Rat Somatic Sensory Cortex Research Articles

Synaptic localization of GLT‐1a in the rat somatic sensory cortex

GLT-1a, the major glutamate transporter, plays an important role in both physiological and pathological conditions. Uncertainty regarding its localization in the cerebral cortex prompted us to re-examine its cellular and subcellular localization in the rat somatic sensory cortex. GLT-1a detection was sensitive to fixation; in optimal conditions approximately 25% of GLT-1a+ profiles were axon terminals. GLT-1a/VGLUT1 double-labeling and pre-embedding electron microscopy studies showed that approximately 50% of GLT-1a+ profiles were in the vicinity of asymmetric synapses. Using pre-embedding electron microscopy, we found that approximately 70% of GLT-1a located in the vicinity of asymmetric synapses was astrocytic and approximately 30% was neuronal. Post-embedding immunogold studies showed that the density of gold particles coding for GLT-1a was much higher in astrocytic processes than in axon terminals, and that in the latter they were never at the active zone. In both astrocytic processes and axon terminals most gold particles were localized in a membrane region extending for about 250 nm from active zone margin, with a peak at 140 nm for astrocytic processes and at 80 for axon terminals. We conclude that, although GLT-1a is expressed by both astrocytes and axon terminals, astrocytic GLT-1a predominates at asymmetric synapses, and that the perisynaptic localization of GLT-1a in cortex is well-suited to modulate Glu concentrations at the cleft and also to restrict Glu spillover.

Postnatal development of the vesicular gaba transporter in rat cerebral cortex

Light and electron microscopic immunocytochemical techniques and Western blotting were used to investigate the postnatal development of the vesicular GABA transporter (VGAT) in the rat somatic sensory cortex. VGAT immunoreactivity was low at birth, it increased gradually through the first and second weeks of life and achieved the adult pattern during the third week. At postnatal day (P)0–P5, VGAT immunoreactivity was associated exclusively to fibers and puncta. Electron microscopic studies performed at P5 showed that all identified synaptic contacts formed by VGAT-positive axonal swellings were of the symmetric type and that a substantial proportion of the boutons appeared not to have formed synapses. From P10 onward, labeled puncta were both scattered in the neuropil and in apposition to unstained cellular profiles; VGAT was also expressed in few GABAergic cell bodies. Western blottings at the same postnatal ages revealed a 55-kDa band whose intensity was weak at P0 (17% of adult), it increased constantly until P15 (P2: 35%; P5: 44%; P10: 68%; P15: 97%), and then leveled off. Overall, the present results show that during neocortical development the expression of VGAT slightly precedes the complete maturation of inhibitory synaptogenesis and suggest that it may contribute to the formation of neocortical GABAergic circuitry.

Substance P and nitric oxide signaling in cerebral cortex: anatomical evidence for reciprocal signaling between two classes of interneurons.

Parvalbumin-containing fast-spiking interneurons in the cerebral cortex exhibit widespread electrical coupling, as do somatostatin-containing low-threshold spiking interneurons. Besides the classical neurotransmitter gamma-aminobutyric acid, these cortical interneurons may also release various neuropeptides including substance P (SP), as well as the freely diffusible messenger nitric oxide (NO). To investigate whether these two networks of interneurons might interact via these nonclassical messengers, we performed immunocytochemistry for SP and NO signaling pathways in rat somatic sensory cortex. SP was found in a subset of parvalbumin-positive cells concentrated in layers IV and V, whereas its receptor, NK1, was found in a subset of somatostatin-containing neurons (and also, at much lower levels, in a disjoint subset of parvalbumin-containing neurons). Only 4% of SP-containing axon terminals were apposed to NK1-positive dendrites, suggesting that in the cerebral cortex, SP may act predominantly as a paracrine neuromediator. Nitric oxide synthase-I (NOS-I), the synthetic enzyme for NO, was found almost exclusively in NK1-positive neurons; 95% of intensely somatostatin/NK1-positive neurons were also positive for NOS-I, and 94% of NOS-positive neurons were also positive for NK1. Immunoreactivity for soluble guanylyl cyclase (the NO receptor) was at high levels in the apical dendrites of layer V pyramidal neurons and in parvalbumin/SP-positive neurons. These data point to a novel reciprocal chemical interaction between two inhibitory networks in the rat neocortex.

Direct inhibition evoked by whisker stimulation in somatic sensory (SI) barrel field cortex of the awake rat.

Whisker deflection typically evokes a transient volley of action potentials in rat somatic sensory (SI) barrel cortex. Postexcitatory inhibition is thought to quickly terminate the cortical cell response to whisker deflection. Using dual electrode extracellular recording in awake rats, we describe an infrequent type of cell response in which stimulation of single hairs consistently blocks the ongoing discharge of neurons without prior excitation (I-only inhibition). Reconstruction of the recording sites indicates that I-only inhibition occurs most frequently when the recording site is clearly in the septum or at the barrel-septum junction. The same cells that respond with I-only inhibition to one whisker can show an excitatory discharge to other whiskers, usually followed by inhibition. Stimulation of either nose hairs or the large mystacial vibrissa can evoke I-only inhibition in SI cortex. I-only inhibition is most commonly observed at low stimulus frequencies ( approximately 1 Hz). At stimulus frequencies of >6 Hz, I-only inhibition typically converts to excitation. We conclude that single whisker low-frequency stimulation can selectively block the spontaneous discharge of neurons in SI barrel field septa. The observation that this cell response is found most often in or at the edge of septa and at relatively long latencies supports the idea that I-only inhibition is mediated through cortical circuits. We propose that in these cells inhibition alone or a combination of inhibition and disfacilitation play a role in suppressing neuronal discharge occasioned by low frequency contact of the whiskers with the environment.

Two systems of giant axon terminals in the cat medial geniculate body: Convergence of cortical and GABAergic inputs

The thalamus plays a critical role in processing sensory information that involves interactions between extrinsic connections and intrinsic circuitry. Little is known regarding how these different systems might interact. We found an unexpected nuclear convergence of two types of giant axon terminals, each of which must have independent origins, in the dorsal division of the cat medial geniculate body. The first class of giant terminal was labeled after injections of biotinylated dextran amines (BDA) in seven auditory cortical areas. A second type was found in sections immunostained for gamma-aminobutyric acid (GABA); these endings had the same nuclear distribution, and they were numerous. The origin of this GABAergic terminal is unknown. The giant corticothalamic terminals were presumably those described in prior accounts using different tracers (Rouiller and de Ribaupierre [1990] Neurosci. Lett. 208:29-35; Ojima [1994] Cerebral Cortex 6:646-663), but with BDA they are labeled more fully. Clusters of such endings were often linked, and hundreds may occur in a single section. Their boutons formed a substantial proportion of the corticothalamic population. Other types of corticogeniculate axon terminals were also labeled, including two kinds that are much smaller and that match closely the classical descriptions of corticothalamic axons. The giant GABAergic endings were found in all dorsal division nuclei and in thalamic visual nuclei such as the lateral posterior nucleus. Like the giant cortical endings, the giant GABAergic terminals often encircled large, pale, immunonegative profiles that may be dendritic. This implies a close spatial, and perhaps a close functional, relationship between the populations of giant axon terminals. Insofar as physiological studies found that pharmacological inactivation of rat somatic sensory cortex suppresses peripheral information transmission through the posterior thalamus, corticofugal input may be essential for normal processing (Diamond et al. [1992] J. Comp. Neurol. 319:66-84). Our findings suggest that the giant corticothalamic endings could play an important role in descending control. Perhaps they are counterbalanced by a GABAergic system and affect thalamic oscillations implicated in shifts in vigilance and attention.

Two systems of giant axon terminals in the cat medial geniculate body: Convergence of cortical and GABAergic inputs

The thalamus plays a critical role in processing sensory information that involves interactions between extrinsic connections and intrinsic circuitry. Little is known regarding how these different systems might interact. We found an unexpected nuclear convergence of two types of giant axon terminals, each of which must have independent origins, in the dorsal division of the cat medial geniculate body. The first class of giant terminal was labeled after injections of biotinylated dextran amines (BDA) in seven auditory cortical areas. A second type was found in sections immunostained for γ-aminobutyric acid (GABA); these endings had the same nuclear distribution, and they were numerous. The origin of this GABAergic terminal is unknown. The giant corticothalamic terminals were presumably those described in prior accounts using different tracers (Rouiller and de Ribaupierre [1990] Neurosci. Lett. 208:29–35; Ojima [1994] Cerebral Cortex 6:646–663), but with BDA they are labeled more fully. Clusters of such endings were often linked, and hundreds may occur in a single section. Their boutons formed a substantial proportion of the corticothalamic population. Other types of corticogeniculate axon terminals were also labeled, including two kinds that are much smaller and that match closely the classical descriptions of corticothalamic axons. The giant GABAergic endings were found in all dorsal division nuclei and in thalamic visual nuclei such as the lateral posterior nucleus. Like the giant cortical endings, the giant GABAergic terminals often encircled large, pale, immunonegative profiles that may be dendritic. This implies a close spatial, and perhaps a close functional, relationship between the populations of giant axon terminals. Insofar as physiological studies found that pharmacological inactivation of rat somatic sensory cortex suppresses peripheral information transmission through the posterior thalamus, corticofugal input may be essential for normal processing (Diamond et al. [1992] J. Comp. Neurol. 319:66–84). Our findings suggest that the giant corticothalamic endings could play an important role in descending control. Perhaps they are counterbalanced by a GABAergic system and affect thalamic oscillations implicated in shifts in vigilance and attention. J. Comp. Neurol. 413:181–197, 1999. © 1999 Wiley-Liss, Inc.

Expression of NR2 receptor subunit in rat somatic sensory cortex: Synaptic distribution and colocalization with NR1 and PSD‐95

Expression of NR2 receptor subunit in rat somatic sensory cortex: Synaptic distribution and colocalization with NR1 and PSD‐95

Expression of NR2 receptor subunit in rat somatic sensory cortex: Synaptic distribution and colocalization with NR1 and PSD-95

Functional N-methyl-D-aspartate (NMDA) receptors comprise heteromeric combinations of NR1 and NR2 subunits. In the present study, we employed light and electron microscopic immunocytochemistry to study the expression of NR2A and NR2B (NR2A/B) protein in somatic sensory cortex of adult rats. To relate this distribution to that of NR1 and to the NMDA receptor anchoring protein PSD-95, we documented extensive cellular colocalization of NR2A/B with NR1 at the light microscopic level. In contrast, PSD-95 exhibited little somatic staining, being restricted mainly to dendrites and neuropil. We employed postembedding immunocytochemistry to study the ultrastructural expression of NR2A/B. Labeling in neuronal perikarya was associated with rough endoplasmic reticulum and Golgi apparatus; in dendrites, gold particles labeled microtubules. The preponderance of labeling was associated with asymmetric synapses. Double immunolabeling revealed that NR2 colocalized in many synapses with NR1 and with PSD-95. Quantitative measurements revealed that density of gold particles coding for both NR2 and PSD-95 was highest just inside the postsynaptic membrane. Tangentially along the membrane, gold particles were concentrated at the synaptic specialization. These data provide structural evidence in neocortex for heteromeric NMDA receptors anchored at the postsynaptic membrane.

Role of the basal forebrain cholinergic projection in somatosensory cortical plasticity.

Trimming all but two whiskers in adult rats produces a predictable change in cortical cell-evoked responses characterized by increased responsiveness to the two intact whiskers and decreased responsiveness to the trimmed whiskers. This type of synaptic plasticity in rat somatic sensory cortex, called "whisker pairing plasticity," first appears in cells above and below the layer IV barrels. These are also the cortical layers that receive the densest cholinergic inputs from the nucleus basalis. The present study assesses whether the cholinergic inputs to cortex have a role in regulating whisker pairing plasticity. To do this, cholinergic basal forebrain fibers were eliminated using an immunotoxin specific for these fibers. A monoclonal antibody to the low-affinity nerve growth factor receptor 192 IgG, conjugated to the cytotoxin saporin, was injected into cortex to eliminate cholinergic fibers in the barrel field. The immunotoxin reduces acetylcholine esterase (AChE)-positive fibers in S1 cortex by >90% by 3 wk after injection. Sham-depleted animals in which either saporin alone or saporin unconjugated to 192 IgG is injected into the cortex produces no decrease in AChE-positive fibers in cortex. Sham-depleted animals show the expected plasticity in barrel column neurons. In contrast, no plasticity develops in the ACh-depleted, 7-day whisker-paired animals. These results support the conclusion that the basal forebrain cholinergic projection to cortex is an important facilitator of synaptic plasticity in mature cortex.

GluR1-immunopositive interneurons in rat neocortex.

Recent in vitro studies suggest that inhibitory interneurons in cortex may express the GluR1 glutamate receptor subunit in the absence of GluR2, leading to calcium-permeable alpha-amino-3-hydroxy-5-methylisoxazole-4-propionate (AMPA) channels. We performed a study of rat somatic sensory cortex to confirm and extend these observations, using quantitative immunocytochemistry for multiple antigens. A morphologically distinct subpopulation of nonpyramidal neurons in neocortex was intensely immunoreactive for GluR1. Electron microscopic analysis of these cells revealed somatic staining for GluR1, mainly in the rough endoplasmic reticulum. Dendritic staining was concentrated at the synaptic active zone and in the adjacent subsynaptic cytoplasm. Double immunostaining revealed that the large majority of intensely GluR1-positive cells contained gamma-aminobutyric acid or its synthetic enzyme, glutamic acid decarboxylase, but little or no GluR2. Thus, AMPA receptors on a subpopulation of inhibitory interneurons in cortex are likely to be calcium permeable. This calcium permeability is likely to influence functional properties of these neurons; it may underlie the high levels of calcium-binding proteins they contain; and may render them liable to excitotoxic injury

Glutamate decarboxylase immunoreactivity in corticocortical projecting neurons of rat somatic sensory cortex.

Combined retrograde tracing and immunocytochemical experiments were carried out on rats to ascertain whether corticocortical projecting neurons in the somatic sensory areas are immunoreactive to an antiserum against glutamate decarboxylase. Injections of a retrograde tracer (colloidal gold-labelled wheat germ agglutinin conjugated to enzymatically inactive horseradish peroxidase) in the first somatic sensory area labelled neurons in the injected area, in the second somatic sensory area, and in the parietoventral area of the ipsilateral hemisphere. The topographical and laminar distribution of these retrogradely-labelled corticocortical neurons in the first and second somatic sensory areas and in the parietoventral area was in line with a previous description (Fabri M. and Burton H. (1991b) J. comp. Neurol. 311, 405-424). In sections processed for the simultaneous visualization of the retrograde tracer and glutamate decarboxylase immunoreactivity, a number of neurons were double-labelled. Double-labelled neurons were most numerous in the first somatic sensory cortex, where they accounted for 5% of all retrogradely-labelled neurons. Outside this region, double-labelled cells were observed in the second somatic sensory cortex and in the parietoventral cortex, where they amounted respectively to 2.8% and 2.3% of all corticocortical neurons labelled in these two areas. Glutamate decarboxylase-immunopositive corticocortical neurons were mainly concentrated in the infragranular layers (73.8% of all double-labelled neurons in the first somatic sensory area, 81.7% in the second somatic sensory area, and 76.5% in the parietoventral area). The results indicate the presence of a small but significant contingent of GABAergic inhibitory neurons in the associative connections of the somatic sensory areas.

Dynamic synaptic modification threshold: computational model of experience-dependent plasticity in adult rat barrel cortex.

Previous electrophysiological experiments have documented the response of neurons in the adult rat somatic sensory ("barrel") cortex to whisker movement after normal experience and after periods of experience with all but two whiskers trimmed close to the face (whisker "pairing"). To better understand how the barrel cortex adapts to changes in the flow of sensory activity, we have developed a computational model of a single representative barrel cell based on the Bienenstock, Cooper, and Munro (BCM) theory of synaptic plasticity. The hallmark of the BCM theory is the dynamic synaptic modification threshold, theta M, which dictates whether a neuron's activity at any given instant will lead to strengthening or weakening of the synapses impinging on it. The threshold theta M is proportional to the neuron's activity averaged over some recent past. Whisker pairing was simulated by setting input activities of the cell to the noise level, except for two inputs that represented untrimmed whiskers. Initially low levels of cell activity, resulting from whisker trimming, led to low values for theta M. As certain synaptic weights potentiated, due to the activity of the paired inputs, the values of theta M increased and after some time their mean reached an asymptotic value. This saturation of theta M led to the depression of some inputs that were originally potentiated. The changes in cell response generated by the model replicated those observed in in vivo experiments. Previously, the BCM theory has explained salient features of developmental experience-dependent plasticity in kitten visual cortex. Our results suggest that the idea of a dynamic synaptic modification threshold, theta M, is general enough to explain plasticity in different species, in different sensory systems, and at different stages of brain maturity.

Glutamate in terminals of thalamocortical fibers in rat somatic sensory cortex

Thalamocortical terminals in layer IV of the somatic sensory cortex (SI) were identified by anterograde transport from ventrobasal thalamus. They were large, contained loosely packed round clear vesicles and abundant mitochondria, and made asymmetric contacts mainly with dendritic spines. Post-embedding immunocytochemistry revealed these terminals to be enriched in glutamate. Levels of glutamate in thalamocortical terminals, and in dendritic spines postsynaptic to them, were significantly higher than in nearby dendrites, astroglia, or in GABAergic terminals (identified by double immunostaining). These results support glutamate as neurotransmitter in thalamocortical fibers, and suggest that dendritic spines may take up glutamate released by these terminals.

Growth of the rat somatic sensory cortex and its constituent parts during postnatal development

We have compared the size and arrangement of the primary somatic sensory cortex (SI) and its constituent parts in juvenile (1 week old) and mature (10-12 weeks old) rats using succinic dehydrogenase histochemistry and digital image analysis. Our goal was to determine whether some regions of the maturing cortex grow more than others. To this end, we examined (1) the growth of barrels and the surrounding (interbarrel) cortex, (2) the growth of the major somatic representations within SI, and (3) the overall growth of SI compared to the neocortex as a whole. With respect to the first of these issues, SI barrels and barrel-like structures grow more than the intervening cortex. The growth of these elements varies according to region: barrels in the head representation more than double in size, whereas the barrel-like structures in the paw representations increase by only about half this amount. The growth of the major somatic representations within SI is also heterogeneous, the representation of the head enlarging to a greater extent than the representations of the paws. Thus, the ratio of the total area of head representation to the combined paw representation is 15% greater in adults than in juveniles. Finally, the primary somatic sensory cortex grows to a somewhat greater extent than the neocortex as a whole. These observations demonstrate that postnatal cortical growth is not uniform; it varies among cortical barrels and the immediately surrounding (interbarrel) cortex, among the representations of different body parts, and between SI and the rest of the neocortex. As an explanation of this differential growth, we suggest that the neuropil of metabolically (and/or electrically) more active cortical regions grows to a greater extent during maturation than that of less active regions.

In situ hybridization reveals VIP precursor mRNA-containing neurons in monkey and rat neocortex

The cDNAs encoding monkey vasoactive intestinal polypeptide (VIP) and PHM-27 and rat VIP and PHI-27 were cloned and used to generate antisense RNA probes. Using in situ hybridization, neurons expressing the VIP/PHM or VIP/PHI precursor mRNAs were localized in monkey and rat somatic sensory and visual cortex. In both neocortical areas of both species, labeled cells were observed in all 6 layers as well as the subcortical white matter.

Maturation of pyramidal cell form in relation to developing afferent and efferent connections of rat somatic sensory cortex

Golgi and axonal labeling methods were used to examine the maturation of pyramidal cells in layers III and V of the rat somatic sensory cortex. The material came from animals late in the gestation period, postnatal, ranging from 0 to 43 days of age and at maturity. Special attention was paid to the period (0–7 days of age) during which it is known that thalamic and callosal fibers grow into the cortex. It is shown that the basic features of the pyramidal cell form are established before the long afferent fibers arrive in layers III and V and before the large number of synapses are established in these layers. Nevertheless, considerable dendritic growth and spine formation occurs after the afferent fibers establish an adult-like pattern of distribution. It is also shown that even at 1 day of age, the axons of pyramidal cells in all layers have reached the vicinity of targets such as the striatum, thalamus, brainstem, spinal cord and contralateral cortex. At 0–1 day the immature pyramidal cells are essentially bipolar in the upper cortical plate, but in the developing infragranular layers they have a few short, almost spine-free, basal dendrites and, rarely, a few oblique branches of the apical dendrite. The apical dendrite extends to the pial surface and the dendritic branches end in growth cones. The dendrites of cells in all layers increase in size and complexity of branching over the first postnatal week; the maturation of dendrites in layer V leads that of dendrites in the supragranular layers by about 2–3 days. As maturation proceeds, basal dendrites acquire secondary and tertiary branches and more oblique branches appear on the apical dendrite. Dendritic spines appear after 4 days of age but remain sparse up to 7–8 days. At 14 days of age, the spine density is much higher than in 7-day-old animals but remains at a much lower density than in 4-week-old, 6-week-old, or adult animals. By 7–14 days, the difference in maturity between superficial (layer III) and deep (layer V) pyramidal cells is difficult to discern qualitatively. All the pyramidal cells now have relatively complex, highly branched dendritic trees when compared to younger cells, but the dendritic tree is still immature in terms of the number, length and complexity of branching of the apical and basal dendritic systems. It can be concluded that the growth of the long axon of cortical pyramidal neurons precedes the acquisition of afferent connections and when these afferent fibers arrive in the cortex the dendritic tree of the pyramidal cell is still highly immature. Thus it remains possible that the finer modeling of the dendritic tree and the formation of spines may be affected by extrinsic influences such as the afferent fibers.

Developmental studies of thalamocortical and commissural connections in the rat somatic sensory cortex.

AbstractAutoradiographic, axonal degeneration, and horseradish peroxidase fiber tracing methods were employed to investigate the organization, development and potential plasticity of the thalamocortical projection to the somatic sensory cortex of the rat. In the adult animal, thalamocortical terminals are concentrated primarily in layers I and IV and in the upper part of layer VI. Fibers terminating in layers IV and VI arise from a different thalamic region than those terminating in layer I. Discrete clusters of fibers and terminals 250–450 μm wide are distributed only to the parts of the SI cortex containing dense aggregates of layer IV granule cells and not to the intervening, less granular and commissurally connected zones.At birth, thalamocortical fibers have invaded the deep part of the developing SI cortex and are concentrated in the upper part of layer VI. Between the age of two and three days, an additional concentration of fibers appears in the part of the cortical plate which will become layer IV. Layer IV is clearly recognizable by three days of age and the dense granule cell aggregates appear in it no more than one day later. The ingrowth of commissural fibers (Wise and Jones, '76) lags behind that of thalamic fibers. The mature commissural fiber pattern is not established until the age of seven days.After removal of the developing thalamocortical system by thalamotomy in newborn rats, subsequent investigation of the commissural system in the adult showed that no commissural fibers or terminals had invaded either laminae or zones of the cortex deprived of thalamic input. Similarly, commissurotomy at birth was not followed by sprouting of thalamic fibers into zones or laminae deprived of commissural connections. The connectional specificity observed in these neocortical fiber systems contrasts markedly with the plasticity of connections reported in allocortical systems.Removal of thalamocortical afferents before they attain their definitive distribution does not radically effect the overall development of the dense granule cell aggregates in layer IV. Within the aggregates, however, subsidiary features such as the “barrels” fail to appear. This finding suggests that certain elements of cortical architecture such as the dense granule cell aggregates are independent of thalamic afferents while others, such as the barrels, result from the interaction of the developing thalamocortical fibers and/or terminals with maturing neurons.

Cells of origin and terminal distribution of descending projections of the rat somatic sensory cortex

The retrograde, horseradish peroxidase technique has been used to demonstrate the cells of origin of corticofugal fiber systems arising in the rat somatic sensory cortex and projecting to the striatum, diencephalon, brainstem, and spinal cord. Correlative experiments conducted with the anterograde, autoradiographic method have been used to confirm the terminal distribution of many of these fiber systems. Corticofugal pathways directed to subcortical structures arise in the first and second somatic sensory areas exclusively from pyramidal cells of the infragranular layers, V and VI. Fibers which descend to the midbrain, pons, medulla and spinal cord arise exclusively from the largest pyramidal cells, the somata of which are found in the deep part of layer V (layer VB). There is some evidence for a sublaminar organization of the different classes of efferent cells within this layer. Fibers projecting to the diencephalon arise from somata situated throughout layer VI and to a lesser extent in layer V. Corticostriatal fibers arise only from cells with somata in layer V, but the somata are more superficially situated than those of the other classes of corticofugal neurons. The laminar distribution of the somata of corticofugal neurons differs considerably from that of commissural and ipsilateral corticocortical neurons.

Somatotopic and columnar organization in the corticotectal projection of the rat somatic sensory cortex

Single injections of tritiated amino acids into the first somatic sensory area (SI) of the rat neocortex result in axoplasmically transported labeling of the stratum griseum intermidiale and stratum griseum profundum of the ipsilateral superior colliculus. The terminal labeling in these layers takes the form of multiple, column-like patches. The SI projection is somatotopically organized with the face and head representations projecting to an extensive anterolateral part of the colliculus and the limb representations projecting to a restricted posterolateral part. Injections of horseradish peroxidase into the superior colliculus result in retrograde labelling of corticotectal cells in the superficial part of layer VB of SI and of the second somatic sensory area (SII).

The organization and postnatal development of the commissural projection of the rat somatic sensory cortex.

Anterograde and retrograde tracing experiments have been used to demonstrate the origin and terminal distribution of commissural fibers in the first somatosensory cortex (SI) of the rat. The commissural fibers originate from pyramidal cells of all layers, but predominantly from layers III and V. The fibers terminate in a series of approximately vertical bands. In each of these there are concentrations of terminals extending from the inner portion of the molecular layer to the deep portion of layer III as well as in the superficial part of layer V, and in layer VI. Discrete vertical bands of cortex are reciprocally connected across the midline to give both the origin and terminal regions of the projection a patchy or "columnar" appearance. The commissural fibers arise from and terminate in areas of the cortex that lie between and alongside the aggregations of granule cells that distinguish SI of the rat. No commissural fibers terminate within the aggregations of layer IV cells themselves but the more superficial terminal ramifications may come to overlie these aggregations. A heterotopic projection to the contralateral second somatosensory cortex has been observed and is similar in form to the homotopic projection to SI. Many commissural fibers have crossed the midline in the corpus callosum by the day of birth but lie in the underlying white matter and do not enter the cortical plate until at least the third postnatal day. During the first postnatal week these fibers grow somewhat diffusely into the maturing cortex and their topographic and laminar pattern of distribution attains its adult characteristics by the end of the first week. Commissural axons, thus, arise from immature cells but the maturation of cell form seems to precede the ingrowth of these axons and the acquisition of commissural synapses.