Dysgranular Zone Research Articles

Metabotropic Glutamate Receptor 5 in the Dysgranular Zone of Primary Somatosensory Cortex Mediates Neuropathic Pain in Rats.

The primary somatosensory cortex (S1) plays a key role in the discrimination of somatic sensations. Among subdivisions in S1, the dysgranular zone of rodent S1 (S1DZ) is homologous to Brodmann’s area 3a of primate S1, which is involved in the processing of noxious signals from the body. However, molecular changes in this region and their role in the pathological pain state have never been studied. In this study, we identified molecular alteration of the S1DZ in a rat model of neuropathic pain induced by right L5 spinal nerve ligation (SNL) surgery and investigated its functional role in pain symptoms. Brain images acquired from SNL group and control group in our previous study were analyzed, and behaviors were measured using the von Frey test, acetone test, and conditioned place preference test. We found that metabotropic glutamate receptor 5 (mGluR5) levels were significantly upregulated in the S1DZ contralateral to the nerve injury in the SNL group compared to the sham group. Pharmacological deactivation of mGluR5 in S1DZ ameliorated symptoms of neuropathic allodynia, which was shown by a significant increase in the mechanical paw withdrawal threshold and a decrease in the behavioral response to cold stimuli. We further confirmed that this treatment induced relief from the tonic-aversive state of chronic neuropathic pain, as a place preference memory associated with the treatment-paired chamber was formed in rats with neuropathic pain. Our data provide evidence that mGluR5 in the S1DZ is involved in the manifestation of abnormal pain sensations in the neuropathic pain state.

Intrinsic functional clustering of ventral premotor F5 in the macaque brain

Neurophysiological and anatomical data suggest the existence of several functionally distinct regions in the lower arcuate sulcus and adjacent postarcuate convexity of the macaque monkey. Ventral premotor F5c lies on the postarcuate convexity and consists of a dorsal hand-related and ventral mouth-related field. The posterior bank of the lower arcuate contains two additional premotor F5 subfields at different anterior-posterior levels, F5a and F5p. Anterior to F5a, area 44 has been described as a dysgranular zone occupying the deepest part of the fundus of the inferior arcuate. Finally, area GrFO occupies the most rostral portion of the fundus and posterior bank of inferior arcuate and extends ventrally onto the frontal operculum. Recently, data-driven exploratory approaches using resting-state fMRI data have been suggested as a promising non-invasive method for examining the functional organization of the primate brain. Here, we examined to what extent partitioning schemes derived from data-driven clustering analysis of resting-state fMRI data correspond with the proposed organization of the fundus and posterior bank of the macaque arcuate sulcus, as suggested by invasive architectonical, connectional and functional investigations. Using a hierarchical clustering analysis, we could retrieve clusters corresponding to the dorsal and ventral portions of F5c on the postarcuate convexity, F5a and F5p at different antero-posterior locations on the posterior bank of the lower arcuate, area 44 in the fundus, as well as part of area GrFO in the most anterior portion of the fundus. Additionally, each of these clusters displayed distinct whole-brain functional connectivity, in line with previous anatomical tracer and seed-based functional connectivity investigations of F5/44 subdivisions. Overall, our data suggests that hierarchical clustering analysis of resting-state fMRI data can retrieve a fine-grained level of cortical organization that resembles detailed parcellation schemes derived from invasive functional and anatomical investigations.

IL-17a promotes sociability in mouse models of neurodevelopmental disorders.

Asubset of children with autism spectrum disorderappear to show an improvement in their behavioural symptoms during the course of a fever, a sign of systemic inflammation1,2. Here we elucidate the molecular and neural mechanisms that underlie the beneficial effects of inflammation on social behaviour deficits in mice. We compared an environmental model of neurodevelopmental disorders in which mice were exposed to maternal immune activation (MIA) during embryogenesis3,4 with mouse models that are genetically deficient for contactin-associatedprotein-like2 (Cntnap2)5, fragileXmentalretardation-1 (Fmr1)6 or Sh3andmultipleankyrinrepeat domains3 (Shank3)7. We establish that the social behaviour deficits in offspring exposed to MIA can be temporarily rescued by the inflammatory response elicited by the administration of lipopolysaccharide (LPS). This behavioural rescue was accompanied by a reduction in neuronal activity in the primary somatosensory cortex dysgranular zone (S1DZ), the hyperactivity of which was previously implicated in the manifestation of behavioural phenotypes associated with offspring exposed to MIA8. By contrast, we did not observe an LPS-induced rescue of social deficits in the monogenic models. We demonstrate that the differences in responsiveness to the LPS treatment between the MIA and the monogenic models emerge from differences in the levels of cytokine production. LPS treatment in monogenic mutant mice did not induce amounts of interleukin-17a (IL-17a) comparable to those induced in MIA offspring; bypassing this difference by directly delivering IL-17a into S1DZ was sufficient to promote sociability in monogenic mutant mice as well as in MIA offspring. Conversely, abrogating the expression of IL-17 receptor subunit a (IL-17Ra) in the neurons of the S1DZ eliminated the ability of LPS to reverse the sociability phenotypes in MIA offspring. Our data support a neuroimmune mechanism that underlies neurodevelopmental disorders in which the production of IL-17a during inflammation can ameliorate the expression of social behaviour deficits by directly affecting neuronal activity in the central nervous system.

Long-Range, Border-Crossing, Horizontal Axon Radiations Are a Common Feature of Rat Neocortical Regions That Differ in Cytoarchitecture.

Employing wide-field optical imaging techniques supported by electrophysiological recordings, previous studies have demonstrated that stimulation of a spatially restricted area (point) in the sensory periphery results in a large evoked neuronal activity spread in mammalian primary cortices. In rats’ primary cortices, such large evoked spreads extend diffusely in multiple directions, cross cortical cytoarchitectural borders and can trespass into other unimodal sensory areas. These point spreads are supported by a spatially matching, diffuse set of long-range horizontal projections within gray matter that extend in multiple directions and cross borders to interconnect different cortical areas. This horizontal projection system is in addition to well-known area-to-area clustered projections to defined targets through white matter. Could similar two-projection cortical systems also be found in cortical regions that differ in their cytoarchitectural structure? To address this question, an adeno-associated viral vector expressing green fluorescent protein (GFP) was injected as an anterograde tract tracer into granular somatosensory cortex (trunk area), dysgranular cortex (somatosensory dysgranular zone and extrastriate cortex) and agranular motor cortex (MCx). Irrespective of the injection site the same two projection systems were found, and their quantification revealed a close similarity to findings in primary sensory cortices. Following detailed reconstruction, the diffuse horizontal axon radiation was found to possess numerous varicosities and to include short, medium and long axons, the latter extending up to 5.2 mm. These “proof of concept” findings suggest that the similarity of the two projection systems among different cortical areas could potentially constitute a canonical motif of neocortical organization.

Diverse Long-Range Axonal Projections of Excitatory Layer 2/3 Neurons in Mouse Barrel Cortex

Excitatory projection neurons of the neocortex are thought to play important roles in perceptual and cognitive functions of the brain by directly connecting diverse cortical and subcortical areas. However, many aspects of the anatomical organization of these inter-areal connections are unknown. Here, we studied long-range axonal projections of excitatory layer 2/3 neurons with cell bodies located in mouse primary somatosensory barrel cortex (wS1). As a population, these neurons densely projected to secondary whisker somatosensory cortex (wS2) and primary/secondary whisker motor cortex (wM1/2), with additional axon in the dysgranular zone surrounding the barrel field, perirhinal temporal association cortex and striatum. In three-dimensional reconstructions of 6 individual wS2-projecting neurons and 9 individual wM1/2-projecting neurons, we found that both classes of neurons had extensive local axon in layers 2/3 and 5 of wS1. Neurons projecting to wS2 did not send axon to wM1/2, whereas a small subset of wM1/2-projecting neurons had relatively weak projections to wS2. A small fraction of projection neurons solely targeted wS2 or wM1/2. However, axon collaterals from wS2-projecting and wM1/2-projecting neurons were typically also found in subsets of various additional areas, including the dysgranular zone, perirhinal temporal association cortex and striatum. Our data suggest extensive diversity in the axonal targets selected by individual nearby cortical long-range projection neurons with somata located in layer 2/3 of wS1.

Reversing behavioural abnormalities in mice exposed to maternal inflammation.

Viral infection during pregnancy is correlated with increased frequency of neurodevelopmental disorders, and this is studied in mice prenatally subjected to maternal immune activation (MIA). We previously showed that maternal T helper 17 cells promote the development of cortical and behavioural abnormalities in MIA-affected offspring. Here we show that cortical abnormalities are preferentially localized to a region encompassing the dysgranular zone of the primary somatosensory cortex (S1DZ). Moreover, activation of pyramidal neurons in this cortical region was sufficient to induce MIA-associated behavioural phenotypes in wild-type animals, whereas reduction in neural activity rescued the behavioural abnormalities in MIA-affected offspring. Sociability and repetitive behavioural phenotypes could be selectively modulated according to the efferent targets of S1DZ. Our work identifies a cortical region primarily, if not exclusively, centred on the S1DZ as the major node of a neural network that mediates behavioural abnormalities observed in offspring exposed to maternal inflammation.

Representation of egomotion in rat's trident and E-row whisker cortices

The whisker trident, a three-whisker array on the rat's chin, has been implicated in egomotion sensing and might function as a tactile speedometer. Here we study the cortical representation of trident whiskers and E-row whiskers in barrel cortex. Neurons identified in trident cortex of anesthetized animals showed sustained velocity-sensitive responses to ground motion. In freely moving animals, about two-thirds of the units in the trident and E-row whisker cortices were tuned to locomotion speed, a larger fraction of speed-tuned cells than in the somatosensory dysgranular zone. Similarly, more units were tuned to acceleration and showed sensitivity to turning in trident and E-row whisker cortices than in the dysgranular zone. Microstimulation in locomoting animals evoked small but significant speed changes, and such changes were larger in the trident and E-row whisker representations than in the dysgranular zone. Thus, activity in trident and E-row cortices represents egomotion information and influences locomotion behavior.

Coalescence of deep and superficial epileptic foci into larger discharge units in adult rat neocortex

Epilepsy is a disease of neuronal hyper-synchrony that can involve both neocortical and hippocampal brain regions. While much is known about the network properties of the hippocampus little is known of how epileptic neocortical hyper-synchrony develops. We aimed at characterizing the properties of epileptic discharges of a neocortical epileptic focus.We established a multi-electrode-array method to record the spatial patterns of epileptiform potentials in acute adult rat brain slices evoked by 4-Aminopyridine in the absence of magnesium.Locations of discharges mapped to two anatomical regions over the somatosensory cortex and over the lateral convexity separated by a gap at a location matching the dysgranular zone. Focal epileptiform discharges were recorded in superficial and deep neocortical layers but over superficial layers, they exhibited larger surface areas. They were often independent even when closely spaced to one another but they became progressively coupled resulting in larger zones of coherent discharge.The gradual coupling of multiple, independent, closely spaced, spatially restricted, focal discharges between deep and superficial neocortical layers represents a possible mechanism of the development of an epileptogenic zone.

Intra‐areal and corticocortical circuits arising in the dysgranular zone of rat primary somatosensory cortex that processes deep somatic input

Somesthesis-guided exploration of the external world requires cortical processing of both cutaneous and proprioceptive information and their integration into motor commands to guide further haptic movement. In the past, attention has been given mostly to the cortical circuits processing cutaneous information for somatic motor integration. By comparison, little has been examined about how cortical circuits are organized for higher order proprioceptive processing. Using the rat cortex as a model, we characterized the intrinsic and corticocortical circuits arising in the major proprioceptive region of the primary somatosensory cortex (SI) that is conventionally referred to as the dysgranular zone (DSZ). We made small injections of biotinylated dextran amine (BDA) as an anterograde tracer in various parts of the DSZ, revealing three distinct principles of its cortical circuit organization. First, its intrinsic circuits extend mainly along the major axis of DSZ to organize multiple patches of interconnections. Second, the central and peripheral regions of DSZ produce differential patterns of intra-areal and corticocortical circuits. Third, the projection fields of DSZ encompass only selective regions of the second somatic (SII), posterior parietal (PPC), and primary motor (MI) cortices. These projection fields are at least partially separated from those of SI cutaneous areas. We hypothesize, based on these observations, that the cortical circuits of DSZ facilitate a modular integration of proprioceptive information along its major axis and disseminate this information to only selective parts of higher order somatic and MI cortices in parallel with cutaneous information.

Transcallosal Pathway of Whisker Information Between Rat Primary Somatosensory Cortices*

It has been suggested that rodent primary somatosensory cortex(SⅠ) can be activated by ipsilateral whisker stimulation,while SⅠ only receives the ascending input from contralateral whiskers.Previous anatomical research revealed two transcallosal pathways transferring whisker information between bilateral cortices: perigranular zone(PGZ) pathway and dysgranular zone(DZ) pathway.But which pathway plays more important role in transferring whisker information remains unknown.We used voltage-sensitive dye(VSD) imaging to visualize the SⅠ activation by stimulating whiskers.We found that the contralateral whisker stimulation first activates the barrel(granular zone,GZ),then the activity forms a propagating wave and spread to the DZ outside the sub-barrel field cortex(BFC).In contrast,ipsilateral whisker stimulation first activates the DZ outside the BFC.The evoked activity in the DZ forms a propagating wave and spreads into the BFC in SⅠ.Inactivating opposite cortex blocks this ipsilateral whisker activation.Electrical stimulus to opposite cortex also first evokes DZ in the imaging cortex.Our results suggested that whisker stimulation first activated the barrel and then DZ in the opposite SⅠ.After a delay of transcallosal transfer,the DZ of the other hemisphere(ipsilateral to the whisker being stimulated) was activated.The callosal fibers connecting the DZs on the two hemispheries play a major role in the SⅠ activation by the ipsilateral whisker stimulation.

Descending projections from the dysgranular zone of rat primary somatosensory cortex processing deep somatic input

In the mammalian somatic system, peripheral inputs from cutaneous and deep receptors ascend via different subcortical channels and terminate in largely separate regions of the primary somatosensory cortex (SI). How these inputs are processed in SI and then projected back to the subcortical relay centers is critical for understanding how SI may regulate somatic information processing in the subcortex. Although it is now relatively well understood how SI cutaneous areas project to the subcortical structures, little is known about the descending projections from SI areas processing deep somatic input. We examined this issue by using the rodent somatic system as a model. In rat SI, deep somatic input is processed mainly in the dysgranular zone (DSZ) enclosed by the cutaneous barrel subfields. By using biotinylated dextran amine (BDA) as anterograde tracer, we characterized the topography of corticostriatal and corticofugal projections arising in the DSZ. The DSZ projections terminate mainly in the lateral subregions of the striatum that are also known as the target of certain SI cutaneous areas. This suggests that SI processing of deep and cutaneous information may be integrated, to a certain degree, in this striatal region. By contrast, at both thalamic and prethalamic levels as far as the spinal cord, descending projections from DSZ terminate in areas largely distinguishable from those that receive input from SI cutaneous areas. These subcortical targets of DSZ include not only the sensory but also motor-related structures, suggesting that SI processing of deep input may engage in regulating somatic and motor information flow between the cortex and periphery.

The Functional Organization and Cortical Connections of Motor Cortex in Squirrels

Despite extraordinary diversity in the rodent order, studies of motor cortex have been limited to only 2 species, rats and mice. Here, we examine the topographic organization of motor cortex in the Eastern gray squirrel (Sciurus carolinensis) and cortical connections of motor cortex in the California ground squirrel (Spermophilus beecheyi). We distinguish a primary motor area, M1, based on intracortical microstimulation (ICMS), myeloarchitecture, and patterns of connectivity. A sensorimotor area between M1 and the primary somatosensory area, S1, was also distinguished based on connections, functional organization, and myeloarchitecture. We term this field 3a based on similarities with area 3a in nonrodent mammals. Movements are evoked with ICMS in both M1 and 3a in a roughly somatotopic pattern. Connections of 3a and M1 are distinct and suggest the presence of a third far rostral field, termed "F," possibly involved in motor processing based on its connections. We hypothesize that 3a is homologous to the dysgranular zone (DZ) in S1 of rats and mice. Our results demonstrate that squirrels have both similar and unique features of M1 organization compared with those described in rats and mice, and that changes in 3a/DZ borders appear to have occurred in both lineages.

Functional connectivity of the insula in the resting brain

The human insula is hidden in the depth of the cerebral hemisphere by the overlying frontal and temporal opercula, and consists of three cytoarchitectonically distinct regions: the anterior agranular area, posterior granular area, and the transitional dysgranular zone; each has distinct histochemical staining patterns and specific connectivity. Even though there are several studies reporting the functional connectivity of the insula with the cingulated cortex, its relationships with other brain areas remain elusive in humans. Therefore, we decided to use resting state functional connectivity to elucidate in details its connectivity, in terms of cortical and subcortical areas, and also of lateralization. We investigated correlations in BOLD fluctuations between specific regions of interest of the insula and other brain areas of right-handed healthy volunteers, on both sides of the brain. Our findings document two major complementary networks involving the ventral-anterior and dorsal-posterior insula: one network links the anterior insula to the middle and inferior temporal cortex and anterior cingulate cortex, and is primarily related to limbic regions which play a role in emotional aspects; the second links the middle-posterior insula to premotor, sensorimotor, supplementary motor and middle-posterior cingulate cortices, indicating a role for the insula in sensorimotor integration. The clear bipartition of the insula was confirmed by negative correlation analysis. Correlation maps are partially lateralized: the salience network, related to the ventral anterior insula, displays stronger connections with the anterior cingulate cortex on the right side, and with the frontal cortex on the left side; the posterior network has stronger connections with the superior temporal cortex and the occipital cortex on the right side. These results are in agreement with connectivity studies in primates, and support the use of resting state functional analysis to investigate connectivity in the living human brain.

Spatiotemporal dynamics of excitation in rat insular cortex: intrinsic corticocortical circuit regulates caudal-rostro excitatory propagation from the insular to frontal cortex

The insular cortex (IC), composing unique anatomical connections, receives multi-modal sensory inputs including visceral, gustatory and somatosensory information from sensory thalamic nuclei. Axonal projections from the limbic structures, which have a profound influence on induction of epileptic activity, also converge onto the IC. However, functional connectivity underlying the physiological and pathological roles characteristic to the IC still remains unclear. The present study sought to elucidate the spatiotemporal dynamics of excitatory propagation and their cellular mechanisms in the IC using optical recording in urethane-anesthetized rats. Repetitive electrical stimulations of the IC at 50 Hz demonstrated characteristic patterns of excitatory propagation depending on the stimulation sites. Stimulation of the granular zone of the IC (GI) and other surrounding cortices such as the motor/primary sensory/secondary sensory cortices evoked round-shaped excitatory propagations, which often extended over the borders of adjacent areas, whereas excitation of the agranular and dysgranular zones in the IC (AI and DI, respectively) spread along the rostrocaudal axis parallel to the rhinal fissure. Stimulation of AI/DI often evoked excitation in the dorsolateral orbital cortex, which exhibited spatially discontinuous topography of excitatory propagation in the IC. Pharmacological manipulations using 6,7-dinitroquinoxaline-2,3(1H,4H)-dione (DNQX), a non-NMDA receptor antagonist, D-2-amino-5-phosphonovaleric acid (D-APV), an NMDA receptor antagonist, and bicuculline methiodide, a GABAA receptor antagonist, indicate that excitatory propagation was primarily regulated by non-NMDA and GABAA receptors. Microinjection of lidocaine or incision of the supragranular layers of the rostrocaudally middle part of excitatory regions suppressed excitation in the remote regions from the stimulation site, suggesting that the excitatory propagation in the IC is largely mediated by cortical local circuits. These features of excitatory propagation in the AI/DI, that is the propagation along the rostrocaudal axis with less propagation in the ventro-dorsal direction, may play an important role for transmitting neural excitation arising from the limbic structures to the frontal and orbital cortices.

Analysis of area-specific expression patterns of RORbeta, ER81 and Nurr1 mRNAs in rat neocortex by double in situ hybridization and cortical box method.

BackgroundThe mammalian neocortex is subdivided into many areas, each of which exhibits distinctive lamina architecture. To investigate such area differences in detail, we chose three genes for comparative analyses, namely, RORbeta, ER81 and Nurr1, mRNAs of which have been reported to be mainly expressed in layers 4, 5 and 6, respectively. To analyze their qualitative and quantitative coexpression profiles in the rat neocortex, we used double in situ hybridization (ISH) histochemistry and cortical box method which we previously developed to integrate the data of different staining and individuals in a standard three-dimensional space.Principal FindingsOur new approach resulted in three main observations. First, the three genes showed unique area distribution patterns that are mostly complementary to one another. The patterns revealed by cortical box method matched well with the cytoarchitectonic areas defined by Nissl staining. Second, at single cell level, RORbeta and ER81 mRNAs were coexpressed in a subpopulation of layer 5 neurons, whereas Nurr1 and ER81 mRNAs were not colocalized. Third, principal component analysis showed that the order of hierarchical processing in the cortex correlates well with the expression profiles of these three genes. Based on this analysis, the dysgranular zone (DZ) in the somatosensory area was considered to exhibit a profile of a higher order area, which is consistent with previous proposal.Conclusions/SignificanceThe tight relationship between the expression of the three layer specific genes and functional areas were revealed, demonstrating the usefulness of cortical box method in the study on the cerebral cortex. In particular, it allowed us to perform statistical evaluation and pattern matching, which would become important in interpreting the ever-increasing data of gene expression in the cortex.

Comparing the Functional Representations of Central and Border Whiskers in Rat Primary Somatosensory Cortex

The anatomical representations of the large facial whiskers, termed barrels, are topographically organized and highly segregated in the posteromedial barrel subfield (PMBSF) of rat layer IV primary somatosensory cortex. Although the functional representations of single whiskers are aligned with their appropriate barrels, their areal extents are rather large, spreading outward from the appropriate barrel along the tangential plane and thereby spanning multiple neighboring and non-neighboring barrels and septal regions. To date, single-whisker functional representations have been characterized primarily for whiskers whose corresponding barrels are located centrally within the PMBSF (central whiskers). Using intrinsic signal imaging verified with post-imaging single-unit recording, we demonstrate that border whiskers, whose barrels are located at the borders of the PMBSF, also evoke large activity areas that are similar in size to those of central whiskers but spread beyond the PMBSF and sometimes beyond primary somatosensory cortex into the neighboring dysgranular zones. This study indicates that the large functional representation of a single whisker is a basic functional feature of the rat whisker-to-barrel system and, combined with results from other studies, suggest that a large functional representation of a small, point-like area on the sensory epithelium may be a functional feature of primary sensory cortex in general.

Parallel Streams for the Relay of Vibrissal Information through Thalamic Barreloids

This study investigated the organization of a vibrissal pathway that arises from the interpolar division of the spinal trigeminal complex (SP5i), transits through the ventral posterior medial nucleus (VPM), and innervates the somatosensory cortical areas in the rat. Using Fluoro-Gold and biotinylated dextran amine, respectively, as retrograde and anterograde tracers, the following organization plan was disclosed. The SP5i projection arises from a population of small-sized neurons that selectively innervate the ventral lateral part of VPM. In cytochrome oxidase-stained material, this region does not display any barreloid arrangement, but Fluoro-Gold injections in single barrel columns labeled rods of cells that extend caudally into the ventral lateral division of VPM. Thus, on the basis of retrograde labeling, barreloids were divided into core and tail compartments, which correspond to the rod segments running across the dorsal and ventral lateral parts of VPM, respectively. Double-labeling experiments revealed that SP5i afferents innervate the tail of barreloids. The anterograde labeling of thalamocortical axons show that most "core cells" project to a single barrel column, whereas some "tail cells" give rise to branching axons that innervate the second somatosensory area and the dysgranular zone of the barrel field. Injections that straddled the transition zone between the core and tail regions disclosed cells projecting to a single barrel column and to the surrounding dysgranular zone. These results suggest that the projection of "barreloids cells" to the granular and/or dysgranular zones relates to the class of prethalamic input(s) they receive.

Regional Differences of Callosal Connections in the Granular Zones of the Primary Somatosensory Cortex in Rats

The primary somatosensory cortex (SI) in rats is cytoarchitectonically divided into three zones: the granular, peri-, and dysgranular zones. To examine callosal connections in the granular zone that bears representation of the body somatosensory map, the distribution of lectin-conjugated horseradish peroxidase labels was explored in the right SI after single or multiple injections into the granular zone of the left SI. After injections in the upper and lower jaw regions, many labeled cell bodies and dense terminal labeling were found in the regions homotopical to the injection sites. Both kinds of labels were densely seen in layers II–III and V, less densely in layer VI. Densely labeled terminals were also observed in layer I. In layer IV, many terminals and a few cell bodies were labeled in the septa, while labeling inside the barrels was sparse or absent. After injections in other regions, i.e., those representing the facial whiskers, fore- and hindlimbs, or trunk in the granular zone, labeled callosal cell bodies and terminals were sparse or absent, except in the septa of the posteromedial barrel subfield representing the facial whiskers. The results clearly show that the density of callosal connections in the granular zone differs in different subfields, and that at least the jaw regions in the granular zones of both hemispheres are directly interconnected, in contrast to the previous assumption that only the dysgranular zone mediates information transmission between the granular zones of both sides.

Cells of origin, thalamic relay and termination of the external cuneothalamocortical tract in the gerbil

The present study is concerned with the connections of the external cuneate nucleus (ECN) in the gerbil following an injection of horseradish peroxidase (HRP) into the ventralis posterior pars oralis (VPLo) or adjacent nuclei of the thalamus. The number, soma size and distribution of the retrograde-labelled ECN neurons were studied and quantified. The application of two retrograde fluorescent tracers was also used to determine whether the ECN neurons would project to the thalamus as well as to the cerebellum through their collaterals. The HRP-positive ECN neurons projecting to the thalamic VPLo were confined to the contralateral caudal half of the ECN, primarily within the intermediate portion represent the forearm and arm territories with a small part of the thoracic and shoulder areas. Labelled neurons were classified into small and medium-sized cells. The majority (96%) of the external cuneothalamic neurons were of the small variety. No double-labelled cells were detected in the ECN following injections of Rhodamine-labelled latex microspheres and Fast blue into the cerebellum and thalamus respectively, suggesting that the ECN neurons projecting to the thalamus form a separate cell group different from those projecting to the cerebellum. The injected HRP into the VPLo was also transported in an anterograde direction by the thalamocortical fibers. The HRP-labelled axonal terminals were distributed within motor area 4 and the dysgranular zones (DZs) of the primary somatosensory cortex (SmI), reaching the deep layers IV and VI as well as superficial layer I. The external cuneothalamocortical pathway shown in the present study may be related to the proprioceptive feedback control of the coordinating motor activity, especially during forelimb muscle movement.

The anatomic organization of evoked potentials in rat parietal cortex: electrically evoked commissural responses

1. Two 8 x 8 channel microelectrode arrays were positioned over 3.5 x 3.5 mm2 areas in homologous regions of right and left parietal cortex of four rats. Potentials were evoked by delivering epicortical electrical stimulation to each electrode on one hemisphere while mapping the commissural response from the contralateral array. Spatial distributions of the electrically evoked potential (EECP) complex were compared directly with cytochrome oxidase-stained sections of the recorded region. 2. Electrode sites most capable of eliciting a commissural EECP were arranged along a diagonal band extending medially from the rostral to caudal region of each electrode array, approximating the pattern of dysgranular cortex separating primary auditory (Te1) from primary somatosensory (Par1) cortex. Electrode sites in the rostromedial and caudolateral region were ineffectual in eliciting an EECP in either hemisphere. Stimulation sites within secondary visual cortex (Oc2L) also produced strong responses. Only weak responses were elicited from stimulation of Te1 and no EECP could be evoked when stimulating within Par1. 3. When an EECP in the maximally sensitive diagonal region was elicited, its spatial distribution was typically asymmetrical throughout the recording array; the response was largest along a diagonal region also extending medially from the rostral to caudal area of each electrode array. Thus the pattern of EECP in each hemisphere closely matched the pattern of electrically excitable regions in the contralateral hemisphere. 4. The EECP was usually heterogeneous. EECP distributions within the strongly responding diagonal area often formed two regions of maximum amplitude separated by a less active zone. Although responses in Te1 were significantly weaker than those in the adjacent dysgranular cortex, they also revealed a heterogeneous spatial distribution with multiple closely spaced maxima. Only responses in Oc2L appeared consistently homogeneous, with a single maximum representing the EECP. 5. These results provide functional evidence supporting a model of parietal cortex in which there are two basic types of recipient regions, densely granular regions, which are the termination sites of specific thalamocortical fibers, and dysgranular or agranular regions, which receive both ipsilateral and contralateral projections. The functional parceling of rodent parietal cortex on the basis of the spatial and temporal distribution of the epicortical evoked potential complex may be superimposed onto the anatomic parceling into granular and dysgranular zones. Implications for stages of sensory information processing are discussed.