[Efferent innervation of pulmonary blood vessels and bronchi in rat (an immunohistochemical study)].

In this investigation the peculiarities of innervation of bronchi and blood vessels of the lung were studied in 20 rats using immunohistochemical demonstration of synaptophysin and alpha-actin. The results obtained have showen that the densest innervation is typical for bronchial walls, particularly, for the muscular lamina. Synaptophysin-immunoreactive terminals (SFIT) were detected in the bronchi in close association with both circular bundles of smooth muscle cells and microganglia. Dense network of SFIT was found in the pulmonary vein--in its middle tunic formed by cardiomyocytes. In contrast to the bronchi and pulmonary vein, large branches of the pulmonary artery contained no SFIT. We briefly discuss the problem of the origin of the nerve fibers described and their functions and suggest that SFIT are formed by efferent fibers (axons) of neurons arising from either the intrapulmonary parasympathetic ganglia.

[Peculiarities of cortico-cortical ipsilateral connections of the primary, secondary and tertiary sensomotor zones of cat cerebral cortex].

Relative distribution of the efferent association fibers of the primary motor--MI (area 4y), secondary--SMII (2pri) and tertiary--SMIII (area 5) sensomotor zones of cerebral cortex were studied in 30 cats using Nauta-Gigax method. The projection of insignificant number of associative fibers to the primary cortical sensory zone--CI (area 1,2, 3a, 3b) was demonstrated. Massive bilateral connections of MI with SMII (2pri) and SMIII (area 5) were shown. It was suggested that the restoration of the motor functions after local destruction of CMI, CMII and CMIII is due to the demonstrated multiple horizontal associative connections between the functional units of the mentioned sensomotor centers.

[Structural basis for the inhibitory function of the parietal cortex efferent systems].

Relative quantitative distribution of all the associative and descending efferent fibers and the ultrastructural organization of the terminals of the parietal cortex areas 5 and 7 in the caudate (NC) and red nucleus (NR) in the cat were analyzed after a local, pointed destruction of the cortex of these areas. The maximal numbers of the associative fibers were found to project to the fundus areas of the motor cortex and to the area of Clare-Bishop; moderate projections were detected to the areas 31, 19 and single degenerating fibers were registered in the areas 1,2, 3a, 3b, 30, and 23. The descending efferents were maximally projecting to NC, NR, reticular nuclei of the thalamus, midbrain, and pons, in all of which, according to the immunocytochemical studies, GABA-ergic terminals are prevalent. On the basis on the electron microscopical studies, it was suggested that the influence of the parietal cortex is mediated by the axo-spinal synapses of the medium shortaxonal spiny cells of the dorsolateral part of NC caput and by the axo-dendritic synapses of Golgi II cells of the parvocellular part of NR. On the basis of the maximal involvement of the fundus areas of the motor cortex, as well as of the inhibitory subcortical (NC) and stem nuclei (NR, reticular nuclei of the thalamus, midbrain, and nuclei pontis), it is suggested that these structures serve as the morphological substrates for the realization of the inhibitory, integrative function of the parietal cortex.

Projections from the ventral nucleus of the lateral lemniscus to the cochlea in the mouse.

Auditory efferents originate in the central auditory system and project to the cochlea. Although the specific anatomy of the olivocochlear (OC) efferents can vary between species, two types of auditory efferents have been identified based upon the general location of their cell bodies and their distinctly different axon terminations in the organ of Corti. In the mouse, the relatively small somata of the lateral (LOC) efferents reside in the lateral superior olive (LSO), have unmyelinated axons, and terminate around ipsilateral inner hair cells (IHCs), primarily against the afferent processes of type I auditory nerve fibers. In contrast, the larger somata of the medial (MOC) efferents are distributed in the ventral nucleus of the trapezoid body (VNTB), have myelinated axons, and terminate bilaterally against the base of multiple outer hair cells (OHCs). Using in vivo retrograde cell body marking, anterograde axon tracing, immunohistochemistry, and electron microscopy, we have identified a group of efferent neurons in mouse, whose cell bodies reside in the ventral nucleus of the lateral lemniscus (VNLL). By virtue of their location, we call them dorsal efferent (DE) neurons. Labeled DE cells were immuno-negative for tyrosine hydroxylase, glycine, and GABA, but immuno-positive for choline acetyltransferase. Morphologically, DEs resembled LOC efferents by their small somata, unmyelinated axons, and ipsilateral projection to IHCs. These three classes of efferent neurons all project axons directly to the cochlea and exhibit cholinergic staining characteristics. The challenge is to discover the contributions of this new population of neurons to auditory efferent function.

Anatomical variations of the levator palpebrae superioris, including observations on its innervation and intramuscular nerves' distribution pattern.

BACKGROUND: The levator palpebrae superioris muscle (LPS) acts as the upper eyelid's major elevator and retractor and is innervated by the oculomotor nerve. The muscle's paralysis is manifested by ptosis. MATERIAL AND METHODS: 70 orbits were dissected. After removing the orbital roof, the LPS' shape and anatomical variations (i.e., the presence of accessory muscular bands or atypical formation of the muscle) were assessed. To visualize the distribution of the oculomotor nerve's intramuscular sub-branches, the isolated levator palpebrae superioris muscles were stained using Sihler's staining technique. RESULTS: Several LPS anatomical variations were observed in the specimens examined, in seven of which (7/70; 10%) additional delicate muscular slips arose from the LPS' lateral border and reached the lacrimal gland. Histological examination confirmed the presence of striated skeletal muscle fibers in all those cases. In three other specimens (3/70; 4.28%), supernumerary muscular bands ("tensor trochleae") were found that linked the levator with the superior oblique muscle's trochlea. In the next case, the LPS' origin was double and the muscle was bipartite on its proximal half. In most cases (55/70; 78.6%), muscular branches formed a single bundle that wrapped around the superior rectus muscle's medial border to reach the levator's inferior surface. Intramuscular sub-branches were distributed largely within the proximal two-thirds of the LPS and formed an irregular, tree-like pattern. However, thin sub-branches and small retrograde sub-branches extended as far as the muscle's insertion. CONCLUSIONS: Plastic surgeons and ophthalmologists should be aware of the levator palpebrae superioris muscle's anatomic variations both in planning and conducting surgeries on the upper eyelid.

Disappearance of afferent and efferent nerve terminals in the inner ear of the chick embryo after chronic treatment with beta-bungarotoxin.

Beta-Bungarotoxin(beta-BT) was applied to chick embryos at 3-day intervals beginning on the 4th day of incubation to see the effect of chronically and massively applied beta-BT, and to investigate the hair cell-nerve relationship in the developing inner ear by electron microscopy. On the 10th day of incubation, nerve terminals had achieved contact with differentiating hair cells, but the acoustico-vestibular ganglion cells of treated animals were decreased in number to one-third of those of the control. By the 14th day, most of the ganglion cells degenerated and disappeared, and only a few nerve terminals were seen in the neuroepithelium. At this time, most of the hair cells lacked synaptic contacts with nerve terminals; but their presynaptic specialization remained intact and they showed evidence of continuing differentiation. On the 17th day, the acoustico-vestibular ganglion cells were completely absent. All the hair cells were devoid of afferent and efferent innervation but were fully differentiated on the 21st day. Beta-BT was found to have a similar destructive effect on cultured spinal ganglion cells. The present study shows that beta-BT kills acoustico-vestibular and spinal nerve cells when applied chronically and massively during development. Furthermore, the differentiation of hair cells proceeds normally, and their presynaptic specializations are maintained when nerve terminals are absent during later developmental stages.

Distribution and structure of efferent synapses in the chicken retina.

The visual system of birds includes an efferent projection from a visual area, the isthmo-optic nucleus in the midbrain, back to the retina. Using a combination of anterograde labeling of efferent fibers, reconstruction of dye-filled neurons, NADPH-diaphorase staining, and transmission electron microscopy, we have examined the distribution of efferent fibers and their synaptic structures in the chicken retina. We show that efferent fibers terminate strictly within the ventral retina. In two completely mapped retinas, only 2 fibers from a total of 15,359 terminated in the dorsal retina. The major synapse made by each efferent fiber is with a single efferent target amacrine cell (TC). This synapse consists of 5-25 boutons of 2 microm diameter, each with multiple active zones, pressed into the TC soma or synapsing with a basketwork of rudimentary TC dendrites in the inner nuclear layer (INL). This basketwork, which is sheathed by Muller cell processes, defines a private neuropil in the INL within which TCs were also seen to receive input from retinal neurons. In addition to the major synapse, efferent fibers typically produce several very thin processes that terminate nearby in single small boutons and for which the soma of a local amacrine cell is one of the likely postsynaptic partners. A minority of efferent fibers also give rise to a thicker process, terminating in a strongly diaphorase-positive ball about 5 microm in diameter.

Forebrain projections to brainstem nuclei involved in the control of mandibular movements in rats.

Mandibular movements occur through the triggering of trigeminal motoneurons. Aberrant movements by orofacial muscles are characteristic of orofacial motor disorders, such as nocturnal bruxism (clenching or grinding of the dentition during sleep). Previous studies have suggested that autonomic changes occur during bruxism episodes. Although it is known that emotional responses increase jaw movement, the brain pathways linking forebrain limbic nuclei and the trigeminal motor nucleus remain unclear. Here we show that neurons in the lateral hypothalamic area, in the central nucleus of the amygdala, and in the parasubthalamic nucleus, project to the trigeminal motor nucleus or to reticular regions around the motor nucleus (Regio h) and in the mesencephalic trigeminal nucleus. We observed orexin co-expression in neurons projecting from the lateral hypothalamic area to the trigeminal motor nucleus. In the central nucleus of the amygdala, neurons projecting to the trigeminal motor nucleus are innervated by corticotrophin-releasing factor immunoreactive fibers. We also observed that the mesencephalic trigeminal nucleus receives dense innervation from orexin and corticotrophin-releasing factor immunoreactive fibers. Therefore, forebrain nuclei related to autonomic control and stress responses might influence the activity of trigeminal motor neurons and consequently play a role in the physiopathology of nocturnal bruxism.

Integration of lobula plate output signals by DNOVS1, an identified premotor descending neuron.

Many motion-sensitive tangential cells of the lobula plate in blowflies are well described with respect to their visual response properties and the connectivity among them. They have large and complex receptive fields with different preferred directions in different parts of their receptive fields matching the optic flow that occurs during various flight maneuvers. However, much less is known about how tangential cells connect to postsynaptic neurons descending to the motor circuits in the thoracic ganglion and how optic flow is represented in these downstream neurons. Here we describe the physiology and the connectivity of a prominent descending neuron called DNOVS1 (for descending neurons of the ocellar and vertical system). We find that DNOVS1 is electrically coupled to a subset of vertical system cells. The specific wiring leads to a preference of DNOVS1 for rotational flow fields around a particular body axis. In addition, DNOVS1 receives input from interneurons connected to the ocelli.

Mechanisms of efferent-mediated responses in the turtle posterior crista.

To study the cellular mechanisms of efferent actions, we recorded from vestibular-nerve afferents close to the turtle posterior crista while efferent fibers were electrically stimulated. Efferent-mediated responses were obtained from calyx-bearing (CD, calyx and dimorphic) afferents and from bouton (B) afferents distinguished by their neuroepithelial locations into BT units near the torus and BM units at intermediate sites. The spike discharge of CD units is strongly excited by efferent stimulation, whereas BT and BM units are inhibited, with BM units also showing a postinhibitory excitation. Synaptic activity was recorded intracellularly after spikes were blocked. Responses of BT/BM units to single efferent shocks consist of a brief depolarization followed by a prolonged hyperpolarization. Both components reflect variations in hair-cell quantal release rates and are eliminated by pharmacological antagonists of alpha9/alpha10 nicotinic receptors. Blocking calcium-dependent SK potassium channels converts the biphasic response into a prolonged depolarization. Results can be explained, as in other hair-cell systems, by the sequential activation of alpha9/alpha10 and SK channels. In BM units, the postinhibitory excitation is based on an increased rate of hair-cell quanta and depends on the preceding inhibition. There is, in addition, an efferent-mediated, direct depolarization of BT/BM and CD fibers. In CD units, it is the exclusive efferent response. Nicotinic antagonists have different effects on hair-cell efferent actions and on the direct depolarization of CD and BT/BM units. Ultrastructural studies, besides confirming the efferent innervation of type II hair cells and calyx endings, show that turtle efferents commonly contact afferent boutons terminating on type II hair cells.

Organization of the basal ganglia: the importance of axonal collateralization.

Recent neuroanatomical data obtained with single-axon or single-cell labeling procedures in both rodents and primates have revealed the presence of various types of projection neurons with profusely collateralized axons within each of the major components of the basal ganglia. Such findings call for a reappraisal of current concepts of the anatomical and functional organization of the basal ganglia,which play such a crucial role in the control of motor behavior. The basal ganglia now stand as a widely distributed neuronal network, whose elements are endowed with a highly patterned set of axon collaterals. The elucidation of this finely tuned network is needed to understand the complex spatiotemporal sequence of neural events that ensures the flow of cortical information through the basal ganglia.

Striatal dopamine- and glutamate-mediated dysregulation in experimental parkinsonism.

Characteristic changes involving interactions between dopamine and glutamate in striatal medium spiny neurons now appear to contribute to symptom production in Parkinson's disease (PD). The balance between kinase and phosphatase signaling modifies the phosphorylation state of glutamate receptors and thus their synaptic strength. Sensitization of spiny-neuron NMDA and AMPA receptors alters cortical glutamatergic input to the striatum and modifies striatal GABAergic output, and thus motor function. Conceivably, the pharmacological targeting of spiny-neuron mechanisms modified in PD will provide a safer and more effective therapy.

Neurofilament proteins are preferentially expressed in descending output neurons of the cat the superior colliculus: a study using SMI-32.

Physiological studies indicate that the output neurons in the multisensory (i.e. intermediate and deep) laminae of the cat superior colliculus receive converging information from widespread regions of the neuraxis, integrate this information, and then relay the product to regions of the brainstem involved in the control of head and eye movements. Yet, an understanding of the neuroanatomy of these converging afferents has been hampered because many terminals contact distal dendrites that are difficult to label with the neurochemical markers generally used to visualize superior colliculus output neurons. Here we show that the SMI-32 antibody, directed at the non-phosphorylated epitopes of high molecular weight neurofilament proteins, is an effective marker for these superior colliculus output neurons. It is also one that can label their distal dendrites. Superior colliculus sections processed for SMI-32 revealed numerous labeled neurons with varying morphologies within the deep laminae. In contrast, few labeled neurons were observed in the superficial laminae. Neurons with large somata in the lateral aspects of the deep superior colliculus were particularly well labeled, and many of their secondary and tertiary dendrites were clearly visible. Injections of the fluorescent biotinylated dextran amine into the pontine reticular formation revealed that approximately 80% of the SMI-32 immunostained neurons also contained retrogradely transported biotinylated dextran amine, indicating that SMI-32 is a common cytoskeletal component expressed in descending output neurons. Superior colliculus output neurons also are known to express the calcium-binding protein parvalbumin, and many SMI-32 immunostained neurons also proved to be parvalbumin immunostained. These studies suggest that SMI-32 can serve as a useful immunohistochemical marker for detailing the somatic and dendritic morphology of superior colliculus output neurons and for facilitating evaluations of their input/output relationships.

Sensory innervation of the hairy skin (light- and electronmicroscopic study.

The sense of touch develops early in phylogeny and is one of the most important senses for the survival of the animal. Touch organs of hairy skin in mammals include the so-called "Haarscheiben" (also Pinkus corpuscles) and all types of hair follicles with their nerve endings. The touch organs of the skin consist of a mechanical transducing component and the sensory component. The epithelium and its derivatives like hair follicles and sebaceous glands are the mechanical transducing component transmitting the mechanical forces like pressure or touch to the second component--the sensory nerve endings. In mammalian hairy skin all sinus and guard hairs and many vellus hairs are touch organs. The sinus hair is a typical example of a touch organ. All mammals except humans are equipped with these highly differentiated touch organs. The hair follicle is almost completely embedded in a blood sinus and equipped with more than 2,000 sensory nerve endings. All sinus and guard hairs are equipped with free nerve endings (nociceptors), Merkel nerve endings (slowly adapting [SA I] mechanoreceptor units-pressure detectors), palisades of lanceolate nerve endings (velocity detectors), and pilo-Ruffini corpuscles (tension receptors). In most of the sinus hairs lamellated corpuscles of Pacini type could be found (rapidly adapting receptors-acceleration detectors). Most vellus hairs are equipped with free and lanceolate nerve endings. Some of the vellus hairs of the upper portion of the body (head, upper extremity) are innervated by Merkel nerve endings. The presence of pilo-Ruffini nerve endings in vellus hairs is very unusual.

The radial fibers in the globus pallidus.

In our Golgi collection of adult monkey brains the striatal efferents, i.e., the radial fibers in the globus pallidus and the "comb" bundle fibers in the internal capsule and in the cerebral peduncle, are well impregnated in the horizontally sectioned brain and in a sagittal sectioned brain. Since collaterals emerging from radial fibers are seen only in the horizontal series and not in the saggittal series, the interpretation is that they proceed anteriorly and posteriorly only, following the curvature of the pallidal segments, and do not run superiorly or inferiorly as they emerge. Although radial fibers emitting collaterals in the lateral segment and in the medial segment of the globus pallidus have been observed, it has not been possible to observe the same radial fiber emitting collaterals in both pallidal segments and the prospects of ever doing so are not good. The radial fibers converging in the globus pallidus pursue many radii and there is little coincidence between the plane of section and the planes in which they travel. At most only severed radial fiber segments 100-150 microns in length can be found in the horizontal sections needed to observe the collaterals. Moreover, sagittal sections trodorsally, as they pass through the internal medullary lamina to enter the medial segment of the globus pallidus. The radial fibers in the medial segment of the globus pallidus are continuous with the "comb" bundle fibers and appear to be thinner than the radial fibers in the lateral segment of the globus pallidus. It is not proved; nonetheless, the view expressed here is that the radial fibers are thinner in the medial segment of the globus pallidus because they may be the same fibers that gave off collaterals in the lateral segment of the globus pallidus. This is discussed in the light of the electrophysiological disclosure of Yoshida et al. ('71, '72) that caudatopallidal fibers are collaterals off caudatonigral fibers. The afferent plexuses of fine, "bouton en passage" fibers, which completely ensheath the long radiating dendrites in the globus pallidus (Fox et al., '66) are well impregnated in the horizontal series. Obviously, they are formed by a number of ultimate branches converging from the collateral brances of a number of different radial fibers. The divergence, too, in this system must be considerable; however, its true extent can only be surmised from the several radial fibers and radial fiber collaterals seen in the incompletely impregnanted Golgi section. Continued.

Ultrastructural study of large efferent neurons in the superior colliculus of the cat after retrograde labeling with horseradish peroxidase.

The ultrastructure of large neurons in the stratum griseum intermedium of the cat superior colliculus was examined following injections of horseradish peroxidase (HRP) into the dorsal tegmental decussation. Four HRP-labeled cells were selected, and the synaptology of their cell bodies and selected regions of proximal and distal dendrites was examined. The four neurons represent four morphologically distinct cell types: multipolar radiating, tufted, large vertical, and medium-sized trapezoid radiating. These four neurons correspond with cell types X1, X2, X3, and T1 respectively, according to the recent classification of neurons in the superior colliculus of the cat by Moschovakis and Karabelas (J. Comp Neurol. 239:276-308, '85). The three X type neurons are similar in having 83% of their somata and over 74% of their proximal dendrites contacted by synaptic profiles. Distal dendrites of the X type neurons, however, receive fewer synaptic contacts. In contrast, in the T1 cell, only 69% of the soma membrane is contacted by synaptic profiles, and the synaptic coverage on proximal and distal dendrites does not vary much from this. Of the eight types of synaptic terminals described in the stratum griseum intermedium of the cat superior colliculus by Norita (J. Comp. Neurol. 190:29-48, '80), only five are found in contact with the X and T type efferent neurons described here. There are some regional differences in terminal distribution, although each terminal is represented on each cell. Type III terminals (small, contain mostly pleomorphic vesicles, and make symmetrical contacts) are the most abundant on cell bodies and dendrites of all four cell types. Terminal types II (medium-sized, containing round and flattened vesicles, and making asymmetrical contacts), and IV (medium to large in size, containing flattened vesicles, and making symmetrical contacts) are well represented. In general, terminal types I (small, containing densely packed round vesicles, and making asymmetrical contacts) and VI (small and irregular in shape, containing flattened vesicles and making symmetrical contacts) are found infrequently. The identity of different types of synaptic terminal is discussed.

Reciprocal synapses between hair cells and first order afferent dendrites in the crista ampullaris of the bullfrog.

Reciprocal synapses were found between afferent dendrites and hair cells of the crista ampullaris in the vestibular system of the bullfrog. The reciprocal synapse consisted of two components forming a dual synaptic complex between a single hair cell and a single dendritic process such that the receptor cell was both presynaptic and postsynaptic to the afferent dendrite. The first component, a typical afferent synaptic complex, consisted of a synaptic sphere surrounded by a single layer of clear vesicles, and was located within the hair cell. In addition, 3-4 arciform structures were located between the sphere and the plasma membrane of the hair cell. The second component of the reciprocal synapse, located approximately 1.5 micron from the first, was recognized by a localized accumulation of clear membrane vesicles within the neuronal process and a short membrane profile within the cytoplasm of the adjacent hair cell. Additional characteristic features included continuity of the arciform structures with the plasma membrane of the hair cell, and within the neuronal process, a greatly reduced number of vesicles when compared with the efferent axonal terminal. This latter was important in distinguishing a first-order afferent dendritic process from an efferent neuronal process. The possible functional significance of these reciprocal synapses has been discussed in terms of the two components having an opposite effect on the membrane polarization of the hair cell.

Anatomy of the posterior lateral line system in young larvae of the zebrafish.

We studied the anatomy of neuromasts, afferent sensory neurons, and efferent neurons of the midbody branch of the posterior lateral line in larvae of the zebrafish (Brachydanio rerio), 5 days after fertilization. This simple sensory system consists of ten or 11 neuromasts, 15-20 sensory neurons, and about nine efferent neurons. The neuromasts are typical free neuromasts and both afferent and efferent synapses are present on hair cells within them. The sensory neurons project into a single longitudinal column of neuropil in the hindbrain. The sensory terminals appear by light microscopy to contact the dorsolateral dendrite of the ipsilateral Mauthner cell. Three types of efferent neurons are present; two types in the hindbrain and one type in the diencephalon. We provide several lines of evidence that demonstrate that these central neurons are efferent to the lateral line. We conclude from this morphology that the larval system includes all of the components of the adult system and is probably functional at this early stage. We also found that larvae have all of the efferent neurons found in adult zebrafish, while the number of neuromasts and sensory neurons will increase during subsequent development.

Chandelier cells in the hippocampal formation of the rat: the entorhinal area and subicular complex.

In the present study we describe the characteristics of the chandelier cells in the rat entorhinal cortex and subicular complex by using the Golgi method and combined Golgi-electron microscopic techniques. In the entorhinal cortex, chandelier cells were frequently stained in layers II/III. Two types of axonal complexes were noted. One had a preferential horizontal orientation and gave rise to terminals located in the upper portion of layers II/III. The second type of chandelier cell axon was observed in the medial entorhinal area, innervating the entire extent of layers II/III. In the subicular complex, chandelier cells were frequently stained in the parasubiculum, whereas only a few cells were found in the presubiculum. In both subfields, chandelier cell axons were restricted to layers II/III. In the subiculum, most chandelier cells were present in the stratum radiatum, giving rise to a descending axon that branched in the stratum pyramidale. Both the size and morphological features of the chandelier cell terminal portions were found to be region-specific. Electron microscopically, the cell body and dendrites of gold-toned chandelier cells displayed typical features of nonpyramidal cells, such as the presence of nuclear infoldings, symmetric and asymmetric synapses on the cell body, and moderate numbers of axon terminals covering the smooth dendritic surface. Five gold-toned chandelier cell axonal complexes were analyzed at the fine structural level. In all parahippocampal regions, gold-labeled axon terminals formed symmetric synaptic contacts with axon initial segments. Our results demonstrate the presence, morphological characteristics, and target selectivity of identified chandelier cells in the parahippocampal region of the rat. Together with previous data, these results suggest a wide distribution of this specialized type of cortical interneuron and indicate that it is a constant and essential component of inhibitory circuits in the cerebral cortex. The possible significance of chandelier cells for the circuits linking several subfields of the hippocampal formation is discussed.

Anatomy and physiology of spiking local and intersegmental interneurons in the median neuroblast lineage of the grasshopper.

The range of anatomical and physiological properties in the adult progeny of an identified neuroblast was investigated. Some 80-90 adult neurons constitute the dorsal unpaired median (DUM) group of the grasshopper metathoracic ganglion. Within the group are efferent, octopaminergic neurons with large cell bodies and overshooting action potentials. Our objective was to determine the properties of the neurons with small cell bodies that make up the majority of the clone, some 60-70 neurons, about which scant information was available. The small DUM neurons have cell body diameters of 10-20 microns and stain with antibodies to GABA (Thompson and Siegler, '89: Proc. Soc. Neurosci. 15:1296 (abstr.); Witten and Truman, '89: Proc. Soc. Neurosci. 15:365 (abstr.)). By employing intracellular electrophysiological and morphological techniques, we have established that the small DUM neurons are spiking interneurons, expressing passively conducted action potentials in the cell body. They fall into two basic classes: local interneurons with bilateral branches in the auditory neuropiles, and intersegmental interneurons with bilateral branches widespread in the methathoracic ganglion and axons traveling in both anterior connectives. The local interneurons typically respond to sound, whereas the intersegmental interneurons selectively respond to wind on the head or to generalized movements by the animal. Primary neurites of small and large DUM neurons enter the neuropil in a bundle, but the neurites of DUM interneurons are more posterior and have a separate trajectory from those of the efferent DUM neurons once in the ganglion core. A model is presented for the sequential development of efferent, local, and intersegmental DUM neurons from the median neuroblast.