculline methiodide (BMI). Even when the neurons were depolarized to perithreshold levels with glutamate, or when local inhibitory influences mediated by GABA were antagonized by BMI, the apparent specificity for one class of afferent input was maintained. Neurons responding to stimulation of either cutaneous or deep receptors maintained their modality specificity, and neurons in cutaneous rapidly adapting regions never took on slowly adapting properties. When ejected at currents that did not elicit action potentials, glutamate lowered the threshold for activation by cutaneous stimuli but did not enlarge the receptive field. With larger ejecting currents, the neurons developed an on-going discharge, but even at these higher doses, glutamate did not produce an increase in the receptive-field size. Some neurons in regions of cortex exhibiting slowly adapting multiunit responses were relatively insensitive to glutamate. These cells required four to five times more glutamate to evoke discharges than did most neurons. Other cells, previously unresponsive to somatic stimuli, could be shown to possess distinct cutaneous receptive fields when either glutamate or BMI was ejected in their vicinity. Iontophoretically administered BMI altered the firing pattern of somatosensory neurons, causing them to discharge in bursts of 3-15 impulses. BMI enlarged the receptive-field size of neurons in regions displaying rapidly adapting multiunit background discharges but not in those regions with slowly adapting multiunit discharges. This differential effect of BMI, suggesting that GABA controls receptive-field size in rapidly adapting regions, also indicates that neurons in rapidly adapting regions differ pharmacologically from those in other submodality regions. In all cortical regions, BMI blocked the poststimulus inhibitory period that normally followed thalamic stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

Functional role of GABA in cat primary somatosensory cortex: shaping receptive fields of cortical neurons.

Responses evoked in neurons of rat sensorimotor cortex upon stimulation of the pyramidal tract and ipsilateral cerebral peduncle were analysed using intracellular recording. Neurons responding antidromically to pyramidal tract stimulation (PT cells) and neurons failing to respond antidromically but exhibiting orthodromic responses were both stained by intracellular injection of horseradish peroxidase (HRP). Layer V pyramidal neurons, including those responding antidromically, exhibited prominent long lasting membrane hyperpolarizations and inhibitions of action potentials following pyramidal tract or cerebral peduncle stimulation. Upon passage of polarizing intracellular current two components were identified within the hyperpolarizing potential. A short duration initial component readily reversed with hyperpolarizing current. Frequently this earlier component overlapped a period of early excitation consisting of action potentials arising from recurrent EPSPs or large slow depolarizing potentials (SDPs). The second, much longer duration hyperpolarizing component did not reverse with passage of hyperpolarizing current and was often followed by a rebound period of depolarization and action potential generation. Both the excitatory and the inhibitory portions of these responses could be demonstrated in animals with acute thalamic transections severing the ascending lemniscal pathway to cortex. Following intracellular staining with HRP, two types of PT cells were identified by their different intracortical axonal arborizations. Most of the injected neurons had local axonal fields extending widely in layers V and VI, but with few or no collaterals extending radially toward the more superficial layers. A second type of PT cell had axon collaterals limited to a narrow zone around the dendritic field but extending radially as far as layer I. Cells of both types were observed to send axon collaterals into neostriatum. Both types of neurons exhibited morphological and physiological characteristics of slow PT cells, and we could find no cells comparable to the fast conducting PT cells observed in other species.

Morphological and electrophysiological characteristics of pyramidal tract neurons in the rat.

Extracellular recordings of 105 neurones in the cat's somatosensory thalamus were obtained with carbon fibre-containing multibarrel micropipettes. The responses of cells to natural stimulation of cutaneous or deep structures were characterized and the responses to electrical stimulation of primary somatosensory cortex were determined. Receptive fields were mapped and the functional properties were examined before and during the microiontophoretic administration of glutamate, γ-aminobutyric acid (GABA) and bicuculline methiodide (BMI). Modality and submodality properties of all cells tested apparently remained unchanged qualitatively, despite all pharmacological interventions. BMI lowered the response threshold of a majority of the 48 cells tested for this variable, although almost 25% responded with elevated thresholds. BMI changed the temporal properties of the responses of both thalamocortical relay neurones and of presumed interneurones. Discharges evoked by natural stimuli and by electrical stimulation of the cortex were prolonged and their pattern was altered. Decreases in the frequency of bursts of discharges were often observed with BMI, and these bursts were invariably prolonged and the interspike interval profiles were altered. Receptive field size changes were observed only in 8 of 48 neurones. For two of these, the field size decreased, while for the others there were small increases.

Bicuculline-induced alterations of response properties in functionally identified ventroposterior thalamic neurones.

Somatosensory cortical neurons with an identifiable electrophysiological signature.

Extracellular and intracellular recordings of corticothalamic (CT) cells were performed in the primary somatosensory cortex of the cat. CT neurons were antidromically activated by electrically stimulating the ventroposterior lateral (VPL) nucleus of the thalamus and were classified into two types according to their physiological properties. Type 1 had no spontaneous activity and no identifiable somatic receptive field. Type 2 fired action potentials spontaneously and responded to mechanical stimulation of the skin or underlying tissues. Axonal conduction velocities were slower for type 1 cells and their cell bodies were located slightly deeper in the cortex than those of type 2 cells. Both types of CT neurons exhibited inhibitory postsynaptic potentials in response to VPL stimulation but an early synaptic excitation and rebound discharge was observed almost exclusively in type 2 cells. These results suggest that only type 2 CT cells can modify the activity of thalamic neurons through a corticothalamic feedback loop.

P. Landry

Papers

Functional role of GABA in cat primary somatosensory cortex: shaping receptive fields of cortical neurons.

Morphological and electrophysiological characteristics of pyramidal tract neurons in the rat.

Bicuculline-induced alterations of response properties in functionally identified ventroposterior thalamic neurones.

Somatosensory cortical neurons with an identifiable electrophysiological signature.

Identification of two populations of corticothalamic neurons in cat primary somatosensory cortex