Nerve fibers innervating the cranial and spinal meninges: morphology of nerve fiber terminals and their structural integration.

Pachymeninx and leptomeninx of cranial cavity and spine are considerably different in their collagenous fiber texture, cellular composition, vascularization, and innervation. The majority of meningeal nerve fibers terminate as free nerve endings whereas encapsulated and lamellated nerve terminals additionally occur in higher vertebrates including man. With respect to nerve fiber classification, arborization pattern, topography, and organization of the microenvironment at the termination site afferent and efferent nerve terminals are differentiated. Only the dura mater and the pial subcompartment of the leptomeninx possess the morphological prerequisites for neurogenic inflammation. In the current review, the results of morphological studies regarding the meningeal innervation including the sites of CSF (cerebrospinal fluid) production and absorption are discussed with emphasis on their structure-function relationships.

Subnuclear organization of the rat habenular complexes.

The habenular complexes represent phylogenetically constant structures in the diencephalon of all vertebrates. Available evidence suggests that this area is engaged in a variety of important biological functions, such as reproductive behaviors, central pain processing, nutrition, sleep-wake cycles, stress responses, and learning. Based on Nissl-stained sections, one medial nucleus and two lateral nuclei (divisions) have been widely accepted in the rat. Cytochemical, hodologic, and functional studies suggest a considerably more complex subnuclear structure. To improve our knowledge of the precise structural composition of the habenular complexes, we have systematically investigated their fine ultrastructure in the rat. Based on the detailed analysis of complete series of large, semithin sections supplemented with electron photomicrographs of selected fields, clear criteria for the delineation of five distinct subnuclei of the medial and ten subnuclei of the lateral habenular complexes were elaborated for the first time. All 15 subnuclei were reconstructed, and their dimensions were determined. A medial and lateral stria medullaris were described. Different roots of the fasciculus retroflexus were differentiated within the medial and lateral habenular complexes. The topographical relationships with respect to the adjacent habenular areas as well as to the neighboring thalamic nuclei were identified and demonstrated. The new understanding of the subnuclear organization of the habenular complexes certainly will facilitate further functional investigations. Whether the newly identified subnuclei finally will be recognized as functionally distinct awaits ongoing immunocytochemical, hodologic, and functional studies.

Topography and immunocytochemical characterization of nerve fibers in the leptomeningeal compartments of the rat. A light- and electron-microscopical study.

The localization of peptidergic, catecholaminergic, and nitroxidergic nerve fibers in the ventral leptomeningeal connective tissue compartment was studied in whole-mount preparations and serial semithin and ultrathin sections. For immunocytochemistry, whole-mount preparations of the leptomeninges and ventral brain slices with the meninges were incubated as free-floating specimens with primary antibodies against protein gene product 9.5 (PGP 9.5), substance P (SP), calcitonin gene-related peptide (CGRP), dopamine beta-hydroxylase (DbetaH), vasoactive intestinal polypeptide (VIP), neuropeptide Y (NPY), and nitric oxide synthase (NOS) using the avidin-biotin-peroxidase method. Based on the regional differences of the connective tissue organization, the leptomeninx is subdivided into the pial, trabecular, and adventitial leptomeninx. The antibody PGP 9.5 stains all unmyelinated nerve fibers in the leptomeninx. Although the highest density of nerve fibers occurs in the adventitial leptomeninx, nerve fibers, and terminals are additionally present in the trabecular and pial leptomeninx. DbetaH-, NPY-, VIP- and NOS-immunoreactive (IR) nerve fibers occur exclusively in the adventitial leptomeninx forming neuromuscular junctions. CGRP- and SP-IR nerve fibers are localized in all three leptomeningeal compartments where they terminate close to the subarachnoid space (type 1) or within the connective tissue (type 2). Due to their morphological and immunocytochemical characterization a possible chemo-, mechano- or nociceptive function is discussed in the context of pathophysiological aspects.

Topography and fine structure of proprioceptors in the hagfish, Myxine glutinosa.

Topography and ultrastructural organization of proprioceptors in the striated parietal muscle of the hagfish, Myxine glutinosa, are described. Due to the topographical location of the afferent nerve terminals four subtypes of proprioceptors are distinguished. Type 1-receptor and type 2-receptor are localized in an encapsulated and lamellated corpuscular organ located along and above the myosepta of the parietal muscle. Type 3-receptor is composed of a complex of free en plaque nerve endings within the dense collagen fibre texture at the site where myoseptum, muscle fascia and body fascia fuse to a raphe. Type 4-receptor resembles type 3 but it is localized in the straight collagen fibre bundles of the myoseptum and the body fascia. A proprioceptive function of the nerve terminals is discussed.

Lamellated receptors in the skin of the hagfish, Myxine glutinosa.

The adult hagfish, Myxine glutinosa, does not exhibit a lateral line system. The hypodermal layer of the dorsal head and body skin contains a prominent receptor system--lamellated corpuscles--arranged in a segmental pattern close to the body fascia. The topography and the structural organization of the lamellated receptors are described at the light- and electronmicroscopical levels. Spinal nerves supply the lamellated receptor organs. A mechanoreceptive function and evolutionary aspects are discussed.

Sensory nerve fiber terminals in the arachnoid granulations of non-human primates.

Myelinated and unmyelinated axons terminate within the arachnoid granulations. Serial section analysis by light and transmission electron microscopy reveals that the nerve fibers terminate at different locations or tissue compartments of the arachnoid granulation. 1. Myelinated axons ramify and terminate as free axon terminals like slowly adapting type II mechanoreceptors in the leptomeningeal connective tissue core of the granulation. 2. Myelinated and unmyelinated axons terminate in the transitional spongy zone which represents a tissue compartment for the diffusion of the cerebrospinal fluid from the subarachnoidal space to the venous sinus. This zone is composed of arachnoidal cells, dural neurothelial cells and fibrocytes. 3. Myelinated and unmyelinated axons terminate within lymphoid cell aggregation associated to the arachnoid granulation.

The anatomy and fine structure of the echidna Tachyglossus aculeatus snout with respect to its different trigeminal sensory receptors including the electroreceptors.

The gross anatomy and nerve supply of the bill of echidna (Tachyglossus aculeatus) is described in relation to its function as an outstanding sensory organ. The sensory innervation of the skin of the echidna snout was investigated by means of frontal serial sections, after decalcification of the specimens. A comprehensive light and electron microscopic description of the location and fine structure of cutaneous sensory receptors of the trigeminal system was made by this means. The encapsulated and non-encapsulated Ruffini receptors, the types of other free receptors in the connective tissue and the Merkel cell receptor do not differ morphologically from those of higher mammals, whereas the pacinian-like corpuscle shows a unique organization of its outer core. This is composed of large perineural cells containing a unique reticulum of parallel-orientated endoplasmic membranes. Lamellated corpuscles, seen in isolation or in association with push rods, are numerous in the snout and in the tip of the tongue of echidna. Push rod receptor organs occur in the hairless skin of the bill with a very dense array at its rostral end and in the pseudopalatal ridges. Gland duct receptors are restricted to the skin adjacent to the nostrils and the mouth opening, including the pseudopalatal plates. Only about one quarter of the total number of 400 seromucous glands receive a sensory innervation of their intraepidermal duct segment. Within each innervated gland two types of receptor terminals are identified. The distributions of the different receptor types are mapped for different regions of the skin, the mucous membrane of the nasal and oral vestibule and the tip of the tongue. The fine structure of nerve terminals is discussed from a comparative anatomical point of view, and some speculations are made about possible transduction processes that underlie the known electrophysiological properties. The sensory organs such as the "push rod" and "gland duct receptor", and most of their sensory terminals, are less differentiated in echidna snout than in the platypus (Ornithorhynchus anatinus) bill.

Neuropeptide Y- and substance P-like immunoreactive nerve fibers in the rat dura mater encephali.

Density and pattern of nerve fibers with neuropeptide Y-like immunoreactivity (NPY-LI) and substance P-like immunoreactivity (SP-LI) in the rat dura mater encephali were investigated by light and electron microscopy using whole-mount preparations. NPY-LI fibers are observed throughout the encephalic dura mater. A remarkable net of NPY-LI nerve fibers is located in the walls of the sagittal and transverse sinuses. Beyond that NPY-LI network, distinct NPY-LI nerve fibers or plexus occur in the rostral falx, parietal dura mater of the olfactory bulb, supratentorial dura mater, parietal dura mater of the cerebellum, tentorium cerebelli and the ventral dura mater. Electron microscopic studies reveal that NPY-LI is exclusively located in unmyelinated axons of small and large nerve fiber bundles, with or without a perineural sheath. Immunopositive C-fibers are predominantly associated with the vascular bed. SP-LI nerve fibers have a moderate and more uniform distribution in the encephalic dura mater. A distinct plexus of SP-LI fibers follows the branches of the middle meningeal artery and the adjacent dura mater. SP-LI fibers are most prominent in the parietal dura mater of the cerebellum. Fine beaded SP-LI fibers, arising from larger SP-LI fiber bundles, are observed in close association to the capillary bed. SP-LI axons are all unmyelinated. They are found in larger nerve fiber bundles with a perineural sheath or in Schwann cells lacking any perineural sheath. The function of NPY-LI and SP-LI nerve fibers in the rat dura mater is discussed in relation to their topography, density and termination.

The fine structure of ampullary and tuberous electroreceptors in the South American blind catfish Pseudocetopsis spec.

Two types of electroreceptors, the ampullary and the tuberous electroreceptor (silurid knollenorgan) in the epidermis of the catfish, Pseudocetopsis spec., were investigated with semithin and ultrathin serial sections. The ampullary organ contains one or two sensory cells which are embedded in supporting cells at the base of open epithelial canals. They bear some slender microvilli on their apical surface and form several synaptic bars. The afferent myelinated nerve fiber arborizes in the connective tissue papilla and looses its myelin sheath about 30 micron below the supporting cell layer. A second thin myelinated axon occur up to the supporting cell layer. The tuberous electroreceptor organ contains one large receptor cell. Most of the cell body is exposed to the lumen of a specialized proximal canal segment and is closely covered with microvilli. A single myelinated axon looses its myelin sheath within the supporting cell layer about 1 micron before terminating as a flat calyx at the base of the sensory cell. A functional significance of the two types of receptors will be discussed.

Nerve fibres and their terminals of the dura mater encephali of the rat.

The dura mater encephali of the rat is richly supplied by myelinated (A-axons) and unmyelinated (C-axons) nerve fibres. For the supratentorial part the main nerve supply stems from all three branches of the trigeminal nerve. Finally, 250 myelinated and 800 unmyelinated nerve fibres innervate one side of the supratentorial part. The vascular bed of the dura mater exhibits long postcapillary venules up to 200 micron in length with segments of endothelial fenestration. Lymphatic vessels occur within the dura mater. They leave the cranial cavity through the openings of the cribriform plate, rostral to the bulla tympani together with the transverse sinus, and the middle meningeal artery. The perineural sheath builds up a tube-like net containing the A- and C-axons. It is spacious in the parietal dura mater and dense at the sagittal sinus along its extension from rostral to caudal and at the confluence of sinuses. Terminals of both the A- and C-axons are of the unencapsulated type. Unencapsulated Ruffini-like receptors stemming from A-axons are found in the dural connective tissue at sites where superficial cerebral veins enter the sagittal sinus and at the confluence of sinuses. The terminations of single A-axons together with C-fibre bundles mix up in their final course in one Schwann cell to build up multiaxonal units or terminations (up to 15 axonal profiles). A morphological differentiation is made due to the topography of these terminations; firstly, in different segments of the vascular bed: postcapillary venule, venule, the sinus wall, lymphatic vessel wall, and secondly, within the dura mater: inner periosteal layer, collagenous fibre bundles of the meningeal layer and at the mesothelial cell layer of the subdural space.

Degeneration patterns of postganglionic fibers following sympathectomy.

In cats the time course of degeneration following lumbal sympathectomy was studied in the ramus communicans griseus (rcg) and in the nerves to the triceps surae muscle using light and electron microscopic methods. The left lumbar sympathetic trunk including its rami communicantes was removed from L2 to S1 using a lateral approach. The animals were sacrificed between 2 and 48 days after the sympathectomy. Tissue samples were taken (a) one cm proximal to the entrance of the rcg into the spinal nerve, and (b) one cm proximal to the entrance of the nerve into the muscle belly. In the rcg signs of degeneration can already be recognized in the myelinated as well as in the unmyelinated axons 48 h after sympathectomy. The degenerative processes in the axons reach their peak activity at about 4 days p.o. They end a week later. Signs of the reactions of the Schwann cells and of the endoneural cells can first be seen 2 days p.o. They are most pronounced around the 8th day p.o., and last at least up to the third week. Thereafter the cicatrization processes settled to a rather steady state (total observation period 7 weeks). In the muscle nerves the first signs of an axonal degeneration of the sympathetic fibers can be recognized 4 days after surgery. The signs of axonal degeneration are most striking about 8 days p.o. They have more or less disappeared another week later. The reactions of the Schwann cells also start on the fourth day but outlast the degenerative processes by some 8 days. Thus the degenerative and reactive processes in the rcg precede those in the muscle nerves by 2 days early after surgery and by 6 days 3 weeks later. Seven weeks after surgery, fragments of folded basement lamella and Remak bundles with condensed cytoplasm and numerous flat processes are persisting signs of the degeneration. In addition to the differences in time course between the proximal and the distal site of observation, it was also noted that both the axonal degeneration and the reactions of the Schwann cells are more pronounced in the rcg than in the muscle nerve. For example there was abundant mitotic activity in the central endoneural and Schwann cells whereas we could not detect such activity in the periphery.(ABSTRACT TRUNCATED AT 400 WORDS)

Sensory innervation of the Achilles tendon by group III and IV afferent fibers.

In sympathectomized cats the innervation of the Achilles tendon by fine afferent nerve fibers was studied with semithin and ultrathin sections. Several different types of sensory endings of group III and group IV nerve fibers were identified. Of the five different types of endings in the group III range (T III endings), two are located within vessel walls. One of them ends in the circumference of the venous vessels (T III/VV). Its lanceolate terminals have characteristic receptor areas at their edges. The second type ends in the adventitia of lymphatic vessels (T III/LV). Its receptive areas are scattered along their terminal course. Two further group III endings ramify within the connective tissue compartments of the vessel-nerve-fascicles of the peritenonium externum and internum. One type is tightly surrounded by collagen fibrils (T III/PTic); the other terminates between the collagen fiber bundles (T III/PTgc). The latter arrangement recalls the ultrastructural relation between nerve terminals and collagen tissue in Golgi tendon organs. The fifth type innervates the endoneural connective tissue of small nerve fiber bundles (T III/EN). At least some of them come into close contact with bundles of collagen fibers which penetrate the perineural sheath to terminate within the endoneurium. The endings of group IV afferents (T IV endings) show a striking topographic relationship to the blood and lymphatic vessels of all connective tissue compartments of the Achilles tendon. They form penicillate endings which may contain granulated vesicles. In any event, they can easily be discriminated from the T III endings in the vessel walls. In close neighborhood to Remak bundles, a cell has been regularly found which fulfilled all ultrastructural criteria for mast cells. But this cell is not a mast cell proper because it is surrounded by a basal lamina (pseudo mast cell).

Sequestration of neuraminidase-treated erythrocytes. Studies on its topographic, morphologic and immunologic aspects.

Scintigraphic experiments and radioactivity measurements of tissues have shown that the radioactivity of 51Cr-labelled and neuraminidase-treated rabbit erythrocytes is rapidly accumulated in liver and spleen. Sequestration of these erythrocytes by liver and spleen was demonstrated by light and electron microscopy of theses tissues after perfusion of the rabbits with solutions for tissue fixation. In liver the phagocytic activity of Kupffer cells was increased after injection of desialylated erythrocytes, while in spleen a significantly enhanced number of erythrocytes was found attached to the sinusoidal walls and in the reticulum of the red pulp. It was shown by scanning electron microscopy that neuraminidase-treatment did not influence the shape of erythrocytes. Desialylated and 51Cr-labelled erythrocytes from the cow are rapidly cleared from the blood-stream with a half-life time of about 3 h. It was shown in an in-vitro test that they adsorb to surviving slices from liver and spleen derived from the same animal. The amount of radioactivity adsorbed is appreciably enhanced in the presence of homologous serum when compared with buffer only. Human neuraminidase-treated erythrocytes are agglutinated in the direct and especially in the indirect Coombs-tests. The involvement of T-antigen in this phenomenon was largely excluded. The in vitro experiments and antibody consumption tests suggest that immunoglobulins (IgG) and complement from serum may be involved in recognition and sequestration of desialylated erythrocytes by macrophages in vivo.

The ultrastructure of taste and touch receptors of the frog's taste organ.

The taste buds from fungiform papillae and the hard palate of frogs were investigated with the scanning and transmission electron microscopes. An immature pre-taste cell and a mature taste cell can be differentiated. Only the mature taste cell exhibits synaptic contact with the afferent taste fibre. Glandular and satellite supporting cells envelop the thin apical processes of the sensory cells. At the base of the taste disc up to 10 Merkel cells form a complex with nerve endings. There are two types of myelinated fibres, large and small. The small fibre innervates the taste cells, the thicker nerve fibre the Merkel cells. The occurrence of two types of receptors explains physiological results.

[New morphologic principles of the physiology of smell and taste]. / Neue morphologische Grundlagen zur Physiologie des Riechens und Schmeckens.

New results as revealed by scanning and transmission electron microscopy have given us further knowledge about the structure of the olfactory region of vertebrates. With comparative studies we are now able to discuss the functional relationship of this region. In all vertebrates the olfactory cell is a primary sensory cell. The apical segment of the olfactory cell with its olfactory vesicle is involved in the formation of the olfactory border. As a rule of the receptor possesses cilia or cilia-like processes. These are absent in the olfactory receptor of the shark, the microvillus receptor of the fish and the olfactory cell of Jabonsons organ of amphibians, reptiles and mammals. The odorous substances in the fish are brought to the receptor membrane by the water flow. In air breathing vertebrates a terminal film is present. This film is a product of secretion from the Bowmans glands. Gasous odorous substances must first be dissolved in the terminal film and penetrate it before reaching the receptor membrane. The cilia-like olfactory process of the fish in the proximal segment is not essentially different from the kinocilia of the supporting cell, except that they are shorter. In contrast the olfactory cell of air-breathing vertebrates form cilia-like processes with a short cilia-like proximal segment and a long and very thin distal end piece. In the amphibians and sauropsidians the end pieces can have a length of up to 150 mu and up to 80 mu in mammals. The olfactory vesicles with its processes undergo continuous regeneration. The olfactory epithelium of man show the same structural formation as observed in other mammals. Regressive changes in the adult can lead to a reduction in the number of sensory cells and also to a flattening of the epithelium. Morphological criteria for regenerative processes in the sensory cell structures are present. A specialized olfactory cell type has been found in some teleosts. This cell is characterized by a small pit below the olfactory border in which the cilia of the olfactory cell are redrawn. There is some evidence that this olfactory cell type may be compared with the olfactory cells in the parafollicular tubes of lamprey. The so called rod-shaped receptor in the olfactory mucosa of fishes has no axon and is therefore no olfactory cell. The same kind of cell is also present in the olfactory mucosa of air-breathing animals. We classify this cell as brush cell.(ABSTRACT TRUNCATED AT 400 WORDS)