Early Forms of Evolution of the Basiepidermal Nerve Plexus of Bilateria as a Possible Evidence for Primary Diversity of Its Initial State

To find out whether there is a real parallelism, as it was understood by Academician A.A. Zavarzin, in the structure of the basiepidermal nerve plexus in primitive representatives of such distant groups of animals, as polychaetes and phoronids, the experimental material obtained earlier on neuronal relations in Myriochele oculata (Polychaeta, Oweniidae) and Phoronopsis harmeri (Tentaculata, Phoronoidea) was analyzed. The similarity of the basiepidermal plexus in the representatives of polychaetes and phoronids has been shown to be merely apparent and has a convergent character, i.e., it does not belong to the type of systemic parallelisms. The structural differences revealed between these two considered evolutionary initial types of nerve plexuses are supposed to be a cause of different directions of differentiation of the Bilateria nervous system. The nerve plexus structure close, by its general organization, to that of oweniids can have many common features with the basiepidermal plexus of especially primitive turbellarias and xenoturbellids, whose neuronal relations are so far non-studied. At the same time, a similar nerve plexus could originate formation of the nervous system of some archiannelids (of the nerillids type), typical polychaetes, and primitive oligochaetes. The neuronal relations similar to those of phoronids probably had more chances to progress in evolution to the direction that is principally close to the neuronal relations in pogonophoras and typical Deuterostomata (including the lower chordates).     

Generation of a locomotory rhythm by a neural network with recurrent cyclic inhibition

An oscillatory intersegmental neuronal network drives the swimming rhythm of the leech. This network consists of interneurons joined via inhibitory connections to form a series of segmentally iterated, concatenated rings. Recurrent cyclic inhibition in these rings produces a multiphasic activity rhythm. By theoretical analysis of such concatenated interneuronal rings and construction of their electronic analogs it is shown that the interneural network identified in the central nervous system of the leech has properties appropriate for generating the observed motor output.     

Local interneurons in the swimmeret system of the crayfish

Summary We describe the structures and physiological properties of thirteen kinds of local interneurons in the swimmeret system of the crayfish,Pacifastacus leniusculus. Eight are unilateral, with processes confined to one side of the midline (Figs. 1, 2); five are bilateral, with processes on both sides of the ganglion (Fig. 6). All have most of their branches in the lateral neuropils. All of the unilateral local interneurons were nonspiking; two of the bilateral interneurons generate action potentials. Three kinds of unilateral interneurons could reset the bursting rhythm or could initiate bursting in quiescent nerve cords. Four others drove tonic firing of motor neurons. Four kinds of bilateral interneurons were premotor, and could affect the period and phase of both pattern generators in their ganglion. One unilateral and one bilateral interneuron were sensory interneurons. At least one bilateral interneuron received input from both pattern generators.Different premotor local interneurons function either in pattern generation, or in hemisegmental coordination of groups of motor neurons, or in bilateral synchronization of the ganglionic pairs of local pattern-generators for the swimmerets.Abbreviations G1. ganglion 1. - LN lateral neuropil - MT miniscule tract     

Differing afferent connections of spiking and nonspiking wind-sensitive local interneurons in the terminal abdominal ganglion of the cricket Gryllus bimaculatus

Fifteen local spiking interneurons (LSIs) and twentyone local non-spiking interneurons (LNIs) were identified in the terminal abdominal ganglion (TAG) of the cricket Gryllus bimaculatus on the basis of intracellular recording and staining (Figs. 1, 5, 6). Although the majority of LNIs showed sharp directionalities (Fig. 7) the LSIs did not (Fig. 3). The directionality of LNIs varied with the recording sites within a single cell (Fig. 8). Electrical stimulations of the cereal sensory nerve suggested that the LNIs are connected monosynaptically with the sensory afferents of both the cerci, and that LSIs may possess a variety of bilateral combinations of polysynaptic connections with the sensory afferents. We found that the spiking and the non-spiking local interneurons in the cereal sensory system differ not only in their membrane properties, but also in their afferent connections, and concluded that their differing connectivity to the sensory afferents will associate them with different roles in signal processing.Abbreviations TAG terminal abdominal ganglion - LSI local spiking interneuron - LNI local non-spiking interneurons - CNS central nervous system - PSP post synaptic potential - GI giant interneuron     

The evolution of the serotonergic nervous system

The pattern of development of the serotonergic nervous system is described from the larvae of ctenophores, platyhelminths, nemerteans, entoprocts, ectoprocts (bryozoans), molluscs, polychaetes, brachiopods, phoronids, echinoderms, enteropneusts and lampreys. The larval brain (apical ganglion) of spiralian protostomes (except nermerteans) generally has three serotonergic neurons and the lateral pair always innervates the ciliary band of the prototroch. In contrast, brachiopods, phoronids, echinoderms and enteropneusts have numerous serotonergic neurons in the apical ganglion from which the ciliary band is innervated. This pattern of development is much like the pattern seen in lamprey embryos and larvae, which leads the author to conclude that the serotonergic raphe system found in vertebrates originated in the larval brain of deuterostome invertebrates. Further, the neural tube of chordates appears to be derived, at least in part, from the ciliary band of deuterostome invertebrate larvae. The evidence shows no sign of a shift in the dorsal ventral orientation within the line leading to the chordates.     

Comparative characterization of the nervous system of the Turbellaria

Initial stages of the centralization of the nervous apparatus in the Turbellaria can be traced through a comparison of the structure of the nervous system in various representatives of the class. The most primitive state, found in the Acoela, is predominantly plexiform with a varying number of longitudinal trunks. Three, and in some cases four, longitudinal trunks are found in the Proseriata and Temnocephalida. Commissures appear in the Macrostomida and all higher orders and form an orthogon. Brain shape varies from ring-shaped in the Acoela to bilobed in the Neorhabdocoela. While the nervous system of the Polycladida is peculiar, having numerous lateral trunks and separation of dorsal and ventral parts of the nervous system, the development of the nervous system in Müller's larvae of polyclads shows it is of an orthogonal type comparable to other platyhelminths. Transition to parasitism is accompanied by some progressive transformations in the structure of the nervous system.     

Microscopic anatomy and ultrastructure of the nervous system of <Emphasis Type="Italic">Phoronopsis harmeri</Emphasis> Pixell, 1912 (Lophophorata: Phoronida)

The microscopic anatomy and ultrastructure of the nervous system of Phoronopsis harmeri was investigated using histological techniques and electron microscopy. The collar nerve ring is basically formed by circular nerve fibers originating from sensitive cells of tentacles. The dorsal nerve plexus principally consists of large motor neurons. It is shown for the first time that the sensitive collar nerve ring immediately passes into the motor dorsal nerve plexus. The basic components of the nervous system have similar cytoarchitectonics and a layered structure. The first layer is formed by numerous nerve fibers surrounded by the processes of glia-like cells. The bodies of glia-like cells constitute the second layer. The third layer consists of neuron bodies overarched by the bodies of epidermal cells. The giant nervous fiber is accompanied by more than one hundred nerve fibers of a common structure and, thus, marks the true longitudinal nerve. The phoronids possess one or two longitudinal nerves. It is supposed that the plexus nature of the nervous system in phoronids may be related to their phylogenesis. A comparison of the nervous system organization and body plans among the Lophophorata suggests that the nervous system of phoronids cannot be considered as a reductive variant of the brachiopod nervous system. At the same time, the structure of the nervous system of bryozoans can be derived from that of phoronids.     

The topological organization of primary afferents in the terminal ganglion of crayfish,Procambarus clarkii

Summary The central projections of primary afferents in the terminal ganglion of the crayfish can be seen when an axonal filling with nickel chloride with subsequent silver intensification was used for identification. We describe here the topological relationships of the projections to the landmark structures of the neuropil.The terminal ganglion has five pairs of sensory nerves associated with the mechanosensory hairs and internal proprioceptors. The projection fields of the primary sensory neurons in the nerves Rl and R2 are almost entirely restricted to the ipsilateral half of the ganglion, whereas those of the nerves R3, R4 and R5 cross the midline to form three sensory commissures, A6SCI, A7SCI and A7SCII. The projection fields are segregated from each other, although all are restricted to the ventral neuropil which lies under the ventral intermediate tract (VIT). The intersegmental projections that ascend via the connective ipsilateral to their origins could be observed. This pattern of projection correlates well with the receptive fields exhibited by several mechanosensory interneurons on the body surface of the final segment.     

Intersegmental sensory tracts and contralateral motor neurons in the leg ganglia of the spider Cupiennius salei Keys

Summary Cobalt filling into spider legs reveals plurisegmental receptor endings and the plurisegmental origin of motor neurons. Motor neuron dendrites are organized into two domains, one interacting with plurisegmental receptors, the other arborizing within the lateral neuropil of the leg neuromere. The intersegmental organization of both motor and sensory elements supports behavioural studies demonstrating inhibitory connections between legs.     

Central afferent pathways of long hair sensilla in the ventral nerve cord of the Indian black scorpion,Heterometrus fulvipes Koch

Histological and electrophysiological studies of identified long hair sensilla (LHS) have provided information on primary afferent fibre pathways in the ventral nerve cord of the Indian black scorpion, Heterometrus fulvipes.Cobalt-filling of single LHS on the metasoma showed that sensory axons enter the respective segmental ganglion, ascend ipsilaterally through the next anterior ganglia and terminate in a 4th ganglion. In each ganglion, these plurisegmental fibres give off collateral branches that terminate in the ganglionic neuropil. Fibres entering heterolateral connectives were not found.Recordings from peripheral nerves after deflections of a hair showed single or multiple spike discharges. A single spike could be recorded from ipsilateral anterior connectives of the ventral nerve cord, indicating a through-conductance of the sensory pathways. Strong deflections of a single hair activated several ipsilateral and fewer contralateral ascending interneurons and some segmentai motor neurons. Behavioral studies demonstrate the mechanoreceptive function of the LHS.The present study provides evidence in support of the notion that sensory afferents of the postabdomen in the scorpion bring about rapid, co-ordinated intersegmental movements of the multisegmented tail of the scorpion.Abbreviations CNS central nervous system - LHS long hair sensillum - TR trichobothria     

Targeting of amacrine cell neurites to appropriate synaptic laminae in the developing zebrafish retina

Cellular mechanisms underlying the precision by which neurons target their synaptic partners have largely been determined based on the study of projection neurons. By contrast, little is known about how interneurons establish their local connections in vivo. Here, we investigated how developing amacrine interneurons selectively innervate the appropriate region of the synaptic neuropil in the inner retina, the inner plexiform layer (IPL). Increases (ON) and decreases (OFF) in light intensity are processed by circuits that are structurally confined to separate ON and OFF synaptic sublaminae within the IPL. Using transgenic zebrafish in which the majority of amacrine cells express fluorescent protein, we determined that the earliest amacrine-derived neuritic plexus formed between two cell populations whose somata, at maturity, resided on opposite sides of this plexus. When we followed the behavior of individual amacrine cells over time, we discovered that they exhibited distinct patterns of structural dynamics at different stages of development. During cellular migration, amacrine cells exhibited an exuberant outgrowth of neurites that was undirected. Upon reaching the forming IPL, neurites extending towards the ganglion cell layer were relatively more stable. Importantly, when an arbor first formed, it preferentially ramified in either the inner or outer IPL corresponding to the future ON and OFF sublaminae, and maintained this stratification pattern. The specificity by which ON and OFF amacrine interneurons innervate their respective sublaminae in the IPL contrasts with that observed for projection neurons in the retina and elsewhere in the central nervous system.     

Investigation of spinal neuronal mechanisms of movement control systems

Morphological and electrophysiological investigations of the means whereby the principal descending motor systems (the cortico-, rubro-, reticulo-, and vestibulo-spinal tracts) are connected with the segmental interneuronal apparatus and motoneurons show that these connections can be based on two different principles. Descending systems either activate motoneurons directly (monosynaptically) or are connected primarily with various interneuron systems, exerting their influence in that case by regulating the activity of simpler or more complex spinal mechanisms. The older descending system (reticulo- and vestibulo-spinal) possess a monosynaptic excitatory action of motoneurons; the evolutionarily newer descending systems, which transmit the most complex motor signals from the cerebral and cerebellar cortex to the spinal cord (cortico- and rubro-spinal), terminate synaptically in every case on interneurons. It is only in primates that a few cortico-spinal fibers form monosynaptic connections with motoneurons. The chief ways of action of the descending systems on interneurons are: control of the afferent inflow into the interneuron system by presynaptic inhibition of the corresponding synapses; control of the interneuron system by postsynaptic interaction with afferent influences; control of motoneurons through the specialized interneuron apparatus. The investigation shows that the last of these mechanisms functions in the cortico- and rubro-spinal, and possibly also in the reticulo- and vestibulo-spinal systems. The functional role of the various means of connection of the descending systems with the spinal neurons in the system of movement control is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 2, pp. 189–202, March–April, 1970.     

Transplantation of neurons reveals processing areas and rules for synaptic connectivity in the cricket nervous system

In order to assess the nature of spatial cues in determining the characteristic projection sites of sensory neurons in the CNS, we have transplanted sensory neurons of the cricket Acheta domesticus to ectopic locations. Thoracic campaniform sensilla (CS) function as proprioceptors and project to an intermediate layer of neuropil in thoracic ganglia while cercal CS transduce tactile information and project into a ventral layer in the terminal abdominal ganglion (TAG). When transplanted to ectopic locations, these afferents retain their modality-specific projection in the host ganglion and terminate in the layer of neuropil homologous to that of their ganglion of origin. Thus, thoracic CS neurons project to intermediate neuropil when transplanted to the abdomen and cercal CS neurons project to a ventral layer of neuropil when transplanted to the thorax. We conclude that CS can be separated into two classes based on their characteristic axonal projections within each segmental ganglion. We also found that the sensory neurons innervating tactile hairs project to ventral neuropil in any ganglion they encounter after transplantation. Ectopic sensory neurons can form functional synaptic connections with identified interneurons located within the host ganglia. The new contacts formed by these ectopic sensory neurons can be with normal targets, which arborize within the same layer of neuropil in each segmental ganglion, or with novel targets, which lack dendrites in the normal ganglion and are thus normally unavailable for synaptogenesis. These observations suggest that a limited set of molecular markers are utilized for cell–cell recognition in each segmentally homologous ganglion. Regenerating sensory neurons can recognize novel postsynaptic neurons if they have dendrites in the appropriate layer of neuropil. We suggest that spatial constraints produced by the segmentation and the modality-specific layering of the nervous system have a pivotal role in determining synaptic specificity. © 1993 John Wiley & Sons, Inc.     

GABA-ergic structures in the intact and chronically isolated cat association cortex (area 5)

The distribution of GABA-ergic structures in the intact and neuronally isolated cat cerebral cortex in area 5 was studied by the histochemical reaction for GABA-transaminase 2 and 3 weeks after isolation. The overwhelming majority of GABA-ergic fibers of the neuropil and of synaptic terminals was shown to be formed by axons of a few GABA-ergic interneurons, and only a small proportion of them belong to afferent axons of extracortical origin. GABA-ergic interneurons were subdivided into short-axonal, forming connections within an isolated area, and long-axonal, forming horizontal connections with more distant cortical neurons. GABA-ergic axons give numerous projections to bodies and proximal segments of dendrites of many pyramidal neurons not containing GABA-transaminase, and of stellate neurons, which include cells with GABA-ergic and non-GABA-ergic mediator nature. It is suggested that the influence of some GABA-ergic neurons on others is responsible for intracortical spatial regulation of inhibition.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 3, pp. 365–371, May–June, 1985.     

The morphology and anatomy of the vestimentiferan worm <Emphasis Type="Italic">Oasisia alvinae</Emphasis> Jones, 1985 (Annelida: Siboglinidae). II. Integument,nervous system and musculature

The histological structure of the epidermis, nervous system, and musculature of the vestimentiferan worm Oasisia alvinae is examined. The differentiation of the integument of different regions of the body and the histological structure of the pyriform glands and papillae with cuticular plaques are described. All the main elements of the nervous system lie in the thickness of the epidermis. The brain structure is examined and an attempt is made to homologize parts of the vestimentiferan brain with the polychaete brain. Part of the vestimentiferan brain above the intrabrain channel corresponds to the supraesophageal ganglion of polychaetes, while part of the vestimentiferan brain under the channel corresponds to the subesophageal ganglion. Two commissures are located in the brain region that corresponds to the supraesophageal ganglion. These commissures could be related to the ventral and dorsal roots of the circumesophageal connectives of sedentary polychaetes. Bundles of fiber occur, which presumably correspond to the circumesophageal connectives and the transverse commissure inside the subesophageal ganglion. The musculature differentiation in the trunk region is a characteristic feature of O. alvinae, i.e., feather-like muscles that occur only in the anterior part of the trunk.     

Modulation of synaptic events by heavy metals in the central nervous system of mollusks

Summary 1. The effects of heavy metals (Pb²⁺, Hg²⁺, and Zn²⁺) on synaptic transmission in the identified neural network ofHelix pomatia L. andLymnaea stagnalis L. (Gastropoda, Mollusca) were studied, with investigation of effects on inputs and outputs as wells as on interneuronal connections.2. The sensory input running from the cardiorenal system to the central nervous system and the synaptic connections between central neurons were affected by heavy metals.3. Lead and mercury (10^–5–10^–3 M) eliminated first the inhibitory, then the excitatory inputs running from the heart to central neurons. At the onset of action lead increased the amplitude of the excitatory postsynaptic potentials, but blockade of sensory information transfer occurred after 10–20 min of treatment.4. The monosynaptic connections between identified interneurons were inhibited by lead and mercury but not by zinc. Motoneurons were found to be less sensitive to heavy metal treatment than interneurons or sensory pathways.5. The treatment with Pb²⁺ and Hg²⁺ often elicited pacemaker and bursting-type firing in central neurons, accompanied by disconnection of synaptic pathways, manifested by insensitivity to sensory synaptic influences.6. Zn²⁺ treatment also sometimes induced pacemaker activity and burst firing but did not cause disconnection of the synaptic transmission between interneurons.7. A network analysis of heavy metal effects can be a useful tool in understanding the connection between their cellular and their behavioral modulatory influences.     

Distribution of NADPH-diaphorase-positive ascending interneurones in the crayfish terminal abdominal ganglion

Previous neuropharmacological studies have described the presence of a nitric oxide-cGMP signalling pathway in the crayfish abdominal nervous system. In this study we have analysed the distribution of putative nitric oxide synthase (NOS)-containing ascending interneurones in the crayfish terminal abdominal ganglion using NADPH-diaphorase (NADPHd) histochemistry. Ascending intersegmental interneurones were stained intracellularly using the fluorescent dye Lucifer yellow and the ganglia containing the stained interneurones subsequently processed for NADPHd activity. Fluorescence persisted throughout histochemical processing. These double-labelling experiments showed that 12 of 18 identified ascending interneurones were NADPHd positive. Thus many ascending interneurones that process mechanosensory signals in the terminal ganglion may contain NOS, and are themselves likely sources of NO which is known to modulate their synaptic inputs. Three clear relationships emerged from our analysis between the effects of NO on the synaptic inputs of interneurones, their output properties and their staining for NADPH-diaphorase. First were class 1 interneurones with no local outputs in the terminal ganglion, the NE type interneurones, which had sensory inputs that were enhanced by NO and were NADPHd positive. Second were class 1 interneurones with local and intersegmental output effects that had sensory inputs that were also enhanced by NO but were NADPHd negative. Third were class 2 interneurones with local and intersegmental outputs that had synaptic inputs that were depressed by the action of NO but were NADPHd positive. These results suggest that NO could selectively enhance specific synaptic connections and sensory processing pathways in local circuits.     

Postural interneurons in the abdominal nervous system of lobster

The nature of the synaptic relationship between 7 identified postural interneurons and 5 pairs of superficial motoneurons was examined by obtaining dual intracellular recordings from interneuron-motoneuron pairs in the lobster 2nd abdominal ganglion. For six different interneuron-motoneuron pairs EPSPs recorded from motoneurons occurred with a short (1 to 3 ms) fixed latency following each presynaptic spike recorded from the interneuron. This suggests that there is a monosynaptic relationship between these interneurons and motoneurons. Monosynaptic pathways accounted for 27% of all excitatory connections. Preliminary evidence indicates that the monosynaptic potentials are mediated by an excitatory chemical synapse since: all IPSPs occurred with latencies greater than 5 ms, there was no evidence for electrical coupling, and one of the interneurons produced facilitating PSPs. A majority of all monosynaptic connections were made by two of the flexion producing interneurons (FPIs), 201 and 301. The synaptic outputs of these FPIs were similar in that both made monosynaptic connections with a different bilaterally homologous pair of motoneurons. Both also produced larger EPSPs and more vigorous spiking in contralateral members of the bilateral motoneuron pairs. A previous study demonstrated that interneurons 201 and 301 are the only postural interneurons yet identified that express motor programs indistinguishable from command neurons. Taken together, these results suggest that certain intersegmental interneurons share properties with command neurons and driver neurons, and that there may not be a sharp morphological or functional distinction between these two cell types.     

The bimodal auditory-vibratory system of the thoracic ventral nerve cord in Locusta migratoria (Acrididae, Locustinae, Oedipodini).

In locusts the auditory receptors of the tympanal organs and many of the vibratory receptors of all 6 legs converge at the level of the thoracic ventral nerve cord, forming a combined auditory-vibratory sensory system; it is represented by the VS-, S-, and V-neurons ascending to the supraesophageal ganglion. The connections between vibratory receptors of the different legs and the dendritic inputs of the bimodal ascending neurons are investigated in this report. As an example, the dendritic branches of the G- and V3-neurons for auditory and vibratory input could be localized by simultaneous recording at 2 different positions of the axon. The vibratory input from the receptors of the different legs was determined. Segmental and/or intersegmental thoracic interneurons are intercalated between the receptors and the ascending auditory-vibratory neurons (G- and V3-neurons). The morphology and function of 2 intersegmental vibratory interneurons (VI1- and VI2-neurons) are described. They probably connect the vibratory receptors of 1 (or 2) leg(s) of 1 thoracic segment with the different bimodal auditory-vibratory neurons. The importance of the anterior Ring Tract for synaptic connection between receptor cells, first order interneurons, and bimodal auditory-vibratory neurons is discussed on the basis of morphological and physiological data.     

Embryonic development of the alimentary canal and ectodermal derivatives in the primitive moth,Neomicropteryx nipponensis issiki (Lepidoptera,Micropterygidae)

The formation of the alimentary canal, nervous system, and of other ectodermal derivatives in the embryo of the primitive moth, Neomicropteryx nipponensis Issiki, is described. The stomodaeum is formed from an invagination in the medioposterior portion of the protocephalon. The proctodaeum arises as an extension of the amnioproctodaeal cavity. The midgut epithelium orginates from anterior and posterior rudiments in blind ends of the stomodaeum and proctodaeum. The decondary dorsal organ is formed in developing midgut. The development of the brain is typical of insects. The ventral nerve cord originates in large part from neuroblasts arising in 3 gnathal, 3 thoracic, and 11 abdominal segments. Intrasegmental median cord cells probably differentiate into both ganglion cells and glial elements of the ventral nerve cord; intersegmental cells appear not to participate in the formation of the nervous system. The stomatogastric nervous system develops from three evaginations in the dorsal wall of the stomodaeum, and consists of the frontal, hypocerebral, and ventricular ganglia, the recurrent nerve, and corpora cardiaca. Five stemmata arise from the epidermis on each side of the head. Five pairs of ectodermal invaginations are formed in the cephalognathal region to produce the tentorium, mandibular apodemes, corpora allata, and silk glands. Prothoracic glands orginate in the prothorax. Mesothoracic spiracles shift anteriorly to the prothorax during development. Oenocytes arise in the first seven abdominal segments. Invaginated pleuropodia are formed in the first abdominal segment.