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[Mechanisms of development of long-periodicity oscillations in activity in nerve nets. Stochastically uniform nerve nets].

The possibility of generation of long-term activity in the stochastic uniform networks consisting of neuron-like threshold elements was analyzed by the mathematical simulation model. It was shown that the long-term (hundreds of milliseconds) self-stopping activity in the networks with a positive feed-back between units can be generated. Termination of the activity was based on the following factors: 1) stochastic properties of network processes which determine the fluctuations of the activity level; 2) neuronal interaction leading either to synchronization of neuronal discharges and respectively to postactivation inhibitory processes, or to reconstruction of the microstructure of the neuronal network activity when weak-connected neurons are predominantly excited. Basing on the data obtained possible participation of the mechanism involved in formation of the long-term self-stopping activity of interneuronal networks of the spinal cord realizing different types of programmed rhythmical activity (generators) is discussed.

Computers↗

Nerve net formation in the primitive nervous system of Hydra--an overview.

Nerve net formation was examined during head-regeneration and budding of Hydra using indirect immunofluorescence on whole mounts. The nerve net was visualized with an antiserum to a neuropeptide, RFamide. The nerve net forms in two steps: the first is the appearance of ganglion cells at the regenerating tip, and the second is the appearance of sensory cells at the apex and the simultaneous disappearance of ganglion cells from the apex. The behavior of epithelial cells during head-regeneration, as defined by monoclonal antibody TS19, corresponded with nerve net formation. We examined nerve net formation in 3 morphogenetic mutants: a head-regeneration deficient mutant (reg-16), a budding deficient mutant (L4) and a multi-headed mutant (mh-1). In addition, we also examined chimeric strains which consist of the epithelial cells from the wild type and nerve cells from a mutant, and vice versa. We obtained clear evidence that nerve net formation is controlled by the environment provided by the epithelial cells.

Animals↗

Phenotypic maturation of neurons and continuous precursor migration in the formation of the peduncle nerve net in Hydra.

Mechanisms of nerve net formation in Hydra were analyzed using a monoclonal antibody (L96) directed against neurons of the peduncle, the basal end of the polyp's body axis. L96+ neurons express RFamide neuropeptides and constitute 70-80% of all ectodermal neurons in the lower peduncle. L96+ neurons arise from neuronal precursors which immigrate from the gastric region into the upper peduncle and first differentiate into neurons lacking the L96 antigen. By tissue movement, these L96- neurons become displaced to the lower peduncle where L96 antigen expression is initiated. The entire L96 neuron differentiation pathway requires about 4 days, but regeneration stimuli shorten it to only 36 hr. Our experiments indicate that local extrinsic signals released by epithelial cells in the peduncle control the L96+ neuron differentiation pathway. Ectopic L96+ neuron differentiation can be induced by LiCl treatment, which also stimulates ectopic feet in the gastric region. Further experiments show that intrinsic signals are also involved in the L96+ neuron differentiation pathway. Neurons of the gastric region become continuously displaced to the peduncle by tissue movement, but these "old" neurons fail to express the L96 antigen in response to the altered epithelial environment. Gastric neurons also fail to express the L96 antigen after LiCl treatment or regeneration in stem cell-depleted polyps. Thus, the competence of neurons to respond to environmental cues with L96 antigen expression is strongly age-dependent. We define this age-dependent acquisition of the neuronal phenotype as phenotypic maturation controlled by the target tissue.

Animals↗

Localisation of WE-14 immunoreactivity in the developing mouse limbo-corneal nerve net.

WE-14 is generated in subpopulations of chromogranin A immunopositive endocrine cells and neurons including those innervating the anterior uvea. This study investigated WE-14 in intact sclero-limbo-corneal tissue from embryonic (E17), neonatal (N0-N16), and adult mice using immunocytochemistry and confocal scanning laser microscopy. Weak WE-14 immunostaining was observed at birth in nerve fibre tracts entering the corneal mid-stroma from the limbo-scleral junction. Immunopositive fibre tracts were evident throughout the cornea at N3; by N5 the mid-stromal plexus had begun to generate fibre populations extending toward the developing corneal epithelium, and some varicose fibres terminated amongst the developing epithelium. Immunostaining was evident at N7 in the developing limbo-scleral nerve net and some fibres exhibited a close association with unidentified vascular elements. By N11 and in subsequent neonates, the cornea had developed a distinct stratified nerve net composed of thick mid-stromal and thinner upper stromal nerve fibre bundles; both possessed populations of varicose WE-14 immunopositive fibres. In the adult, a sub-epithelial network of varicose WE-14 immunopositive fibres were evident at the limbo-scleral junction. Some fibres exhibited a close association with unidentified vascular elements, while others extended into the upper peripheral corneal stroma. WE-14 was evident in leashes throughout the basal corneal epithelium and generated fibres ramifying between the stratified epithelium with some fibres terminating amongst the outermost corneal epithelia. This study has demonstrated that WE-14 was evident in the limbo-corneal nerve net at birth and that its detection parallels corneal development to adulthood, where WE-14 is evident in a subpopulation of nerve fibres.

Animals↗

Action potential in neurons of motor nerve net of Cyanea (Coelenterata).

Neurons of the motor nerve net of the jellyfish Cyanea were impaled with microelectrodes for intracellular recordings. The cells have conventional, negative resting potentials and produce variable-amplitude action potentials with complex waveforms. The variability and complexity of these spikes is due to the superimposition of two classes of Ca2+-dependent potentials on an otherwise fast, clean action potential. Repetitive stimulation and ionic manipulation reveal that most superimposed potentials are chemically induced excitatory postsynaptic potentials (EPSPs). These account for the complexity and variability of the action potential. The remaining potential is interpreted as a Ca2+ component of the action potential. The action potential is a Na+-dependent but tetrodotoxin- (TTX) insensitive event. Repolarization is achieved by two pharmacologically distinct mechanisms: a tetraethylammonium- (TEA) and 4-amino-pyridine- (4-AP) sensitive K+ efflux and a delayed, Ca2+-activated, K+ efflux. The latter is responsible for the afterhyperpolarization that follows the action potential. The results indicated that these neurons are physiologically conventional. This is interesting in view of the phylogenetic primitiveness of the preparation and important, since it means that this preparation can provide generally useful information on chemical synaptic physiology.

4-Aminopyridine↗

Structure, development, and maintenance of the nerve net of the body column in Hydra.

The anatomy and developmental dynamics of the nerve net in the body column of Hydra viridissima were examined immunocytochemically with a monoclonal antibody (CC04) that recognizes an antigen in nerve cells and with an antiserum against vasopressin. CC04+ neuron cell bodies, their neurites, and vasopressin-like-immunoreactive (VLI+) neurites could be clearly visualized on whole-mount preparations. All neurites of the CC04+ neurons in the body column were VLI+. However, only half of the VLI+ neurites in the body column were CC04+. Immunocytochemical analysis of macerated preparations showed that half of the neurons in the gastric region of the body column were CC04+. These results suggest that most of the neurons in the gastric region are VLI+. The density of the VLI+ neurites was uniform along the entire length of the body column. The CC04+ neuron density in the gastric region remained constant at all stages of asexual development and during foot regeneration. After pulse-labeling with 5-bromo-2'-deoxyuridine (BrdU), CC04+ neurons with labeled nuclei appeared in the body column. We conclude that neuron density in the gastric region is maintained at a constant value by insertion of new neurons in parallel with continuous epithelial cell division.

Animals↗

Three digestive movements in Hydra regulated by the diffuse nerve net in the body column.

The mammalian digestive tract undergoes various digestive movements such as peristalsis and segmentation movement. How those digestive movements and the underlying mechanisms appeared in evolution remains unraveled. A widely accepted view has been that, early in evolution, the digestive process was static based upon diffusion, and later it became dynamic involving digestive movements. Here, we report digestive movements which occur in Hydra, a member of the phylum Cnidaria. We find that the body column of Hydra undergoes a series of movements when fed with Artemia. Comparison of the movements to those in mammals showed similarities in appearance to esophageal reflex, segmentation movement, and defecation reflex. When nerve cells were eliminated, polyps showed only a weak segmentation movement, demonstrating that the diffuse nerve net in the body column of Hydra primarily regulates the movements just as the netlike enteric nervous system does in mammals. Elimination of both secretory gland cells and nerve cells resulted in the complete loss of movement, suggesting that the gland cells are involved in the weak movement. Overall, these observations suggest that the digestive process in Hydra is dynamic and that the diffuse nerve net regulates the digestive movements as a primitive form of enteric nervous system.

Animals↗

A method for recording electrical activity from the body wall nerve nets in sea anemones

Glass microelectrodes were used to record electrical activity from thin rings cut from the column of the sea anemone Calliactis parasitica. This is the first time that pulses have been recorded from the nervous system in the column. Three pulse types were detected, types A, B and C. Type A pulses are probably associated with neurones of the through-conducting nerve net. Type B pulses may be from the endodermal slow conduction system (SS2). Type C pulses have not previously been recorded and are thought to represent activity in a local nerve net. At this stage we cannot positively state whether the recordings are intracellular from endodermal myoepithelial cells or are extracellular from the sub-epithelial region.

Journal Article↗

Electrophysiology of the peripheral nerve net in the polyclad flatworm Freemania litoricola.

1. A diffuse-conducting system close to the dorsal epithelium of the polyclad flatworm Freemania litoricola is described. Tactile stimuli elicit small action potentials which can be conducted around lesions through the body wall. The potentials can occur in bursts or barrages. 2. This conducting system appears to be insensitive to Mg2+ ions. 3. Conduction velocities (0-26--71 m/sec) vary over the animal. Conduction spread in the anterior half of the animal appears to be greater than that in the posterior portion. 4. Response decrement to repeated stimulation can be recorded in the peripheral system but it is not clear if this is due to habituation or fatigue. 5. Conduction from the peripheral net to the brain occurs. Some central units appear to pick up information only, or mainly, through the anterior nerves, while other units can respond to information conducted through the network to nerves of the contralateral side. 6. Different possibilities to account for this system are discussed, and it is suggested that the animals either possess a unique Mg2+ insensitive synaptic nerve-net or else the network is electrically coupled.

Action Potentials↗

A giant nerve net with multi-effector synapses underlying epithelial adhesive strips in the mouth of Beroë (Ctenophora).

We present ultrastructural evidence for the first known example of a giant nerve net in the phylum Ctenophora. The giant fibre system in Beroë underlies paired strips of adherent epithelial cells that run inside the lips. Interlocking actin-lined cell junctions between opposing adhesive strips keep Beroë's large mouth closed while the ctenophore searches for prey. The giant neurons, up to 6-8 microns in diameter, form a continuous lattice-like plexus rich in vesicles, microtubules, and 'presynaptic triads'. A novel feature is that individual giant axons make synaptic contacts with more than one type of effector, i.e. longitudinal muscle fibres and epithelial adhesive cells. Contact of prey with sensory receptors on the lips of Beroë induces rapid disappearance of the actin-lined adhesive cell junctions, and muscular opening of the mouth to ingest prey. Electron microscopy of food-opened mouths shows local thickening of longitudinal muscles and widening of the basal ends of epithelial cells in the adhesive strip, correlated with retraction of the adhesive epithelium into the mesoglea. Addition of 1% Triton X-100 to formaldehyde fixative in the absence of prey also elicits regional thickening of longitudinal muscles at the location of the adhesive strips (visualized by rhodamine-phalloidin staining). The giant neuron system may serve as a final common pathway to rapidly signal disassembly of actin-based junctions between adhesive cells as well as contractions of longitudinal muscles underlying the adhesive strips, thereby enabling Beroë to open its mouth rapidly to engulf prey.

Animals↗

Effect of high-level oxygen exposure on the peroxidase activity and the neuromelanin-like pigment content of the nerve net in the earthworm, Lumbricus terrestris.

Histochemical examination of 1-microm tissue sections from the dorsal nerve plexus of the earthworm, Lumbricus terrestris, reveals multiple brown intraneuronal granules. These granules contain material morphologically and histochemically consistent with neuromelanin. When viewed with transmission electron microscopy, these were seen as single membrane-enclosed biphasic granules with diameters of 370-730 nm. Exposure of L. terrestris to high-level environmental oxygen resulted in an increase in the number of neuromelanin-like pigment granules within the neurons of the circular muscle layer. As measured by ortho-phenylenediamine hydrochloride, the endogenous peroxidase activity of extracts from worms incubated in high-level environmental oxygen was 51% more than controls. The endogenous peroxidase activity was localized in situ with 3,3-diaminobenzidine (DAB) and was found to increase in and around the neuromelanin-like pigment-containing neurons within the circular muscle layer. These studies suggest that the nerve net of L. terrestris may serve as a model to study the role of neuromelanin production in oxidative stress and its relationship to endogenous peroxidases.

Animals↗

Effect of structure on function in model nerve nets.

A theoretical analysis has been made on the effect of the pattern of interneuronal connectivity in model nerve nets on the activity of these nets. Two types of nets have been investigated: one in which the likelihood of a connection between a given neuron and any other element in the net is given by a Poisson probability distribution, and a second type in which the pattern of interconnection follows a Gaussian distribution. An analytical treatment is presented of the equations for noiseless nets in these two conditions. The principal result is that nets with Poisson connectivity law are activated by extraneous firing of a single neuron and continue in spontaneous activity indefinitely. On the other hand, similar nets in which the connections are, however, distributed according to a normal connectivity law, exhibit a definite threshold and produce spontaneous activity only subsequent to extraneous activation of a substantial fraction of the population. Moreover, spontaneous activity in Gaussian nets, but not in Poisson nets, becomes extinguished if the number of active neurons falls below the critical threshold. Some neuroanatomical implications are discussed which suggest that the pyramidal system of the cerebral cortex and other neuronal systems histologically characterized by large numbers of synapses per neuron may incorporate a Gaussian connectivity law, whereas a Poisson law may be characteristic of these cortical layers and nuclei primarily containing granule cells.

Electric Conductivity↗

A subset of cells in the nerve net of Hydra oligactis defined by a monoclonal antibody: its arrangement and development.

A monoclonal antibody, termed JD1, was generated that bound to a subset of the nerve cells in the hypostome and tentacles of Hydra oligactis. Using a whole-mount technique the spatial pattern of the subset of nerve cells and their processes could be clearly visualized using indirect immunofluorescence. The subset largely corresponds to the epidermal sensory cells. Using the same technique the development of the pattern during head regeneration and budding was examined. The appearance of the nerve cells coincides with the formation of both the tentacles and hypostome. When head regeneration does not occur, JD1+ cells do not appear suggesting the differentiation of JD1+ cells is an integral event in head formation dependent on antecedent patterning processes.

Animals↗

Characterization of a RFamide-positive subset of ganglionic cells in the hydrozoan planular nerve net.

The complexity of the hydrozoan planular nervous system was examined. Using a whole-mount technique with indirect immunofluorescence, the spatial pattern of ganglionic cells showing RFamide-like immunoreactivity was visualized. RFamide antiserum bound a subset of ganglionic cells in the anterior and upper middle regions of the planula and a few ganglionic cells in the upper tail region. Labeled cells consisted of bipolar and multipolar neurons. Stained processes from these cells formed a three-dimensional nerve net that followed the contour of the mesoglea; such fibers were striking in terms of their large numbers, long lengths, and organization into distinct bundles. Labeled fibers were seen to contact other ganglionic cells, sensory cells, epithelio-muscle cells, the mesoglea, and the outside free surface. All stained cell bodies and fibers were found in the ectoderm. Using the same technique the reappearance of RFamide-positive ganglionic cells in epithelial tissue of chimeric grafts of planulae was observed. Interstitial cells capable of forming RFamide-positive ganglionic cells underwent extensive anterior-posterior migrations in the grafts, moved into the epithelial tissue, and differentiated into RFamide-positive ganglionic cells. Stained repopulated ganglionic cells always formed in the same position in the epithelial tissue as was observed in control planulae suggesting that the expression of RFamide-like substances may be position dependent in the planula.

Animals↗