Presynaptic inhibition of olfactory receptor neurons in crustaceans

Presynaptic inhibition of transmitter release from primary sensory afferents is a common strategy for regulating sensory input to the arthropod central nervous system. In the olfactory system, presynaptic inhibition of olfactory receptor neurons has been long suspected, but until recently could not be demonstrated directly because of the difficulty in recording from the afferent nerve terminals. A preparation using the isolated but intact brain of the spiny lobster in combination with voltage-sensitive dye staining has allowed stimulus-evoked responses of olfactory receptor axons to be recorded selectively with optical imaging methods. This approach has provided the first direct physiological evidence for presynaptic inhibition of olfactory receptor neurons. As in other arthropod sensory systems, the cellular mechanism underlying presynaptic afferent inhibition appears to be a reduction of action potential amplitude in the axon terminal. In the spiny lobster, two inhibitory transmitters, GABA and histamine, can independently mediate presynaptic inhibition. GABA- and histaminergic interneurons in the lobster olfactory lobe (the target of olfactory receptor neurons) constitute dual, functionally distinct inhibitory pathways that are likely to play different roles in regulating primary olfactory input to the CNS. Presynaptic inhibition in the vertebrate olfactory system is also mediated by dual inhibitory pathways, but via a different cellular mechanism. Thus, it is possible that presynaptic inhibition of primary olfactory afferents evolved independently in vertebrates and invertebrates to fill a common, fundamental role in processing olfactory information.     

Neuromodulation of arthropod mechanosensory neurons

Arthropod mechanosensory afferents have long been known to receive efferent synaptic connections onto their centrally located axon terminals. These connections cause presynaptic inhibition by attenuating the action potentials arriving at the axon terminals, thus reducing the synaptic potentials in the postsynaptic neurons. This type of inhibition can specifically reduce the excitation of selected postsynaptic neurons while leaving others unaffected. However, recent research has demonstrated that sensory signals detected by arthropod mechanosensory neurons can also be synaptically modulated before they ever arrive at the axon terminals. In arachnids and crustaceans, wide and complex networks of synapses on all parts of the afferent neurons, including the somata and dendrites, provide mechanisms to inhibit or enhance the responses to mechanical stimuli as they are being detected. This modulation will affect the signal transmission to all axonal branches and postsynaptic cells of the affected receptor neuron. In addition to the increased complexity of mechanosensory information transmission produced by these synapses, a variety of circulating neuroactive substances also modulate these neurons by acting on their postsynaptic receptors.     

Synaptic morphology of inner and outer hair cells of the human organ of Corti

Innervations of inner and outer hair cells of the organ of Corti of the human cochlea were studied by serial section electron microscopy. At the base of inner hair cells, presumed afferent fibers were of varying size and demonstrated synaptic specialization consisting of a presynaptic body, vesicles, and asymmetrical synaptic membrane specialization. Two types of neurons, vesiculated presumably efferent and nonvesiculated presumably afferent, synapsed at the base of outer hair cells. The synaptic specialization of afferent fibers included presynaptic body, vesicles, and asymmetrical membrane thickening, whereas efferent synapses demonstrated presynaptic vesicles and a subsynaptic cisterna. Some presumably afferent nerve terminals formed a reciprocal synapse with outer hair cells in both the human and the chimpanzee. Such a synaptic relationship demonstrated morphologic specialization consistent with both hair cell-to-neuron and neuron-to-hair cell transmission between the same outer hair cell and nerve terminal. The innervation density of inner and outer hair cells and the comparative anatomy of the afferent and efferent innervation are discussed.     

Freeze-fracture organization of hair cell synapses in the sensory epithelium of guinea pig organ of Corti

The fine structure of both the afferent and efferent hair cell synapses in the sensory epithelium of guinea pig organ of Corti was examined by freeze-fracture electron microscopy. In the afferent synapse, barlike aggregates of intramembrane particles (IMPs) of about 10 nm in diameter were seen on the P-face of the afferent presynaptic membrane directly beneath the presynaptic dense projection which is located in the active zone of the presynaptic membrane. Small and large depressions have been seen on the presynaptic membrane. The former were observed in the proximity of the barlike aggregates, while the latter were observed some distance from the aggregate. In outer hair cells, IMPs of about 10 nm in diameter were seen on the P-face of the afferent postsynaptic membrane at a density of 3,000/μm². In the efferent synapse, many aggregates composed of from several to tens of large IMPs of 13 nm in diameter were observed on the presynaptic membrane. These aggregates were localized to small membrane depressions, which tended to be deeper as particle number per aggregate increased. Dense populations of IMPs of about 9 nm in diameter were observed on the P-face of the efferent postsynaptic membrane at a density of 4,000/μm². A fenestrated subsynaptic cistern completely covers the efferent postsynaptic membrane. Moreover, the subsynaptic cistern spans several efferent postsynaptic membranes when efferent synapses are gathered in a group. In the afferent and efferent synapses of hair cells, specializations of the synaptic membranes were represented by marked aggregates characteristic of IMPs. In the efferent synapse, IMP movement inside the synaptic membrane was proposed in relationship to retrival of synaptic vesicle membrane. Structural relationship between the subsynaptic cistern and efferent postsynaptic membrane was revealed.     

GABAergic synapses in the brain identified with antisera to GABA and its synthesizing enzyme,glutamate decarboxylase

GABA is a known inhibitory neurotransmitter in the mammalian brain. The site of GABAergic synapses can be determined with immunocytochemical methods that localize either GABA or its synthesizing enzyme, glutamate decarboxylase (GAD). In general, GABAergic axon terminals contain pleomorphic synaptic vesicles and form symmetric synapses. However, a small number of GABAergic axon terminals in selected brain regions (spinal cord, cerebellum, superior colliculus, striatum, globus pallidus, inferior olive, and substantia nigra) form asymmetric synapses. GAD- and GABA-immunoreactive processes that contain synaptic vesicles participate in every known morphological type of chemical synapse. These include axosomatic, axodendritic, axospinous, initial segment, axoaxonic, dendrodendritic, serial, reciprocal, and ribbon synapses. Although GABAergic synapses form a heterogeneous group, they most commonly form axosomatic, axodendritic, and initial segment synapses in the brain and spinal cord. These findings provide helpful guidelines for the identification of GABAergic synapses in future studies.     

Synaptic structure, distribution, and circuitry in the central nervous system of the locust and related insects.

The Orthopteran central nervous system has proved a fertile substrate for combined morphological and physiological studies of identified neurons. Electron microscopy reveals two major types of synaptic contacts between nerve fibres: chemical synapses (which predominate) and electrotonic (gap) junctions. The chemical synapses are characterized by a structural asymmetry between the pre- and postsynaptic electron dense paramembranous structures. The postsynaptic paramembranous density defines the extent of a synaptic contact that varies according to synaptic type and location in single identified neurons. Synaptic bars are the most prominent presynaptic element at both monadic and dyadic (divergent) synapses. These are associated with small electron lucent synaptic vesicles in neurons that are cholinergic or glutamatergic (round vesicles) or GABAergic (pleomorphic vesicles). Dense core vesicles of different sizes are indicative of the presence of peptide or amine transmitters. Synapses are mostly found on small-diameter neuropilar branches and the number of synaptic contacts constituting a single physiological synapse ranges from a few tens to several thousand depending on the neurones involved. Some principles of synaptic circuitry can be deduced from the analysis of highly ordered brain neuropiles. With the light microscope, synaptic location can be inferred from the distribution of the presynaptic protein synapsin I. In the ventral nerve cord, identified neurons that are components of circuits subserving known behaviours, have been studied using electrophysiology in combination with light and electron microscopy and immunocytochemistry of neuroactive compounds. This has allowed the synaptic distribution of the major classes of neurone in the ventral nerve cord to be analysed within a functional context.     

Distribution of GABA and glycine receptors on bipolar and ganglion cells in the mammalian retina

The amino acids GABA and glycine mediate synaptic transmission via specific neurotransmitter receptors. Molecular cloning studies have shown that there is a great diversity of GABA and glycine receptors. In the present article, the distribution of GABA and glycine receptors on identified bipolar and ganglion cell types in the mammalian retina is reviewed. Immunofluorescence obtained with antibodies against GABA and glycine receptors is punctate. Electron microscopy shows that the puncta represent a cluster of receptors at synaptic sites. Bipolar cell types were identified with immunohistochemical markers. Double immunofluorescence with subunit-specific antibodies was used to analyze the distribution of receptor clusters on bipolar axon terminals. The OFF cone bipolar cells seem to be dominated by glycinergic input, whereas the ON cone bipolar and rod bipolar cells are dominated by GABAergic input. Ganglion cells were intracellularly injected with Neurobiotin, visualized with Streptavidin coupled to FITC, and subsequently stained with subunit specific antibodies. The distribution and density of receptor clusters containing the alpha1 subunit of the GABA(A) receptor and the alpha1 subunit of the glycine receptor, respectively, were analyzed on midget and parasol cells in the marmoset (a New World monkey). Both GABA(A) and glycine receptors are distributed uniformly along the dendrites of ON and OFF types of parasol and midget ganglion cells, indicating that functional differences between these subtypes of ganglion cells are not determined by GABA or glycinergic input.     

Immunoelectron microscopic localization of neurotransmitters in the cochlea

This paper presents the works and methods of our respective laboratories using electron microscopic immunocytochemistry to identify and localize cochlear neurotransmitters. Antibodies to various prospective neurotransmitters and associated enzymes have been used to study the ultrastructural localization of several candidates for olivocochlear efferent neurotransmitters previously suggested by light microscopic immunocytochemistry. Antibodies against enkephalins label lateral olivocochlear efferent fibers. Antibodies against choline acetyltransferase (ChAT) (an enzyme marker for acetylcholine) label a major population of both lateral and medial efferent fibers and terminals, whereas antibodies to γ-aminobutyric acid (GABA) label what might be a small subpopulation of both the lateral and medial efferent systems. The GABA-like immunostained medial efferent fibers are preferentially located in the upper turns of the guinea pig cochlea, particularly the third turn. Immunoelectron microscopy shows that neither GABA nor ChAT immunolabels all medial efferent terminals, regardless of cochlear turn. All the different types of immunolabeled efferent terminals have been observed to make characteristic synaptic contacts; lateral efferent terminals on afferent dendrites and medial efferent terminals on outer hair cells and occasionally on type II afferent dendrites. Other types of contacts involving GABA-like, and sometimes met-enkephalin-like, immunostained fibers are occasionally seen particularly in the upper turns of the cochlea. Immunoelectron microscopic results suggest that both medial and lateral efferent systems might be further subdivided on the basis of differences in neurotransmitters. Future trends of immunocytochemical research on cochlear neurotransmitters are proposed, particularly colocalization studies, which show a complex pattern of coexistence of neurotransmitters in the lateral efferent system.     

Application of the Golgi/electron microscopy technique for cell identification in immunocytochemical, retrograde labeling, and developmental studies of hippocampal neurons.

In this study the Golgi/electron microscopy (EM) technique has been used for an analysis of the fine structure, specific synaptic connections, and differentiation of neurons in the hippocampus and fascia dentata of rodents. In a first series of experiments the specific synaptic contacts formed between cholinergic terminals and identified hippocampal neurons were studied. By means of a variant of the section Golgi impregnation procedure, Vibratome sections immunostained for choline acetyltransferase, the acetylcholine-synthesizing enzyme, were Golgi-impregnated in order to identify the target neurons of cholinergic terminals in the hippocampus. It could be shown with this combined approach that cholinergic septohippocampal fibers form a variety of synapses with different target structures of the Golgi-impregnated and gold-toned hippocampal neurons. In this report cholinergic synapses on the heads of small spines, the necks of large complex spines, dendritic shafts, and cell bodies of identified dentate granule cells are described. The variety of cholinergic synapses suggests that cholinergic transmission in the fascia dentata is a complex event. Next, the Golgi/EM technique was applied to Vibratome sections that contained retrogradely labeled neurons in the hilar region of the fascia dentata following horseradish peroxidase (HRP) injection into the contralateral hippocampus. With this combined approach some of the hilar cells projecting to the contralateral side were identified as mossy cells by the presence of retrogradely transported HRP in thin sections through these Golgi-impregnated and gold-toned neurons. Our findings suggest that the mossy cells are part of the commissural/associational system terminating in the inner molecular layer of the fascia dentata. They are mainly driven by hilar collaterals of granule cell axons that form giant synapses on their dendrites. Finally, the Golgi/EM procedure was used to study the differentiation and developmental plasticity of hippocampal and dentate neurons in transplants and slice cultures of hippocampus. Under both experimental conditions, the differentiating neurons are deprived of their normal laminated afferent innervation but develop their major cell-specific characteristics including a large number of postsynaptic structures (spines). As revealed in thin sections of gold-toned identified cells, all these spines formed synapses with presynaptic boutons suggesting sprouting of the transplanted and cultured neurons, respectively. Altogether, the present report demonstrates the usefulness of the Golgi/EM technique, particularly of the section impregnation procedure, for a variety of studies requiring the identification of individual neurons at the ultrastructural level.     

Serial reconstruction of inner hair cell afferent innervation using semithick sections

The afferent innervation pattern of inner hair cells in the apex of the guinea pig cochlea was studied using serial reconstruction of semithick (0.25–μm) sections and high-voltage electron microscopy (HVEM). This thickness produced a good compromise between the ability to resolve details of the synaptic contacts between the hair cells and sensory neurons and the number of sections required to reconstruct the nerve terminals within the receptor organ. The use of a goniometer allowed the sections to be tilted to angles optimum for viewing either the synaptic membrane specializations or the presynaptic bodies. Reasonably good images of 0.25-μm sections could be obtained using a conventional 120-keV microscope, but the images produced by the HVEM were clearly superior. The sensory nerve terminals and hair cells were reconstructed using a microcomputer-based computer-aided-design system. Nerve terminals with complex shapes could be successfully rendered as surface models viewed as stereo pairs. The advantages and limitations of the techniques used are discussed.     

Efferent controls in crustacean mechanoreceptors

Since the 1960s it has been known that central neural networks can elaborate motor patterns in the absence of any sensory feedback. However, sensory and neuromodulatory inputs allow the animal to adapt the motor command to the actual mechanical configuration or changing needs. Many studies in invertebrates, particularly in crustacea, have described several mechanisms of sensory-motor integration and have shown that part of this integration was supported by the efferent control of the mechanosensory neurons themselves. In this article, we review the findings that support such an efferent control of mechanosensory neurons in crustacea. Various types of crustacean proprioceptors feeding information about joint movements and strains to central neural networks are considered, together with evidence of efferent controls exerted on their sensory neurons. These efferent controls comprise (1) the neurohormonal modulation of the coding properties of sensory neurons by bioamines and peptides; (2) the presynaptic inhibition of sensory neurons by GABA, glutamate and histamine; and (3) the long-term potentiation of sensory-motor synapses by glutamate. Several of these mechanisms can coexist on the same sensory neuron, and the functional significance of such multiple modulations is discussed.     

Presynaptic proteins of ribbon synapses in the retina

The synapses of photoreceptors and bipolar cells in the retina are characterized ultrastructurally by the presence of an electron-dense bar, the synaptic ribbon, lying perpendicular to the plasma membrane at the active zone and extending about 0.5 microm into the cytoplasm. Hence, these synapses are known as ribbon synapses. All neurons that make ribbon synapses release neurotransmitter tonically. That is, neurotransmitter is released continuously from these neurons and the rate of release is modulated in response to graded changes in the membrane potential. This contrasts with action potential-driven, phasic release from other neurons. Similar to other synapses, neurotransmitter is released at ribbon synapses by the calcium-dependent exocytosis of synaptic vesicles. Most components of the molecular machinery governing transmitter release are conserved between ribbon and conventional synapses, but several differences that may be important determinants of tonic transmitter release have been identified in the retina by immunohistochemistry. For example, the presynaptic calcium channels of bipolar cells and photoreceptors are different from those elsewhere in the brain. Differences have also been found in the proteins involved in synaptic vesicle recruitment to the active zone and in synaptic vesicle fusion. These differences and others are discussed in terms of their implications for neurotransmitter release from photoreceptors and bipolar cells in the retina.     

Technique to study three-dimensional spatial arrangement of synaptic vesicles using data from single sections

Synaptic vesicles are membrane-bound organelles storing neurotransmitters in presynaptic terminals and releasing them into the synaptic cleft. Coordinated movements of synaptic vesicles relate to synaptic function and their spatial arrangement can provide useful information about the activity of a synapse. This article presents a technique to extract quantitative information about three-dimensional (3D) spatial arrangement of synaptic vesicles from measurements performed on single ultrathin random sections of a presynaptic terminal. The technique presumes quantification of a 2D density as well as 2D spatial pattern formed by vesicle profiles using a minimum spanning tree (MST) algorithm, in digitized micrographs of a presynaptic terminal. Further, original software was used to simulate a 3D spatial arrangement of synaptic vesicles and their random sectioning. A 3D density and pattern of synaptic vesicles were used as basic input parameters of the model, while a 2D density and MST quantities for vesicle profiles served as output, model-derived parameters allowing one to compare and fit simulated distributions to experimental ones. Pilot simulations performed to check the validity of the technique have shown that a 2D density and MST quantities of vesicle profiles closely relate to a 3D density and spatial pattern of vesicles. The technique was demonstrated in the analysis of spatial distribution of synaptic vesicles in axonal terminals forming asymmetric synaptic densities in the stratum radiatum of the CA1 subfield of the murine hippocampus.     

Electrical properties of chemoreceptor elements in the carotid body

The electrical properties of chemoreceptor afferent nerve fibers and glomus cells and the behavior of cytosolic Ca(2+) in glomus cells are reviewed. While this has not been confirmed, spontaneously depolarizing potentials (SDPs) recorded in a chemoreceptor afferent terminal may be the postsynaptic expression of presynaptic events. Glomus cells, which are presynaptic elements, either depolarized or hyperpolarized in response to natural and chemical stimulation. After-hyperpolarization following an initial depolarization and after-depolarization following an initial hyperpolarization were often seen. When a glomus cell depolarizes, voltage noise increases despite a decrease in input resistance in both intact and denervated carotid bodies. The voltage noise may be "receptor noise" generated in the glomus cell itself. The electrical properties of glomus cells change in the denervated carotid body, which suggests that the chemoreceptor afferent nerve exerts some trophic effect(s) on glomus cells. Hypoxia either increases or decreases cytosolic Ca(2+), while ACh or NaCN induces either an increase or no change in cytosolic Ca(2+) in glomus cells. There are at least two possible explanations for voltage changes in glomus cells: a chemical stimulus first depolarizes the glomus cell and induces Ca(2+) influx to release chemical substances, or a chemical stimulus induces an increase in [Ca(2+)](i) and then hyperpolarizes the glomus cell via potassium influx.     

Motion detectors in the locust visual system: From biology to robot sensors

Motion detectors in the locust optic lobe and brain fall into two categories: neurones that respond selectively to approaching vs. receding objects and neurones that respond selectively to a particular pattern of image motion over a substantial part of the eye, generated by the locust's own movements through its environment. Neurones from the two categories can be differentiated on the basis of their response to motion at a constant velocity at a fixed distance from the locust: neurones of the first category respond equally well to motion in any direction whereas neurones in the second category respond selectively to one preferred direction of motion. Several of the motion detectors of the first category, responding to approaching objects, share the same input organisation, suggesting that it is important in generating a tuning for approaching objects. Anatomical, physiological, and modelling studies have revealed how the selectivity of the response is generated. The selectivity arises as a result of a critical race between excitation, generated when image edges move out over the eye and delayed inhibition, generated by the same edge movements. For excitation to build up, the velocity and extent of edge motion over the eye must increase rapidly. The ultrastructure of the afferent inputs onto the dendrites of collision sensitive neurones reveals a possible substrate for the interaction between excitation and inhibition. This interpretation is supported by both physiological and immunocytochemical evidence. The input organisation of these neurones has been incorporated into the control structure of a small mobile robot, which successfully avoids collisions with looming objects. The ecological role of motion detectors of the second category that respond to image motion over a substantial part of the visual field, is discussed as is the input organisation that generates this selective response. The broad tuning of these neurones, particularly at low velocities (<0.02 degree/s), suggests they may have a role in navigation during migratory flights at altitude. By contrast, their optimum tuning to high-image velocities suggests these motion detectors are adapted for use in a fast flying insect, which does not spend significant time hovering.     

Neural input and neural control of the subcommissural organ

The neural control of the subcommissural organ (SCO) has been partially characterized. The best known input is an important serotonergic innervation in the SCO of several mammals. In the rat, this innervation comes from raphe nuclei and appears to exert an inhibitory effect on the SCO activity. A GABAergic innervation has also been shown in the SCO of the rat and frog Rana perezi. In the rat, GABA and the enzyme glutamate decarboxylase are involved in the SCO innervation. GABA is taken up by some secretory ependymocytes and nerve terminals, coexisting with serotonin in a population of synaptic terminals. Dopamine, noradrenaline, and different neuropeptides such as LH-RH, vasopressin, vasotocin, oxytocin, mesotocin, substance P, alpha-neoendorphin, and galanin are also involved in SCO innervation. In the bovine SCO, an important number of fibers containing tyrosine hydroxylase are present, indicating that in this species dopamine and/or noradrenaline-containing fibers are an important neural input. In Rana perezi, a GABAergic innervation of pineal origin could explain the influence of light on the SCO secretory activity in frogs. A general conclusion is that the SCO cells receive neural inputs from different neurotransmitter systems. In addition, the possibility that neurotransmitters and neuropeptides present in the cerebrospinal fluid may also affect the SCO activity, is discussed.     

The ultrastructure of neuronal-pinealocytic interconnections in the monkey pineal.

Recent ultrastructural studies of neuronal-pinealocytic interconnections in the monkey pineal are reviewed. The pinealocytes in the adult monkey show almost all of the cytological specializations known in subprimate mammals. Adjacent pinealocytes are functionally coupled through ribbon synapses on cell bodies and gap junctions on cell bodies and cell processes. The pinealocytes receive direct synpatic contacts of nerve fibers with cholinergic terminal morphology. Nerve cells restricted to the central portion of the pineal receive synaptic contacts with more than three different morphologically defined types of nerve terminals. In addition to nerve terminals containing small clear vesicles or vesicles of pleomorphic morphology, a pinealocyte's terminal process containing the synaptic ribbon forms a true synaptic contact on the nerve cell body. The diversity of synapses on these nerve cells strongly suggests multiple origins of these neurons rather than a single peripheral parasympathetic origin. The possible involvement of pineal neurons in an intrinsic circuit that regulates the function of pinealocytes and integrates the neural input from the central as well as the peripheral nervous systems is discussed.     

Ultrastructural description of glutamate-, aspartate-, taurine-, and glycine-like immunoreactive terminals from five rat brain regions

The ultrastructural localization of putative excitatory (glutamate, aspartate) and inhibitory (taurine, glycine) amino acid neurotransmitters is described in several selected rat brain regions. In general, axon terminal profiles immunoreactive for excitatory amino acids formed asymmetric synapses with non-immunoreactive small diameter dendritic profiles or dendritic spines. In the cerebellum, both mossy fiber terminals and parallel fiber terminals were immunoreactive for glutamate and aspartate. In the hippocampus, mossy fiber terminals within the stratum lucidum of the CA3 region were immunoreactive for glutamate. Localization of glutamate and aspartate to cerebellar parallel and mossy fibers, as well as the identification of glutamate in hippocampal mossy fibers, is consistent with the excitatory nature of these fibers as described in previous physiological studies. Glutamate-like immunoreactive terminals were also identified in subnucleus caudalis of the spinal trigeminal nucleus and in the dorsal horn of the spinal cord. Immunoreactive axon terminals for two putative inhibitory neurotransmitters, glycine and taurine, displayed a greater number of morphological variations in synaptic structure. In the cerebellum, taurine-like immunoreactivity was present in both basket cell axon terminals which formed symmetric synapses with Purkinje cell neurons, and in a few mossy fiber terminals which formed asymmetric synapses with dendritic spines. In the area dentata of the hippocampus, taurine-like immunoreactive profiles formed asymmetric synapses with dendritic elements. Glycine-like immunoreactive terminals formed symmetric synapses with cell perikarya in both the ventral horn of the spinal cord and in the cochlear nuclei, and on axon terminals in the spinal trigeminal and cochlear nuclei. In contrast, some glycine-like immunoreactive terminals formed asymmetric synapses with distal dendritic profiles in the spinal cord and spinal trigeminal nucleus. The localization of taurine to cerebellar basket cell axons and glycine to axon terminals that synapse on ventral horn motor neuron perikarya is consistent with the hypothesis that these amino acids are functioning as inhibitory neurotransmitters at these synapses. Taurine localization to cerebellar mossy fibers and to fibers in the molecular layer of the dentate gyrus may be more consistent with a proposed neuromodulator role of taurine.     

A review of recent observations concerning the synaptic organization of the basilar pontine nuclei

Ultrastructural studies are described that have identified in the basilar pontine nuclei (BPN), the synaptic boutons formed by the corticopontine, cerebellopontine, tectopontine, and dorsal column nucleipontine afferent projection systems. In addition, immunocytochemical studies visualized neuronal somata, dendrites, and synaptic boutons that contain immunoreactivity for GABA or the synthesizing enzyme glutamic acid decarboxylase (GAD). Based upon differences in the mode of degeneration and postsynaptic locus of degenerative synaptic boutons in the BPN, it is suggested that two types of cortical neurons and three classes of deep cerebellar nuclear cells project to the BPN. For similar reasons, it appears that two types of neurons in the dorsal column nuclei project to the BPN while only one type of afferent synaptic bouton takes origin from the superior colliculus. Furthermore it appears that the population of BPN neurons projecting to the paramedian lobule receives convergent inputs from the cutaneous periphery and the corresponding region of sensorimotor cortex. Studies employing GAD immunohistochemistry indicate that GABA-ergic neurons and axon terminals are present in the BPN and thus support the suggestion that a local inhibitory interneuron is present within the BPN. Taken together these observations suggest that basilar pontine neurons might play a more active role in the integration of various types of information destined for the cerebellar cortex than has previously been recognized.     

Peripheral synapses and giant neurons in whip spiders

Among invertebrates the synapses between neurons are generally restricted to ganglia, i.e., to the central nervous system (CNS). As an exception, synapses occur in the sensory nerves of arachnid legs, indicating that some nervous integration is already taking place far out in the periphery. In the antenniform legs of whip spiders (Amblypygi), a very special synaptic circuit is present. These highly modified legs contain several large interneurons (giant neurons) that receive mechanosensory input from 700-1,500 tarsal bristles. Some of the sensory cell axons contact a giant neuron at its short, branched dendrite, a few at the soma, but most synapse onto the long giant axon. The fine structure of these synapses resembles that of typical chemical synapses in other arthropods. Although thousands of sensory fibers converge on a single giant neuron, there is no reduction in the actual number of sensory fibers, because these afferent fibers continue their course to the CNS after having made several en passant synapses onto the giant neuron. Touching a single tarsal bristle is sufficient to elicit action potentials in a giant neuron. Owing to the large diameter of the giant axon (10-20 microm), the action potentials reach the CNS within 55 ms, at conduction velocities of up to 7 m/s. However, mechanical stimulation of the tarsal bristles does not elicit a fast escape response, in contrast to giant fiber systems in earthworms, certain insects, and crayfishes. A quick escape is observed in whip spiders, but only after stimulation of the filiform hairs (trichobothria) on the regular walking legs. Although the giant fiber system in the antenniform legs undoubtedly provides a fast sensory pathway, its biological significance is not clearly understood at the moment.