Morphology, distribution, and density of sensory receptors in the glabrous skin of the cat rhinarium

The sensory organization of the cat rhinarium has been investigated. Individual rete pegs were found to contain a triad of receptors comprising free nerve endings ascending in the peg to terminate in close proximity to the skin surface, a basally situated layer of Merkel corpuscles, and an abundance of encapsulated receptors lying at the base and to one side of the rete peg. Neither the Merkel corpuscles nor the encapsulated receptors were evenly distributed. Merkel corpuscles were more abundant dorsally; ventrally they were fewer and asymmetrically arranged within individual rete pegs. The encapsulated corpuscles were more evenly distributed, but dorsally they were consistently present as encapsulated clusters of up to nine corpuscles.     

Distribution of the intensity-characteristic parameters of cat rapidly adapting mechanoreceptive fibers

Modeling population responses of nerve fibers requires statistical characterization of fiber-response properties. The rate/intensity characteristics of cat rapidly adapting (RA) fibers were fitted by four-parameter, piece-wise linear functions using nonlinear regression (n = 14; R2 > 0.958). The parameters were tested against the null hypothesis that they are log normally distributed. The test fail to reject this hypothesis (Kolmogorov-Smirnov p>0.477). However, a significant statistical difference was found between the specific lognormal distributions obtained from monkey (Johnson, J Neurophysiol 37: 48-72, 1974) and cat for all four parameters (Kolmogorov-Smirnov, p<0.0075, p<0.05, p<0.0001, p<0.00007). Although the stimulus contactor size was not the same in monkey and cat studies, the differences between monkey and cat fibers are attributed to anatomical differences in the glabrous sin of both species. Modeling studies suggest that the absolute firing thresholds of RA fibers have a right-skewed distribution because of the anatomical constraints present in both species' skin. Meissner corpuscles, which are the sensory end-organs of RA fibers, are likely to be found deeper in the skin within dermal papilla, therefore, the thresholds can be elevated. However, the thresholds are bounded at lower end, probably due to the epidermal junction that acts as a superficial mechanical barrier for these corpuscles.     

Localization of met-enkephalin like immunoreactivity in the glabrous skin of the cat rhinarium

The presence of met-enkephalin like immunoreactivity (MEL IR) was investigated immunohisto-chemically in the glabrous skin of the cat rhinarium using the peroxidase-antiperoxidase Sternberger's method. Neither sensory corpuscles nor nerve bundles show MEL IR. MEL IR was found in the epidermal Merkel cells, as well as in Langerhans cells and/or melanocytes. In dermal papillae the reaction results positive in a number of cells which could be identified as Schwann or pigmentary cells.     

The skin of primates. XXXIX. The skin of the white-browed capuchin (Cebus albifrons)

The skin of the white-browed capuchin (Cebus albifrons), although basically similar to that of the squirrel monkey (Saimiri sciureus), contains several outstanding peculiarities: (1) both the epidermis and dermis of the general body surface are devoid of melanotic melanocytes; (2) the skin of the prehensile tail has no modified, glabrous friction surface; (3) the prehensile surface of the tail has no specialized nerve end-organs; (4) cholinesterase-positive, papillary nerve end-organs rest beneath the epidermal ridges of volar skin; and (5) both the clear cells and dark cells of the eccrine glands contain glycogen but neither shows phosphorylase activity.     

A fine structural localization of the non-specific cholinesterase activity in glomerular nerve formations (endings)

Snout glabrous skin (rhinarium) of the cat is innervated not only by typical simple lamellar corpuscles but also glomerular formations. In contrast to simple lamellar corpuscles, glomerular nerve formations are located away the dermal papillae. In cross sections, glomerular nerve formation consists of several axonal profiles enveloped by 1-2 cytoplasmic lamellae of Schwann cells. The space among them is filled by collagenous microfibrils and the basal lamina-like material. Capsule was composed from fibroblast-like cells without definite basal lamina. An electron-dense reaction product due to non-specific cholinesterase activity was associated with Schwann cells and their processes surrounding unmyelinated terminal portion of the sensory axons. Abundant reaction product was bound to the collagenous microfibrils and was deposited in extracellular matrix between Schwann cell processes. These results are further evidence for the presence of the non-specific cholinesterase molecules as integral component of the extracellular matrix in sensory corpuscles. On the basis of histochemical study two possible explanation are considered for functional involving of this enzyme in sensory nerve formations.     

Mesenteric and tactile Pacinian corpuscles are anatomically and physiologically comparable

Pacinian corpuscles (PCs) in cat mesentery have been studied extensively to help determine the structural and functional bases of tactile mechanotransduction. Although we, like many other investigators, have found that the mesenteric receptors are anatomically very similar to those found in mammalian skin, few physiological characteristics of the mesenteric PCs and those of the skin have been compared. Action-potential rate-amplitude and frequency characteristics (10 Hz–1 KHz), as well as interval (IH) and peri-stimulus-time (PSTH) histograms in response to sinusoidal displacements were obtained from nerve fibers innervating mesenteric PCs and from PC fibers innervating cat glabrous skin. The intensity characteristics obtained on both preparations showed similar response profiles, including equal slopes for low stimulus intensities (approximately 10, with impulse ratios/20 dB displacement) and one and two impulse/cycle entrainment. The frequency characteristics of both groups were U-shaped with similar low-frequency slopes (?12.5 dB/octave) and bandwidths (Q_3dB = 1.4). The best frequency for both the tactile PCs' and mesenteric PCs was 250 Hz, which is in the expected range. The IHs showed entrainment and the PSTHs showed neither transient responses nor adaptation to steady-state sinusoidal stimuli. The functional similarity between mesenteric PCs' nerve responses and those of tactile PC afferents, as well as the receptors’ anatomical similarity, lead us to suggest that the mesenteric PC can act as a model for those in the skin. Furthermore, since the frequency characteristics of the two PC types are similar, it is concluded that the skin, while attenuating stimulus intensity, does not impart temporal filtering of vibratory stimuli.     

The skin of primates. XXX. The skin of the woolly monkey. (Lagothrix lagotricha)

The anatomical and histochemical features of the skin of the woolly monkey are intermediate between those of the Cercopithecoidea and the Pithecoidea. The animal has a prehensile tail, the glabrous, friction surface of which is similar to that of the fingers. The epidermis is heavily pigmented. The dermal vascularization is relatively well-developed and similar to that of the skin of the Cercopithecoidea. Hair follicles grow in groups of 4 to 15, as in the skin of the Pithecoidea. In the hairy skin, eccrine sweat glands occur only in the tail and genitalia. The woolly monkey, like the green monkey, possesses only acetylcholinesterase-containing nerve fibers around its eccrine sweat glands.     

人胎儿真皮乳头和皮纹发生的研究

本文用扫描电镜法研究了入胎儿的皮纹发生过程,包括初级真皮嵴、次级真皮嵴、真皮乳头和表皮隆线的发生。为研究人皮纹的发生和皮肤的异常提供了皮肤正常发育的形态学依据。共观察111例从第6周到第9个月胎儿的皮纹区皮肤,表明第3个月末胎儿开始形成初级真皮嵴,以后逐渐加深,至第16周嵴的顶端中央产生纵沟形成两条平行的次级真皮嵴;自19周后,次级真皮嵴局部隆起,由波浪形逐渐形成乳头。至30周乳头呈犬牙状。表皮隆线于第4—5月形成,随真皮乳头的增高而渐趋明显。至第6个月,全部皮纹图样已可辨认。本文还讨论了真皮乳头发生的过程。     

Localization of Merkel cells in the monkey skin: an anatomical model

The Merkel cell-neurite complex is considered to be one class of mechanoreceptors in the skin. Merkel cells are innervated by slowly adapting type I (SAI) tactile nerve fibers. In this paper, the detailed distribution of Merkel cells is studied by immunohistochemical labeling of the monkey (Macaca fascicularis) digital glabrous skin. Specific morphometric variables (density of intermediate epidermal ridges and Merkel cells, distance between skin surface and ridge tips and bases, maximum and average cell counts per ridge, distance between cells and ridges) were measured by a combination of light/fluorescence microscopy and computer-image analysis. The morphometric results were similar for each digit of the monkey's hand. Next, the anatomical data were used to form a three-dimensional reconstruction of the Merkel-cell distribution in the fingertip skin. A patch of the distal-pad surface was then computationally flattened to obtain the two-dimensional distribution of Merkel cells. Based on previous anatomical and physiological data, SAI fibers were simulated to innervate clusters of Merkel cells in the distal-pad surface. On average, 28 cells were innervated by a single fiber. The resulting anatomical model may be used to estimate the population response of SAI fibers by incorporating spike generation.     

Time Course and Characteristics of the Capacity of Sensory Nerves to Reinnervate Skin Territories outside Their Normal Innervation Zones

Factors involved in the outcome of regeneration of the saphenous nerve after a cut or crush lesion were studied in adult rats with electrophysiological recordings of low-threshold mechanoreceptor activity and plasma extravasation of Evans blue after electrical nerve stimulation that activated C fibers.

In the first series of experiments, saphenous and sciatic nerve section was combined with anastomosis of the transected proximal end of the saphenous nerve to the distal end of the cut tibial nerve. Regeneration of saphenous nerve fibers involved in plasma extravasation and low-threshold mechanoreceptor activity in the glabrous skin was observed 13 weeks after nerve anastomosis. Substance P-, calcitonin gene-related peptide-, and protein gene product 9.5 (PGP-9.5)-immunoreactive (IR) thin epidermal and dermal nerve endings, as well as coarse dermal PGP-9.5-IR nerve fibers and Meissner corpuscles and Merkel cell-neurite-like complexes, were observed in the reinnervated glabrous skin at this time.

In a second series of experiments, the time course of the regeneration of saphenous nerve axons to the permanently sciatic-nerve-denervated foot sole was examined. Saphenous-nerve-induced plasma extravasation and low-threshold mechanoreceptor activity in the saphenous nerve were found in the normal saphenous nerve territory 2, 3, 4, and 6 weeks after sciatic nerve cut combined with saphenous nerve crush in the left hindlimb. Saphenous-nerve-induced plasma extravasation was also present in the glabrous skin normally innervated by the sciatic nerve 3, 4, and 6 weeks after the sciatic cut/saphenous crush lesion. However, no low-threshold mechanoreceptor activity was detected in the saphenous nerve when the glabrous skin area was stimulated.

In a third series of experiments, the fate of the expansion of the saphenous nerve territory after saphenous nerve crush was examined when the crushed sciatic nerve had been allowed to regenerate. Nerve fibers involved in plasma extravasation were observed in the glabrous skin of the hindpaw after saphenous nerve, as well as after tibial nerve, C-fiber stimulation 3, 12, and 43 weeks after the saphenous crush/sciatic crush lesion.

Low-threshold mechanoreceptors from the regenerated saphenous nerve, which primarily innervates hairy skin, seem to be functional in the glabrous skin if the axons are guided by the transected tibial nerve by anastomosis. Furthermore, the results indicate that fibers from the regenerating saphenous nerve that have extended into denervated glabrous skin areas can exist even if sciatic nerve axons are allowed to grow back to their original territory.     

Calcitonin gene-related peptide (CGRP) immunoreactivity in the neuroendocrine Merkel cells and nerve fibres of pig and human skin

The presence of calcitonin gene-related peptide (CGRP) in the skin of pig snout and human fingertip was investigated using immunohistochemical techniques. CGRP immunoreactivity was found in Merkel cells and nerve fibres of both species. In pig snout skin, Merkel cells containing CGRP were seen forming clusters at the tips of rete ridge epidermis and in the external root sheath of sinus hair follicles (vibrissae). Human Merkel cells immunostained for CGRP were found isolated or forming small groups in the basal layer of glandular epidermal ridges. In all cases, immunoreactivity was more intense on the side of the Merkel cell facing the associated nerve terminal (which was never positive for CGRP). This part of the Merkel cell has the greatest density of dense-cored granules, suggesting that CGRP must be stored in these granules. Nerve bundles containing CGRP-immunoreactive fibres were found at dermal and hypodermal level, and blood vessels were often surrounded by CGRP nerve fibres. In pig snout skin some nerve fibres containing CGRP penetrated the epidermis and terminated as free endings, and in the human fingertip a small number of CGRP-immunoreactive nerve fibres were seen in Meissner's corpuscles.     

Mesenteric and tactile Pacinian corpuscles are anatomically and physiologically comparable

Pacinian corpuscles (PCs) in cat mesentery have been studied extensively to help determine the structural and functional bases of tactile mechanotransduction. Although we, like many other investigators, have found that the mesenteric receptors are anatomically very similar to those found in mammalian skin, few physiological characteristics of the mesenteric PCs and those of the skin have been compared. Action-potential rate-amplitude and frequency characteristics (10 Hz-1 KHz), as well as interval (IH) and peri-stimulus-time (PSTH) histograms in response to sinusoidal displacements were obtained from nerve fibers innervating mesenteric PCs and from PC fibers innervating cat glabrous skin. The intensity characteristics obtained on both preparations showed similar response profiles, including equal slopes for low stimulus intensities (approximately 10, with impulse ratios/20 dB displacement) and one and two impulse/cycle entrainment. The frequency characteristics of both groups were U-shaped with similar low-frequency slopes (-12.5 dB/octave) and bandwidths (Q(3dB) = 1.4). The best frequency for both the tactile PCs' and mesenteric PCs was 250 Hz, which is in the expected range. The IHs showed entrainment and the PSTHs showed neither transient responses nor adaptation to steady-state sinusoidal stimuli. The functional similarity between mesenteric PCs' nerve responses and those of tactile PC afferents, as well as the receptors' anatomical similarity, lead us to suggest that the mesenteric PC can act as a model for those in the skin. Furthermore, since the frequency characteristics of the two PC types are similar, it is concluded that the skin, while attenuating stimulus intensity, does not impart temporal filtering of vibratory stimuli.     

Calcitonin gene-related peptide (CGRP) immunoreactivity in the neuroendocrine Merkel cells and nerve fibres of pig and human skin

Summary The presence of calcitonin gene-related peptide (CGRP) in the skin of pig snout and human fingertip was investigated using immunohistochemical techniques. CGRP immunoreactivity was found in Merkel cells and nerve fibres of both species. In pig snout skin, Merkel cells containing CGRP were seen forming clusters at the tips of rete ridge epidermis and in the external root sheath of sinus hair follicles (vibrissae). Human Merkel cells immunostained for CGRP were found isolated or forming small groups in the basal layer of glandular epidermal ridges. In all cases, immunoreactivity was more intense on the side of the Merkel cell facing the associated nerve terminal (which was never positive for CGRP). This part of the Merkel cell has the greatest density of dense-cored granules, suggesting that CGRP must be stored in these granules. Nerve, bundles containing CGRP-immunoreactive fibres were found at dermal and hypodermal level, and blood vessels were often surrounded by CGRP nerve fibres. In pig snout skin some nerve fibres containing CGRP penetrated the epidermis and terminated as free endings, and in the human fingertip a small number of CGRP-immunoreactive nerve fibres were seen in Meissner's corpuscles.     

Dendritic cell migrations involving the pilosebaceous unit in the development of murine skin

Over the lint week of postnatal life, dermal dendritic cells stream upwards to invade the epidermis of the mouse tail and back skin. Their migrations seem associated with the development of distinct types of epidermal physiology:ortho- and parakeratosis. Changes from neonatal epidermal morphology occur at similar times in both back and tail skin. The hairv mouse back skin is alwavs orthokeratotic, but the initially orthokeratotic tail epidermis later becomes parakeratolir in the scale regions, remaining orthokeratotic in areas of hair production.
Dermal cells studied were adenosine triphosphatase (ATPase)-, non-specific esterase (NSE)-, naphthvl AS-D chloroacetate-, and dihydroxyphenvlalanine (dopa)-positive dendritic cells. The results are discussed in connection with hair growth and glabrous epidermal kcratinization. Dendritic cell regulation of epidermal physiology involving the dermis and pilosebaceous unit is discussed in relation to reviewed work on mesenchymal-epithelial interactions in animal and human skin.     

Zu den Nervenendigungen (Merkeische Endigungen) in der haarlosen Nasenhaut der Katze

Zusammenfassung Die Nervenendigungen in der Epidermis der haarlosen Haut der Katzennase wurden licht- und elektronenmikroskopisch untersucht. Im Stratum germinativum werden Merkeische Endigungen gefunden. Sie bestehen aus spezialisierten Zellen und je einer anliegenden verdickten Nervenfaser. Diese Rezeptoren werden in zwei Formen beobachtet. Einzelne Zell-Neurit-Komplexe liegen in der Basalschicht der Epidermis über den bindegewebigen Papillen und Gruppen solcher Komplexe im Stratum germinativum der Epidermalleisten (Epithelialzapfen). Die spezialisierten Zellen sind mit den umliegenden Epidermiszellen durch Desmosomen verbunden.Freie Nervenendigungen konnten in der Epidermis der haarlosen Nasenhaut nicht beobachtet werden.

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Nerve endings (Merkel's corpuscles) in hairless skin of the nose of cat

Summary The nerve endings in the epidermis of the hairless skin of the nose of the cat have been studied by means of light and electron microscopy. In the stratum germinativum there are Merkel's corpuscles. They consist of specialized cells with a thickened nerve fibre adjacent to each one of them. These receptors have been observed in two forms. Single cell-neurit-complexes are lying in the basal layer of the epidermis above the connective tissue papillae, and groups of these complexes are situated in the interpapillary pegs of the epidermis. The specialized cells are connected by desmosomes with the neighbouring cells of the epidermis. There are no free nerve endings in the hairless skin of the nose.

     

Localization of the lamellated corpuscle neurons of the cat large intestine mesentery

Electrical activity and ultrastructure of lamellated corpuscles (Pacinian corpuscles) of the cat large intestine mesentery were studied 10 days after uni- and bilateral extirpation of the lumbar spinal ganglia. Terminal degenerated in all the receptors, and there were no electrical reactions at bilateral extripation of ganglia L3-L4, they were preserved in some receptors at bilateral extirpation of ganglia L2-L3 and L4-L5 and at unilateral extirpation of ganglia L3-L4. Structure of the receptors and electrical reactions were preserved in all the lamellated corpuscles at bilateral extirpation at the level of L1-L2 and L5-L6. After enucleation of the corpuscles from the large intestine mesentery, in some neurons of ganglia L3-L4, chromatolysis was observed. After the caudal mesenteric nerve was sectioned, there were no electrical reactions in the lamellated corpuscles of the large intestine mesentery. Hence, the sensitive neurons-theri peripheral processes participate in the formation of the lamellated corpuscles of the cat large intestine mesentery-are situated in the lumbar ganglia L3 and L4.     

Embryonic dermal condensation and adult dermal papilla induce hair follicles in adult glabrous epidermis through different mechanisms

Hair induction in the adult glabrous epidermis by the embryonic dermis was compared with that by the adult dermis. Recombinant skin, composed of the adult sole epidermis and the embryonic dermis containing dermal condensations (DC), was transplanted onto the back of nude mice. The epidermis of transplants formed hairs. Histology on the induction process demonstrated the formation of placode-like tissues, indicating that the transplant produces hair follicles through a mechanism similar to that underlying hair follicle development in the embryonic skin. An isolated adult rat sole skin piece, inserted with either an aggregate of cultured dermal papilla (DP) cells or an intact DP between its epidermis and dermis, was similarly transplanted. The transplant produced hair follicles. Histology showed that the epidermis in both cases surrounded the aggregates of DP cells. The epidermis never formed placode-like tissues. Thus, it was concluded that the adult epidermal cells recapitulate the embryonic process of hair follicle development when exposed to DC, whereas they get directly into the anagen of the hair cycle when exposed to DP. The expression pattern of Edar and Shh genes, and P-cadherin protein during the hair follicle development in the two types of transplants supported the above conclusion.     

Development of dermal ridges in the volar skin of fetal pigtailed macaques (Macaca nemestrina)

Development of the dermal ridges in volar skin was investigated in 28 pigtailed macaque (Macaca nemestrina) fetuses of known gestational age, ranging from 51 days postconception to newborn. Histology, scanning electron microscopy, and staining of the abraded dermal surface were used in the study. Morphological features of the dermal-epidermal system and their changes with advancing age are described. Chronology was established for stages in the development of the volar skin, i.e., the differentiation of the primary and secondary epidermal ridges (PER and SER) at the undersurface of the epidermis corresponding to the formation of primary and secondary dermal ridges (PDR and SDR) and the development of the dermal papillae. PDRs were first seen at 55 gestational days and SDRs at 93 days. Differentiation of sweat ducts occurred over the period between 60 and 119 gestational days. A regional sequence of differentiation starting with the digital apices of the hand and ending in the calcar area and the phalanges of the foot was documented. Generally, morphogenesis in the macaque was accelerated relative to that in the human fetus by approximately 3 weeks.     

Identification of murine type I keratin 9 (73 kDa) and its immunolocalization in neonatal and adult mouse foot sole epidermis

The foot sole epidermis of the fore and hind feet of the adult mouse contains an acidic (type I) mRNA-encoded 73-kDa keratin polypeptide which cannot be detected in any other skin site of the mouse integument. Western blot analysis using an antibody specific for the 64-kDa keratin 9 of human and bovine callus-forming epidermis [A. C. Knapp et al. (1986) J. Cell Biol. 103, 657-667] demonstrates that the 73-kDa keratin represents the murine analog of keratin 9 of man and cow. Concomitant investigations in two related rodent species indicate that the size of this keratin varies more among species than that of any other orthologous keratin. Histological examination of adult mouse foot sole skin reveals an extremely thick and undulated epidermis covering the apical portion of the six footpads, whereas the epidermal-dermal junction of the lateral walls of these nodular protuberances as well as that of the remainder of the foot sole skin is essentially flat. If sections of adult foot sole skin are investigated by indirect immunofluorescence with the keratin 9-specific antibody, intense cytoplasmic staining is restricted to the apical rete pegs of the footpad epidermis in which virtually all suprabasal cells express keratin 9. However, we also observed keratin 9-negative cell columns ascending straight above the tips of the dermal papillae and separating the keratin 9-positive rete pegs from each other. At the transition from the strongly undulated apical epidermis to the flat epidermis of the lateral walls of the footpads, keratin 9-positive cells loose their coherence and gradually disappear toward the inter-footpad epidermis. This intimate relationship between the morphogenesis of epidermal ridges and inter-ridges and the expression of keratin 9 is also visible in foot sole epidermis of neonatal mice. Here we observed the appearance of keratin 9-positive suprabasal cells concomitant with the onset of pronounced folding of the apical footpad epidermis by about Day 3 after birth. Our findings confirm the view that the expression of keratin 9 is characteristic of a highly specialized pathway of epidermal differentiation. We propose a hypothesis for keratin expression in skin sites which are subject to pronounced mechanical wear and tear.     

Adaptive differentiations of the skin of the head in a subterranean rodent,Spalax ehrenbergi

The skin of macroscopically distinct regions (hairy skin, vibrissal fields, buccal ridge, and rhinarium) of the head of the blind mole-rat, Spalax ehrenbergi, was studied by routine histological methods. Few guard and several soft vellus hairs are organized into tufts that grow from a group of hair follicles localized in an invaginated compound cavity. We suggest that this hair arrangement may be a burrowing adaptation to match frictional resistance. The follicles and the compound cavity possess either well developed complex striated musculature or errector pili muscles. There are no structural specializations (sweat glands, glomus bodies) to enhance thermo-regulatory (heat dissipative) capacities in the hairy skin of the head. Vibrissae penetrate the epidermal surface as single hairs. They are microscopically normally developed are arranged in vibrissal fields according to a basal mammalian pattern. Most of them are, however, relatively short and inconspicuous. The mystical vibrissal field is horizontally divided by a prominent buccal ridge which is probably involved in bulldozing. The hairs in the ridge leave the compound cavity singularly. The follicles of guard hairs and bristles are equipped with well developed pilo-Ruffini complexes indicating that the buccal ridge may serve also as a tactile organ. The glabrous skin of the rhinarium has a highly interdigitated dermal-epidermal interface. The dermal papillae possess simple lamellated and/or simple Meissner's corpuscles and few Merkel cell-axon-complexes indicating that the skin of the rhinarium may be particularly sensitive to perception of vibrations. J Morphol 233:53–66, 1997. © 1997 Wiley-Liss, Inc.