Pedal neuron 3 serves a significant role in effecting turning during crawling by the marine slug <Emphasis Type="Italic">Tritonia diomedea</Emphasis> (Bergh)

The marine nudibranch Tritonia diomedea crawls using its ciliated foot surface as the sole means of propulsion. Turning while crawling involves raising a small portion of the lateral foot margin on the side of the turn. The cilia in the lifted area no longer contribute to propulsion, and this asymmetry in thrust turns the animal towards the lifted side. Neurons located in the pedal ganglia of the brain contribute to these foot margin contractions. T. diomedea has a natural tendency to turn upstream (rheotaxis), and pedal flexion neuron Pedal 3 elicits foot margin lift and receives modulatory input from flow receptors. To assess the contribution of this single cell in turning behavior, two fine wires were glued to the surface of the brain over left and right Pedal 3. We determined that Pedal 3 activity is correlated with subsequent ipsilateral turns, preceding the lift of the foot margin and the change in orientation by a consistent interval. Both Pedal 3 cells show synchronous bursts of activity, and the firing frequency of the ipsilateral Pedal 3 increased before turns were observed to that side. Stimulation of the electrode over Pedal 3 proved sufficient to elicit an ipsilateral turn in Tritonia.     

Immunochemical and electrophysiological analyses of magnetically responsive neurons in the mollusc Tritonia diomedea

Tritonia diomedea uses the Earth’s magnetic field as an orientation cue, but little is known about the neural mechanisms that underlie magnetic orientation behavior in this or other animals. Six large, individually identifiable neurons in the brain of Tritonia (left and right Pd5, Pd6, Pd7) are known to respond with altered electrical activity to changes in earth-strength magnetic fields. In this study we used immunochemical, electrophysiological, and neuroanatomical techniques to investigate the function of the Pd5 neurons, the largest magnetically responsive cells. Immunocytochemical studies localized TPeps, neuropeptides isolated from Pd5, to dense-cored vesicles within the Pd5 somata and within neurites adjacent to ciliated foot epithelial cells. Anatomical analyses revealed that neurites from Pd5 are located within nerves innervating the ipsilateral foot and body wall. These results imply that Pd5 project to the foot and regulate ciliary beating through paracrine release. Electrophysiological recordings indicated that, although both LPd5 and RPd5 responded to the same magnetic stimuli, the pattern of spiking in the two cells differed. Given that TPeps increase ciliary beating and Tritonia locomotes using pedal cilia, our results are consistent with the hypothesis that Pd5 neurons control or modulate the ciliary activity involved in crawling during orientation behavior.     

Conservation of a Tritonia Pedal peptides network in gastropods

Adults of the nudibranch mollusc Tritonia diomedea crawl using mucociliary locomotion. Crawling is controlled in part by the large Pedal 5 (Pd5) and Pedal 6 (Pd6) neurons that produce Tritonia Pedal peptides (TPeps). TPeps elicit an increase in ciliary beat frequency, thereby increasing crawling speed. In adults of T. diomedea, an extensive network of TPep‐containing neurites adjacent to the basement membrane of the pedal epithelium delivers TPeps to the ciliated cells. In this study, we show that diverse nudibranchs all have a pattern of TPep‐like immunoreactivity similar to that of T. diomedea, with thin tracts of TPep‐like immunoreactive (TPep‐LIR) neurites projecting to the epithelial layer. We also show that members of two non‐nudibranch gastropod species have a pattern of TPep‐innervation similar to that of the nudibranchs. In addition, we characterized two pairs of motor neurons in adults of the nudibranch Armina californica that are possible homologues of the Pd5 and Pd6 cells in T. diomedea. Activity in one of these pairs, the Pedal Peptidergic Dorsal 1 (PPD1) cells, was correlated with mucociliary locomotion. The second pair, the Pedal Peptidergic Ventral 1 cells, shared synchronous synaptic input with the PPD1 cells, a pattern consistent with the shared synaptic input of the T. diomedea Pd5 and Pd6 cells. These findings suggest that the roles of the Pd5 and Pd6 cells as mucociliary motor neurons in nudibranchs are conserved evolutionarily. Additionally, the extensive network of TPep‐LIR neurites seen in the foot of T. diomedea appears likely to be a common feature among gastropods.     

Physiological basis of feeding behavior in Tritonia diomedea. III. Role of depolarizing afterpotentials

The nature and role of the depolarizing afterpotentials (DAPs) of buccal motoneurons of Tritonia diomedea were examined. Neuron B5 exhibits a DAP whose ionic dependence and modifiability by TEA and 4-AP suggest a similarity to the DAP previously described in pleural pacemaker neurons. Reduction of the DAP severely reduces the ability of these neurons to generate bursts of action potentials. Certain other motoneurons (B1 and B6) are reexcited by a slow DAP (SDAP) which appears to be of synaptic origin. It is concluded that DAPs, which are dependent upon motoneuron activity, contribute to the synthesis of motor output by the buccal ganglion.     

Shoreward orientation involving geomagnetic cues in the nudibranch mollusc Tritonia diomedea

The marine nudibranch mollusc Tritonia diotnedea orients to the geomagnetic field in the laboratory, and has identifiable brain cells (Pd5, 6) which respond electrically when the ambient magnetic field is rotated artificially. Field studies reported here seek to determine if and why Tritonia diomedea uses geomagnetic cues to orient in the field. Animals were collected in their natural habitat using SCUBA, and placed on lines with magnetic headings parallel to the shore, at different locations with respect to their site of origin. Observations made at two or more tidal cycles later indicate that most animals move from the line in a direction corresponding to the original shoreward direction, regardless of the actual shoreward direction at the site of release, suggesting guidance by geomagnetic cues. Tritonia diomedea are close to neutrally buoyant, and subject to transport over great distances when dislodged by tidal currents or during escape swimming behavior. Since the natural distribution of food and mates is along the shoreline, shoreward orientation using geomagnetic cues, particularly when other cues are weak or ambiguous, may have adaptive value.     

Function of identified nerves in orientation to water flow in Tritonia diomedea

We determined which sensory and motor nerves mediate orientation to flow in the marine slug Tritonia diomedea, and tested the hypothesis that the slug orients to water flow by comparing the intensities of water flow stimulation on each side of its body. Lesion experiments revealed which nerves carried information necessary for flow orientation. The lateral branches of Cerebral Nerve # 2 were the only cerebral nerves necessary for flow orientation. Cutting all cerebral nerves except the lateral branches of Cerebral Nerve # 2 did not eliminate flow orientation. Thus, the lateral branches of Cerebral Nerve # 2 were both necessary and sufficient (among the cerebral nerves) for flow orientation. Denervation of one side of the head by cutting Cerebral Nerves # 1–4 on one side did not eliminate normal flow orientation. We have revised our model of how Tritonia diomedea orients to flow to allow for this unilateral determination of flow direction. Unilaterally cutting Pedal Nerve # 3, which contains many pedal motor axons, reduced turning toward that side, but did not affect final orientation to flow. The ability to detect flow direction was not compro mised by the inability to initially turn towards flow.Abbreviations CeN cerebral nerve - PeN pedal nerve - PlN pleural nerve     

About possible functional interaction between serotonin and neuropeptides in control processes of embryogenesis (Experiments on embryos ofTritonia diomedea)

Ritanserin and inmecarb hydrochloride, antagonists of serotonin, act cytostatically and teratogenically on early embryos ofTritonia diomedea, a nudibranch mollusk. On the basis of a pharmacological analysis and the type of developmental abnormalities observed, this action appears to be due to disturbances in the functional activity of endogenous serotonin and is associated with damage to the cytoskeleton. The effects of ritanserin and inmecarb are prevented or attenuated by lipophilic serotonin analogs (serotoninamides of polyenoic fatty acids), as well as by polypeptides isolated from neurons Pd5 and Pd6 of the pedal ganglia of the adultTritonia. In late embryos (stage of veligers), serotonin and to a lesser extent its lipophilic analogs strongly increase embryonic motility. This effect of serotonin is potentiated by some neuropeptides and inhibited by others. These results provide evidence for functional interaction between serotonin and neuropeptides in the control processes of embryogenesis.     

Homologues of serotonergic central pattern generator neurons in related nudibranch molluscs with divergent behaviors

Homologues of a neuron that contributes to a species-specific behavior were identified and characterized in species lacking that behavior. The nudibranch Tritonia diomedea swims by flexing its body dorsally and ventrally. The dorsal swim interneurons (DSIs) are components of the central pattern generator (CPG) underlying this rhythmic motor pattern and also activate crawling. Homologues of the DSIs were identified in six nudibranchs that do not exhibit dorsal–ventral swimming: Tochuina tetraquetra, Melibe leonina, Dendronotus iris, D. frondosus, Armina californica, and Triopha catalinae. Homology was based upon shared features that distinguish the DSIs from all other neurons: (1) serotonin immunoreactivity, (2) location in the Cerebral serotonergic posterior (CeSP) cluster, and (3) axon projection to the contralateral pedal ganglion. The DSI homologues, named CeSP-A neurons, share additional features with the DSIs: irregular basal firing, synchronous inputs, electrical coupling, and reciprocal inhibition. Unlike the DSIs, the CeSP-A neurons were not rhythmically active in response to nerve stimulation. The CeSP-A neurons in Tochuina and Triopha also excited homologues of the Tritonia Pd5 neuron, a crawling efferent. Thus, the CeSP-A neurons and the DSIs may be part of a conserved network related to crawling that may have been co-opted into a rhythmic swim CPG in Tritonia. This material is based upon work supported by the National Science Foundation, under Grant No. 0445768, and a GSU Research Program Enhancement grant to PSK.     

Magnetic orientation and navigation in marine turtles, lobsters, and molluscs: concepts and conundrums

The Earth's magnetic field provides a pervasive source of directionalinformation used by phylogenetically diverse marine animals.Behavioral experiments with sea turtles, spiny lobsters, andsea slugs have revealed that all have a magnetic compass sense,despite vast differences in the environment each inhabits andthe spatial scale over which each moves. For two of these animals,the Earth's field also serves as a source of positional information.Hatchling loggerhead sea turtles from Florida responded to themagnetic fields found in three widely separated regions of theAtlantic Ocean by swimming in directions that would, in eachcase, facilitate movement along the migratory route. Thus, foryoung loggerheads, regional magnetic fields function as navigationalmarkers and elicit changes in swimming direction at crucialgeographic boundaries. Older turtles, as well as spiny lobsters,apparently acquire a "magnetic map" that enables them to usemagnetic topography to determine their position relative tospecific goals. Relatively little is known about the neuralmechanisms that underlie magnetic orientation and navigation.A promising model system is the marine mollusc Tritonia diomedea,which possesses both a magnetic compass and a relatively simplenervous system. Six neurons in the brain of T. diomedea havebeen identified that respond to changes in magnetic fields.At least some of these appear to be ciliary motor neurons thatgenerate or modulate the final behavioral output of the orientationcircuitry. These findings represent an encouraging step towarda holistic understanding of the cells and circuitry that underliemagnetic orientation behavior in one model organism.     

Variable neuronal participation in stereotypic motor programs

To what extent are motor networks underlying rhythmic behaviors rigidly hard-wired versus fluid and dynamic entities? Do the members of motor networks change from moment-to-moment or from motor program episode-to-episode? These are questions that can only be addressed in systems where it is possible to monitor the spiking activity of networks of neurons during the production of motor programs. We used large-scale voltage-sensitive dye (VSD) imaging followed by Independent Component Analysis spike-sorting to examine the extent to which the neuronal network underlying the escape swim behavior of Tritonia diomedea is hard-wired versus fluid from a moment-to-moment perspective. We found that while most neurons were dedicated to the swim network, a small but significant proportion of neurons participated in a surprisingly variable manner. These neurons joined the swim motor program late, left early, burst only on some cycles or skipped cycles of the motor program. We confirmed that this variable neuronal participation was not due to effects of the VSD by finding such neurons with intracellular recording in dye-free saline. Further, these neurons markedly varied their level of participation in the network from swim episode-to-episode. The generality of such unreliably bursting neurons was confirmed by their presence in the rhythmic escape networks of two other molluscan species, Tritonia festiva and Aplysia californica. Our observations support a view that neuronal networks, even those underlying rhythmic and stereotyped motor programs, may be more variable in structure than widely appreciated.     

Characterization of the synaptic actions of an interneuron in the central nervous system of Tritonia

The motor program that drives the swimming behavior of the marine mollusk Tritonia diomedea is generated by three interneuronal populations in the cerebral ganglia. One of these populations, the pair of C2 neurons, is shown to also exert powerful synaptic actions upon most cells in the contralateral pedal ganglion. Intracellular staining with Co²⁺ showed that the C2 neurons projected to the contralateral pedal ganglion as a single unbranched axon, and nearly all contralateral pedal neurons received monosynaptic input from C2. Orthodromic stimulation of most peripheral nerves caused monosynaptic excitation of C2 by afferent sensory cells and, in some cases, monosynaptic inhibition from an unidentified source. C2 neurons produced four types of postsynaptic potential (PSP) on pedal neurons: (1) a fast, Cl^?-mediated inhibition (FIPSP); (2) a fast, Na⁺-mediated excitation (FEPSP); (3) a slow, K⁺-mediated inhibition (SIPSP); and (4) a slow, conductance-decrease excitation (SEPSP). All four could be recorded simultaneously in some pedal neurons. The C2 neurons appear to be high-order, multiaction neurons involved in both the generation of a complex motor program and the coordination of ancillary neuronal activity.     

Evidence for homologous peptidergic neurons in the buccal ganglia of diverse nudibranch mollusks

The buccal ganglia of seven nudibranches (Aeolidia papillosa, Armina californica, Dirona albolineata, D. picta, Hermissenda crassicornis, Melibe leonina, and Tritonia diomedea) were examined to explore possible homologies between large cells that reacted with antibodies directed against small cardioactive peptide B (SCP_B). The buccal ganglion of each species possessed a pair of large, dorsal–lateral, whitish neurons that contained an SCP_B-like peptide. We refer to these neurons as the SLB (SCP_B-immunoreactive Large Buccal) cells. In all species examined, the SLB cells project out the gastroesophageal nerves and appear to innervate the esophagus. In each species, an apparent rhythmic feeding motor program (FMP) was observed by intracellular recording from both SLB neurons and other neurons in isolated preparations of the buccal ganglia. SLB cells often fire at a high frequency, and usually burst in a specific phase relation to the FMP activity. Stimulation of SLB cells enhances expression of the feeding motor program, either by potentiating existing activity or eliciting the FMP in quiescent preparations. Finally, perfusion of isolated buccal ganglia with SCP_B excites the SLB cells and activates FMPs. Thus, both the immunohistochemical and electrophysiological data suggest that the SLB cells within three suborders of the opistobranchia (Dendronotacea, Arminacea, and Aeolidacea) are homologous. A comparison of our data with previously published studies indicates that SLB cell homologs may exist in other gastropods as well.     

Detection of substrate-associated odour cues versus prey-associated cues by the oral veil in Tritonia diomedea

Our goal was to test two potential sensory roles for the oral veil in the nudibranch Tritonia diomedea (now synonymous with T. tetraquetra). First, we hypothesized this cephalic sensory organ could detect substrate-associated odours left behind by an odour plume flowing across sediment. In two experiments in a laboratory flow tank, however, T. diomedea did not show consistent crawling headings in response to either prey or predator odours associated with sediment substrate. In one of the experiments, the slugs did significantly decrease crawling speed in response to prey odours. Although slugs could thus detect at least some substrate-associated odours, these results suggest such cues are not used for navigation. We next considered the oral veil’s potential role in behaviours requiring responses to nearby cues. Our observations of animals before and after denervation of the oral veil suggest that, unsurprisingly, predatory bite-strikes do rely on sensory input from the oral veil. Overall, these data, combined with the results of earlier studies, are consistent with the oral veil detecting cues primarily from nearby stimuli (including both chemical and mechanical modalities), while having little or no role in detecting and responding to odour cues originating from distant sources used for navigation behaviour.     

Commissural ring nerve: A female-specific neurosecretory tract supplied by bifurcating median neurons in the cockroach Periplaneta americana (L.) and the cricket Teleogryllus commodus (Walker)

Summary In the American cockroach, Periplaneta americana, and the Australian field cricket, Teleogryllus commodus, the two nerves supplying the bases of the cerci are joined by a branch that crosses behind the last abdominal ganglion. This commissural ring nerve is restricted to females, and it contains many axons filled with granular and agranular vesicles. The axons stem from somata located within the ganglion. There are one (Periplaneta) or two (Teleogryllus) groups of median neurons with bilaterally symmetrical bifurcations, and a group of postero-ventral neurons on each side. In T. commodus, these neurons are distinct from others associated with the cerci. In the two species, the ring nerve neurons contribute to a neuropile near the root of each cereal nerve. The bifurcating median neurons arborize on both sides before entering the ring nerve, while the postero-ventral ones branch more extensively ipsilateral to their somata. The possibilities are discussed that the bifurcating neurons may be homologous to dorsal unpaired median neurons, and that the ring nerve may be a neurohemal area.     

Properties of central motor neurons exciting locomotory cilia inTritonia diomedea

Summary A pair of large, identifiable neurons (Pd 21), one in each pedal ganglion, can excite previously inactive locomotory cilia on the sole of the foot ofTritonia diomedea (Audesirk, 1978; Fig. 3). These neurons exert their effect via axons which innervate the foot and are probably central motor neurons for pedal cilia. IntactTritonia are stimulated to crawl by the application of 1.5 M NaCl to the tail, and conversely usually stop crawling when the chemosensitive oral veil is touched with food (sea whip,Virgularia sp.). The Pd 21 neurons are excited by 1.5 M NaCl applied externally to the tail, and are inhibited by sea whip touch to the oral veil (Figs. 4 and 5). When aTritonia performs its escape swim, the cilia move strongly, and the Pd 21 neurons fire bursts of spikes in phase with dorsal flexions (Figs. 6 and 7). After a swim, aTritonia rapidly crawls along the substrate; during this time the spiking rate of the Pd 21s is greatly accelerated. Interneurons thought to drive swim bursts produce monosynaptic EPSPs in the Pd 21s (Fig. 8). The Pd 21s are coordinated in their spike activity by synaptic activity which is synchronous in the two neurons regardless of the site of external stimulation, and by electrical coupling between the two cells via axons in a pedal commissure (Figs. 9 and 10). The coupling coefficient for passively conducted potentials is quite high, about 0.15, despite an axon 8 to 12 mm long separating the two cells.Abbreviations BPSP biphasic postsynaptic potential - SW sea water     

The organization of plurisegmental mechanosensitive interneurons in the central nervous system of the wandering spider Cupiennius salei

Summary In spiders the bulk of the central nervous system (CNS) consists of fused segmental ganglia traversed by longitudinal tracts, which have precise relationships with sensory neuropils and which contain the fibers of large plurisegmental interneurons. The responses of these interneurons to various mechanical stimuli were studied electrophysiologically, and their unilateral or bilateral structure was revealed by intracellular staining. Unilateral interneurons visit all the neuromeres on one side of the CNS. They receive mechanosensory input either from a single leg or from all ipsilateral legs via sensory neurons that invade leg neuromeres and project into specific longitudinal tracts. The anatomical organization of unilateral interneurons suggests that their axons impart their information to all ipsilateral leg neuromeres. Bilateral interneurons are of two kinds, symmetric and asymmetric neurons. The latter respond to stimulation of all legs on one side of the body, having their dendrites amongst sensory tracts of the same side of the CNS. Anatomical evidence suggests that their terminals invade all four contralateral leg neuromeres. Bilaterally symmetrical plurisegmental interneurons have dendritic arborizations in both halves of the fused ventral ganglia. They respond to the stimulation of any of the 8 legs. A third class of cells, the ascending neurons have unilateral or bilateral dendritic arborizations in the fused ventral ganglia and show blebbed axons in postero-ventral regions of the brain. Their response characteristics are similar to those of other plurisegmental interneurons. Descending neurons have opposite structural polarity, arising in the brain and terminating in segmental regions of the fused ventral ganglia. Descending neurons show strong responses to visual stimulation. Approximately 50% of all the recorded neurons respond exclusively to stimulation of a single type of mechanoreceptor (either tactile hairs, or trichobothria, or slit sensilla), while the rest respond to stimulation of a variety of sensilla. However, these functional differences are not obviously reflected by the anatomy. The functional significance of plurisegmental interneurons is discussed with respect to sensory convergence and the coordination of motor output to the legs. A comparison between the response properties of certain plurisegmental interneurons and their parent longitudinal tracts suggests that the tracts themselves do not reflect a modality-specific organization.Abbreviations BPI bilateral plurisegmental interneuron - CNS central nervous system - FVG fused ventral ganglia - LT longitudinal tract - PI plurisegmental interneuron - PSTH peristimulus timehistogram - UPI unilateral plurisegmental interneuron     

Leucokinin and diuretic hormone immunoreactivity of neurons in the tobacco hornworm,Manduca sexta,and co-localization of this immunoreactivity in lateral neurosecretory cells of abdominal ganglia

Because leucokinins stimulate diuresis in some insects, we wished to identify the neurosecretory cells in Manduca sexta that might be a source of leucokinin-like neurohormones. Immunostaining was done at various stages of development, using an antiserum to leucokinin IV. Bilateral pairs of neurosecretory cells in abdominal ganglia 3–7 of larvae and adults are immunoreactive; these cells project via the ipsilateral ventral nerves to the neurohemal transverse nerves. The immunoreactivity and size of these lateral cells greatly increases in the pharate adult, and this change appears to be related to a period of intensive diuresis occurring a few days before adult eclosion. Relationships of these neurons to cells that are immunoreactive to a M. sexta diuretic hormone were also investigated. Diuretic hormone and leucokinin immunoreactivity are co-localized in the lateral neurosecretory cells and their neurohemal projections. A median pair of leucokinin-immunoreactive, and a lateral pair of diuretic hormone-immunoreactive neurons in the larval terminal abdominal ganglion project to neurohemal release sites within the cryptonephridium. The immunoreactivity of these cells is lost as the cryptonephridium is eliminated during metamorphosis. This loss appears to be related to the change from the larval to adult pattern of diuresis.     

Spatial tuning of neurons in the inferior colliculus of the big brown bat: effects of sound level, stimulus type and multiple sound sources

We examined factors that affect spatial receptive fields of single units in the central nucleus of the inferior colliculus of Eptesicus fuscus. Pure tones, frequency- or amplitude-modulated sounds, or noise bursts were presented in the free-field, and responses were recorded extracellularly. For 58 neurons that were tested over a 30 dB range of sound levels, 7 (12%) exhibited a change of less than 10° in the center point and medial border of their receptive field. For 28 neurons that were tested with more than one stimulus type, 5 (18%) exhibited a change of less than 10° in the center point and medial border of their receptive field.The azimuthal response ranges of 19 neurons were measured in the presence of a continuous broadband noise presented from a second loudspeaker set at different fixed azimuthal positions. For 3 neurons driven by a contralateral stimulus only, the effect of the noise was simple masking. For 11 neurons driven by sound at either side, 8 were unaffected by the noise and 1 showed a simple masking effect. For the remaining 2, as well as for 5 neurons that were excited by contralateral sound and inhibited by ipsilateral sound, the peak of the azimuthal response range shifted toward the direction of the noise.Abbreviations E/E excitation at either ear - I/E inhibition at the ipsilateral ear, excitation at the contralateral ear - O/E no effect from the ipsilateral ear, excitation at the contralateral ear - FM downward frequency modulation - FM upward frequency modulation - IC inferior colliculus - ICC central nucleus of the inferior colliculus - ILD interaural level difference - ITD interaural time difference - PT pure tone - SAM sinusoidally amplitude modulated sounds - SFM sinusoidally frequency modulated sounds     

The oculomotor system of Daphnia magna

Summary The highly mobile cyclopic compound eye of Daphnia magna is rotated by six muscles arranged as three bilateral pairs. The three muscles on each side of the head share a common origin on the carapace and insert dorsally, laterally and ventrally on the eye. The dorsal and ventral muscles are each composed of two muscle fibers and the lateral muscle is composed of from two to five fibers, with three the most common number. Individual muscle fibers are spindle-shaped mononucleated cells with organized bundles of myofilaments. Lateral eye-muscle fibers are thinner than those of the other muscles but are otherwise similar in ultrastructure. Two motor neurons innervate each dorsal and each ventral muscle and one motor neuron innervates each lateral muscle. The cell bodies of the motor neurons are situated dorsally in the supraesophageal ganglion (SEG) and are ipsilateral to the muscles they innervate. The dendritic fields of the dorsal-muscle motor neurons are ipsilateral to their cell bodies; those of the ventral-muscle motor neurons are bilateral though predominantly contralateral. The central projections of the lateral-muscle motor neurons are unknown. In the dorsal and ventral muscles one motor axon synapses principally with one muscle fiber; in each lateral muscle the single motor axon branches to, and forms synapses with, all the fibers. The neuromuscular junctions, characterized by pre- and postsynaptic densities and clear vesicles, are similar in all the eye muscles.