The structural organization of layer IV in the somatosensory region (SI) of mouse cerebral cortex. The description of a cortical field composed of discrete cytoarchitectonic units.

Functional architecture of the mystacial vibrissae.

Our confocal three-dimensional analyses revealed substantial differences in the innervation to vibrissal follicle-sinus complexes (FSCs) in the rat and cat. This is the first study using anti-protein gene product 9.5 (PGP9.5) immunolabeling and confocal microscopy on thick sections to examine systematically the terminal arborizations of the various FSC endings and to compare them between two species, the rat and the cat, that have similar-appearing FSCs but different exploratory behaviors, such as existence or absence of whisking. At least eight distinct endings were clearly discriminated three dimensionally in this study: 1) Merkel endings at the rete ridge collar, 2) circumferentially oriented lanceolate endings, 3) Merkel endings at the level of the ring sinus, 4) longitudinally oriented lanceolate endings, 5) club-like ringwulst endings, 6) reticular endings, 7) spiny endings, and 8) encapsulated endings. Of particular contrast, each nerve fiber that innervates Merkel cells at the level of the ring sinus in the rat usually terminates as a single, relatively small cluster of endings, whereas in the cat they terminate en passant as several large clusters of endings. Also, individual arbors of reticular endings in the rat ramify parallel to the vibrissae and distribute over wide, overlapping territories, whereas those in the cat ramify perpendicular and terminate in tightly circumscribed territories. Otherwise, the inner conical body of rat FSCs contains en passant, circumferentially oriented lanceolate endings that are lacking in the cat, whereas the cavernous sinus of the cat has en passant corpuscular endings that are lacking in the rat. Surprisingly, the one type of innervation that is the most similar in both species is a major set of simple, club-like endings, located at the attachment of the ringwulst, that had not previously been recognized as a morphologically unique type of innervation. Although the basic structure of the FSCs is similar in the rat and cat, the numerous differences in innervation suggest that these species would have different tactile capabilities and perceptions possibly related to their different vibrissa-related exploratory behaviors.

Similarities and differences in the innervation of mystacial vibrissal follicle-sinus complexes in the rat and cat: a confocal microscopic study.

This comparative study was conducted to provide a detailed, comprehensive description of the innervation to the follicle-sinus complex (F-SC) of mystacial vibrissae and to determine if interspecies variability in the innervation of the F-SCs may be related to differences in the structure or existence of barrels in the primary somatosensory (SI) cortex. Two silver techniques (Winkelmann on 100 micron-thick-frozen sections and Sevier-Munger on 8 micron-thick paraffin sections) were applied to comparable mystacial skin samples from adult hamsters, mice, rats, gerbils, rabbits, guinea pigs and cats. The basic structure and innervation of the F-SCs is the same in all species. Six distinct populations of sensory receptors are identified at consistent locations: Merkel endings in the epidermal rete ridge collar at the mouth of the follicle; circularly disposed presumptive lanceolate, Ruffini, and free nerve endings (FNE) in the inner conical body; longitudinal lanceolate endings in a dense palisade in the mesenchymal sheath at the level of the ring sinus; Merkel endings in the external root sheath at the level of the ring sinus; scattered corpuscular and FNEs (possibly lanceolate or Ruffini endings) in the cavernous sinus; and a few FNEs in the dermal papilla. In each F-SC, the first two locations are supplied by several superficial vibrissal nerves that arise from several small nerves that also innervate the skin between the vibrissae. These superficial nerves may innervate more than one F-SC. The next three locations are supplied by a single large deep vibrissal nerve that is derived directly from a row fascicle of the infraorbital nerve. Each deep nerve innervates a single F-SC. The source of the papilla innervation was not found. The ring sinus locations are consistently the most heavily innervated in all species. The number of axons in comparable deep vibrissal nerves is similar among the rodents, higher in the cat, and lower in the rabbit. Innervation of the inner conical body varies considerably, being dense in species that vigorously whisk their vibrissae (hamster, mouse, rat, and gerbil) and sparse or absent in species that minimally or never whisk (guinea pig, rabbit, and cat). Innervation to the cavernous sinus is sparse particularly in hamsters and gerbils. The innervation to the rete ridge is uniquely absent in the rabbit.(ABSTRACT TRUNCATED AT 400 WORDS)

A comparative light microscopic analysis of the sensory innervation of the mystacial pad. I. Innervation of vibrissal follicle-sinus complexes.

The present review of cutaneous sensory receptors begins with a consideration of free nerve endings (FNEs) that can be considered as sensory terminals evidencing the least structural specialization of the axon and associated cells. Using the criteria established by KRUGER et al (1981), FNEs of both A delta and C fibers can be identified on the basis of ultrastructural characteristics that include an intimate relationship between axons and the associated epithelium, the lack of a complete Schwann cell investment, the accumulation of numerous vesicles and other cytoplasmic organelles, and for A delta terminals a 1:1 relationship between axon and investing Schwann cell. Using these criteria, the so-called genital end bulbs of the human glans penis are merely a skein of FNEs based on the ultrastructural study of HALATA and MUNGER (1986).Hair follicles of most species studied to date (the exception being the rabbit and to some extent the guinea pig) are multiply innervated with lanceolate, Ruffini and FNEs. The lanceolate terminals are the rapidly adapting terminals that are numerous in guard hairs. Ruffini terminals of hairs resemble those of the periodontal ligament or joint capsules and both are remarkably similar to Golgi tendon organs in terms of ultrastructural characteristics. The key ultrastructural characteristic is the encircling of collagen bundles by axons and associated Schwann and connective tissue cells.Axons frequently enter the epidermis either to terminate as FNEs or become associated with Merkel cells in glabrous skin at the base of the papillary ridges or in clusters of Merkel cells in hairy skin in touch domes or Haarscheiben. Merkel cells have clusters of apparent secretory granules polarized toward the axon and the axon is typically a slowly adapting mechanoreceptor. The function of the granules is not known.Pacinian corpuscles are the largest of the corpuscular receptors of the dermis and are characterized by an elaborate inner core of stacks of numerous thin lamellae arranged in a bilaterally symmetrical manner. Based on the fact that the lamellae are coupled with gap junctions and the outer core lamellae isolated by numerous tight junctions, the authors have proposed that the unique ionic environment may be in part responsible for the remarkable sensitivity of Pacinian corpuscles (MUNGER and IDE, 1987).Meissner corpuscles are a typical corpuscular receptor of murine (IDE, 1976, 1977), marsupial and primate glabrous skin (MUNGER, 1971). The axons typically weave back and forth between stacks of lamellae. Simple corpuscles are common in many mammals especially in the glabrous snout skin (MUNGER, 1965, HALATA and MUNGER, 1983). The lamellae around the axon are typically arranged circumferentially around the axon.The authors have previously noted that axons in mechanoreceptors including lanceolate terminals, Pacinian and Meissner corpuscles, and muscle spindles frequently have axonal spines projecting from the elongated or Y-axis of the axon (MUNGER and IDE, 1987). Axonal spines typically are filled with filaments and lack other cytoplasmic organelles. The common presence of axonal spines provided the basis for the speculation that they may be involved in mechano-electric transduction.

The Structure and Function of Cutaneous Sensory Receptors

The innervation of rat vibrissae has been studied by light and electron microscopy. Nerve fibers enter the external root sheath and become associated with Merkel cells. Merkel cells are specialized sheath cells containing many secretory granules apposed to the neurite. These secretory granules are PAS-positive and diastase-resistant in light microscopic preparations. The neurite at the level of the Merkel cell is expanded and contains masses of mitochondria. As contrasted to nerve fibers in glabrous skin, the neurite does not penetrate deeply into the external root sheath but appears to terminate at the level of the Merkel cells. The neurite-Merkel cell complex is interpreted as representing a mechanoreceptor.

The ultrastructure and innervation of rat vibrissae.

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The ultrastructure and innervation of rat vibrissae.