Cerebrospinal fluid-contacting neurons in the regenerating spinal cord of lizards and amphibians are likely mechanoreceptors

During spinal cord (SC) regeneration in the tail of amphibians and lizards, small neurons in contact with the central canal and cerebrospinal fluid (CSF) are formed. The present review summarizes previous and recent studies that have characterized most of these neurons as cerebrospinal fluid-contacting neurons (CSFCNs), especially in the regenerating caudal SC of lizards. CSFCNs form tufts of stereocilia immersed in the CSF, secrete exosomes, and are often in contact with a secreted protein-rod indicated as Reissner fiber. Ultrastructural, autoradiographic, immunohistochemical, and behavioral studies strongly indicate that most of these cells are mechanoreceptors that differentiate from ependymal cells within 20–30 days after SC amputation. Numerous CSFCNs are gamma amino-butyric acid (GABA)-ergic, uptake amino acids, receive few synaptic boutons, and contain neurofilaments, fibroblast growth factor (FGFs), and other signaling proteins, the latter likely secreted into the central canal. Similar neurons are formed in the SC of the tuatara (Sphenodon puctatus), anurans, and urodeles during tail regeneration. In lizard, most of their projection remains in the SC close to the regenerated tail, but they form synapses with neurons that receive descending nerves from the brainstem, including vestibular nuclei. CSFCNs, aside a possible neurosecretory activity, might sense liquor movements for maintenance of balance, a role that is supported from recent studies on other caudate vertebrates. The regeneration of these cells also in the nervous system of other vertebrates remains unknown.     

Cerebrospinal fluid-contacting neurons in the paraventricular organ and in the spinal cord of the quail embryo: a fluorescence-histochemical study

Although the cerebrospinal fluid-contacting neurons of the avian paraventricular organ exhibit considerable amounts of catecholamines, they show no tyrosine hydroxylase immunoreactivity. In the quail embryo, the development of these neurons has been studied using the paraformaldeyde-glutaraldeyde method for the fluorescence-histochemical localization of catecholamines. The timing of the appearance of catecholamine fluorescence in cerebrospinal fluid-contacting neurons and that in catecholamine-containing neurons of the brainstem have been compared. The first neurons displaying catecholamine fluorescence are found within the locus coeruleus and the nucleus subcoeruleus ventralis on the 5.5th day of incubation. Catecholaminergic neuronal groups of the medulla and mesencephalon can be identified by embryonic day 7, and fluorescent cerebrospinal fluid-contacting neurons of the hypothalamic paraventricular organ can be first recognized at the 8th day of incubation. If the catecholamine content of cerebrospinal fluid-contacting neurons that lack tyrosine hydroxylase depends upon an uptake mechanism, it may be significant that, in fluorescence-histochemical preparations, these neurons can be identified 1–3 days later than those in which catecholamines are synthesized and from which catecholamines are released at an earlier developmental stage. Moreover, cerebrospinal fluid-contacting neurons that have previously been shown to be tyrosine-hydroxylase immunoreactive, and that lie at the spinal-medullary junction display a different developmental pattern. By fluorescence histochemistry, they can be detected only by embryonic day 10.5. The chemical, developmental and topographical differences suggest that the catecholamine-containing cerebrospinal fluid-contacting elements of the paraventricular organ and those of the spinal cord represent two different subsets of cerebrospinal fluid-contacting neurons whose respective functional roles remain to be investigated.     

Morphometric studies of prepyramidal and pyramidal neurons of various vertebrates]

Praepyramidal and pyramidal neurons were demonstrated in homologous telencephalic areas of Salmo irideus (Gibbons 1855), Rana temporaria L., Rattus norvegicus, forma alba, by means of the Golgi-technique. A comparative morphometrical analysis was made concerning the following parameters: main dendritic length, length of the pericaryon, number of dendritic spines in the first 50-micrometer-dendritic segment and length of the spines-free zone. From these data a quotient was calculated and expressing the theoretical dendritic length per ten spines. There is an increase of length of the pericaryon, length of the spines-free zone, main dendritic length and a pseudo-decrease of the number of dendritic spines in the first 50 micrometer-dendritic segment in rats, as compared with Rana and Salmo. The results are discussed with respect to following problems: homology, phylogenesis and corticalisation.     

Cerebrospinal fluid-contacting neurons,sensory pinealocytes and Landolt's clubs of the retina as revealed by means of an electron-microscopic immunoreaction against opsin

Opsin-immunoreactive sites of hypothalamic cerebrospinal fluid (CSF)-contacting neurons, pinealocytes and retinal cells were studied in various vertebrates (Carassius auratus, Phoxinus phoxinus, Triturus cristatus, Bombina bombina, Rana esculenta) by means of postembedding immuno-electron microscopy with the use of the protein A-gold labeling method. The retina of the rat served as a general reference tissue for the quality of the immunocytochemical reaction. A strong opsin immunoreaction (rat-antibovine opsin serum) was obtained in the rod-type outer segments of photoreceptors in the retina of all species studied. Cone-type outer segments exhibited only very few antigenic binding sites. In the pineal organ of the goldfish and the frog, outer segments of the photoreceptor cells displayed strong immunoreactivity. No immunoreaction was found in hypothalamic CSF-contacting neurons and Landolt's clubs of nerve cells of the bipolar layer of the retina. The morphological similarity between the ciliated dendritic terminal of the Landolt's club and the intraventricular dendritic ending of the CSF-contacting neurons is emphasized.     

Vasoactive intestinal peptide-immunoreactive cerebrospinal fluid-contacting neurons in the reptilian lateral septum nucleus accumbens

By means of immunocytochemical demonstration of vasoactive intestinal peptide (VIP) an accumulation of cerebrospinal fluid (CSF)-contacting neurons was found in a circumscribed region of the nucleus accumbens/lateral septum of eleven reptilian (chelonian, lacertilian, ophidian, crocodilian) species. Basal processes of these cells contribute to a subependymal plexus whose density displays considerable interspecific variation. VIP-immunoreactive nerve fibers occur also in the lateral septum and the nucleus accumbens where they encompass immunonegative cells in a basket-like pattern. The CSF-contacting neurons are surrounded by columnar ependymocytes frequently arranged in a pseudostratified manner. These specialized arrays of ependymal cells, however, occupy a more extended area than the VIP-immunoreactive CSF-contacting neurons and can be traced from the rostro-ventral pole of the lateral ventricle to the interventricular foramen. These observations suggest the existence of a telencephalic site of CSF-contacting neurons which may be more widespread than hitherto thought and which may participate in a circumventricular system of the lateral ventricle. Previous studies mainly performed with birds indicate that the VIP-immunoreactive CSF-contacting neurons of the nucleus accumbens might form a part of the encephalic (extraretinal and extrapineal) photoreceptor. However, further experiments are required to test this supposition since the VIP-immunoreactive neurons of the nucleus accumbens remained unlabeled by antibodies against bovine rodopsin and chicken cone-opsin in all eleven species analysed in this investigation.     

Distribution of plasma magnesium in various vertebrates

Plasma magnesium in at least five mammalian species (humans, rat, dog, sheep, cattle) is in the form of a complex, separable from ionic magnesium and plasma protein by size exclusion chromatography on Sephadex G-10. Plasma magnesium in three non-mammalian vertebrates (toads, trout, chicken) behaves similarly to ionic magnesium or as a very small magnesium complex on Sephadex G-10.     

Neuromodulation of central pattern generators in invertebrates and vertebrates

Central pattern generators are subject to extensive modulation that generates flexibility in the rhythmic outputs of these neural networks. The effects of neuromodulators interact with one another, and modulatory neurons are themselves often subject to modulation, enabling both higher order control and indirect interactions among central pattern generators. In addition, modulators often directly mediate the interactions between functionally related central pattern generators. In systems such as the vertebrate respiratory central pattern generator, multiple pacemaker types interact to produce rhythmic output. Modulators can then alter the relative contributions of the different pacemakers, leading to substantial changes in motor output and hence to different behaviors. Surprisingly, substantial changes in some aspects of the circuitry of a central pattern generator, such as a several-fold increase in synaptic strength, can sometimes have little effect on the output of the CPG, whereas other changes have profound effects.     

Aldehyde dehydrogenase activity in blood from various vertebrates

1. Aldehyde dehydrogenase activity was determined in whole blood samples from 17 selected vertebrates of 5 classes, using 3,4-dihydroxyphenylacetaldehyde (the aldehyde derived from dopamine) as substrate. 2. Aldehyde dehydrogenase activity in blood was widely but unevenly distributed among the species studied. 3. Mean aldehyde dehydrogenase activities in the range of 40-140 nmol/min.ml blood (measured at 37 degrees C, pH 8.8) were found in blood from man, monkey, rabbit, guinea pig and mouse (C57BL and NMRI strains), with the highest activity in rabbit blood. 4. Much lower aldehyde dehydrogenase activities (0.5-7.5 nmol/min.ml blood) were found in blood from Sprague-Dawley and Wistar rat, dog, cat, horse, pig, chicken, caiman, frog and rainbow trout, whereas the activities in blood from DBA mouse, cow, sheep and crucian carp were close to the detection limit.     

Phylogenetic study of serotonin-immunoreactive structures in the pancreas of various vertebrates

Summary The distribution pattern of serotonin (5HT) in the pancreas was studied immunohistochemically by using a 5HT monoclonal antibody in various vertebrates including the eel, bullfrog, South African clawed toad, turtle, chicken, mouse, rat, guinea-pig, cat, dog and human. In all species examined, except the bullfrog, 5HT-like immunoreactivity was observed in nerve fibers, in endocrine cells, or in both. Positive nerve fibers were found in the eel, turtle, mouse, rat and guinea-pig. These fibers ran mainly along the blood vessels and partly through the gap between the exocrine glands. In the eel and guinea-pig, positive fibers invaded the pancreatic islet. Occasionally, these positive fibers were found adjacent to the surface of both exocrine and endocrine cells, suggesting a regulatory role of 5HT in pancreatic function. 5HT-positive endocrine cells were observed in the pancreas of all species except for the bullfrog and rat. In the eel and in mammals such as the mouse, guinea-pig, cat, dog and human, 5HT-positive cells were mainly observed within the pancreatic islet. In the South African clawed toad, turtle and chicken, the positive cells were mainly in the exocrine region. The present study indicates that the distribution patterns of 5HT in the pancreas varies considerably among different species.     

The system of cerebrospinal fluid-contacting neurons. Its supposed role in the nonsynaptic signal transmission of the brain

Recent investigations confirm the importance of nonsynaptic signal transmission in several functions of the nervous tissue. Present in various periventricular brain regions of vertebrates, the system of cerebrospinal fluid (CSF)-contacting neurons seems to have a special role in taking up, transforming and emitting nonsynaptic signals mediated by the internal and external CSF and intercellular fluid of the brain. Most of the CSF-contacting nerve cells send dendritic processes into the internal CSF of the brain ventricles or central canal where they form terminals bearing stereocilia and a 9+0-, or 9+2-type cilium. Some of these neurons resemble known sensory cells of chemoreceptor-type, others may be sensitive to the pressure or flow of the CSF, or to the illumination of the brain tissue. The axons of the CSF-contacting neurons transmit information taken up by dendrites and perikarya to synaptic zones of various brain areas. By forming neurohormonal terminals, axons also contact the external CSF space and release various bioactive substances there. Some perikarya send their axons into the internal CSF, and form free endings there, or synapses on intraventricular dendrites, perikarya and/or on the ventricular surface of ependymal cells. Contacting the intercellular space, sensory-type cilia were also demonstrated on nerve cells situated in the brain tissue subependymally or farther away from the ventricles. Among neuronal elements entering the internal CSF-space, the hypothalamic CSF-contacting neurons are present in the magnocellular and parvicellular nuclei and in some circumventricular organs like the paraventricular organ and the vascular sac. The CSF-contacting dendrites of all these areas bear a solitary 9 x 2+0-type cilium and resemble chemoreceptors cytologically. In electrophysiological experiments, the neurons of the paraventricular organ are highly sensitive to the composition of the ventricular CSF. The axons of the CSF-contacting neurons terminate not only in the hypothalamic synaptic zones but also in tel-, mes- and rhombencephalic nuclei and reach the spinal cord as well. The supposed chemical information taken up by the CSF-contacting neurons from the ventricular CSF may influence the function of these areas of the central nervous system. Some nerve cells of the photoreceptor areas form sensory terminals similar to those of the hypothalamic CSF-contacting neurons. Special secondary neurons of the retina and pineal organ contact the retinal photoreceptor space and pineal recess respectively, both cavities being embryologically derived from the 3rd ventricle. The composition of these photoreceptor spaces is important in the photochemical transduction and may modify the activity of the secondary neurons. Septal and preoptic CSF-contacting neurons contain various opsins and other compounds of the phototransduction cascade and represent deep encephalic photoreceptors detecting the illumination of the brain tissue and play a role in the regulation of circadian and reproductive responses to light. The medullo-spinal CSF-contacting neurons present in the oblongate medulla, spinal cord and terminal filum, send their dendrites into the fourth ventricle and central canal. Resembling mechanoreceptors of the lateral line organ, the spinal CSF-contacting neurons may be sensitive to the pressure or flow of the CSF. The axons of these neurons terminate at the external CSF-space of the oblongate medulla and spinal cord and form neurohormonal nerve endings. Based on information taken up from the CSF, a regulatory effect on the production or composition of CSF was supposed for bioactive materials released by these terminals. Most of the axons of the medullospinal CSF-contacting neurons and the magno- and parvicellular neurosecretory nuclei running to neurohemal areas (neurohypophysis, median eminence, terminal lamina, vascular sac and urophysis) do not terminate directly on vessels, instead they form neurohormonal nerve terminals attached by half-desmosomes on the basal lamina of the external and vascular surface of the brain tissue. Therefore, the bioactive materials released from these terminals primarily enter the external CSF and secondarily, by diffusion into vessels and the composition of the external CSF, may have a modulatory effect on the bioactive substances released by the neurohormonal terminals. Contacting the intercellular space, sensory-type cilia were also demonstrated on nerve cells situated subependymally or farther away from the ventricles, among others in the neurosecretory nuclei. Since tight-junctions are lacking between ependymal cells of the ventricular wall, not only CSF-contacting but also subependymal ciliated neurons may be influenced by the actual composition of the CSF besides that of the intercellular fluid of the brain tissue. According to the comparative histological data summarised in this review, the ventricular CSF-contacting neurons represent the phylogenetically oldest component detecting the internal fluid milieu of the brain. The neurohormonal terminals on the external surface of the brain equally represent an ancient form of nonsynaptic signal transmission.     

Comparison of various approaches to calculating the optimal hematocrit in vertebrates

An interesting problem in hemorheology is to calculate that volume fraction of erythrocytes (hematocrit) that is optimal for transporting a maximum amount of oxygen. If the hematocrit is too low, too few erythrocytes are present to transport oxygen. If it is too high, the blood is very viscous and cannot flow quickly, so that oxygen supply to the tissues is again reduced. These considerations are very important, since oxygen transport is an important factor for physical performance. Here, we derive theoretical optimal values of hematocrit in vertebrates and collect, from the literature, experimentally observed values for 57 animal species. It is an interesting question whether optimal hematocrit theory allows one to calculate hematocrit values that are in agreement with the observed values in various vertebrate species. For this, we first briefly review previous approaches in that theory. Then we check which empirical or theoretically derived formulas describing the dependence of viscosity on concentration in a suspension lead to the best agreement between the theoretical and observed values. We consider both spatially homogeneous and heterogeneous distributions of erythrocytes in the blood and also possible extensions, like the influence of defective erythrocytes and cases where some substances are transported in the plasma. By discussing the results, we critically assess the power and limitations of optimal hematocrit theory. One of our goals is to provide a systematic overview of different approaches in optimal hematocrit theory.