Adjacent Whiskers Research Articles

Responses of barrel cortex neurons in awake rats and effects of urethane anesthesia.

A "barrel" is an interconnected network of layer IV neurons that is an important component of a functional cortical column in the whisker area of the rodent primary somatosensory cortex. The present study was undertaken in order to resolve apparently conflicting findings from single-unit studies of barrel neurons conducted in rats maintained under different anesthetic conditions. Multiunit responses to controlled deflections of mystacial vibrissae were recorded from the whisker/barrel cortex of awake, undrugged rats, and responses at the same recording site were reexamined after the animal was anesthetized with urethane. In contrast to the awake condition, stimulus-evoked responses under urethane were characterized by a large late component. Such effects were more pronounced for deflections of noncolumnar or "adjacent" whiskers than for the columnar whisker. Latencies to peak responses were virtually identical for the columnar whisker in awake and urethane states (11.9 vs 11.8 ms) but were considerably longer for adjacent whisker deflections in urethane-anesthetized animals (15.5 vs 29.0 ms). The magnitudes of adjacent whisker responses, relative to the response evoked by the columnar whisker, varied with the laminar location of the recording site in awake but not in urethane-anesthetized animals; in awake rats, receptive fields were clearly smallest in the layer IV barrels. Results in the awake condition confirm those of previous studies conducted in unanesthetized or lightly sedated animals, and data obtained with urethane are comparable to others' results in urethane-anesthetized rats. The former have important implications for how barrel cortex processes information in behaving animals.

Functional Asymmetries in the Rodent Barrel Cortex

Neurophysiological and 2-deoxyglucose (2DG) studies of the rodent whisker barrel cortex have demonstrated asymmetries in its functional organization. To examine the possibility that the activity gradients observed in metabolic studies can be attributed to subtle rostral-caudal and dorsal-ventral asymmetries in electrophysiologically measured surround or cross-whisker inhibition, we compared 2DG results with predictions generated from quantitative single-cell receptive field data. Despite differences in the two experimental approaches, there is remarkable agreement between the findings. (1) The distribution of 2DG activity declines across the barrel cortex of the behaving animal from anteromedial barrels to posterolateral barrels, and is qualitatively and quantitatively similar to the values predicted from neurophysiology. (2) The strength of surround inhibition in barrel neurons predicts the twofold increase in activation of the C3 barrel following acute clipping of adjacent whiskers. And (3) within a cortical column, the decrease in metabolic activity associated with adjacent whisker stimulation is greatest in layer IV and least in the infragranular layers; this corresponds to the laminar distribution of inhibitory interactions observed electrophysiologically.

Thalamocortical response transformation in the rat vibrissa/barrel system.

1. Extracellular single-unit recordings and controlled whisker stimuli were used to compare response properties between cells in the "barreloids" of the thalamic ventrobasal complex and those in the cytochrome oxidase-rich centers of the "barrels" in the first somatic sensory cortex. Individual vibrissae were deflected alone or in paired combination involving the neuron's maximally excitatory whisker and an adjacent one in the same or neighboring whisker rows. Quantitative data were derived from 135 thalamocortical unit's (TCUs), 242 "regular-spike" barrel units (RSUs), and 16 "fast-spike" barrel units (FSUs) recorded in 26 normal adult rats. 2. Compared with TCUs, RSUs displayed lower rates of spontaneous activity and responded less vigorously to whisker stimuli. Proportionally, more than twice as many TCUs as RSUs responded in slowly adapting fashion to at least one angular direction of whisker displacement. Discharges of slowly adapting TCUs were approximately 3.5 times greater than those of slowly adapting RSUs. 3. Proportionally, about twice as many TCUs than RSUs responded selectively to whisker movements in different angular directions. 4. Cells in the thalamus responded more vigorously to a larger number of whiskers than RSUs in the cortex. Depending on the stimulus conditions, two to three times more TCUs than RSUs were excited by two or more whiskers. 5. Following displacement of an adjacent whisker, unit discharges to subsequent deflections of the maximally excitatory whisker were reduced in a time-dependent fashion. The time course of response suppression was similar in TCUs and RSUs, but inhibitory interactions between adjacent whiskers were observed much less often in the thalamus. A cyclic pattern of stimulus-evoked excitation/inhibition characterizes responses in the cortical barrels but is considerably less pronounced in the thalamic barreloids. 6. The presence and/or degree of response suppression depended on which adjacent whisker was stimulated and on the angular direction of that whisker's movement. For individual TCUs, some adjacent whiskers evoked inhibition, others did not. The vast majority of RSUs displayed response suppression to all adjacent whiskers. Unlike receptive fields of TCUs, those of RSUs have small--i.e., single-whisker--excitatory centers with potent and symmetrical inhibitory surrounds. 7. Fast-spike units in the barrels displayed the greatest spontaneous and stimulus-evoked activities and were the least selective for whisker movements at different angular directions. FSUs had the largest excitatory receptive fields; 100% responded to two or more vibrissae.(ABSTRACT TRUNCATED AT 400 WORDS)

Somatotopic organization of the second somatosensory area (SII) in the cerebral cortex of the mouse.

The somatotopic organization of the parietal cortex of barbiturate-anesthetized, adult mice was studied using tungsten microelectrodes. A complete representation of the contralateral face and body occupying approximately 4.0-4.5 mm2 was found immediately posterior and lateral to the representation of the face in the first somatosensory area (SI). Within this second somatosensory area (SII), the following findings were made: A relatively large region is devoted to representations of the paws and face, especially the sinus hairs associated with the anterior upper lip and mystacial vibrissae. Receptive fields on these body regions are among the smallest found in SII, though larger than corresponding receptive fields in SI. In particular, vibrissae receptive fields always include at least several adjacent whiskers, and paw receptive fields always include at least two adjacent digits. In regions representing proximal body parts, receptive fields are considerably larger, may include both contralateral and ipsilateral limb or trunk surfaces, and sometimes include the entire body and face. Responses to both somatosensory and auditory stimulation were consistently found in the body (i.e., trunk and limb) representation, but rarely found in the face region. The face is represented most anteriorly, and the hindlimb and tail most posteriorly. Forepaw and hindpaw digits and anterior aspects of the face (e.g., perioral sinus hairs and the incisors) are represented laterally, while the back, caudal head, and mystacial vibrissae are represented medially. Within SII, therefore, a "musculus" can be viewed as having an upright body orientation with the face area bordering the face representation within SI. By comparison with SI, SII is characterized by a less pronounced layer IV, which is of irregular thickness and packing density, and by less uniformity in the layering of pyramidal cells in lamina V. In addition, SII is generally thicker from pia to white matter than SI. These results are in general accord with earlier findings from evoked potential studies in mice, but are at variance with recent reports in mice and rats that the mystacial vibrissae have only a minimal, or no, representation within SII. Indeed, the present findings suggest that the representation of the whiskers in SII may have a specialized function paralleling that in SI.

Temporal and spatial integration in the rat SI vibrissa cortex.

Glass micropipettes were used to record the activity of 124 single units in the somatosensory vibrissa cortex (SI) of 16 rats in response to combined deflections of contralateral vibrissae. Compact multiangular electromechanical stimulators were used to stimulate individual vibrissal hairs alone or in combinations of two or three adjacent whiskers. Each whisker was stimulated independently to produce controlled temporal and spatial patterns of mechanical stimuli. Following displacement of a vibrissa, unit discharges to subsequent deflections of adjacent whiskers are reduced in a time-dependent fashion. Response suppression is strongest at short interdeflection intervals, i.e., 10-20 ms and decreases progressively during the 50-100 ms following the first deflection. In many cases this period also corresponds with a reduction in ongoing unit discharges. Response suppression was not observed for first-order neurons recorded in the trigeminal ganglion of barbiturate-anesthetized rats. In the cortex, the presence and/or degree of response suppression depends on a number of spatial factors. These include 1) the angular direction(s) in which the individual hairs are moved, 2) the sequence in which two whiskers are deflected, that is, which one is deflected first, 3) the particular combination of whiskers stimulated, and 4) the number (2 or 3) of vibrissae comprising the multiwhisker stimulus. Within a vertical electrode penetration, one particular whisker typically elicits the strongest excitatory and inhibitory effects; other, nearby vibrissae elicit variable (or no) excitation or inhibition. Excitatory and inhibitory subregions of a receptive field could thus be distributed asymmetrically around the maximally effective whisker. In these cases, the receptive fields displayed spatial orientations. Quantitative criteria were used to classify 30 cortical units on the basis of the distribution of inhibitory subregions on either side of the maximally effective whisker. Twenty-one of these cells had receptive fields (RFs) with symmetrical inhibitory side regions. Responses of the other nine units were strongly suppressed by a preceding deflection of a vibrissa on one side but relatively unaffected, or even slightly facilitated, by preceding deflection of the whisker on the other side.(ABSTRACT TRUNCATED AT 400 WORDS)