Modification of statocyst input to local interneurons by behavioral condition in the crayfish brain

Posture control by statocysts is affected by leg condition in decapod crustaceans. We investigated how, in the crayfish brain, the synaptic response of local interneurons to statocyst stimulation was affected by leg movements on and off a substratum. The magnetic field stimulation method permitted sustained stimulation of statocyst receptors by mimicking body rolling. The statocyst-driven local interneurons were classified into four morphological groups (Type-I–IV). All interneurons except Type-IV projected their dendritic branches to the parolfactory lobe of the deutocerebrum where statocyst afferents project directly. Type-I interneurons having somata in the ventral-paired lateral cluster responded invariably to statocyst stimulation regardless of the leg condition, whereas others having somata in the ventral-unpaired posterior cluster showed response enhancement or suppression, depending on the cell, during leg movements on a substratum, but no response change during free leg movements off the substratum. The synaptic responses of Type-II and IV interneurons were also affected differently by leg movements depending on the substratum condition, whereas those of Type-III remained unaffected. These findings suggest that the statocyst pathway in the crayfish brain is organized in parallel with local circuits that are affected by leg condition and those not affected.     

Multiple gate control of the descending statocyst-motor pathway in the crayfish Procambarus clarkii Girard

1. Synaptic responses of uropod motoneurons and interneurons to magnetic field stimulation of the statocyst were studied in a whole animal preparation using intracellular recording and staining techniques to characterize the descending statocyst pathways controlling uropod steering behavior. 2. When the animal was engaged in abdominal postural movement, all uropod motoneurons received sustained excitatory input. Motoneurons which were to be activated during steering behavior showed excitatory responses to the stimulus superimposed on the sustained excitation. In the resting state, they showed weaker responses or no visible responses to the same stimulation. 3. Motoneurons to be suppressed during steering showed inhibitory responses to the stimulus only during abdominal movement. These included both active inhibition as well as disfacilitatory suppression of excitatory input to the motoneurons. 4. Premotor nonspiking interneurons, like motoneurons, showed greater responses to the stimulus during abdominal movement than at rest. Unlike motoneurons, however, they did not always receive sustained input during abdominal movement. 5. Descending axons which responded to statocyst stimulation independent of abdominal movement were found in the 4th and 5th abdominal ganglia. Other axons showed greater responses during abdominal movement than at rest. 6. A number of intersegmental descending interneurons with cell bodies and dendrites in the 4th or 5th ganglion were found to receive excitatory inputs from both the statocyst and the motor system controlling abdominal posture. These responses were found to summate with each other to generate spikes. 7. Statocyst signals are thus transmitted to uropod motoneurons by two types of descending pathways: one whose operation is affected by the abdominal system and the other which operates independently. The former pathway functions by recruiting intersegmental abdominal interneurons and makes stronger connections with motoneurons than the latter.     

Identified nonspiking interneurons in leg reflexes and during walking in the stick insect

In the stick insect Carausius morosus identified nonspiking interneurons (type E4) were investigated in the mesothoracic ganglion during intraand intersegmental reflexes and during searching and walking.In the standing and in the actively moving animal interneurons of type E4 drive the excitatory extensor tibiae motoneurons, up to four excitatory protractor coxae motoneurons, and the common inhibitor 1 motoneuron (Figs. 1–4).In the standing animal a depolarization of this type of interneuron is induced by tactile stimuli to the tarsi of the ipsilateral front, middle and hind legs (Fig. 5). This response precedes and accompanies the observed activation of the affected middle leg motoneurons. The same is true when compensatory leg placement reflexes are elicited by tactile stimuli given to the tarsi of the legs (Fig. 6).During forward walking the membrane potential of interneurons of type E4 is strongly modulated in the step-cycle (Figs.8–10). The peak depolarization occurs at the transition from stance to swing. The oscillations in membrane potential are correlated with the activity profile of the extensor motoneurons and the common inhibitor 1 (Fig. 9).The described properties of interneuron type E4 in the actively behaving animal show that these interneurons are involved in the organization and coordination of the motor output of the proximal leg joints during reflex movements and during walking.Abbreviations CLP reflex, compensatory leg placement reflex - CI1 common inhibitor I motoneuron - fCO femoral chordotonal organ - FETi fast extensor tibiae motoneuron - FT femur-tibia - SETi slow extensor tibiae motoneuron     

Morphology of descending statocyst interneurons in the crayfish Procambarus clarkii Girard

Summary The morphological features of descending interneurons that responded to the artificial bending of statolith hairs were assessed with intracellular recording and staining techniques. Seven statocyst interneurons were identified on the basis of their structure and response characteristics and designated as interneurons S1 to S7. All seven identified interneurons project to the optic lobe, where the optic nerve also projects, and to the dorsal part of the tritocerebrum, where the eyestalk motoneurons originate. All except interneuron S6 also extend their major branches to other neuropilar regions. S2 projects to the dorsal part of the deutocerebrum, where the statocyst nerve terminates, and S3 to the dorsal part of deutocerebrum and the antennal lobe. Four other interneurons (S1, S4, S5, S7) also extend their branches to the parolfactory lobe to which the statocyst nerve projects as well as to the deutocerebrum and antennal lobe. The extensive dendritic projections of S1–S7 suggest that they are complex multimodal interneurons rather than simple relay interneurons, receiving at least visual and statocyst sensory information. The function of the antennal lobe branches, however, has yet to be determined since the functional role of antennal input in equilibrium control is unknown.     

Central organization of common inhibitory motoneurons in the locust: role of afferent signals from leg mechanoreceptors

Intracellular recordings of mesothoracic common inhibitory neurons (CI₁, CI₂ and CI₃) were made while tactile hairs of the middle legs of locusts (Locusta migratoria) were mechanically stimulated. Generally the three common inhibitory neurons were excited by stimulation of tactile hairs on the ventral and dorsal surface of femur and tibia. The response pattern of all three CI neurons was similar suggesting that they work as a functional unit. Touching hairs on the dorsal surface of tibia and tarsus in some cases led to inhibition of CIs. The connection between sensory cells of tactile hairs and common inhibitory neurons is polysynaptic.To identify interneurons which mediate afferent signals, simultaneous intracellular recordings from CIs and interneurons were made. Different spiking interneurons were identified which made excitatory or inhibitory monosynaptic connections with CIs. Interneurons with inhibitory input to CIs belonged to the ventral midline group of spiking local interneurons. Behavioral and electrophysiological results indicate that reflex movements of the leg are accompanied by activity of CI neurons. Further it appears that CI activity is inhibited when reflex movements of the leg are actively suppressed by the animal.Abbreviations CI common inhibitor - IN interneuron - LY Lucifer Yellow     

How morphology of semicircular canals affects transduction,as shown by response characteristics of statocyst interneurons in the crabCarcinus maenas (L.)

Summary Action potentials in directional statocyst interneurons are measured during sinusoidal oscillation of the crabCarcinus around horizontal axes. Peak response of the interneurons precede peak position of the animal by 120 ° (Fig. 3). Optimum response of an interneuron occurs when the plane of oscillation of the crab coincides with the circumferential plane of the vertical canal of the statocyst providing input to the interneuron in question (Fig. 6). Directionality is explained in terms of unidirectional sensory receptors inCarcinus statocyst. Increased phase lead of response over peak position of the crab is explained in terms of reduced viscous damping in the open statocyst.The author is grateful to the Science Research Council for financial assistance.     

Local spiking interneurons controlling the equilibrium response in the crayfish Procambarus clarkii

1. In the crayfish brain, the responses of local spiking interneurons to body roll simulated by bending of statocyst hairs, were investigated with intracellular recording and staining techniques. The neurons had two separate branching portions in the protocerebrum and the deutocerebrum. They were named as type-I local neurons and further classified into 5 types (ac-U, vplc-U, vplc-B, vupc-U, vupc-B). 2. Vupc-U neurons showed excitatory responses and vplc-U neurons showed inhibitory responses to inward hair deflection of the statocyst ipsilateral to their deutocerebral branches. The other 3 types were of mixed populations of the interneurons showing either excitatory or inhibitory responses to the stimulation. 3. Of 10 type-I local neurons showing excitatory responses to inward hair deflection, 6 interneurons had output effects on oculomotor and/or descending neurons. All these 6 interneurons showed large EPSPs and much higher frequency of spikes to the hair stimulation than those of the other 4. All 8 type-I local neurons that showed inhibitory responses had no output effects. 4. Type-I local neurons controlled two equilibrium responses, compensatory eye movement and righting reflex, either simultaneously or independently.     

On the control of stridulation in the acridid grasshopper Omocestus viridulus L.

Summary In tethered, minimally dissected grasshoppers stridulation was elicited by DC brain stimulation. Intracellular recording, stimulation and staining of interneurons in the metathoracic ganglion complex were performed simultaneously with measurements of hindleg movements. The functional significance of interneurons for generation of the stridulation rhythm and bilateral coordination of the hindlegs was tested and quantitatively analysed. Interneurons involved in stridulation were found in the metathoracic and abdominal neuromeres. Typically they included arborizations in a dorsal neuropil parallel to the long axis of the ganglion. Interneuron T3-LI-3 started and drove the ipsilateral rhythm generator. The stridulation rhythm was reset by interneuron A1-AC-1. It was stopped if interneuron T3-AC-1 was stimulated. Bilateral coordination of stridulation movements was based on excitatory and inhibitory pathways between the hemiganglionic networks. Switching in the coordination of hindleg movement patterns was induced by changes in the discharge rate of the bilateral arborizing interneuron T3-LC-4. The hemiganglionic interneurons T3-LI-3 and T3-LI-1 influenced coordination via the activity level within the networks.     

Synaptic connections between sensory afferents and the common inhibitory motoneuron in crayfish

The sensory inputs to the common inhibitory motoneuron that innervates every leg muscle of the crayfish Procambarus clarkii (Girard) were analyzed by performing intracellular recordings from its neurite within the neuropil of the 5th thoracic ganglion. Two types of sensory inputs involved in locomotion were studied, those from a movement coding proprioceptor (the coxobasal chordotonal organ) and those from sensory neu rons coding contact forces exerted at the tip of the leg on the substrate (the dactyl sensory afferents). Sinusoidal movements applied to the chordotonal organ strand induced a stable biphasic response in the common inhibitory motoneuron that consisted of bursts of spikes during release and stretch of the strand, corresponding to raising and lowering of the leg, respectively. Using ramp movements imposed on the chordotonal strand, we demonstrated that only movement-coding chordotonal afferents produce excitatory post-synaptic potentials in the common inhibitory motoneuron; these connections are monosynaptic. Mechanical or electrical stimulation of the dactyl sensory afferents resulted in an increase in the tonic discharge of the common inhibitory motoneuron through polysynaptic excitatory pathways. These two types of sensory cues reinforce the central command of the common inhibitory motoneuron and contribute to enhancing its activity during leg movements, and thus facilitate the relaxation of tonic muscle fibres during locomotion.Abbreviations ADR anterior distal root - A Lev anterior levator nerve - CB coxo-basipodite joint - CBCO coxo-basal chordotonal organ - CI common inhibitory motoneuron - Dep depressor nerve - DSA dactyl sensory afferents - EPSP excitatory post-synaptic potential - IN interneuron - MN motoneuron - PDR posterior distal root - P Lev posterior levator nerve - Pro promotor nerve - Rem remotor nerve     

Local interneurons and information processing in the olfactory glomeruli of the moth Manduca sexta

Intracellular recordings were made from the major neurites of local interneurons in the moth antennal lobe. Antennal nerve stimulation evoked 3 patterns of postsynaptic activity: (i) a short-latency compound excitatory postsynaptic potential that, based on electrical stimulation of the antennal nerve and stimulation of the antenna with odors, represents a monosynaptic input from olfactory afferent axons (71 out of 86 neurons), (ii) a delayed activation of firing in response to both electrical- and odor-driven input (11 neurons), and (iii) a delayed membrane hyperpolarization in response to antennal nerve input (4 neurons).Simultaneous intracellular recordings from a local interneuron with short-latency responses and a projection (output) neuron revealed unidirectional synaptic interactions between these two cell types. In 20% of the 30 pairs studied, spontaneous and current-induced spiking activity in a local interneuron correlated with hyperpolarization and suppression of firing in a projection neuron. No evidence for recurrent or feedback inhibition of projection neurons was found. Furthermore, suppression of firing in an inhibitory local interneuron led to an increase in firing in the normally quiescent projection neuron, suggesting that a disinhibitory pathway may mediate excitation in projection neurons. This is the first direct evidence of an inhibitory role for local interneurons in olfactory information processing in insects. Through different types of multisynaptic interactions with projection neurons, local interneurons help to generate and shape the output from olfactory glomeruli in the antennal lobe.Abbreviations AL antennal lobe - EPSP excitatory postsynaptic potential - GABA -aminobutyric acid - IPSP inhibitory postsynaptic potential - LN local interneuron - MGC macroglomerular complex - OB olfactory bulb - PN projection neuron - TES N-tris[hydroxymethyl]methyl-2-aminoethane-sulfonic acid     

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     

Consistency of interneuronal group formation in response to stimulation of identified cells

In crayfish stimulation of abdominal positioning interneurons (APIs) recruits other interneurons producing various abdominal movements. We investigated whether: (1) the same API from different preparations activated a similar number or group of interneurons, (2) different APIs activated different groups, and (3) repeated stimulation of an API consistently affected a similar set of interneurons. To quantify the similarities and differences of the recruited interneuronal groups we compared the number of interneurons affected, their firing frequencies, and motor outputs. Three types of APIs (Curly Q, L and T) were identified and each type was stimulated in three preparations. Our results showed that for the Curly Q and L cells, each cell type activated interneuronal groups that were statistically similar in number and firing frequency. The T cell activated interneuronal groups that were more variable. Some APIs generally provided a repeatable motor output; all did not. The interneuronal groups activated by the Curly Q, L and T cells were very different from each other. Repeated stimulation of one Curly Q cell affected similar although not identical sets of interneurons. These data suggest that repeated motor outputs could be produced by a similar but not identical group of cells. Accepted: 29 September 1997     

Intersegmental transfer of sensory signals in the stick insect leg muscle control system

Intersegmental coordination during locomotion in legged animals arises from mechanical couplings and the exchange of neuronal information between legs. Here, the information flow from a single leg sense organ of the stick insect Cuniculina impigra onto motoneurons and interneurons of other legs was investigated. The femoral chordotonal organ (fCO) of the right middle leg, which measures posture and movement of the femur-tibia joint, was stimulated, and the responses of the tibial motoneuron pools of the other legs were recorded. In resting animals, fCO signals did not affect motoneuronal activity in neighboring legs. When the locomotor system was activated and antagonistic motoneurons were bursting in alternation, fCO stimuli facilitated transitions from flexor to extensor activity and vice versa in the contralateral leg. Following pharmacological treatment with picrotoxin, a blocker of GABA-ergic inhibition, the tibial motoneurons of all legs showed specific responses to signals from the middle leg fCO. For the contralateral middle leg we show that fCO signals encoding velocity and position of the tibia were processed by those identified local premotor nonspiking interneurons known to contribute to posture and movement control during standing and voluntary leg movements. Interneurons received both excitatory and inhibitory inputs, so that the response of some interneurons supported the motoneuronal output, while others opposed it. Our results demonstrate that sensory information from the fCO specifically affects the motoneuronal activity of other legs and that the layer of premotor nonspiking interneurons is a site of interaction between local proprioceptive sensory signals and proprioceptive signals from other legs.     

Synchronous firing by specific pairs of cercal giant interneurons in crickets encodes wind direction

One prominent stimulus to evoke an escape response in crickets is the detection of air movement, such as would result from an attacking predator. Wind is detected by the cercal sensory system that consists of hundreds of sensory cells at the base of filiform hairs. These sensory cells relay information to about a dozen cercal giant and non-giant interneurons. The response of cercal sensory cells depends both, on the intensity and the direction of the wind. Spike trains of cercal giant interneurons then convey the information about wind direction and intensity to the central nervous system. Extracellular recording of multiple cercal giant interneurons shows that certain interneuron pairs fire synchronously if a wind comes from a particular direction. We demonstrate here that directional tuning curves of synchronously firing pairs of interneurons are sharper than those of single interneurons. Moreover, the sum total of all synchronously firing pairs eventually covers all wind directions. The sharpness of the tuning curves in synchronously firing pairs results from excitatory and inhibitory input from the cercal sensory neurons. Our results suggest, that synchronous firing of specific pairs of cercal giant interneurons encodes the wind direction. This was further supported by behavioral analyses.     

Quantitative analysis of inputs to somatostatin-immunoreactive descending interneurons in the myenteric plexus of the guinea-pig small intestine

Somatostatin immunoreactivity occurs in a specific subgroup of cholinergic descending interneurons in the myenteric plexus of the guinea-pig small intestine. In the present work, we made light- and electron-microscopic investigations of chemically defined inputs to these neurons, in order that the origins of the connections of other neurons with them could be deduced. Somatostatin-immunoreactive synapses and close contacts were found on the cell bodies and filamentous processes of somatostatin neurons; these were 84% of all inputs. It is thus confirmed that this class of interneuron forms chains that project anally. Descending interneurons with immunoreactivity for nitric oxide synthase provided 14% of inputs to somatostatin-immunoreactive descending interneurons. An antiserum against a calcium-binding protein, calbindin, was used as marker for the majority of intrinsic primary afferent neurons, AH/Dogiel type II neurons; this class of neurons provided only 2.5% of the inputs to somatostatin-immunoreactive descending interneurons. We conclude that somatostatin-immunoreactive descending interneurons are involved in the conduction of impulses distally along the full length of the small intestine, but receive only a minor input from calbindin-immunoreactive primary afferent neurons.     

Premotor interneurons in generation of adaptive leg reflexes and voluntary movements in stick insects

We investigated the role of local nonspiking interneurons involved in motor control of legs in the stick insect, Carausius morosus. In a preparation that allowed the animals to perform active leg movements such as adaptive tactile reflexes, proprioceptive reflexes, and walking, we gathered the following results. Almost all tested nonspiking interneurons that provide synaptic drive onto motoneurons of the proximal leg muscles contribute to all of the motor programs underlying tactile reflexes and voluntary leg movements such as walking, searching, and rocking. Most of them are also involved in the generation of proprioceptive reflexes. All motor programs for coactivation, avoidance reflexes, resistance reflexes, and voluntary leg movements result from parallel pathways including nonspiking interneurons that support and others that oppose the motoneuronal activity. The contribution of a single interneuron to the different motor programs is specific: it can be supporting for one motor program but opposing for the other. Even for the same motor program, for example, coactivation, the contribution of an individual interneuron can depend on the stimulus site from where the response is elicited. Our results support the idea that the different motor patterns for adaptive tactile reflexes, resistance reflexes, and voluntary leg movements emerge from a multifunctional neuronal circuit that is reorganized corresponding to the motor behavior performed. The actual motor pattern is then shaped by distributed information processing in parallel supporting and opposing pathways. © 1996 John Wiley & Sons, Inc.     

Cerebral-buccal pathways in Aplysia californica: synaptic connections, cooperative interneuronal effects and feedback during buccal motor programs

Ingestion of seaweed by Aplysia is in part mediated by cerebral-buccal interneurons that drive rhythmic motor output from the buccal ganglia and in some cases cerebral-buccal interneurons act as members of the feeding central pattern generator. Here we document cooperative interactions between cerebral-buccal interneuron 2 and cerebral-buccal interneuron 12, characterize synaptic input to cerebral-buccal interneuron 2 and cerebral-buccal interneuron 12 from buccal peripheral nerve 2,3, describe a synaptic connection between cerebral-buccal interneuron 1 and buccal neuron B34, further characterize connections made by cerebral-buccal interneurons 2 and -12 with B34 and B61/62, and describe a novel, inhibitory connection made by cerebral-buccal interneuron 2 with a buccal neuron. When cerebral-buccal interneurons 2 and 12 were driven synchronously at low frequencies, ingestion-like buccal motor programs were elicited, and if either was driven alone, indirect synaptic input was recruited in the other cerebral-buccal interneuron. Stimulation of BN2,3 recruited both ingestion and rejection-like motor programs without firing in cerebral-buccal interneurons 2 or 12. During motor programs elicited by cerebral-buccal interneurons 2 or 12, high-voltage stimulation of BN2,3 inhibited firing in both cerebral-buccal interneurons. Our results suggest that cerebral-buccal interneurons 2 and 12 use cooperative interactions to modulate buccal motor programs, yet firing in cerebral-buccal interneurons 2 or 12 is not necessary for recruiting motor programs by buccal peripheral nerve BN2,3, even in preparations with intact cerebral-buccal pathways.     

Modulatory effects of parallel fiber and molecular layer interneuron synaptic activity on purkinje cell responses to ascending segment input: a modeling study

Based on anatomical, physiological, and model-based studies, it has been proposed that synapses associated with the ascending segment of granule cell axons provide the principle excitatory drive on Purkinje cells which is then modulated by the more numerous parallel fiber synapses. In this study we have evaluated this idea using a detailed compartmental model of a cerebellar Purkinje cell by providing identical ascending segment synaptic inputs during different levels of random parallel fiber and molecular interneuron input. Results suggest that background inputs from parallel fibers and molecular layer interneurons can have a substantial effect on the response of Purkinje cells to ascending segment inputs. Interestingly, these effects are not reflected in the average firing rate of the Purkinje cell and are thus entirely dendritic in effect. These results are considered in the context of the known segregated spatial distribution of the parallel fibers and ascending segment synapses and a new hypothesis concerning the functional organization of cerebellar cortical circuitry.     

Excitatory influence of wind-sensitive local interneurons on an ascending interneuron in the cricket cercal sensory system

This study examined the effects of a set of identified wind-sensitive local interneurons (9DL interneurons) on the wind-evoked spike output and directional sensitivity of an ascending interneuron (10-3) in the cricket (Acheta domesticus) cercal sensory system. Comparison of the directional sensitivities of the 9DL interneurons and 10-3 revealed that 3 of the 9DL interneurons have a large degree of overlap in their excitatory receptive fields with that of 10-3. Photoinactivation of any one of these 3 9DL interneurons resulted in a significant decrease in the spike output of 10-3 at its optimal excitatory wind stimulus positions. However, the overall directional sensitivity of 10-3 remained essentially unchanged. Photoinactivation of the one 9DL interneuron which had no overlap in its excitatory receptive field with 10-3 did not affect 10-3's responsiveness to wind stimuli. Results from simultaneous intracellular recordings of 10-3 and one of the 9DL interneurons which had an excitatory influence on 10-3 showed that depolarization of the local interneuron produced an epsp in 10-3, and could elicit several action potentials. Comparison of the morphologies of the 9DL interneurons and 10-3 revealed that the 3 9DL interneurons which had an excitatory influence on 10-3 all had regions of dendritic overlap with this ascending interneuron.Abbreviations ANOVA analysis of variance - Contra contralateral - epsp excitatory post-synaptic potential - Ipsi ipsilateral - LGI lateral giant interneuron - MGI medial giant interneuron     

Synaptic mechanisms of monaural and binaural processing in the locust

(1) Responses of auditory interneurones were recorded intracellularly within the metathoracic ganglion of the locust when stimulating each tympanic membrane with a piezoelectric transducer. Thus, in contrast to conventional sound stimulation, each of the two ears could be activated independently from the other at variable intensities, duration and stimulus onsets. By means of this ‘earphone-like’ stimulation technique the binaural integration properties of auditory interneurons could be analysed. (2) A minority of units (3 out of 43) was affected by input from one side only. Their synaptic input was purely excitatory and the intensity characteristics reflected those of auditory receptor fibres. (3) Most interneurones received input from both ears, each being excitatory or one excitatory or one excitatory and one inhibitory. In some units the unilateral synaptic response already included both an EPSP and an IPSP. As a result of varying temporal interactions between the EPSP and the IPSP within the unilaterally evoked complex response the intensity characteristics differed widely from unit to unit. (4) With binaural simultaneous stimulation the complexity of the postsynaptic responses of most interneurones increased as the synaptic input from both ears coincided at the level of the recorded interneurone. Although both ears were stimulated symmetrically (at the same time and intensity), units were recorded where the latencies of ipsilateral and contralateral synaptic input were different. Contralateral inhibition could either follow or precede ipsilateral excitation and in some cases both EPSP and IPSP had the same latency. On the basis of these findings the binaural synaptic mechanisms of directional coding are discussed and compared with corresponding results under free field stimulus conditions.