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Distributed processing of sensory information in the leech. II. Identification of interneurons contributing to the local bending reflex.

Isolated midbody ganglia of the leech Hirudo medicinalis were surveyed for interneurons contributing to the dorsal component of the local bending reflex, i.e., to the excitation of dorsal excitatory motor neurons that follows stimulation of dorsal mechanoreceptors responsive to pressure (P cells). Nine types of local bending interneuron could be distinguished on physiological and morphological grounds--8 paired and 1 unpaired cell per ganglion. Synaptic latencies from sensory neurons to interneurons were consistent with a direct or possibly disynaptic pathway. Connections between interneurons appeared to be rare and hyperpolarization of individual interneurons during local bending produced small but reliable decrements in motor neuron response, suggesting that multiple parallel pathways contribute to the behavior. Paradoxically, most interneurons received substantial inputs from ventral as well as dorsal mechanoreceptors, indicating that interneurons that were distinguished by their contribution to dorsal local bending were, in fact, active in ventral and lateral bends as well. The capacity to detect a particular stimulus and produce the appropriate response cannot be localized to particular types of interneuron; rather, it appears to be a distributed property of the entire local bending network.

Animals↗

Parallel processing of proprioceptive signals by spiking local interneurons and motor neurons in the locust.

The connections made by afferents from a proprioceptor at the femorotibial joint in a hind leg of a locust, the femoral chordotonal organ (FCO), were determined by making intracellular recordings from motor neurons and spiking local interneurons in the central nervous system and from afferent cell bodies in the periphery. Staining the central projections of the afferent neurons with dye introduced into their axons at the receptor, and the intracellular injection of dye into motor neurons and interneurons, shows that the branches of all 3 types of neuron overlap in specific regions of neuropile. Afferents excited by a movement of the receptor apodeme that is equivalent to an imposed extension of the femorotibial joint excite flexor tibiae motor neurons and some spiking local interneurons with cell bodies at the ventral midline of the metathoracic ganglion. The opposite movement excites extensor tibiae motor neurons and a different set of spiking local interneurons. Spikes in afferents that excite flexor motor neurons evoke depolarizing potentials that follow each spike with a consistent central latency of approximately 1.5 msec. The amplitude of the depolarizing potentials is dependent upon the membrane potential of the motor neuron. This evidence points to the connection being direct and to the potentials' being EPSPs. Simultaneous recordings from certain spiking local interneurons and certain flexor motor neurons show that they receive many synaptic potentials in common and are driven in a parallel fashion by movements of the receptor apodeme. Spikes of some afferents evoke EPSPs in both neurons with the same consistency and latency. An afferent can therefore synapse directly upon a motor neuron and a spiking local interneuron. Each afferent synapses on several motor neurons and possibly upon several interneurons. In turn, each motor neuron and each interneuron receives inputs from several afferents.

Animals↗

Expression of NGF and NT3 mRNAs in hippocampal interneurons innervated by the GABAergic septohippocampal pathway.

We used in situ hybridization for the detection of nerve growth factor (NGF), brain-derived neurotrophic factor (BDNF), and neurotrophin 3 (NT3) mRNAs combined with immunocytochemistry against the calcium-binding proteins parvalbumin (PARV), calbindin 28k (CALB), and calretinin (CALR) to determine the expression of neurotrophins in functionally distinct subsets of hippocampal interneurons. Most PARV-immunoreactive neurons in the hippocampus were NGF mRNA-positive (82%), which corresponds to 71% of NGF-positive neurons in the hippocampus proper and in the dentate gyrus (excluding granule cells). In contrast, only a subset of CALB- and CALR-immunoreactive interneurons (24% and 23%, respectively) displayed hybridization signals for NGF. Small subsets of PARV- and CALR-positive cells expressed NT3 mRNA, but we did not find hippocampal interneurons expressing BDNF mRNA. These results show that NGF and NT3 genes are differentially regulated in distinct subsets of GABAergic cells, and these interneurons are a major source of NGF production in the hippocampus. We also addressed whether hippocampal interneurons expressing neurotrophins were targets of the GABAergic septohippocampal pathway. We developed a triple-labeling method that combines anterograde tracing of this pathway by means of Phaseolus vulgaris leucoagglutinin injections, with in situ hybridization for the detection of neurotrophins, and immunocytochemistry for calcium-binding proteins. Virtually every PARV-positive neuron innervated by GABAergic septohippocampal baskets expressed NGF mRNA (86%), whereas 39-59% of CALR- and CALB-positive interneurons that were contacted by GABAergic septohippocampal axons showed NGF gene expression. A small subset of NT3 mRNA-expressing interneurons was also innervated by septohippocampal baskets. These findings show that the GABAergic septohippocampal pathway preferentially terminates on interneurons expressing NGF mRNA, suggesting that this neurotrophic factor might be involved in the specification of this connection and in its maintenance and normal function in the adult brain.

Afferent Pathways↗

Estimation of the size and directional output of functional groups of interneurons underlying abdominal positioning behaviors in crayfish.

Quantitative studies were made of a large population of interneurons that controls postural flexion and extension of the crayfish abdomen. The number of interneurons needed to produce a motor program was estimated by stimulating a single abdominal positioning interneuron and recording interneuronal activity that was evoked from rostral and caudal connectives in an isolated abdominal nerve cord. We also examined the role that these functional groups have in producing a stronger motor output in either a rostral or caudal direction and thus specifying various abdominal geometrics. The average number of interneurons responding to stimulation of a single abdominal positioning interneuron was 32 (range: 3-50; n = 27). The average number of interneurons that decreased activity was 10 (range: 2-32). Of 653 activated interneurons from 20 preparations, approximately 43% fired between 2 and 5 Hz, 33% fired between 6 and 15 Hz, and 25% fired > 15 Hz. The size of a recruited group was usually but not always correlated with the strength of its motor response or with the direction of motor bias. Therefore, the contribution of a group may depend upon the number of active elements as well as synaptic efficacy.

Abdomen↗

A novel type of GABAergic interneuron connecting the input and the output regions of the hippocampus.

The main excitatory pathway of the hippocampal formation is controlled by a network of morphologically distinct populations of GABAergic interneurons. Here we describe a novel type of GABAergic interneuron located in the outer molecular layer (OML) of the rat dentate gyrus with a long-range forward projection from the dentate gyrus to the subiculum across the hippocampal fissure. OML interneurons were recorded in hippocampal slices by using the whole-cell patch-clamp configuration. During recording, cells were filled with biocytin for subsequent light and electron microscopic analysis. Neurons projecting to the subiculum were distributed throughout the entire OML. They had round or ovoid somata and a multipolar dendritic morphology. Two axonal domains could be distinguished: an extensive, tangential distribution within the OML and a long-range vertical and tangential projection to layer 1 and stratum pyramidale of the subiculum. Symmetric synaptic contacts were established by these interneurons on dendritic shafts in the OML and subiculum. OML interneurons were characterized physiologically by short action potential duration and marked afterhyperpolarization that followed the spike. On sustained current injection, they generated high-frequency (up to 130 Hz, 34 degrees C) trains of action potentials with only little adaptation. In situ hybridization and single-cell RT-PCR analysis for GAD67 mRNA confirmed the GABAergic nature of OML interneurons. GABAergic interneurons in the OML projecting to the subiculum connect the input and output regions of the hippocampus. Hence, they could mediate long-range feed-forward inhibition and may participate in an oscillating cross-regional interneuron network that may synchronize the activity of spatially distributed principal neurons in the dentate gyrus and the subiculum.

Animals↗

Up and down states in striatal medium spiny neurons simultaneously recorded with spontaneous activity in fast-spiking interneurons studied in cortex-striatum-substantia nigra organotypic cultures.

In vivo intracellular spontaneous activity in striatal medium spiny (MS) projection neurons is characterized by "up" and "down" states. How this type of activity relates to the neuronal activity of striatal fast-spiking (FS) interneurons was examined in the presence of nigral and cortical inputs using cortex-striatum-substantia nigra organotypic cultures grown for 45 +/- 4 d. The nigrostriatal projection was confirmed by tyrosine hydroxylase immunoreactivity. Corticostriatal (CS) projection neurons, striatal MS neurons, and FS neurons were intracellularly recorded and morphologically and electrophysiologically characterized. Intracellular spontaneous activity in the cultures consisted of intermittent depolarized periods of 0.5-1 sec duration. Spontaneous depolarizations in MS neurons were restricted to a narrow membrane potential range (up state) during which they occasionally fired single spikes. These up states were completely blocked by the glutamate antagonist CNQX. In FS interneurons, depolarized periods were characterized by large membrane potential fluctuations that occupied a wide range between rest and spike threshold. Also, FS interneurons spontaneously fired at much higher rates than did MS neurons. Simultaneous intracellular recordings established that during spontaneous depolarizations MS neurons and FS interneurons displayed correlated subthreshold neuronal activity in the low frequency range. These results indicate that (1) the CS projection neurons, striatal MS neurons, and FS interneurons grown in cortex-striatum-substantia nigra organotypic cultures show morphological and electrophysiological characteristics similar to those seen in vivo; (2) striatal MS neurons but not FS interneurons show an up state; (3) striatal MS neurons and FS interneurons receive common, presumably cortical inputs in the low frequency range. Our results support the view that the cortex provides a feedforward inhibition of MS neuron activity during the up state via FS interneurons.

6-Cyano-7-nitroquinoxaline-2,3-dione↗

Functionally distinct groups of interneurons identified during rhythmic carbachol oscillations in hippocampus in vitro.

During distinct behavioral states, the hippocampus exhibits characteristic rhythmic electrical activity. Evidence in vivo suggests that both principal pyramidal cells and GABAergic interneurons participate in generating oscillations. We found that during rhythmic oscillations in area CA3, functionally distinct classes of interneurons could be identified, although all recorded interneurons had similar dendritic and axonal arbors. One group of interneurons was powerfully excited by CA3 pyramidal cells, whereas two other interneuron groups were relatively unaffected by pyramidal cell firing. One of these groups of interneurons was potently inhibited by other local interneurons during the pyramidal cell bursts. Our findings emphasize that morphologically similar cells are wired together very differently within the local circuit. The classes of hippocampal interneurons we have tentatively defined may be used during distinct behavioral states to switch the local network from one oscillatory state to another.

Animals↗

[Interneurons of the motor region of the neocortex].

Interneurons of motor area in the brain cortex have been studied in cats and monkeys. The greatest attention has been paid to pyramidal interneurons, among which six cell types have been described according to their axonal composition. Unlike stellate interneurons, all types of pyramidal interneurons possess less developed axonal collaterals. Interneuronal contacts are situated on dendrites or cell bodies of middle and large long-axonal pyramids. Functional role of cortical interneurons seems to be different. Some of them are of inhibitory nature (basket cells and, perhaps, other types of long-axonal stellate neurons), others are exciting elements. The latter include short-axonal stellate neurons and, perhaps, pyramidal interneurons. While comparing the cortex in cats and monkeys, it is evident that the neocortex in monkeys, especially its lower layers, is rich in pyramidal interneurons.

Animals↗

Pyramidal neurons proportionately alter the identity and survival of specific cortical interneuron subtypes.

The mammalian cerebral cortex comprises a complex neuronal network that maintains a precise balance between excitatory pyramidal neurons and inhibitory interneurons. Accumulating evidence indicates that specific interneuron subtypes form stereotyped microcircuits with distinct pyramidal neuron classes. Here we show that pyramidal neurons play an active role in this process by promoting the survival and terminal differentiation of their associated interneuron subtypes. In wild-type cortex, interneuron subtype abundance mirrors the prevalence of their pyramidal neuron partners. In Fezf2 mutants, which lack layer 5b pyramidal neurons and are expanded in layer 6 intratelencephalic neurons, corresponding subtype-specific shifts occur through two distinct mechanisms: somatostatin interneurons adjust their programmed cell death, whereas parvalbumin interneurons switch their subtype identity. Silencing neuronal activity or blocking vesicular release in L5b pyramidal neurons revealed that their communication with interneurons does not require voltage-gated synaptic activity and engages both tetanus toxin-sensitive and -insensitive pathways. Moreover, a targeted bioinformatic screen for ligand-receptor pairs displaying subtype-specific expression and reduced expression of pyramidal neuron-derived ligand in Fezf2 mutants identified candidate secreted factors and adhesion molecules. These findings reveal distinct, pyramidal neuron-driven mechanisms for sculpting interneuron diversity and integrating them into local cortical circuits.

Journal Article↗

Interneurons of the ganglionic layer in the mormyrid electrosensory lateral line lobe: morphology, immunohistochemistry, and synaptology.

This is the second paper in a series that describes the morphology, immunohistochemistry, and synaptology of the mormyrid electrosensory lateral line lobe (ELL). The ELL is a highly laminated cerebellum-like structure in the rhombencephalon that subserves an active electric sense: Objects in the nearby environment of the fish are detected on the basis of changes in the reafferent electrosensory signals that are generated by the animal's own electric organ discharge. The present paper describes interneurons in the superficial (molecular, ganglionic, and plexiform) layers of the ELL cortex that were analyzed in the light and electron microscopes after Golgi impregnation, intracellular labeling, neuroanatomical tracing, and gamma-aminobutyric acid (GABA) immunohistochemistry. The most numerous interneurons in the ganglionic layer are GABAergic medium-sized ganglionic (MG) cells and small ganglionic (SG) cells. MG cells have 10-20 spiny apical dendrites in the molecular layer, a cell body of 10-12 microns diameter in the ganglionic layer, a single basal dendrite that gives rise to fine, beaded, axon-like branches in either the plexiform layer (MG1 subtype) or the deeper granular layer (MG2 subtype), and an axon that terminates in the plexiform layer. Their apical dendritic tree has 12,000-22,000 spines that are contacted by GABA-negative terminals, and it receives, 1,250-2,500 GABA-positive contacts on the smooth dendritic surface between the spines. The average ratio of GABA-negative to GABA-positive contacts on the interneuron apical dendrites (14:1) is significantly higher than that for the efferent projection cells that have been described previously (Grant et al. [1996] J. Comp. Neurol., this issue). The somata and basal dendrites of MG cells receive a low to moderate density of GABAergic synaptic input, and their axons make GABAergic synaptic contacts with the somata and cell bodies of MG as well as with large ganglionic (LG) cells. SG cells probably represent immature, growing MG cells. Other interneurons in the superficial ELL layers include GABAergic stellate cells in the molecular layer, two types of non-GABAergic cells with smooth dendrites in the deep molecular layer that are named thick-smooth dendrite cells and deep molecular layer cells, and horizontal cells that are encountered particularly in the plexiform layer. Comparison with the ELL of waveform gymnotiform fish, which is another group of active electrolocating teleosts that has been investigated thoroughly, shows striking differences. In these fish, no GABAergic interneurons are found in the ganglionic (pyramidal) layer of the ELL, and GABA-negative interneurons with smooth dendrites in the molecular layer also seem to be lacking. At present, the phylogenetic origin of the described superficial interneurons in the mormyrid ELL is uncertain.

Animals↗

Y retinal terminals contact interneurons in the cat dorsal lateral geniculate nucleus.

One of the largest influences on dorsal lateral geniculate nucleus (dLGN) activity comes from interneurons, which use the neurotransmitter gamma-aminobutyric acid (GABA). It is well established that X retinogeniculate terminals contact interneurons and thalamocortical cells in complex synaptic arrangements known as glomeruli. However, there is little anatomical evidence for the involvement of dLGN interneurons in the Y pathway. To determine whether Y retinogeniculate axons contact interneurons, we injected the superior colliculus (SC) with biotinylated dextran amine (BDA) to backfill retinal axons, which also project to the SC. Within the A lamina of the dLGN, this BDA labeling allowed us to distinguish Y retinogeniculate axons from X retinogeniculate axons, which do not project to the SC. In BDA-labeled tissue prepared for electron microscopic analysis, we subsequently used postembedding immunocytochemical staining for GABA to distinguish interneurons from thalamocortical cells. We found that the majority of profiles postsynaptic to Y retinal axons were GABA-negative dendrites of thalamocortical cells (117/200 or 58.5%). The remainder (83/200 or 41.5%) were GABA-positive dendrites, many of which contained vesicles (59/200 or 29.5%). Thus, Y retinogeniculate axons do contact interneurons. However, these contacts differed from X retinogeniculate axons, in that triadic arrangements were rare. This indicates that the X and Y pathways participate in unique circuitries but that interneurons are involved in the modulation of both pathways.

Animals↗

Cortical inputs to m2-immunoreactive striatal interneurons in rat and monkey.

Previous anatomical studies have been unsuccessful in demonstrating significant cortical inputs to cholinergic and somatostatinergic striatal interneurons in rats. On the other hand, electrophysiological studies have shown that cortical stimulation induces monosynaptic EPSPs in cholinergic interneurons. It has been proposed that the negative anatomical findings might have been the result of incomplete labeling of distal dendrites. In the present study, we reinvestigated this issue using m2 muscarinic receptor antibodies as a selective marker for cholinergic and somatostatinergic interneurons in the striatum. This was combined with injections of either the anterograde tracer biotinylated dextran amine (BDA) in the monkey prefrontal cortex or aspiration lesion of the sensorimotor cortex in rats. The results showed that, in both species, a small percentage (1-2%) of cortical terminals make asymmetric synaptic contacts with m2-immunoreactive interneurons in the striatum. Interestingly, the majority of these synapses are onto small dendritic spines or spine-like appendages, as opposed to dendritic shafts and/or cell bodies. Thus, m2-containing striatal interneurons do receive direct cortical inputs and can, therefore, integrate and modulate cortical information flow through the striatum. Although the density of cortical terminals in contact with individual striatal interneurons is likely to be relatively low compared to the massive cortical input to projection neurons, both cholinergic and somatostatinergic interneurons display intrinsic properties that allow even small and distal inputs to influence their overall state of neuronal activity.

Animals↗

Afferents from the auditory thalamus synapse on inhibitory interneurons in the lateral nucleus of the amygdala.

Physiological studies suggest that afferents to the lateral nucleus of the amygdala (LA) from the auditory thalamus initiate feedforward inhibition [Li et al. (1996b)]. This model of neural processing requires that thalamic afferents synapse directly onto inhibitory interneurons. To determine whether such synaptic contacts occur, we combined anterograde tract tracing with interneuron immunocytochemistry. The anterograde tracer biotinylated dextran amine (BDA) was injected into the auditory thalamus. Inhibitory interneurons in the LA were identified using antibodies directed against gamma aminobutyric acid (GABA) or one of the calcium binding proteins (CBPs), parvalbumin (PARV), calbindin (CALB), or calretinin (CALR), since CBPs identify distinct populations of GABAergic cells within the amygdala. The distribution of GABAergic and CBP interneurons in each subregion of the LA was examined by light microscopy and the relationships between thalamo-amygdala terminals and interneurons were examined by confocal and electron microscopy. Immunoreactive cells were distributed in all three subdivisions of LA, except for CALR-ir neurons, which were sparse in the dorsal subregion and were found mainly in the ventromedial and ventrolateral subregions. Confocal microscopy revealed some thalamo-amygdala terminals in close proximity to LA interneurons, while electron microscopy showed that thalamo-amygdala terminals made direct synaptic contacts onto distal dendritic processes of inhibitory neurons. These data provide morphological evidence that thalamic afferents synapse directly onto inhibitory interneurons in LA, and are consistent with the possibility that inputs from the auditory thalamus initiate feedforward inhibition in LA. This architecture could play an important role in the suppression of background neural noise, thereby enhancing the response of LA cells to incoming auditory stimuli.

Afferent Pathways↗

Apoptosis of spinal interneurons induced by sciatic nerve axotomy in the neonatal rat is counteracted by nerve growth factor and ciliary neurotrophic factor.

We have previously shown that not only motoneurons and dorsal root ganglion cells but also small neurons, presumably interneurons in the spinal cord, may undergo apoptotic cell death as a result of neonatal peripheral nerve transection in the rat. With the aid of electron microscopy, we have here demonstrated that apoptosis in the spinal cord is confined to neurons and does not involve glial cells at the survival time studied (24 hours). To define the relative importance of the loss of a potential target (motoneuron) and a potential afferent input (dorsal root ganglion cell) for the induction of apoptosis in interneurons in this situation, we have compared the distributions and time courses for TUNEL labeling, which detects apoptotic cell nuclei, in the L5 segment of the spinal cord and the L5 dorsal root ganglion after sciatic nerve transection in the neonatal (P2) rat. In additional experiments, we studied the effects on TUNEL labeling of interneurons after treatment of the cut sciatic nerve with either ciliary neurotrophic factor (CNTF) to rescue motoneurons or nerve growth factor (NGF) to rescue dorsal root ganglion cells. The time courses of the TUNEL labeling in motoneurons and interneurons induced by the lesion show great similarities (peak at 8-48 hours postoperatively), whereas the labeling in dorsal root ganglion cells occurs later (24-72 hours). Both CNTF and NGF decrease the number of TUNEL-labeled interneurons, but there is a regional difference, in that CNTF preferentially saves interneurons in deep dorsal and ventral parts of the spinal cord, whereas the rescuing effects of NGF are seen mainly in the superficial dorsal horn. The results are interpreted as signs of a trophic dependence on both the target and the afferent input for the survival of interneurons neonatally.

Afferent Pathways↗

Synaptic connections of distinct interneuronal subpopulations in the rat basolateral amygdalar nucleus.

Although it is well established that the activity of pyramidal projection neurons in the basolateral amygdala (ABL) is controlled by gamma-aminobutyric acid (GABA)ergic inhibitory interneurons, very little is known about the connections of specific interneuronal subpopulations in this region. In the present study, immunohistochemical techniques were used at the light and electron microscopic levels to identify specific populations of interneurons and to analyze their connections with each other and with unlabeled presumptive pyramidal neurons. Double-labeling immunofluorescence experiments revealed that antibodies to vasoactive intestinal peptide (VIP) and calbindin-D28K (CB) labeled two separate interneuronal subpopulations in the ABL. Light microscopic double-labeling immunoperoxidase experiments demonstrated that many VIP-positive (VIP+) axon terminals formed intimate synaptic-like contacts with the CB-positive (CB+) neurons and that both CB+ and VIP+ terminals often contributed to the formation of pericellular baskets that surrounded unlabeled perikarya of pyramidal neurons. By using a dual immunoperoxidase/immunogold-silver procedure at the ultrastructural level, it was found that 30% of VIP+ terminals in the anterior subdivision of the basolateral nucleus innervated interneurons that were either CB+ (25%) or VIP+ (5%). A smaller percentage (15%) of CB+ terminals formed synapses with labeled interneurons. Both VIP+ and CB+ terminals also innervated unlabeled perikarya, dendrites, and spines, most of which probably belonged to pyramidal neurons. The interconnections between interneurons may be important for disinhibitory mechanisms and the mediation of rhythmic oscillations in the ABL.

Amygdala↗

Commissurally projecting inhibitory interneurons of the rat hippocampal dentate gyrus: a colocalization study of neuronal markers and the retrograde tracer Fluoro-gold.

Improved methods for detecting neuronal markers and the retrograde tracer Fluoro-Gold (FG) were used to identify commissurally projecting neurons of the rat hippocampus. In addition to the dentate hilar mossy cells and CA3 pyramidal cells shown previously to transport retrograde tracers after injection into the dorsal hippocampus, FG-positive interneurons of the dentate granule cell layer and hilus were detected in numbers greater than previously reported. FG labeling of interneurons was variable among animals, but was as high as 96% of hilar somatostatin-positive interneurons, 84% of parvalbumin-positive cells of the granule cell layer and hilus combined, and 33% of hilar calretinin-positive cells. By comparison, interneurons of the dentate molecular layer and all hippocampal subregions were conspicuously FG-negative. Whereas hilar mossy cells and CA3 pyramidal cells were FG-labeled throughout the longitudinal axis, FG-positive interneurons exhibited a relatively homotopic distribution. "Control" injections of FG into the neocortex, septum, and ventral hippocampus demonstrated that the homotopic labeling of dentate interneurons was injection site-specific, and that the CA1-CA3 interneurons unlabeled by contralateral hippocampal FG injection were nonetheless able to transport FG from the septum. These data suggest a hippocampal organizing principle according to which virtually all commissurally projecting hippocampal neurons share the property of being monosynaptic targets of dentate granule cells. Because granule cells innervate their exclusively ipsilateral target cells in a highly lamellar pattern, these results suggest that focal granule cell excitation may result in commissural inhibition of the corresponding "twin" granule cell lamella, thereby lateralizing and amplifying the influence of the initiating discharge.

Animals↗

The morphology of two groups of spiking local interneurons in the metathoracic ganglion of the locust.

Two bilaterally symmetrical groups of spiking local interneurons are described in a segmental ganglion of the locust. Interneurons in both groups are excited by specific sets of sensory receptors on one leg. The cell bodies of the anterior-lateral group lie amongst approximately 40 small cell bodies at the anterior of the ganglion, close to the lateral edge of an anterior connective. Interneurons in this group have primary neurites in Ventral Commissure I ( VCI ), and dorsoventral processes in the Oblique Tract, which divide the extensive neuropilar branches into distinct ventral and dorsal regions. Cell bodies of the midline group lie amongst a group of approximately 100 small cell bodies near the ventral midline. Interneurons in this group have primary neurites in Ventral Commissure II ( VCII ), and dorsoventral processes in the Perpendicular Tract, which divide the neuropilar branches into dorsal and ventral regions. The ventral branches of interneurons in both groups are numerous and of a uniform texture, whereas the dorsal branches are sparse and varicose. The ventral branches project to the same ventral areas of neuropil as the afferents from some hairs on a hind leg. The dorsal branches of a midline interneuron and the branches of a leg motor neuron that it affects project to the same dorsal area of neuropil. Some midline interneurons receive direct inputs from leg hair afferents and make direct connections with leg motor neurons.

Animals↗

Regional specialization in synaptic input and output in an identified local nonspiking interneuron of the crayfish revealed by light and electron microscopy.

Lateral inhibition of several mechanosensory interneurons in the crayfish terminal ganglion is mediated by a pair of identified local nonspiking interneurons called local directionally selective (LDS) interneurons. The ultrastructure and synaptic distribution of these interneurons were investigated by using intracellular labeling with horseradish peroxidase for electron microscopy. The LDS interneuron has bilateral, asymmetric arborizations. They are connected by a thick transverse neurite on which no synapses are made. The neurites on the side ipsilateral to the cell body are smooth and receive almost entirely input synapses. On the other hand, the contralateral neurites have prominent numerous swellings or varicosities from which many short fine spines arise, with intermingled input and output synapses. Both synapses, however, are not always associated with varicosities and spines. Presynaptic neurites of the LDS interneuron contain only round agranular synaptic vesicles in the vicinity of synaptic specializations. These morphological findings correlate well with the physiology of this interneuron: it is functionally polarized to receive input on one side (ipsilateral to the cell body) of the ganglion and convey it to the other side.

Afferent Pathways↗