Morphological reproductive characteristics of testes and fertilization capacity of cryopreserved sperm after the Fukushima accident in raccoon (Procyon lotor).

Since the Fukushima Daiichi Nuclear Power Plant (FDNPP) accident, we have established an archive system of livestock and wild animals from the surrounding ex-evacuation zone. Wildlife within the alert zone have been exposed to low-dose-rate (LDR) radiation for a long continuous time. In this study, we analysed the morphological characteristics of the testes and in vitro fertilization (IVF) capacity of cryopreserved sperm of racoons from the ex-evacuation zone of the FDNPP accident. The radioactivity of caesium-137 (137 Cs) was measured by gamma-ray spectrometry, and the measured radioactivity concentration was 300-6,630 Bq/kg in the Fukushima raccoons. Notably, normal spermatogenesis was observed in the seminiferous tubules of the testes, with the germinal epithelium composed of a spermatogenic cell lineage with no evident ultrastructural alterations; freeze-thawing sperm penetration ability was confirmed using the interspecific zona pellucida-free mouse oocytes IVF assays. This study revealed that the chronic and LDR radiation exposure associated with the FDNPP accident had no adverse effect on the reproductive characteristics and functions of male raccoons.

Elliptical Fourier descriptors of outline and morphological analysis in caudal view of foramen magnum of the tropical raccoon (Procyon cancrivorus) (Linnaeus, 1758).

OBJECTIVES: To evaluate sexual-size dimorphism and attempt at categorization of inter-individual shapes of foramen magnum outlines using Fourier descriptors which allow for shape outline evaluations with a resultant specimen character definition. MATERIALS AND METHODS: Individual characterization and quantification of foramen magnum shapes in direct caudal view based on elliptical Fourier technique was applied to 46 tropical raccoon skulls (26 females, 20 males). RESULTS: Incremental number of harmonics demonstrates morphological contributions of such descriptors with their relations to specific anatomical constructions established. The initial harmonics (1st to 3rd) described the general foramen shapes while the second (4th to 12th) demonstrated fine morphological details. Sexual-size dimorphism was observed in females (87.1%) and 91.7% in males, normalization of size produces 75% in females and 83% in males. With respect to foramen magnum dimorphism analysis, the result obtained through elliptic Fourier analysis was comparatively better in detail information of outline contours than earlier classical methods. The first four effective principal components defined 70.63% of its shape properties while the rest (22.51%) constituted fine details of morphology. CONCLUSION: Both size and shape seems important in sexual dimorphisms in this species, this investigation suggest clinical implications, taxonomic and anthropologic perspectives in foramen characterization magnum characterization and further postulates an increased possibility of volume reduction cerebellar protrusion, ontogenic magnum shape irregularities in the sample population with neurologic consequences especially among females.

Light and scanning electron microscopic study on the tongue and lingual papillae of the common raccoon, Procyon lotor.

We observed the external surface and connective tissue cores (CTCs), after exfoliation of the epithelium of the lingual papillae (filiform, fungiform, foliate and vallate papillae) of the common raccoon (Procyon lotor) using scanning electron microscopy and light microscopy. The tongue was elongated and their two-third width was almost fixed. Numerous filiform papillae were distributed along the anterior two-thirds of the tongue and fungiform papillae were distributed between the filiform papillae. Eight vallate papillae that had a weak circumferential ridge were distributed in a V-shape in the posterior part of the tongue and numerous taste buds were observable in the circumferential furrows of vallate papillae. Weak fold-like foliate papillae were observable at the lateral edge in the posterior part of the tongue and a few salivary duct orifices were observable beneath the foliate papillae. An islet-like structure with numerous taste buds, was observable on the deep part of the salivary duct of foliate papillae. Large conical papillae were distributed at the posterior part and root of the tongue. After removal of epithelium, filiform papillae of CTCs were appeared to be a thumb or cone-like main core and associating several finger-like short accessory cores. These cores were surrounded an oval concavity. The main core was situated behind the concavity and associated with accessory cores. CTCs of fungiform papillae were cylinder-like with numerous vertically running ridges and with a few concavities seen at the top of the cores. CTCs of vallate papillae and their surrounded circumferential ridge were covered with numerous pimple-like protrusions. The lingual papillae of Common raccoon's tongue had morphological feature of carnivore species.

Morphology and glycoconjugate histochemistry of the eccrine glands in the snout skin of the North American raccoon (Procyon lotor).

The eccrine nasolabial glands were found in the hypodermis of the nasal plane in the North American raccoon (Procyon lotor). In addition to light and electron microscopic observations, the distribution and selectivity of complex glycoconjugates in the eccrine tubular glands of the raccoon snout skin were studied using various histochemical methods, particularly lectin staining. The secretory epithelium and the luminal secretions exhibited high amounts of glycoconjugates with various saccharide residues (alpha-D: -mannose, alpha-L: -fucose, beta-D: -galactose, beta-N-acetyl-D: -glucosamine, sialic acid). The excretory duct cells also showed positive reactions with most of the histochemical methods applied. The results are discussed with regard to possible functions of the glandular secretions. The complex glycoconjugates that are produced by the eccrine nasolabial glands may be related to moistening of the skin surface as well as protecting the epidermis against physical damage or microbial contamination. This is the first report on the glands in the snout skin of carnivores.

Microvasculature of the retina, ciliary processes and choroid in the North American raccoon (Procyon lotor) eye: a scanning electron microscopic study of corrosion casts.

We have studied the vasculature of the retina, ciliary processes and choroid in the North American raccoon (Procyon lotor), a nocturnal mammal, using light and scanning electron microscopic examination of corrosion casts. We carried out an identical study in the crab-eating macaque (Macaca fascicularis), which forages only during the daytime, in order to compare the ocular vasculature with that of nocturnal mammals. Our observations in raccoons demonstrated a photoreceptor layer associated with rich lymph and a poorly vascularized retina. The meridian region of the eye, which lies in the horizontal plane and pass around the optic disc, had a markedly sparse capillary network. This horizontal sparse vascular band may correspond to a visual streak. Ciliary process capillaries were delicate, and formed a well-developed and compact network. Choriocapillaries were quite thin and formed a coarse capillary network. This contrasted with the dense retinal and well-extended choroidal capillary networks noted in the macaques. Our findings suggest that the sparse retinal capillary network in raccoons is extremely beneficial for photon capture, thereby allowing the raccoon to see well at night, as the retinal vessels restrict the inflow of photons toward the photoreceptors. The well-developed lymph probably compensates for the sparse retinal capillaries and choriocapillaries and nourishes the retina in the nocturnal raccoon.

Projections of digit afferents to the cuneate nucleus in the raccoon before and after partial deafferentation.

Within the cuneate nucleus of the raccoon, the representations of individual forepaw digits are anatomically separated by densely myelinated laminae. This unique arrangement was utilized to determine whether the terminations of cutaneous afferents from individual digits are precisely restricted to the appropriate region of the cuneate nucleus or overlap with afferents from adjacent digits. By using the transganglionic transport of horseradish peroxidase (HRP), it was found that, for each digit, the terminal labeling was restricted to the appropriate 150-250-micron-wide column that extended rostrocaudally throughout the nucleus. The topographical arrangement of digit input corresponded to the known electrophysiology, with the terminal column for the fifth digit located most medially within the nucleus and those for digits 4 to 1 successively more laterally. Within a column, the density of labeling was greater over cell clusters than between clusters. These results indicate that afferents from adjacent digits do not overlap in the cuneate nucleus. In six animals, the fifth digit was amputated, and 2-4 months later, HRP was injected into the nerves of the fourth digit to determine whether its afferents had sprouted into the denervated fifth-digit column. The projection pattern from the fourth digit in each of these animals was the same as in normal animals and the same as in the intact contralateral side. These results indicate that the reorganization seen in the cerebral cortex following peripheral deafferentation cannot be attributed to changes in the afferent fiber projections to the cuneate nucleus.

Somatosensory thalamocortical connections in the raccoon: an HRP study.

In the North American raccoon (Procyon lotor), representations of the glabrous surfaces of the hand digits are found within separate subnuclei of the thalamic ventrobasal complex (VB) and on separate subgyri of the somatosensory cortex (SmI). In the present study, the retrograde transport of horseradish peroxidase from SmI to VB was utilized to study relationships between physiologically identified cortical subgyri and somatotopically corresponding thalamic subnuclei. Single large or multiple small injections confined to a single gyral crown led to retrograde labeling of large groupings of cells filling the entire VB subnucleus for the appropriate digit. In the aggregate, the regions of label appeared as thin, wedge-shaped sheets extended in the dorsoventral and anteroposterior dimensions, but flattened mediolaterally, and curving to form a laterally directed convexity; these appear to correspond to the lamellae of monkey VB described by others. These large injections led to labeling of approximately 80% of all large (18-30-micron diameter) cells within the lamella. Single, small, focal injections of a gyral crown led to variable amounts of labeling, ranging from an entire digital lamella to only a small focal cluster of cells. No evidence was obtained for the existence of anteroposteriorly extending "rods" of cells, as reported in primates. Finally, there was a sparse, but consistent labeling of cells of the posterior nuclear group (Po) following gyral crown injections. These results are in agreement with expectations based on prior electrophysiological studies of raccoon VB and SmI, as well as prior anatomical studies of thalamocortical relationships.

Distribution of cells projecting to thalamus vs. those projecting to cerebellum in subdivisions of the dorsal column nuclei in raccoons.

To learn the distribution of cells projecting to the thalamus, as opposed to the cerebellum, in the mechanosensory nuclei of the dorsal medulla of raccoons, we analyzed the retrograde transport of horseradish peroxidase from the ventrobasal complex of the thalamus and from the cerebellum. We found six nuclear regions projecting heavily to the thalamus with very small projections to the cerebellum: Bischoff's, central cuneate, central gracile, rostral cuneate, rostral gracile nuclei, and cell group z. Two regions showed heavy projections to the cerebellum with no projections to the thalamus: the lateral portion of the external cuneate nucleus and the compact portion of cell group x. Four regions showed more equivalent projections to both target regions: basal cuneate, medial portion of the external cuneate nucleus, medial tongue extension of the external cuneate nucleus, and reticular portion of cell group x. Three more ventral regions were labeled: lateral cervical nucleus from thalamic injections but not from cerebellar injections; central cervical nucleus from cerebellar injections, which crossed the midline, but not from thalamic injections; and lateral reticular nucleus from both target regions. In most medullary regions, most cells project to one target and very few project to the other; we suggest that the cells projecting to the minor target convey samples of the information going to the major target.

Medullary sources of projections to the kinesthetic thalamus in raccoons: external and basal cuneate nuclei and cell groups x and z.

In raccoons and other mammals, a pathway for kinesthetic sensation (from muscles, fascia, tendons, and joints) reaches the anterodorsal cap of the ventrobasal thalamus and the anteriormost part of the somatic sensory cerebral cortex. To find the medullary component of this kinesthetic pathway in raccoons, small injections of horseradish peroxidase were made in the thalamus under guidance of simultaneous electrophysiological recording from kinesthetic projections. As determined by retrograde labeling following these injections, kinesthetic thalamic subregions receive projections as follows: caudomedial from cells in the external cuneate nucleus and its medial tongue, rostromedial from cells in basal cuneate nucleus, and rostrolateral from cells in cell group z and the reticular division of cell group x. Electrophysiological recording showed kinesthetic representations in each of these medullary regions. Labeled cells were also observed in the infratrigeminal subnucleus of the lateral reticular nucleus. Cats have kinesthetic projections to the thalamus from the basal cuneate and cell group z; raccoons (and monkeys) have these plus projections from the external cuneate and cell group x. This suggests that the kinesthetic projection system in raccoons and monkeys is expanded in correlation with their more dextrous use of the hand.

Somatosensory receptive field properties of corpus callosum fibres in the raccoon.

Anatomical studies in a number of species have shown that most areas of the somatosensory cortex are callosally interconnected. This is also true for the raccoon, at least for those parts representing proximal and axial body regions. Electrophysiologically, studies carried out in cats and monkeys have demonstrated that all sensory sub-modalities cross in the callosum. Moreover, cells representing the paws and fingers, though occupying a large portion of areas SI and SII, seem to send proportionately fewer axons through the callosum than axial structures. No comparable study has been carried out in the raccoon. The purpose of the present experiment was therefore to investigate the functional organization of the callosal system in this animal by examining the receptive field properties of the somatosensory fibres crossing in the callosum. Axonal activity was recorded directly through tungsten microelectrodes in the corpus callosum of eight raccoons. Results indicated that somatosensory information is transmitted in its rostral portion. Most receptive fields concerned axial and proximal body regions and the head and face. Some receptive fields represented para-axial regions of the body and a few concerned the hands and fingers. Slowly and rapidly adapting fibres were found, as were all the sensory sub-modalities tested. A substantial proportion of the axons had bilateral receptive fields. These results are discussed in relation to those obtained in other species, with particular reference to: (1) the midline fusion hypothesis of callosal function; (2) the representation within this structure of the distal extremities, and (3) the origin of the bilateral receptive fields.

A horseradish peroxidase study of projections from the main and external cuneate nuclei to the cerebellum of the North American raccoon.

Neurons of the raccoon main (MCN) and external (ECN) cuneate nuclei having terminal fields in the anterior lobe and paramedian lobule of the cerebellum were studied by means of the retrograde transport of horseradish peroxidase (HRP). In the MCN, neurons of the polymorphic regions, but not of the round cell clusters, were labeled following HRP injections of the anterior lobe. HRP-labeled fusiform, triangular, and stellate cells were observed from 3-4 mm caudal to the obex to 2-3 mm rostral to it. The main body of labeled MCN neurons was located in the 1.25 mm immediately rostral to the obex, there forming a recognizable subnucleus in the ventral MCN. The ECN displayed uniform labelling from the anterior lobe with no particular cell type or nuclear region being labeled preferentially. A minor projection to the anterior lobe appeared to originate in the contralateral MCN and ECN. Injections of the paramedian lobule produced MCN labeling similar to that observed after anterior lobe injections, but the majority of labeled ECN neurons were found in the dorsomedial part of the nucleus. In lobule V of the anterior lobe, sagittal zone c contained the greatest density of cuneocerebellar terminals as judged by the degree of retrograde labeling seen in neurons of both the MCN and ECN. This observation was consistent with evoked potentials recorded in the anterior lobe of the raccoon cerebellum. The position of cuneocerebellar neurons in the ventrolateral MCN suggests that the bulk of the information being transferred to the cerebellum by the raccoon MCN is related to deep, rather than cutaneous, modalities.

Organization of intracortical and commissural connections in somatosensory cortical areas I and II in the raccoon.

The organization of intracortical and callosal projecting cell bodies was examined in somatosensory representation areas I (SI) and II (SII) of the raccoon by use of horseradish peroxidase (HRP) or horseradish peroxidase-wheat germ agglutin (HRP-WHA). HRP and HRP-WHA were injected into commissurally and noncommissurally connected subdivisions of SI and SII. Injection sites in SII were identified electrophysiologically. Results were obtained from transverse sections in which the HRP was visualized with the aid of the substrates dihydrochlorobenzidine or tetramethyl benzidine in the presence of hydrogen peroxidase. The principal findings were the following: (1) there are reciprocal connections between SI and SII; (2) in SI the intracortically projecting cell bodies and terminals are located primarily in sulcal cortex; (3) intracortically projecting neurons in SI are located primarily in layers III whereas in SII they are located principally in layers III and V; (4) there are connections between disparate areas within SI; and (5) there are intracortical connections between callosum-connected and acallosal regions in SII. These results are discussed with regard to the results of mapping studies of the SI, the significance of intracortical connections to the formation of sulci in SI, and the possible roles of nonhomotopic connections in the intermanual transfer of learning.

Demarcations of the mechanosensory projection zones in the raccoon thalamus, shown by cytochrome oxidase, acetylcholinesterase, and Nissl stains.

To determine anatomically the boundaries and internal organization of the kinesthetic and cutaneous mechanosensory regions of the ventrobasal thalamus, alternate section series from electrophysiologically mapped tissues from 14 raccoons were stained for cytochrome oxidase, myelinated fibers, acetylcholinesterase, and Nissl substance. Microelectrode tracks, along with electrolytic lesions placed as tissue markers, reveal that the mechanoreceptor projection zones have higher cytochrome oxidase and lower acetylcholinesterase staining than some neighboring regions. Both these enzymatic stains reveal particularly sharp boundaries separating the mechanoresponsive region, from the lateral posterior nucleus dorsally and from the ventroposterior inferior nucleus ventrally. The kinesthetic projection zone is often separated from other mechanoreceptor projections by bundles as well as laminae of myelinated fibers, similar to those separating cutaneous projections from distinct body parts. These subdivisions are particularly well marked by the cytochrome oxidase stain. The combination, in neighboring sections, of the use of the several stains adds considerably to the visible delineation of these functionally distinct regions, beyond what can be seen in Nissl-stained sections.

Distribution of neurons immunoreactive for parvalbumin and calbindin in the somatosensory thalamus of the raccoon.

The aim of this study was to assess the distribution of neurons immunoreactive for parvalbumin (PV), calbindin (CaBP), glutamic acid decarboxylase (GAD), and gamma-aminobutyric acid (GABA) in the somatosensory thalamus of the raccoon and to compare these features to those of other species, especially primates. Immunohistochemistry was used to study the location of these neurons in the ventroposterior nucleus (VP), ventroposterior inferior nucleus (VPI), posterior group of nuclei (Po), and reticular nucleus (Rt). A consistent differential pattern of PV-positive (PV+) and CaBP-positive (CaBP+) cells was found in the somatosensory thalamus. Many PV+ neurons were observed in VP and Rt, but very few were found in VPI or Po. In contrast, CaBP+ neurons were distributed throughout VP, VPI, and Po but were very sparse or absent in Rt. In the VP nucleus, PV+ cells were distributed in clusters separated by interclusteral regions with a sparse distribution of PV+ cell bodies. The distributions of PV+ and CaBP+ cells tended to be complementary to each other in VP; regions with a high density of PV+ neurons had a low density of CaBP+ cell bodies. Double-labeling experiments revealed very few neurons in which PV and CaBP immunoreactivities were colocalized. Cells immunoreactive for GAD or GABA were found in PV-dense clusters of VP; fewer GABAergic neurons were present in the CaBP-dense interclusteral regions of VP and in VPI and Po. GAD+ and GABA+ neurons were most prominently distributed in Rt. We conclude that the distributions of PV+ and CaBP+ cell bodies in the raccoon somatosensory thalamus are very similar to those in primates. The density of GABAergic neurons in the somatosensory thalamus of the raccoon is less than that in the cat and monkey, but the relative distribution of GABAergic neurons in the different somatosensory nuclei is very similar to that in the cat and monkey. These results are discussed in relation to findings in other species and are related to the functions of lemniscal and nonlemniscal somatosensory pathways.

Somatotopic organization of mechanosensory projections to SII cerebral neocortex in the raccoon (Procyon lotor).

The pattern of projections of peripheral receptors to the neocortex in the second somesthetic receiving area (SII) was mapped in raccoons. The purpose was to determine if the projection area of peripheral receptors to the forepaw area in the SII region is disproportionally enlarged as it is in SI. Tungsten microelectrode recording procedures were used to map thoroughly the inferior wall of the suprasylvian sulcus for regions responsive to mechanical stimulation of peripheral receptors. The results show that: 1. The forepaw area in SII shows an enlargement commensurate with that found in the SI. This suggests that those factors that are selective for tactile acuity of the raccoon forepaw were operating in the evolution of SII as they were in SI. 2. The somatotopic organization of mechanoreceptive projections to SII is reversed mediolaterally compared to previous descriptions of this arrangement in other mammals: projections form axial structures lie medially and those from apical structures lie laterally along the inferior bank of the suprasylvian sulcus in the raccoon.

Innervation of the digit on the forepaw of the raccoon.

The innervation of the digits on the raccoon forepaw was examined by using immunochemistry for protein gene product 9.5, calcitonin-gene related peptide, substance P, neuropeptide-Y, tyrosine hydroxylase, and neurofilament protein. The larger-caliber axons in the ventral glabrous skin terminate as Pacinian corpuscles deep in the dermis, small corpuscles and Merkel endings around the base of dermal papillae, and Merkel endings on rete pegs in dermal papillae. Extensive fine-caliber innervation terminates in the epidermis and on the microvasculature. The innervation is more dense in the distal than in the proximal volar pads. Pacinian endings are also concentrated in the transverse crease separating the distal and proximal pads. In the dorsal hairy skin, hair follicles are well innervated with piloneural complexes. Merkel innervation is located under slight epidermal elevations and in some large Merkel rete pegs located at the apex of transverse skin folds just proximal to the claw. No cutaneous Ruffini corpuscles were found anywhere on the digit. The claw is affiliated with dense medial and lateral beds of Pacinian endings, bouquets of highly branched Ruffini-like endings at the transition from the distal phalanx and unmyelinated innervation in the skin around the perimeter. Encapsulated endings are located at the lateral edge of the articular surface of the distal phalanx. Extensive fine-caliber innervation is affiliated with sweat glands and with the vasculature and is especially dense at presumptive arteriovenous sphincters. Virtually all of the sweat gland and vascular innervation is peptidergic, whereas most of the unmyelinated epidermal innervation is nonpeptidergic.

Species differences in mechanosensory projections from the mouth to the ventrobasal thalamus.

To determine whether the largely ipsilateral, inverted representation of mouth parts in the ventrobasal thalamus of sheep was unique to that species or an expansion of a general mammalian pattern, the corresponding thalamic projections were mapped electrophysiologically in a selected series of mammals (oppossums, agoutis, squirrel monkeys, cats, raccoons, and sheep) representing major branches of evolution among therian mammals. In mapping, tungsten microelectrodes were used to record multi-unit discharges in the thalamus in response to mechanical stimulation of oral surfaces. The pattern of projections seen in sheep is not a general mammalian pattern; there is extensive variability among mammals in the laterality and internal orgainzation of the projections from the mouth. In spite of the great variability, the results suggest an hypothesis concerning phylogenetic trends: descendants of palaeoryctoid insectivores (cats, raccoons, and sheep in our sample) have extensive ipsilateral projections from the mouth, in other therian mammals (opossums, agoutis, and squirrel monkeys in our sample) the ipsilateral component is small or absent.

Variation and evolution of mammalian corticospinal somata with special reference to primates.

The morphology of the somata originating the corticospinal tract was examined in 24 species of mammals to identify commonalities and major sources of variation among the different species. Horseradish peroxidase was applied to a hemisection of the spinal cord at the C1-C2 junction. After tetramethylbenzidine processing, the labeled somata throughout the cerebral cortex were plotted and counted. Then, 23 morphological characteristics of the corticospinal somata were examined, including their number, size, and density across the cortical surface. The results show that morphological characteristics of corticospinal somata are closely related to an animal's body, brain, and cerebral cortex size. That is, mammals with large neocortical surfaces tend to have larger as well as more corticospinal somata; mammals with large bodies tend to have corticospinal somata that are less densely distributed. Moreover, the probable increase in the ratio of local noncorticospinal somata to corticospinal somata implies that the evolution of the corticospinal tract was accomplished by an increase in "support" or "server" cells as well as an increase in the size of the tract itself. The results also show that several characteristics are reliably related to an animal's taxonomic classification and hence its ancestry. Comparisons among three mammalian lineages indicate that some characteristics may have changed uniquely in the anthropoid primate lineage, and thus, presumably, in the human lineage. The results suggest that if morphological characteristics of the corticospinal tract important in the evolution of the specialized motor abilities in anthropoid primates are sought, then examination of the role of changes in soma diameter, rostral (motor)/caudal (sensory) ratios of density, concentration, surface density, and volume density may be more instructive than examination of the total number of corticospinal neurons alone.

The connections of cortical somatosensory areas I and II with separate nuclei in the ventroposterior thalamus in the raccoon.

The thalamocortical afferents to cortical somatosensory areas I (SI) and II (SII) were investigated in the raccoon using the horseradish peroxidase technique. The purpose of this study was to determine if the cell bodies of origin for thalamocortical afferents to these cortical regions were localized in the same or different nuclei in the ventroposterior region of the thalamus. Horseradish peroxidase was injected into subdivisions of SI or SII and after post-injection survival periods of 12-72 hours the horseradish peroxidase in the tissue was reacted with the chromogens dihydrochlorobenzidine or tetramethylbenzidine in the presence of hydrogen peroxide. The results show that SI and SII receive projections from neurons in separate and distinct nuclei in the ventroposterior thalamus. Following injections into subdivisions of area I, a topographical distribution of retrogradely-labelled cell bodies was observed in the ventrobasal complex. Following injections of horseradish peroxidase into subdivisions of area II, a topographical distribution of labelled cell bodies was observed in the ventroposterior inferior nucleus. No labelled cell bodies were observed in the ventrobasal complex. The thalamocortical connections of somatosensory cortices I and II in raccoon are compared with those in other animals and it is suggested that these two cortical areas may be involved in differential processing of tactile information.

The anterior border zones of primary somatic sensory (S1) neocortex and their relation to cerebral convolutions, shown by micromapping of peripheral projections to the region of the fourth forepaw digit representation in raccoons.

In raccoons the somatic sensory neocortex is greatly expanded, with separate gyral crowns devoted to and intervening sulci separating, sensory representations of separate body parts, most strikingly those of the volar surfaces of individual forepaw digits. Most of the cortex in this region is buried in widely ramifying sulcal walls, wherein sensory projections have not been studied. We have determined mechanosensory projections to the fourth digit representation region including all neighboring sulcal walls, using tungsten microelectrodes for 3-dimensional micromapping. We found no significant alteration in the location and pattern of projections when the following different anesthetics were used: dial-urethane, chloralose, or methoxyflurane with nitrous oxide. The precisely organized somatotopic representation of the distal volar surface of the fourth digit, on the causal aspect of its gyral crown, continues down the anterior bank of the triradiate sulcus. This meets, at the fundus, projections from the proximal volar surface of the digit which occupy the posterior sulcus wall; they in turn meet projections from the volar palm at the gyral crown. In the anterior part of the crown containing the representation of the distal volar digit, across the crown. In the anterior part of the crown containing the representation of the distal volar digit, across the crown of the gyral bridge intervening between the medial and lateral segments of the central sulcus, throughout the posterior walls of the central sulci, and in the walls of the interbrachial sulcus, we found a distinctive border-zone of projections from heterogeneous receptive fields. Within a roughly somatotopic basic pattern of organization we found intermingled projections from single and multiple claws and dorsal hairy surfaces of digits and proximal hand, along with additional projections from volar surfaces. These projections can be construed as forming something of a distorted mirror-image of the representation of the volar hand. Beyond this was a second zone of distinctive projections from afferents of the forelimb muscles, in the anterior walls of the central sulci. These projections are interrupted where the sulci are interrupted. The zone of muscle afferent projections corresponds to those seen between sensory and motor regions in other species; its strict association with sulcal folding here and in other species suggests a general relationship of these projections to central sulci. The zone of heterogeneous projections resembles similar zones seen at other levels of this system in raccoons, in the cortex of other species, and it may relate to some of the multiple representation reported in other species. It also may be related to the formation of sulci in this region and may be a specialized zone for cortico-cortical connections.