Forepaw Barrel Research Articles

Lower jaw-to-forepaw rapid and delayed reorganization in the rat forepaw barrel subfield in primary somatosensory cortex.

We used the forepaw barrel subfield (FBS), that normally receives input from the forepaw skin surface, in rat primary somatosensory cortex as a model system to study rapid and delayed lower jaw-to-forepaw cortical reorganization. Single and multi-unit recording from FBS neurons was used to examine the FBS for the presence of "new" lower jaw input following deafferentations that include forelimb amputation, brachial plexus nerve cut, and brachial plexus anesthesia. The major findings are as follows: (1) immediately following forelimb deafferentations, new input from the lower jaw becomes expressed in the anterior FBS; (2) 7-27weeks after forelimb amputation, new input from the lower jaw is expressed in both anterior and posterior FBS; (3) evoked response latencies recorded in the deafferented FBS following electrical stimulation of the lower jaw skin surface are significantly longer in both rapid and delayed deafferents compared to control latencies for input from the forepaw to reach the FBS or for input from lower jaw to reach the LJBSF; (4) the longer latencies suggest that an additional relay site is imposed along the somatosensory pathway for lower jaw input to access the deafferented FBS. We conclude that different sources of input and different mechanisms underlie rapid and delayed reorganization in the FBS and suggest that these findings are relevant, as an initial step, for developing a rodent animal model to investigate phantom limb phenomena.

Functional connectivity in rat brain at 200 μm resolution.

The somatosensory functional magnetic resonance imaging (fMRI) response to electrical stimulation of the middle phalange of the second digit of four rats at a spatial resolution of 200 μm cubic at 9.4 T is reported. At high threshold (p<0.002), activated voxels encompass a penetrating vein that passes across the gray matter. These voxels lie mostly in three contiguous slices perpendicular to the pial surface. This activation is assigned to the representation in the forepaw barrel subfield (FBS) of a single cortical column of this phalange. In addition, the activation of the indusium griseum (IG) is visualized robustly. Voxels revealed by fMRI were used to observe functional connectivity to other voxels of the sensorimotor cortex using fcMRI. Results of this experiment were analyzed as a function of decreasing threshold, which exhibited spreading connectivity that revealed S2, M1/M2, and contralateral S1. Noting that every cubic millimeter of tissue contains 125 voxels, connectivity patterns are complex. It is hypothesized that they reflect connections within gray matter by association fibers. S2 and IG revealed connectivities with many voxels across the sensorimotor cortex. These regions also showed subregional variation of connectivity. A 1-cm-diameter surface coil with a local low-noise RF amplifier was used in these studies. The usual region of sensitivity (ROS) of such a coil is 1 cm diameter by 0.5 cm depth. Significant connectivity was observed between time courses of voxels that were within the ROS and voxels that were outside, which extends the volume of tissue that can be observed by the methods of this article.

Forelimb amputation-induced reorganization in the ventral posterior lateral nucleus (VPL) provides a substrate for large-scale cortical reorganization in rat forepaw barrel subfield (FBS)

In this study, we examined the role of the ventral posterior lateral nucleus (VPL) as a possible substrate for large-scale cortical reorganization in the forepaw barrel subfield (FBS) of primary somatosensory cortex (SI) that follows forelimb amputation. Previously, we reported that, 6 weeks after forelimb amputation in young adult rats, new input from the shoulder becomes expressed throughout the FBS that quite likely has a subcortical origin. Subsequent examination of the cuneate nucleus (CN) 1 to 30 weeks following forelimb amputation showed that CN played an insignificant role in cortical reorganization and led to the present investigation of VPL. As a first step, we used electrophysiological recordings in forelimb intact adult rats (n=8) to map the body representation in VPL with particular emphasis on the forepaw and shoulder representations and showed that VPL was somatotopically organized. We next used stimulation and recording techniques in forelimb intact rats (n=5) and examined the pattern of projection (a) from the forelimb and shoulder to SI, (b) from the forepaw and shoulder to VPL, and (c) from sites in the forepaw and shoulder representation in VPL to forelimb and shoulder sites in SI. The results showed that the projections were narrowly focused and homotopic. Electrophysiological recordings were then used to map the former forepaw representation in forelimb amputated young adult rats (n=5) at 7 to 24 weeks after amputation. At each time period, new input from the shoulder was observed in the deafferented forepaw region in VPL. To determine whether the new shoulder input in the deafferented forepaw VPL projected to a new shoulder site in the deafferented FBS, we examined the thalamocortical pathway in 2 forelimb-amputated rats. Stimulation of a new shoulder site in deafferented FBS antidromically-activated a cell in the former forepaw territory in VPL; however, similar stimulation from a site in the original shoulder representation, outside the deafferented region, in SI did not activate cells in the former forepaw VPL. These results suggest that the new shoulder input in deafferented FBS is relayed from cells in the former forepaw region in VPL. In the last step, we used anatomical tracing and stimulation and recording techniques in forelimb intact rats (n=9) to examine the cuneothalamic pathway from shoulder and forepaw receptive field zones in CN to determine whether projections from the shoulder zone might provide a possible source of shoulder input to forepaw VPL. Injection of biotinylated dextran amine (BDA) into physiologically identified shoulder responsive sites in CN densely labeled axon terminals in the shoulder representation in VPL, but also gave off small collateral branches into forepaw VPL. In addition, microstimulation delivered to forepaw VPL antidromically-activated cells in shoulder receptive field sites in CN. These results suggest that forepaw VPL also receives input from shoulder receptive sites in CN that are latent or subthreshold in forelimb intact rats. However, we speculate that following amputation these latent shoulder inputs become expressed, possibly as a down-regulation of GABA inhibition from the reticular nucleus (RTN). These results, taken together, suggest that VPL provides a substrate for large-scale cortical reorganization that follows forelimb amputation.

Forelimb amputation-induced reorganization in the cuneate nucleus (CN) is not reflected in large-scale reorganization in rat forepaw barrel subfield cortex (FBS)

We examined reorganization in cuneate nucleus (CN) in juvenile rat following forelimb amputation (n=34) and in intact controls (n=5) to determine whether CN forms a substrate for large-scale reorganization in forepaw barrel subfield (FBS) cortex. New input from the shoulder first appears in the FBS 4 weeks after amputation, and by 6 weeks, the new shoulder input comes to occupy most of the FBS. Electrophysiological recording was used to map CN in controls and in forelimb amputees during the first 12 weeks following deafferentation and at 26 and 30 weeks post-amputation. Mapping was confined to a location 300μm anterior to the obex where a medial-to-lateral row of electrode penetrations traversed through a complete complement of cytochrome-oxidase stained clusters (called barrelettes) that are associated with the representation of the glabrous forepaw digits and pads and adjacent non-cluster zones that are associated with the representation of the wrist, arm, and shoulder. Following amputation, non-cluster zones became occupied with new input from the body/chest and head/neck, while the cluster zone remained largely devoid of new input except at the border. A regression analysis comparing controls and amputees over the first 12 weeks post-amputation found significant differences for the total area of new input from the body/chest and head/neck in the non-cluster zones, while no significant differences were found for any new input into the cluster zone. When the averaged areas of a body-part representation were re-examined as a percentage of the averaged zonal area, a non-significant increase in new input from the body was observed within the cluster zone during post-amputation weeks 2–3 that returned to baseline in the subsequent weeks. In contrast, significant differences in averaged area of body-part representations for body/chest and head/neck were found in non-cluster zones over the first 12 weeks post-amputation. The present findings suggest that reorganization occurs only within the non-cluster zones whereby new input from the body/chest and head/neck moves in and occupies the deafferented territory immediately after amputation. Additionally, the lack of significant differences in new shoulder input in either cluster or non-cluster zones over the first 12 weeks after amputation suggests that CN provides an unlikely substrate for large-scale reorganization in the FBS.

Spike-Timing-Dependent Potentiation of Sensory Surround in the Somatosensory Cortex Is Facilitated by Deprivation-Mediated Disinhibition

Functional maps in the cerebral cortex reorganize in response to changes in experience, but the synaptic underpinnings remain uncertain. Here, we demonstrate that layer (L) 2/3 pyramidal cell synapses in mouse barrel cortex can be potentiated upon pairing of whisker-evoked postsynaptic potentials (PSPs) with action potentials (APs). This spike-timing-dependent long-term potentiation (STD-LTP) was only effective for PSPs evoked by deflections of a whisker in the neuron's receptive field center, and not its surround. Trimming of all except two whiskers rapidly opened the possibility to drive STD-LTP by the spared surround whisker. This facilitated STD-LTP was associated with a strong decrease in the surrounding whisker-evoked inhibitory conductance and partially occluded picrotoxin-mediated LTP facilitation. Taken together, our data demonstrate that sensory deprivation-mediated disinhibition facilitates STD-LTP from the sensory surround, which may promote correlation- and experience-dependent expansion of receptive fields.

Amputation with Median Nerve Redirection (Targeted Reinnervation) Reactivates Forepaw Barrel Subfield in Rats

Prosthetic limbs are difficult to control and do not provide sensory feedback. Targeted reinnervation was developed as a neural-machine interface for amputees to address these issues. In targeted reinnervation, amputated nerves are redirected to proximal muscles and skin, creating nerve interfaces for prosthesis control and sensory feedback. Touching the reinnervated skin causes sensation to be projected to the missing limb. Here we use electrophysiological brain recording in the Sprague Dawley rat to investigate the changes to somatosensory cortex (S1) following amputation and nerve redirection with the intent to provide insight into the sensory phenomena observed in human targeted reinnervation amputees. Recordings revealed that redirected nerves established an expanded representation in S1, which may help to explain the projected sensations that encompass large areas of the hand in targeted reinnervation amputees. These results also provide evidence that the reinnervated target skin could serve as a line of communication from a prosthesis to cortical hand processing regions. S1 border regions were simultaneously responsive to reinnervated input and also vibrissae, lower lip, and hindfoot, suggesting competition for deactivated cortical territory. Electrically evoked potential latencies from reinnervated skin to cortex suggest direct connection of the redirected afferents to the forepaw processing region of S1. Latencies also provide evidence that the widespread reactivation of S1 cortex may arise from central anatomical interconnectivity. Targeted reinnervation offers the opportunity to examine the cortical plasticity effects when behaviorally important sensory afferents are redirected from their original location to a new skin surface on a different part of the body.

Mapping plasticity in the forepaw digit barrel subfield of rat brains using functional MRI

The topographic organization of the forepaw barrel subfield in layer IV of rat primary somatosensory cortex (S1) is a good model for studying neural function and plasticity. The goal of this study was to test the feasibility of functional MRI (fMRI) to map the forepaw digit representations in the S1 of the rat and its plasticity after digit amputation. Three dimensional echo-planar imaging with 300micron isotropic resolution at 11.7T was used to achieve high signal-to-noise ratios and laminar layer resolution. By alternating electrical stimulation of the 2nd (D2) and 4th (D4) digits, functional activation in layer IV of the barrel subfields could be distinguished using a differential analysis. Furthermore, 2 and a half months after the amputation of the 3rd digit in baby rats, the overlapping area between D2 and D4 representations was increased. This indicates that the forepaw barrel subfield previously associated with the ablated digit is now associated with the representation of nearby digits, which is consistent with studies using electrophysiology and cytochrome oxidase staining.

Prenatal alcohol exposure (PAE) reduces the size of the forepaw representation in forepaw barrel subfield (FBS) cortex in neonatal rats: relationship between periphery and central representation

Prenatal alcohol exposure (PAE) alters limb development that may lead to structural and functional abnormalities of the limb reported in children diagnosed with Fetal Alcohol Spectrum Disorder. To determine whether PAE alters the central representation of the forelimb we used the rodent barrel cortex as our model system where it was possible to visualize and quantitatively measure the size of the forepaw representation in the forepaw barrel subfield (FBS) in first somatosensory cortex. In the present study, we examined the effects of PAE on pattern and size of the forepaw and forepaw representation in FBS in neonatal rats at gestational day 32 that corresponds to postnatal day 9. Pregnant Sprague-Dawley rats were chronically intubated with binge doses of ethanol (6 g/kg) from gestational day 1 through gestational day 20. The offspring of the ethanol treated dams comprised the ethanol (EtOH) group. The effect of PAE on the EtOH group was compared with a nutritional-controlled pairfed (PF) group and a normal chowfed (CF) group. The ventral (glabrous) surface area of the forepaw digits, length of digit 2 through digit 5, and the corresponding glabrous forepaw digit representations in the FBS were measured and compared between treatment groups. In rats exposed to in utero alcohol, the sizes of the overall glabrous forepaw and forepaw digits were significantly reduced in EtOH pups compared to CF and PF pups; overall glabrous forepaw area was 11% smaller than CF controls. Glabrous digit lengths were also smaller in EtOH rats compared to CF controls and significantly smaller in digit 2 through digit 4. The glabrous digit representation in FBS was 18% smaller in the EtOH group when compared to the CF treatment. However, PAE did not produce malformations in the forepaw or alter the pattern of the forepaw representation in FBS; instead, PAE significantly reduced both body and brain weights compared to controls. Unexpectedly, little or no correlation was observed between the size of the glabrous forepaw compared to the size of the glabrous forepaw representation in the FBS for any of the treatment groups. The present findings of PAE-related alterations in sensory periphery and the central cortical representation may underlie deficits in sensorimotor integration reported among children with Fetal Alcohol Spectrum Disorder.

Delayed reorganization of the shoulder representation in forepaw barrel subfield (FBS) in first somatosensory cortex (SI) following forelimb deafferentation in adult rats.

We previously reported that 6-16 weeks after forelimb amputation in adult rats, neurons in layer IV of rat first somatosensory cortex (SI) in the forepaw barrel subfield (FBS) associated with the representation of the forepaw became responsive to new input from the shoulder (Pearson et al. 1999). These new shoulder-responsive sites in deafferented FBS had longer evoked response latencies than did sites in the shoulder representation located in the posterior part of the trunk subfield, hereafter referred to as the original shoulder representation. Furthermore, projection neurons in the original shoulder representation in both intact and deafferented adults did not extend their axons into the FBS, and ablation of the original shoulder representation cortex and/or the second somatosensory cortex (SII) failed to eliminate new shoulder input in the deafferented FBS (Pearson et al. 2001). These results led us to conclude that large-scale reorganization in FBS quite likely involved a subcortical substrate. In addition, the time course for large-scale cortical reorganization following forelimb amputation was unknown, and this information could shed light on potential mechanisms for large-scale cortical reorganization. In the present study, we extended our previous findings of large-scale cortical reorganization in the FBS by investigating the time course for reorganization following forelimb amputation. The major findings are: a) deafferented forelimb cortex remained unresponsive to shoulder stimulation during the 1st week following forelimb amputation; b) new responses to shoulder stimulation were first observed in deafferented forelimb cortex 2-3 weeks after forelimb amputation; however, the new shoulder input was restricted to locations in the former forearm cortex; c) islet(s) of new shoulder representation were first observed in deafferented FBS 4 weeks after amputation; these islets occupied a larger percentage of FBS in subsequent weeks; d) portions of FBS remained unresponsive as many as 4 months after deafferentation (maximum time examined between amputation and recording); and e) the increase in total size of the shoulder representation appeared to result from the establishment of new shoulder representations that were often discontinuous from the original shoulder representation. These findings provide evidence that forelimb amputation results in delayed reorganization of the FBS and we describe possible mechanisms and substrates underlying the reorganization.

Removal of GABAergic inhibition alters subthreshold input in neurons in forepaw barrel subfield (FBS) in rat first somatosensory cortex (SI) after digit stimulation.

Our objective was to test the hypothesis that suppression of GABAergic inhibition results in an enhancement of responses to stimulation of the surround receptive field. Neurons in the forepaw barrel subfield (FBS) in rat first somatosensory cortex (SI) receive short latency suprathreshold input from a principal location on the forepaw and longer latency subthreshold input from surrounding forepaw skin regions. Input from principal and surround receptive field sites was examined before, during, and after administration of the GABA(A) receptor blocker bicuculline methiodide (BMI) (in 165 mM NaCl at pH 3.3-3.5). In vivo extracellular recording was used to first identify the location of the glabrous forepaw digit representation within the FBS. In vivo intracellular recording and labeling techniques were then used to impale single FBS neurons in layer IV as well as neurons in layers III and V, determine the receptive field of the cell, and fill the cell with biocytin for subsequent morphological identification. The intracellular recording electrode was fastened with dental wax to a double-barrel pipette for BMI iontophoresis and current balance. A stimulating probe, placed on the glabrous forepaw skin surface, was used to identify principal and surround components of the receptive field. Once a cell was impaled and a stable recording was obtained, a stimulating probe was placed at a selected site within the surround receptive field. Single-pulse stimulation (1 Hz) was then delivered through the skin probe and the percentage of spikes occurring in 1-min intervals before BMI onset was used as a baseline measure. BMI was then iontophoresed while the periphery was simultaneously stimulated, and spike percentage measured during and after BMI ejection was compared with the pre-BMI baseline. The major findings are: (1) suppression of GABAergic inhibition enhanced evoked responses to firing level from sites in surround receptive fields in 65% of the cells ( n=17); (2) evoked responses were rapidly elevated (within 1 min) to suprathreshold firing in the presence of BMI in 31% of the cells; (3) GABAergic inhibition was reversible [suprathreshold spiking gradually reversed to subthreshold excitatory postsynaptic potentials (EPSPs) in 45% of the cells tested]; (4) BMI altered the stimulus-evoked and non-stimulus-evoked firing pattern in SI neurons from single spikes to burst patterns in all tested cells; and (5) iontophoresis of NaCl (165 mM) without BMI was ineffective in altering evoked responses in control cells ( n=4). The present findings support the notion that subthreshold input from surround receptive fields is one possible mechanism for rapid cortical reorganization in barrel cortex and that GABAergic inhibition may regulate its expression. Possible corticocortical and thalamocortical substrates for subthreshold input to reach barrel neurons are discussed.

Large-scale cortical reorganization following forelimb deafferentation in rat does not involve plasticity of intracortical connections.

Physiological mapping of the body representation 1 month or longer after forelimb removal in adult rats revealed new pockets of shoulder representation in the forepaw barrel subfield (FBS) in the first somatosensory cortex (SI). These "new" shoulder representations have longer evoked response latencies than sites in the shoulder representation within the trunk subfield, hereafter referred to as the "original" shoulder representation. We postulated that the "new" shoulder representations in the FBS were relayed from the "original" shoulder representation. We investigated this hypothesis by studying anatomical connectivity between the "original" shoulder representation and the FBS in intact control and forelimb deafferented adult rats using Phaseolus vulgaris leucoagglutinin (PHA-L), biocytin, and biotin dextran-amine (BDA) as anterograde tracers. The retrograde tracer cholera toxin beta subunit (CT-B) injected into the FBS was also used to study connectivity between the "original" shoulder representation and the FBS. Using these anatomical tracing techniques, we were unable to show the existence of a direct corticocortical connection between the "original" shoulder representation in the trunk subfield and the FBS in either intact or deafferented rats. Functional connectivity between the two cortical regions was studied by ablating the "original" shoulder representation alone or in combination with the shoulder representation in the second somatosensory cortex (SII) while recording evoked responses in the FBS following electrical stimulation of the shoulder. Both ablations failed to eliminate the evoked responses at the "new" shoulder sites in the FBS, suggesting that SI and SII are not necessary for "new" shoulder input in the FBS. It is suggested that subcortical sites may play a major role in large-scale cortical reorganization. Results of projections from the "original" shoulder representation to parietal medial (PM), parietal lateral (PL), SII, parietal ventral (PV), and parietal rhinal (PR) sensory fields and agranular lateral (AgL) and agranular medial (AgM) motor fields are also described.

Effects of large-scale limb deafferentation on the morphological and physiological organization of the forepaw barrel subfield (FBS) in somatosensory cortex (SI) in adult and neonatal rats.

The physiological representation of the shoulder and surrounding body was examined in layer IV of somatosensory cortex (SI) in rats that had underground removal of the forelimb, either as newborns on postnatal day three (PND-3) or as adults (at least 8 weeks of age). Electrophysiological recordings were used to map the shoulder and body representations (physiological map), and the mitochondria marker, cytochrome oxidase (CO), was used to visualize recording sites in barrel and barrel-like structures (morphological map) in layer IV of deafferents and intact controls. The SI shoulder representation lies in a nebulously stained region that lies posterior to the forearm, wrist, and forepaw representations; the latter region is associated with the well-defined forepaw barrel subfield (FBS). The major findings are: (1) the shoulder is represented as a single zone located at the posterior extent of the SI body map in intact rats; (2) limb deafferentation in adult or neonatal rats that were physiologically mapped 6-16 weeks post-amputation resulted in two or more islets of "new" representation of the shoulder in the FBS in addition to the representation of the "original" shoulder in the posterior part of the body map; (3) deafferentations made in neonatal rats, physiologically mapped as adults, had a significantly greater (Mann-Whitney U) amount of "new" cortical representation within the FBS than did rats deafferented as adults; (4) fewer unresponsive sites in the FBS were found for neonate deafferents than for adult deafferents; (5) evoked response latencies following electrical stimulation of the shoulder were shortest for cortical sites within the "original" shoulder representation in intact controls, and latencies recorded at the "original" shoulder representation in deafferents were also shorter than latencies recorded in "new" shoulder representations in both groups of deafferents; and (6) morphological maps of the FBS were altered in neonate deafferents to the extent that the barrel structure was poorly formed, as exemplified by the absence of the four mediolateral running bands; however, the overall ovoid shape of the FBS was still apparent, but not as sharply defined as for intact controls or adult deafferents. Possible mechanisms for reorganization following large-scale deafferentation are discussed.

In vivo intracellular recording and labeling of neurons in the forepaw barrel subfield (FBS) of rat somatosensory cortex: possible physiological and morphological substrates for reorganization.

We examined the physiological properties and morphology of neurons in the forepaw barrel subfield (FBS) in somatosensory cortex (SI) of adult rats using in vivo intracellular recording and biocytin labeling techniques. Our results show that both pyramidal and non-pyramidal type spiny neurons can be activated with short latency by peripheral stimulation. FBS neurons within individual barrels receive both suprathreshold and subthreshold convergent input from one or more forepaw digits and pads. We hypothesize that some of these subthreshold inputs may be elevated to firing level by some, as yet unknown, mechanism(s) following peripheral deafferentation. Examination of the relationship between the dendritic pattern of labeled neurons and individual barrels within the FBS suggests that neurons with dendrites extending into neighboring barrel bands may serve as a possible morphological substrate for immediate reorganization.

Relationship between representation of hindpaw and hindpaw barrel subfield (HBS) in layer IV of rat somatosensory cortex.

We describe the organization of the hindpaw barrel subfield (HBS) in layer IV of rat somatosensory cortex (SI) and relate this organization to the representation of the hindpaw. The ovoid-shaped, HBS is oriented anterior to posterior and comprises barrels and barrel-like structures, the most prominent of which consist of at least five anteriorly-located elongated barrel bands. Posterior to these elongated bands is a cluster of four barrels. Two additional barrels are found, one lateral, the other medial. The lateral border is formed by a nearly continuous band that overlaps portions of the anterior elongated bands and posterior barrels. The HBS shows considerable variability in size and shape; nevertheless, the overall pattern reflects a common plan of organization. Electrophysiological mapping confirmed that hindpaw representation is somatotopically organized. The glabrous toes are represented anteriorly, the pads posteriorly, and the dorsal hairy skin of the toes and hindpaw laterally. By aligning physiological and morphological (HBS) maps according to lesion sites, our data suggest that the elongated anteriorly-located barrel bands represent the hindpaw toes, the four toe pads are represented immediately posterior followed by barrels representing the plantar pads. The representations of dorsal hairy skin of toe and dorsal hindpaw form the lateral border; the heel and ankle are represented most posterior. We interpret our findings as support that individual barrels in the HBS are associated with discrete regions of the hindpaw; however, the precise relationship of structure and function reported between the vibrissae and posteromedial barrel subfield (PMBSF) and between the forepaw and the forepaw barrel subfield (FBS) were not observed.

Digit removal leads to discrepancies between the structural and functional organization of the forepaw barrel subfield in layer IV of rat primary somatosensory cortex.

The physiological representation of the forepaw in rat primary somatosensory cortex (SI) is topographically organized. This representation is associated with the unique arrangement of barrels in layer IV of the forepaw barrel subfield (FBS) in SI and provides an example of a relationship between cortical structure and function. It has been reported that removal of peripheral afferent input to the FBS prior to postnatal day 5 or 6 results in a disorganized FBS, while deafferentation at later times produces little or no alteration of the FBS. Therefore, restricted deafferentations of individual digits in adult rats should result in little, if any, disruption of the FBS, while at the same time eliminating afferent input to the FBS from a localized region of the periphery. This manipulation is likely to create a mismatch between structure and function and offer insight into what barrels actually represent in the adult deafferent. In the present study, we amputated digit three (D3) in eight adult rats, allowed a 1-month survival time, physiologically mapped the representation of D2, D4, and the stump, and compared this physiological map to the underlying barrels in the FBS. Our results showed that FBS barrels formerly associated with the representation of D3 were now associated with the representation of surrounding digits D2 and D4, as well as the remaining stump. By superimposing the morphological and physiological map upon one another, it was clear that the D2 and D4 representations expanded into the former D3 barrel territory and septae between the barrels. The reorganized physiological map was somatotopically organized, even though the general configuration of the morphological map remained unaltered, as visualized with cytochrome oxidase staining. These results suggest that in the deafferent, neurons within FBS barrels previously associated with the representation of punctate regions of skin become associated with neighboring regions of skin. A morphological substrate to account for this cortical reorganization is described.

Relationship between the organization of the forepaw barrel subfield and the representation of the forepaw in layer IV of rat somatosensory cortex.

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.