Intracellular recordings and organic and inorganic Ca2+ channel blockers were used in a neocortical brain slice preparation to test whether high-voltage-activated (HVA) Ca2+ channels are differentially coupled to Ca2+-dependent afterhyperpolarizations (AHPs) in sensorimotor neocortical pyramidal neurons. For the most part, spike repolarization was not Ca2+ dependent in these cells, although the final phase of repolarization (after the fast AHP) was sensitive to block of N-type current. Between 30 and 60% of the medium afterhyperpolarization (mAHP) and between approximately 80 and 90% of the slow AHP (sAHP) were Ca2+ dependent. Based on the effects of specific organic Ca2+ channel blockers (dihydropyridines, omega-conotoxin GVIA, omega-agatoxin IVA, and omega-conotoxin MVIIC), the sAHP is coupled to N-, P-, and Q-type currents. P-type currents were coupled to the mAHP. L-type current was not involved in the generation of either AHP but (with other HVA currents) contributes to the inward currents that regulate interspike intervals during repetitive firing. These data suggest different functional consequences for modulation of Ca2+ current subtypes.

Following amputation, most amputees still report feeling the missing limb and often describe these feelings as excruciatingly painful. Phantom limb sensations (PLS) are useful while controlling a prosthesis; however, phantom limb pain (PLP) is a debilitating condition that drastically hinders quality of life. Although such experiences have been reported since the early 16th century, the etiology remains unknown. Debate continues regarding the roles of the central and peripheral nervous systems. Currently, the most posited mechanistic theories rely on neuronal network reorganization; however, greater consideration should be given to the role of the dorsal root ganglion within the peripheral nervous system. This Review provides an overview of the proposed mechanistic theories as well as an overview of various treatments for PLP.

ABSTRACT:The motor representation of the body musculature was studied in 11 adult mice by using ICMS. The motor responses elicited from both granular and agranular cortical fields showed that the mouse motor cortex is topographically organized; however, within the representation of individual body-parts the movements are multiply represented. In addition, several sites were encountered where more than one movement was elicited at the same stimulus threshold. The locations of pyramidal cells contributing axons to the pyramidal tract were examined by means of retrograde tracing with HRP injected into the cervical enlargement. This procedure labeled neurons only in lamina V in granular and agranular cortical fields. The similarities between the organization of motor cortex demonstrated in this study and the organization in the rat suggest that the rat and mouse share a common plan of rodent motor cortical organization. RESUME: Etude de I'organisation du cortex moteur de la souris par tracage retrograde et cartographic par microstimulation intracorticale (MSIC). La representation motrice de la musculature corporelle a 6t6 etudiee chez 11 souris adultes au moyen de la MSIC. Les reponses motrices elicitees a partir des champs corticaux de cellules granuleuses et non-granuleuses ont montre que le cortex moteur de la souris a une organisation topographique; cependant, la representation des mouvements est multiple au sein de la representation de parties individuelles du corps. De plus, plusieurs sites ont 6te repeY6s ou plus d'un mouvement etaient provoques pour un meme seuil de stimulation. La localisation des cellules pyramidales contribuant des axones au faisceau pyramidal a ete examinee au moyen du tracage retrograde en injectant du HRP dans le renflement cervical. Cette technique marquait seulement les neurones de la coucheV dans les champs corticaux granuleux et agranuleux. Les similitudes entre I'organisation du cortex moteur d6montr6es par cette etude et I'organisation chez le rat suggerent que le rat et la souris ont un plan commun d'organisation motrice corticale en tant que rongeurs. Can. J. Neurol. Sci. 199J; 18: 28-38The functional organization of the motor cortex in rat has been extensively studied 1 * 6 by systematically sampling a large number of cortical sites using intracortical microstimulation (ICMS), a technique developed by Asanuma and colleagues. These maps of motor cortex describe a complete representation of movements for all body parts. Often the results of these studies were related to the cytoarchitecture of the region studied. Although some ambiguities remain concerning the definition of primary motor cortex, the size and location of individual body part representations, and the extent of cytoarchitectonic boundaries, these studies have established some general parameters of motor organization of rat cortex.In contrast, knowledge of the functional organization of mouse motor cortex lags considerably behind that of the rat even though the mouse somatosensory cortex and the barrel field in part...

We studied the organization of the forepaw barrel subfield (FBS) in layer IV of adult rat somatosensory cortex using the mitochondrial marker cytochrome oxidase and related this organization to the representation of the forepaw. The FBS is an ovoid structure consisting of barrels and barrel-like structures, the most conspicuous of which form four centrally located medio lateral running bands. Each band contains three to four barrels. These centrally located bands are bordered along their entire lateral side by a nebulous zone of undifferentiated labeling. At the anterior border, two small barrels are located laterally and one or two larger barrels are located medially. Medial to the central zone are three well-defined barrels. The posterior border consists of a nebulous field of labeling and occasional barrel-like structures. The results from our electrophysiological recording and mapping revealed that the forepaw representation was topographically organized into a single map and that the forepaw map matches almost precisely with individual barrels and barrel-like structures in the FBS. Each of the four central bands is associated with the representation of a single glabrous digit. Digit two (D2) is represented anteriorly and followed posteriorly by D3 through D5. Within each digit band the digit is somatotopically organized, with the skin over the distal phalanx represented in the two lateral barrels and the middle and proximal phalanges represented in the medial barrel(s). The dorsal hairy digit skin and dorsal hand are represented in the lateral zone. D1 is represented by two small anteriorly located barrels. Medial to the representation of the glabrous digits is the representation of the palmar pads. The representation of these pads, in turn, lies between the representations of the thenar (located anteriorly) and hypothenar (located posteriorly) pads. Posterior to the hypothenar pad representation lie the representations of the wrist and forearm. While the present results support the conclusion that individual barrels are associated with discrete locations on the forepaw, examples were found where the recording site was not precisely located within the predicted barrel. Some of these errors may be accounted for by limitations in the mapping techniques; nevertheless, the FBS offers an excellent model system to study relationships between cortical structure and function.

Neurons in layer IV of rat somatosensory (SI) barrel cortex receive punctate somatic input from well-defined regions of the periphery. Following peripheral deafferentation, SI neurons in deafferented cortex respond to new input from neighboring regions of the skin surface. The precise mechanism(s) through which this occurs is unknown, although corticocortical and barreloid to barrel connections have been suggested as possible substrates. Because layer-IV barrels receive a strong afferent input from ventroposterior (VP) thalamic projection neurons, any divergence in the thalamocortical (TC) projection to multiple cortical barrels could also provide an anatomical substrate for rapid cortical reorganization. We used in-vivo intracellular recording methods to record and physiologically identify neurons in rat VP and to label those neurons with an intracellular tracer. Thalamic neurons (n=117) were impaled with sharp intracellular electrodes, and the receptive field(s) and firing pattern were measured. Cells were then injected with biocytin or biotinylated dextran amine (BDA). A total of 38 labeled TC neurons were quantitatively analyzed for soma size and dendritic arborization size; quantitative analysis of TC-axon arborizations in layer IV of barrel cortex was carried out in a total of 13 TC neurons. Two different axon-arborization patterns were identified in SI cortex: direct-projecting axons (n=6) were observed to project to and arborize within a single cortical barrel as well as extend their fibers into adjacent barrels; bifurcating-type axons (n=7) were seen to bifurcate in the subcortical white matter or in layer VI and then project to multiple barrel columns, where they arborized in layer IV. Axon fibers were always observed in three or more cortical barrels (mean=5, range=3-7). The mean mediolateral extent of arborizations in layer IV for the direct-projecting and bifurcating type axons were 458 microm and 1,302 microm, respectively, and these were significantly different (t=3.78, P<0.01). Axon-fiber length within cortical laminae was measured for each arborization pattern in relationship to the total fiber length within a cortical column. Direct-projecting axons always had greater than 50% of their fiber length within layer IV. Bifurcating-type axons were differentially distributed within multiple columns and always had less than 50% of their total column fiber length in layer IV. Morphological analysis of TC somata and dendrites revealed no correlation between local neuron morphology and axonal-arborization patterns. All intracellularly recorded TC neurons had similar adapting firing patterns when injected with a long-duration pulse. Our results showed that TC neurons project to multiple cortical barrels with one barrel receiving the principal input. This divergent TC projection pattern in SI cortex may provide an anatomical substrate for cortical plasticity and must be considered in any mechanism of rapid cortical reorganization.