A simple method for overcoming some problems when observing thick reflective biological samples with a confocal scanning laser microscope.

A simple device is described, which allows the range of depth of scanning to be reduced when observing thick reflecting biological samples with a confocal scanning laser microscope (CSLM). Thick histological sections of human skin and rat brain stem were mounted between two coverslips ('sandwich' style) and the optical tomography was performed from both sides by turning the 'sandwich' upside-down. The samples were impregnated using standard Golgi-Cox, 'rapid Golgi' or other silver methods. The ability to turn the 'sandwich' upside-down is particularly useful when the reflective structure inspected is deep inside the section, i.e., near the lower surface of the specimen, or when it is opaque to the laser beam or excessively reflective.

A quantitative ultrastructural investigation of tyrosine hydroxylase-immunoreactive axons in the hairy skin of the guinea pig.

Besides its thermoregulatory role, the sympathetic innervation of the skin is involved in a modulation of sensory processing and trophic functions that has not been fully characterized. To investigate possible sites at which such sympathosensory interactions might occur, a quantitative ultrastructural study of the sympathetic innervation of the skin was attempted. The hairy skin of the guinea pig was studied because the sympathetic and sensory nerve axons in this species can easily be discriminated by the presence of immunoreactivity to the catecholamine-synthesizing enzyme, tyrosine hydroxylase (TH). The thermoregulatory role of the sympathetic skin innervation was highlighted by the almost exclusive sympathetic innervation of piloarrector muscles which contained 62% (n = 195) of randomly selected TH-immunoreactive (TH-IR) axon profiles. Of TH-IR pilomotor axons, 53% were filled with vesicles. Vesicle-containing axonal profiles were equally frequent around dermal arterial blood vessels (partly associated with mast cells), hair follicles, and within nerve fibre bundles surrounded by a perineural sheath, in each case accounting for about 3% of all dermal TH-IR axonal profiles. In contrast to piloarrector muscles, at these locations TH-IR (sympathetic) and non-reactive (sensory) axons were found in close association. These findings are in line with the previously reported inhibitory influence of sympathetic stimulation upon hair follicle afferents and perivascular sensory nerve terminals. In addition, they point to a yet underestimated target of sympathetic axon terminals, i.e. preterminal nerve fibre bundles.

SP- and CGRP-immunoreactive axons differ in their ability to reinnervate the skin of the rat tail.

The reinnervation of cutaneous targets was studied in the rat tail after proximal lesions of all collector nerves. The distribution of immunofluorescent nerve fibres stained for calcitonin-gene-related peptide (CGRP) and substance P (SP) was examined after 110-210 days. Targets at all sites were reinnervated by CGRP-immunoreactive (IR) fibres. However, SP-IR terminals were rare, particularly distally, despite staining within subdermal nerve trunks.

Central terminations of low-threshold mechanoreceptive afferents in the trigeminal nuclei interpolaris and caudalis of the cat.

Previous studies indicate that vibrissa, nonvibrissa, guard hair, hairy skin, and periodontal ligament afferents give rise to morphologically distinct terminal arbors in the trigeminal nuclei principalis (Vp) and oralis (Vo) in the cat. The present study describes the extent to which morphological and functional relationships exist in the nuclei interpolaris (Vi) and caudalis (Vc). Twenty-two fibers were physiologically characterized and stained by intra-axonal HRP injection techniques. The fast adapting (FA) vibrissa (VF) afferents gave rise to compact and circumscribed arbors in Vi and Vc. These tended to be larger in Vc than in Vi. The slowly adapting (SA) vibrissa (VS) afferents in Vi and Vc had more widespread and larger arbors than those of the VF afferents. The VS arbors in Vc tended to be larger and less circular than those in Vi. Guard hair (GH) afferents gave rise to circumscribed arbors in both nuclei, but those in Vc tended to have larger and more circular arbors than those in Vi. Down hair (DH) afferents gave rise to small, circumscribed arbors or a few distinct patches of boutons within a small area in Vi; arbors in Vc were less extensive and "stringy." Unlike other afferents, DH arbors were larger in Vi than in Vc, but smaller than those of GH afferents in either nuclei. The SA hairy skin (SS) afferents had arbors that were widespread with a few distinct patches of boutons; the arbors in Vc were larger than those in Vi. The arbors of SS afferents were smaller than those of VS and GH afferents in both nuclei. Like GH afferents, FA periodontal ligament (PF) afferents gave rise to large and circumscribed arbors in Vi, although the arbors in Vc were smaller and less dense. The present study demonstrated significant functional and morphological relationships for primary afferents in Vi and Vc, thus suggesting that sensory information from each of the distinct fiber or functional classes is processed in a characteristic manner in the individual nuclei.

Differential ascending projections from neurons in the cat's lateral cervical nucleus.

Extracellular microelectrode recordings were made from single cells of the lateral cervical nucleus (LCN) in cats anaesthetized with chloralose and paralysed with gallamine triethiodide. The cells were tested for antidromic activation from the contralateral medial lemniscus and the contralateral tectum. Seventy-two LCN units were recorded which projected to one or both targets. Sixty (83%) projected through the medial lemniscus, and of these 36 (50% of the total) also projected to the tectum, whereas 24 (33%) projected through the medial lemniscus only; 12 (17%) projected only to the tectum. Twenty-nine units (40%) were excited by moving hairs of the coat but not by pinch of the skin, and 9 (31%) of these projected to the tectum, 11 (38%) through the medial lemniscus and 9 (31%) to both targets. Forty units (56%) were excited by hair movement and noxious pinch, and 3 (7%) of these projected to the tectum, 10 (25%) through the medial lemniscus and 27 (68%) to both targets. Three units (4%) had no discernible receptive fields and they all projected through the medial lemniscus, but not to the tectum. Of the 12 units projecting only to the tectum, 11 had receptive fields completely or partially on the trunk. Units projecting either through the medial lemniscus only, or through the medial lemniscus and also into the tectum, had receptive fields more widely distributed: these included small fields on the fore- and hind feet, on the limbs and also, a minority, on the trunk. Units with glove- or stocking-like receptive fields projected through the medial lemniscus. The results show that while most LCN cells project through the medial lemniscus, those excited by hair movement alone preferentially project either to the tectum or through the medial lemniscus, but not by both routes. The differences in receptive field properties of the differently projecting units are discussed in terms of the possible functions of the spinocervical system.

Receptive field properties of trigeminothalamic neurons in the rostral trigeminal sensory nuclei of cats.

This study described topographic and receptive field representation in the region of the rostral trigeminal nuclei, and evaluated whether thalamic neurons from the principal sensory nucleus relay muscle afferent information to the thalamus. Extracellular single-unit activity was recorded from anesthetized cats. Units were tested for responses to natural stimuli (i.e., air bursts, brushing, light pressure, and pinch) applied to the face and oral cavity, electrical stimulation of the masseter nerve, and ramp-and-hold movements of the jaw. The receptive fields and physiological properties for 110 units were studied; we were able to verify the recording site for 96 of these units. Most of the units had discrete receptive fields in the oral cavity, skin, hair, and masseter muscle. Only 2 units received convergent inputs. Stimulation of the ipsilateral and contralateral ventroposteromedial nucleus of the thalamus was performed to identify antidromically activated units. The results showed that the dorsal principal sensory nucleus received its input primarily from the oral cavity. Most of the units (85%) that were activated by antidromic stimulation from the ipsilateral thalamus were located in this nucleus. In contrast, 82% of the units that projected to the contralateral thalamus were located in the ventral principal sensory nucleus. A complete somatotopic representation of the ipsilateral face and oral cavity was observed in the ventral principal sensory nucleus. Although 24 units had muscle receptive fields, none were activated by stimulation of the ipsilateral thalamus, and only 1 responded to stimulation of the contralateral thalamus. Most of the units that were not antidromically driven were recorded outside of the cytoarchitectural boundaries of the principal sensory nucleus. Retrograde labeling of the rostral trigeminal nuclei indicated that most of the neurons in the dorsal principal sensory nucleus projected to the ipsilateral thalamus, whereas those in the ventral principal sensory nucleus projected to the contralateral thalamus. Taken together, these observations do not support the presence of a substantial relay for muscle afferent input from the dorsal principal sensory nucleus to the ventrobasal thalamus in cats.

Hairy skin: psychophysical channels and their physiological substrates.

Experiments were conducted in which threshold-frequency characteristics were measured on the hairy skin of the forearm of human observers. Thresholds were measured with two stimulus probe areas (2.9 and 0.008 cm2) at three skin-surface temperatures (15 degrees, 30 degrees, and 40 degrees C). The results suggest that whereas glabrous skin uses four distinct channels of information, only three channels may be involved in mediating the sense of touch for hairy skin. The three channels are defined as Ph, (Pacinian, hairy skin), NPh low (non-Pacinian, hairy skin, low frequencies) and NPh mid (non-Pacinian, hairy skin, middle frequencies). In addition, it is proposed that the neural substrates for the three psychophysically characterized channels are, respectively, the Pacinian corpuscle (PC) nerve fibers, the slowly adapting type II (SAII) fibers, and the rapidly adapting (RA) fibers.

Receptive field reorganization in dorsal column nuclei during temporary denervation.

Altered sensory input can result in the reorganization of somatosensory maps in the cerebral cortex and thalamus, but the extent to which reorganization occurs at lower levels of the somatosensory system is unknown. In cat dorsal column nuclei (DCN), the injection of local anesthetic into the receptive fields of DCN neurons resulted in the emergence of a new receptive field in all 13 neurons studied. New receptive fields emerged rapidly (within minutes), sometimes accompanied by changes in adaptation rates and stimulus selectivity, suggesting that the new fields arose from the unmasking of previously ineffective inputs. Receptive field reorganization was not imposed by descending cortical inputs to the DCN, because comparable results were obtained in 10 additional cells when the somatosensory and motor cortex were removed before recording. These results suggest that mechanisms underlying somatotopic reorganization exist at the earliest stages of somatosensory processing. Such mechanisms may participate in adaptive responses of the nervous system to injury or continuously changing sensory stimulation.

Morphology of A beta hair follicle afferent collaterals in dorsal horn of cats with neonatal chronic denervation of digits.

1. The effect of neonatal denervation of two adjacent hindpaw digits (digits 3 and 4) in the cat on the morphology of spinal collaterals of identified A beta hair follicle afferents (HFAs) with receptive fields (RFs) on the neighbouring digits has been investigated using intra-axonal injection of horseradish peroxidase. 2. The morphology of collaterals of HFAs innervating intact digits (digit 2 or digit 5) was examined within the neonatally and chronically deprived region of dorsal horn by selectively injecting the afferents directly over this region. 3. Of a total of nine injected axons, seven had RFs on the distal part of digit 2, whereas two had RFs on the distal part of digit 5. With the exception of one of the digit 5 afferents, all injected afferents gave rise to stained collaterals throughout the chronically deprived digit representation. 4. Many collaterals within the chronically deprived region of dorsal horn gave rise to arborizations having relatively large numbers of boutons when compared with previously described somatotopically inappropriate (SIA) collaterals in the normal adult cat. 5. It is suggested that neonatal denervation of digits in the cat may lead to collateral sprouting and bouton formation by SIA collaterals of A beta HFAs innervating adjacent intact digits. This morphological plasticity could explain the previously reported functional reorganization of the RFs of spinocervical tract neurons that occurs in these neonatally deafferented animals.

Differential activation and classification of cutaneous afferents in the rat.

1. A total of 312 cutaneous afferent units identified in the rat foot as belonging to one of nine major types of sensory receptors were included in the present study. A natural stimulus set was defined to differentiate optimally among those receptor types according to the distinguishing response patterns that it produced. It included air puffs, 30- and 300-Hz sinusoids, 200-mN force indentation of the skin, 1.2- and 6-N compressions of a skin fold, cooling the skin by 5 and 20 degrees C, warming by 5 degrees C, and heating by 15 degrees C. 2. The responses to predefined stimuli of 188 units were subjected to multivariate statistical analyses. The responses of an individual unit were measured as the number of impulses evoked by 10 stimuli, each lasting 10 s. Additionally, the number of impulses occurring for 5 s after withdrawal of a 200-mN indentation (1 of the 10 stimuli) was counted. 3. In discriminant analysis, the 11 stimulus variables predicted fairly correctly the grouping of afferent units into nine predetermined receptor categories (175 of 188, 93.1%), indicating a powerful ability to discriminate among different receptor types. Using hierarchical cluster analysis, afferent unit data described by 11 variables were divided into clusters that well represented prior receptor categories (170 of 188, 90.4%), suggesting the reliable application of this procedure to the classification of newly recorded cutaneous sensory receptors. 4. Eleven variables were then reduced to 7 on the basis of the results of factor analysis (95% of variance accounted for). The seven variables corresponded to 1.2-N compression, heating the skin by 15 degrees C, cooling the skin by 20 degrees C, 30- and 300-Hz sinusoids, withdrawal of a 200-mN indentation, and air puffs. 5. The seven selected variables correctly assigned afferent units into five modality-based categories in the discriminant solution (177 of 188, 94.1%). In the cluster solution, afferent units described by the seven selected variables were divided into clusters, most of whose members were modality specific (176 of 188, 93.6%). 6. The results indicate that cutaneous receptors can be divided into modality-specific groups according to similarities in their responses to seven stimulus variables. It is proposed that the stimulus set developed here and multivariate statistical methods can be used as powerful tools for the functional classification of central somatosensory neurons.

Morphology and topography of identified primary afferents in trigeminal subnuclei principalis and oralis.

1. Intra-axonal recording, receptive field mapping, horseradish peroxidase injection, cytochrome oxidase staining, and computer-assisted reconstruction/morphometric methods were used to elucidate the structure and topography of trigeminal primary afferent collaterals in the normal adult rat. Prior studies focused on trigeminal brain stem subnuclei interpolaris and caudalis. This work is extended here to the remaining 2 subnuclei, principalis (PrV) and oralis (SpVo), where collaterals from 66 axons in 37 adult rats were studied. In nine rats, three to five axons were stained for within-nucleus comparisons of different fibers. Quantitative analyses were restricted to vibrissa sensitive fibers. 2. All of the axons conducted rapidly with small, low-threshold receptive fields. The majority responded to vibrissa deflection (n = 47); the remainder responded to guard hair deflection; gentle pressure applied to hairy skin, glabrous skin, lingual mucosa, or an incisor; or jaw movement. All descended in the trigeminal sensory root where some bifurcated into ascending and descending branches. Each well-stained fiber gave rise to transversely oriented collaterals in PrV and SpVo. 3. Within PrV and SpVo, fibers with differing adaptation properties and receptive fields had indistinguishable collateral morphologies. Arbors from single axons were rostrocaudally discontinuous, small relative to collaterals in subnuclei interpolaris and caudalis, circumscribed and topographically organized in a manner consistent with cytochrome oxidase and bulk-labeled primary afferent staining patterns. In SpVo and caudal PrV, the map is inverted with the nose pointing medially. In rostral PrV, the map turns 90 degrees such that the nose points dorsally. 4. Axons had different quantitative properties along the rostrocaudal axis of the trigeminal brain stem complex. Whereas arbors subtended similar transverse areas throughout PrV and SpVo, collaterals in the rostral third of PrV had a relatively low bouton density. Arbors in the caudal two thirds of PrV had the highest bouton density. Arbors in SpVo tended to be more variable in size and shape than those of caudal PrV, and their bouton numbers were significantly lower than in PrV. 5. In PrV, arbors were largely confined to somatotopically corresponding cytochrome oxidase patches, precluding significant overlap of neighboring whisker projections. In SpVo, termination sites were not as strictly confined and numerous examples of within- and between-row overlap were obtained for whisker afferents in cases where multiple axons were stained.(ABSTRACT TRUNCATED AT 400 WORDS)

Sensory innervation of the hairy skin (light- and electronmicroscopic study.

The sense of touch develops early in phylogeny and is one of the most important senses for the survival of the animal. Touch organs of hairy skin in mammals include the so-called "Haarscheiben" (also Pinkus corpuscles) and all types of hair follicles with their nerve endings. The touch organs of the skin consist of a mechanical transducing component and the sensory component. The epithelium and its derivatives like hair follicles and sebaceous glands are the mechanical transducing component transmitting the mechanical forces like pressure or touch to the second component--the sensory nerve endings. In mammalian hairy skin all sinus and guard hairs and many vellus hairs are touch organs. The sinus hair is a typical example of a touch organ. All mammals except humans are equipped with these highly differentiated touch organs. The hair follicle is almost completely embedded in a blood sinus and equipped with more than 2,000 sensory nerve endings. All sinus and guard hairs are equipped with free nerve endings (nociceptors), Merkel nerve endings (slowly adapting [SA I] mechanoreceptor units-pressure detectors), palisades of lanceolate nerve endings (velocity detectors), and pilo-Ruffini corpuscles (tension receptors). In most of the sinus hairs lamellated corpuscles of Pacini type could be found (rapidly adapting receptors-acceleration detectors). Most vellus hairs are equipped with free and lanceolate nerve endings. Some of the vellus hairs of the upper portion of the body (head, upper extremity) are innervated by Merkel nerve endings. The presence of pilo-Ruffini nerve endings in vellus hairs is very unusual.

Cutaneous sensory receptors in the rat foot.

1. A total of 574 cutaneous afferent units in the sural and plantar nerves supplying the skin of the rat foot was examined: 399 A beta-units, 55 A delta-units, and 120 C-units. Their receptive-field (RF) properties were similar to those described in other mammals. However, the receptor type composition of units was different between the two nerves. 2. The sural A beta-fiber sample (n = 160) consisted of G-hair (41%), field (11%), rapidly adapting (RA; 6%), slowly adapting type I (SA-I; 7%), and type II (SA-II; 35%) mechanoreceptors. The plantar A beta-fiber sample (n = 239) was composed of G-hair (3%), RA (35%), SA-I (30%), SA-II (24%), and Pacinian corpuscle (PC; 8%) mechanoreceptors. 3. The RFs of SA-II units were located on both hairy and glabrous skin overlying the foot joints. Many of the SA-II units responded to movement of the foot joints. The RFs of both SA-I and RA units were small in size and located in high density on the toe tips and footpads. PC units were very sensitive to vibration and had extremely large RFs as in other species, although they were rare and found only in the plantar nerve. Field units were similar to SA-II units in response properties and RF distribution. 4. The sural A delta-fiber sample (n = 44) included nociceptors (68%), D-hair (27%), and cold (5%) receptors. All sampled plantar A delta-fibers (n = 11) were nociceptors. Of A delta-nociceptor units, A delta-mechanical nociceptors (73%) were dominant. 5. The sural C-fiber sample (n = 85) included nociceptors (44%), C-mechanoreceptors (33%), and cold receptors (21%). The plantar C-fiber sample (n = 35) included nociceptors (77%) and cold receptors (23%). No warm units were found among either the sural or plantar nerve fibers. Of C-nociceptors, C-mechanoheat nociceptors (80%) were dominant. 6. The results indicate that all well-known types of cutaneous receptors, except warm receptors, exist in the foot skin of the rat. On the basis of the fact that RFs of RA and SA-I units are in high density on the toe tips and footpads, it is suggested that those regions may have a spatial discriminating capacity. It is also suggested that SA-II receptors may play a role in proprioception, because they have RFs on the skin over foot joints and respond to joint movement.(ABSTRACT TRUNCATED AT 400 WORDS)

Identified hair follicle afferent boutons in the spinal cord of the cat are enriched with L-glutamate-like immunoreactivity.

Hair follicle afferent boutons in the spinal dorsal horn of the cat were examined for L-glutamate enrichment. Two hair follicle afferent axons were labelled intra-axonally with horseradish peroxidase, and post-embedding immunogold reactions of L-glutamate were performed on thin sections containing horseradish peroxidase-labelled boutons. Quantitative analysis showed that hair follicle boutons were associated with immunogold reactions for L-glutamate which were almost twice as dense as average values for dorsal horn tissue. Further analysis revealed that hair afferent boutons displayed 2.3-times the average gold particle density associated with post-synaptic dendrites and 3-times the average immunogold density for L-glutamate of structures which were known to be immunoreactive for GABA. This enrichment of L-glutamate in identified hair afferent terminals supports the idea that the amino acid is a neurotransmitter of hair follicle primary afferent axons.

Slow potentials activated by afferent and direct stimulation in spinal interneurons of laminae III-V.

Stimulation of hairy skin afferent fibers and/or direct depolarizing current injection initiates persistent changes in membrane potential (slow potentials), lasting up to 1 min or longer, in some deep dorsal horn neurons. Intracellular staining with horseradish peroxidase shows that these cells are interneurons, suggesting that slow potentials modulate moment-to-moment impulse transmission in local spinal circuits.

[Peptidergic innervation in the sinus hair follicles of several mammalian species].

In this study, the detailed distribution of peptidergic nerve fibers in the sinus hair follicle was immunohistochemically investigated by the avidin biotin-complex method using antibodies against calcitonin gene-related peptide (CGRP), substance P (SP), vasoactive intestinal polypeptide (VIP) and neuropeptide Y (NPY) in the cat, dog, hamster, rat, mouse and guinea pig. The peptide-immunoreactive (IR) nerve fibers presented a fine varicose appearance. They entered the sinus hair follicle after penetrating the capsule (Cap) at various levels. Peptide-IR fibers in the sinus hair were classified into the 3 types described below based on their course of entry and terminal distribution. Type A: these fibers arrive at the orifice of the sinus hair follicle from superficial dermis and innervate the upper portion of the follicle. Most of them form a network around the vibrissal shaft (VS) in the outer and inner conical body (OCB and ICB, respectively), and some are distributed around the rete ridge collar (RRC) and the sebaceous gland (SG). Type B: these fibers enter the lower third of the sinus hair follicle after forming a nerve bundle together with myelinated fibers, or accompany an artery. After distribution in the trabeculae (Trab) of the cavernous sinus (CS), they form a dense plexus in the connective tissue follicle (CTF) at the level of the CS. Some ascend through the CTF and terminate at the level of the ring sinus (RS). Type C: these fibers enter the sinus hair follicle at its base. They innervate the hair papilla (HP) and the CTF of the hair bulb. CGRP and SP were detected in all types of nerve fibers in all species investigated. These peptidergic nerve fibers showed the same distribution pattern, but CGRP fibers were more numerous than SP fibers. They were distributed at high density in the OCB, ICB, CTF, ringwulst (Rw), the Trab of the CS and the HP. A moderate number of VIP- and NPY-IR fibers, mainly types B and C, were detected in these portions in the cat, dog and hamster, but few fibers were observed in other portions or in other species. Although the basic structures of the sinus hair follicle presented almost the same features as in the mammalian species, the connective tissue of the HP extended to the level of the RS in the cat, rat, hamster, and guinea pig. In these species, CGRP-, SP- and/or VIP-IR fibers extended to the top of the papillary connective tissue, and in the cat, were especially well developed.(ABSTRACT TRUNCATED AT 400 WORDS)

Mechanisms underlying somatosensory cortical dynamics: I. In vivo studies.

The experiments of this study demonstrate that relatively modest rates of repetitive tactile stimulation are accompanied by rapid and reversible modifications (either increases or decreases) in the response of SI neurons. Complete recovery occurs in a few minutes following cessation of stimulation. The modifications are reproducible (1) if stimulus parameters remain the same and (2) if time for recovery is provided between successive exposures. In contrast, repetitive tactile stimuli identical to those that modify SI neuron response rarely lead to changes in the response of cutaneous mechanoreceptive afferents. SI neuron functional properties conventionally regarded as immutable [e.g., directional selectivity, and distribution of sensitivity within the receptive field (RF)] also modify with repetitive stimulation. While the changes in RF organization differ in detail from one neuron to the next, they are similar in form: the response generated by stimulus contact with one (or more rarely, several) RF region(s) becomes enhanced relative to the response the same stimulus evokes from neighboring regions. Neurons in the same column (sampled in the same radial penetration) exhibit very similar changes in the distribution of sensitivity within the RF, whereas neurons sampled in tangential penetrations exhibit diverse, apparently unrelated changes in RF organization in response to the same repetitive stimulus. Simultaneous multichannel recordings reveal that a repetitive tactile stimulus exerts similar effects on the response and RFs of the neurons within local (no more than 100 microns) neuron groupings. A model that incorporates a manner of SI topographical organization (segregate organization) and well-known aspects of neocortical cellular, neurotransmitter/receptor, and connectional architecture accounts for the changes in SI neuron behavior observed during repetitive stimulation.

Receptive field properties of somatosensory neurons in the cat superior colliculus.

In general, knowledge of the internal organization of receptive fields has played an important role in shaping current understanding of sensory physiology. Such knowledge is particularly important for understanding the function of the superior colliculus, since this structure is at once implicated in spatial localization and has relatively large receptive fields. While this issue has been addressed in the visual and auditory modalities represented in the superior colliculus, there are no previous studies of its somatosensory receptive field organization. Here, the properties of somatosensory receptive fields in the cat superior colliculus were studied quantitatively to determine whether they contain internal non-homogeneities that might aid in the determination of stimulus detail. Of special interest was the possibility that these comparatively large receptive fields would contain areas of differential excitability that could aid in spatial resolution, that within-field spatial summation and/or inhibition would be exhibited, and that the borders of the excitatory receptive field would be flanked by inhibitory regions. The data demonstrate that while inhibition beyond the receptive field borders is a rarity, these somatosensory receptive fields nearly always contain a well-defined area of maximal sensitivity within which the size of the stimulus is a critical feature in determining the magnitude of the response. These best areas are systematically distributed across receptive fields as a function of their location in the structure, and indicate that the resolution of stimulus location and size may be greater than expected on the basis of receptive field size alone.

Innervation of hairs in the facial skin of marsupial mammals.

The innervation of pelage, guard hairs and vibrissae hairs was studied in five species of marsupial mammals by means of electron microscopy for the first time. This study showed that members of different superfamilies in marsupial mammals displayed the same pattern of hair innervation. This also resembled the pattern seen in the placental mammals. All types of hairs had both longitudinal and transverse lanceolate nerve terminals. Pelage hairs did not have any Merkel cells. Guard hairs were very richly innervated and had free nerve endings, lanceolate nerve endings, many Merkel cells with their associated nerve endings and pilo-Ruffini nerve endings. Vibrissae hairs had free nerve endings, Merkel nerve endings and lamellated corpuscles, but pilo-Ruffini nerve endings were not seen in this investigation. Because of the profusion and variety of innervation in guard hairs of the marsupial mammals, these hairs may have a similar function to vibrissae hairs in placental mammals.