Spatial propagation of interictal discharges along the cortex

Interictal discharges (IIDs) accompany epileptic seizures and highlight the mechanisms of pathological activity. The propagation of IIDs along the neural tissue is not well understood. To simulate IID propagation, this study proposes a new mathematical model that uses the conductance-based refractory density approach for glutamatergic and GABAergic neuronal populations. The mathematical model is found to be consistent with experimental double-patch registrations in the 4-aminopyridine in vitro model of epilepsy. In slices, the spontaneous activity of interneurons leads to their synchronization by means of the depolarizing GABAmediated response, thus initiating IIDs. Modeling reveals a clustering of interneuronal synchronization followed by IIDs with activity fronts that propagate along the cortex. The GABA-mediated depolarization either remains to be subthreshold for the principal neurons and thus results in pure GABAergic IIDs (IID1s) or leads to glutamatergic excitation, thus resulting in another type of IIDs (IID2s). In both the model and experiment, IIDs propagate as waves, with constant activity profiles and velocity. The speed of IIDs is of the order of tens of mm/s and is larger for IID2s than for IID1s (40 and 20?mm/s, respectively). The simulations, consistent with experimental observations, show that the wavelike propagation of IIDs initiated by interneurons is determined by local synaptic connectivity under the conditions of depolarizing GABA.     

Spatial orientation in the bushcricket <Emphasis Type="Italic">Leptophyes punctatissima</Emphasis> (Phaneropterinae; Orthoptera): III. Peripheral directionality and central nervous processing of spatial cues

We examined peripheral and central nervous cues underlying the ability of the bushcricket Leptophyes punctatissima to orient to elevated and depressed sound sources broadcasting the female acoustic reply. The peripheral spatial directionality of the ear was measured physiologically using monaural preparations of an auditory interneuron (T-fibre). In the azimuth, maximal interaural intensity differences of 18 dB occur between ipsi- and contralateral stimulation. With increasing elevation or depression of the sound sources, IIDs decrease systematically and reach zero with the source exactly above or below the preparation. Bilateral, simultaneous recordings of the activity of the pair of interneurons allowed determining the binaural discharge differences which occur in response to the extremely short (1 ms) female reply. These discharge differences are large (four action potentials/stimulus) and reliable in the azimuth with lateral stimulation, and decrease gradually with more frontal stimulation. With elevation and depression of sound sources these differences again decrease to one action potential/stimulus at 60° or 75° elevation, and lateral stimulus angles of about 60°. We also calculated the reliability with which a receiver could correctly determine the location of the sound source. We discuss these quantitative measures in relation to the spatial phonotactic behaviour of male L. punctatissima.     

Out of phase: relevance of the medial septum for directional hearing and phonotaxis in the natural habitat of field crickets

A modified tracheal system is the anatomical basis for a pressure difference receiver in field crickets, where sound has access to the inner and outer side of the tympanum of the ear in the forelegs. A thin septum in the midline of a connecting trachea coupling both ears is regarded to be important in producing frequency-dependent interaural intensity differences (IIDs) for sound localization. However, the fundamental role of the septum in directional hearing has recently been challenged by the finding that the localization ability is ensured even with a perforated septum, at least under controlled laboratory conditions. Here, we investigated the influence of the medial septum on phonotaxis of female Gryllus bimaculatus under natural conditions. Surprisingly, even with a perforated septum, females reliably tracked a male calling song in the field. Although reduced by 5.2 dB, IIDs still averaged at 7.9 dB and provided a reliable proximate basis for the observed behavioural performance of operated females in the field. In contrast, in the closely related species Gryllus campestris the same septum perforation caused a dramatic decline in IIDs over all frequencies tested. We discuss this discrepancy with respect to a difference in the phenotype of their tracheal systems.     

Brain interneurons in noctuoid moths: Binaural excitation and slow potentials

A method is described for measuring small differences in the acoustic sensitivity of protocerebral interneurons on one side of the noctuid brain when ultrasonic pulses are directed first at one tympanic organ and then at the other. In 23 preparations ipsilateral sensitivity of the brain interneurons was consistently greater by 3 to 4 dB (range 0–7 dB). The spike response of protocerebral interneurons to tympanic stimulation was accompanied by a negative potential having a time course of 40 to 50 msec in response to a 10 msec stimulus pulse. The consistent positive ipsilateral bias in the sensitivity of brain interneurons is much less than the increased sensitivity of the tympanic organ when sounds originate on the ipsilateral side as compared with its sensitivity to sounds directed at the contralateral side. A possible neural mechanism and the behavioural significance of this arrangement are discussed.     

硕螽(Deracantha onos)听觉中间神经元的声反应特征

本文报道了硕螽听通路单个听觉中间神经元的声反应特征。依据动作电位发放模式的不同,听觉中间神经元可分为两类,即紧张型与相位型。紧张型听觉中间神经元属于窄凋谐带神经元,敏感的频率范围8—18千赫,反应最佳频率在12千赫附近,与同种雄硕螽叫声的主能峰相匹配。相位型听觉中间神经元属于宽调谐带神经元,有二个敏感频率范围,分别为5—8千赫和12—18千赫。它们对声强度的编码方式也不一样:分别以动作电位的数目与反应潜伏期对声强编码。本文还讨论了不同类型听觉中间神经元的功能意义。     

Silencing synaptic communication between random interneurons during Drosophila larval locomotion

Genetic manipulation of individual neurons provides a powerful approach toward understanding their contribution to stereotypic behaviors. We describe and evaluate a method for identifying candidate interneurons and associated neuropile compartments that mediate Drosophila larval locomotion. We created Drosophila larvae that express green fluorescent protein (GFP) and a shibire(ts1) (shi(ts1)) transgene (a temperature-sensitive neuronal silencer) in small numbers of randomly selected cholinergic neurons. These larvae were screened for aberrant behavior at an elevated temperature (31-32°C). Among larvae with abnormal locomotion or sensory-motor responses, some had very small numbers of GFP-labeled temperature-sensitive interneurons. Labeled ascending interneurons projecting from the abdominal ganglia to specific brain neuropile compartments emerged as candidates for mediation of larval locomotion. Random targeting of small sets of neurons for functional evaluation, together with anatomical mapping of their processes, provides a tool for identifying the regions of the central nervous system that are required for normal locomotion. We discuss the limitations and advantages of this approach to discovery of interneurons that regulate motor behavior.     

EPSP amplification and the precision of spike timing in hippocampal neurons

The temporal precision with which EPSPs initiate action potentials in postsynaptic cells determines how activity spreads in neuronal networks. We found that small EPSPs evoked from just subthreshold potentials initiated firing with short latencies in most CA1 hippocampal inhibitory cells, while action potential timing in pyramidal cells was more variable due to plateau potentials that amplified and prolonged EPSPs. Action potential timing apparently depends on the balance of subthreshold intrinsic currents. In interneurons, outward currents dominate responses to somatically injected EPSP waveforms, while inward currents are larger than outward currents close to threshold in pyramidal cells. Suppressing outward potassium currents increases the variability in latency of synaptically induced firing in interneurons. These differences in precision of EPSP-spike coupling in inhibitory and pyramidal cells will enhance inhibitory control of the spread of excitation in the hippocampus.     

Hearing and frequency dependence of auditory interneurons in the parasitoid fly <Emphasis Type="Italic">Homotrixa alleni</Emphasis> (Tachinidae: Ormiini)

The parasitoid tachinid fly Homotrixa alleni detects its hosts by their acoustic signals. The tympanal organ of the fly is located at the prothorax and contains scolopidial sensory units of different size and orientation. The tympanal membrane vibrates in the frequency range of approximately 4–35 kHz, which is also reflected in the hearing threshold measured at the neck connective. The auditory organ is not tuned to the peak frequency (5 kHz) of the main host, the bush cricket Sciarasaga quadrata. Auditory afferents project in the three thoracic neuromeres. Most of the ascending interneurons branch in all thoracic neuromeres and terminate in the deutocerebrum of the brain. The interneurons do not differ considerably in frequency tuning, but in their sensitivity with lowest thresholds around 30 dB SPL. Suprathreshold responses of most neurons depend on frequency and intensity, indicating inhibitory influence at higher intensities. Some neurons respond particularly well at low frequency sounds (around 5 kHz) and high intensities (80–90 dB SPL), and thus may be involved in detection of the primary host, S. quadrata. The auditory system of H. alleni contains auditory interneurons reacting in a wide range of temporal patterns from strictly phasic to tonic and with clear differences in frequency responses.     

Audition in the praying mantis,Mantis religiosa L.: identification of an interneuron mediating ultrasonic hearing

1. The praying mantis possesses a single ear located in the ventral midline of the metathorax. We have studied the mantis' auditory nervous system using both extracellular and intracellular techniques and have identified anatomically and physiologically a mirror-image pair of interneurons (MR-501-T3) in the metathoracic ganglion that mediates ultrasonic hearing. 2. MR-501-T3 is tuned broadly to ultrasound with best sensitivity (55-60 dB SPL) between 25 and 45 kHz. Its tuning matches closely that of the whole tympanal nerve. 3. The physiological responses of MR-501-T3 are characterized by: (1) a phasic-tonic firing pattern with a distinctive initial burst at 500-800 spikes/s; (2) minimum latencies of 8-12 ms; (3) no spontaneous activity; (4) sigmoid intensity response curves with a small (10 dB) dynamic range; (5) accurate coding of stimulus duration and of repetition rates up to 60 pps. 4. The ascending axon of MR-501-T3 conducts action potentials at 4 m/s, a rate comparable with some giant fiber systems. 5. MR-501-T3 shows no directional capability. Sound from right and left produce identical responses in both cells of the pair. Neither cutting one tympanal nerve nor removing one hemi-ear leads to different responses in the two cells indicating that they must receive a common input, either from the auditory afferents or from interneurons. We present evidence that the two cells are not directly connected. 6. MR-501-T3 is a large, symmetrical cell with its processes primarily in the intermediate neuropil (lateral ring tract). Its integration segment crosses the midline in the supramedian commissure, and the cell body lies dorsally near the entrance of the leg nerve. The axon travels in the dorsal lateral tract and is one of the largest (17 microns) in the connective. 7. Given the strong anatomical similarities between MR-501-T3 and the G and B cells of the locust, these cells may be homologous. 8. We present arguments based on our physiological results and existing behavioral data that MR-501-T3 is part of an ultrasonic warning/escape system in the mantis. As in moths, lacewings, and crickets, this system may provide a defense against nocturnally foraging bats.     

Phase resetting curves and oscillatory stability in interneurons of rat somatosensory cortex

Synchronous oscillations in neural activity are found over wide areas of the cortex. Specific populations of interneurons are believed to play a significant role in generating these synchronized oscillations through mutual synaptic and gap-junctional interactions. Little is known, though, about the mechanism of how oscillations are maintained stably by particular types of interneurons and by their local networks. To obtain more insight into this, we measured membrane-potential responses to small current-pulse perturbations during regular firing, to construct phase resetting curves (PRCs) for three types of interneurons: nonpyramidal regular-spiking (NPRS), low-threshold spiking (LTS), and fast-spiking (FS) cells. Within each cell type, both monophasic and biphasic PRCs were observed, but the proportions and sensitivities to perturbation amplitude were clearly correlated to cell type. We then analyzed the experimentally measured PRCs to predict oscillation stability, or firing reliability, of cells for a complex stochastic input, as occurs in vivo. To do this, we used a method from random dynamical system theory to estimate Lyapunov exponents of the simplified phase model on the circle. The results indicated that LTS and NPRS cells have greater oscillatory stability (are more reliably entrained) in small noisy inputs than FS cells, which is consistent with their distinct types of threshold dynamics.     

Response properties of visual interneurons to motion stimuli in the praying mantis,Tenodera aridifolia

Intracellular responses of motion-sensitive visual interneurons were recorded from the lobula complex of the mantis, Tenodera aridifolia. The interneurons were divided into four classes according to the response polarity, spatial tuning, and directional selectivity. Neurons of the first class had small, medium, or large receptive fields and showed a strong excitation in response to a small-field motion such as a small square moving in any direction (SF neurons). The second class neurons showed non-directionally selective responses: an excitation to a large-field motion of gratings in any direction (ND neurons). Most ND neurons had small or medium-size receptive fields. Neurons of the third class had large receptive fields and exhibited directionally selective responses: an excitation to a large-field motion of gratings in preferred direction and an inhibition to a motion in opposite, null direction (DS neurons). The last class neurons had small receptive fields and showed inhibitory responses to a moving square and gratings (I neurons). The functional roles of these neurons in prey recognition and optomotor response were discussed.     

Pedal serotonergic neurons modulate the synaptic input of withdrawal interneurons ofHelix

A group of serotonergic cells, located in the pedal ganglia ofHelix lucorum, modulates synaptic responses of neurons involved in withdrawal behavior. Extracellular or intracellular stimulation of these serotonergic cells leads to facilitation of spike responses to noxious stimuli in the putative command neurons for withdrawal behavior. Noxious tactile stimuli elicit an increase in background spiking frequency in the modulatory neurons and a corresponding increase in stimulus-evoked spike responses in withdrawal interneurons. The serotonergic neurons have processes in the neuropil of the parieto-visceral ganglia complex, consistent with their putative role in modulating the activity of giant parietal interneurons, which send processes to the same neuropil and to the pedal ganglia. The serotonergic cells respond to noxious tactile and chemical stimuli. Although the group as a whole respond to noxious stimuli applied to any part of the body, most cells respond more to ipsilateral than contralateral stimulation, and exhibit differences in receptive areas. Intracellular investigation revealed electrical coupling between serotonergic neurons which could underlie the recruitment of members of the group not responding to a given noxious stimulus.     

Binaural disparity cues available to the barn owl for sound localization

1. Bilateral recording of cochlear potentials was used to measure the variations in interaural time differences (ITDs) and interaural intensity differences (IIDs) as a free-field auditory stimulus was moved to different positions around a barn owl's head. 2. ITD varied smoothly with stimulus azimuth across a broad frequency range. 3. ITD varied minimally with stimulus elevation, except at extreme angles from the horizontal. 4. IID varied with both stimulus elevation and stimulus azimuth. Lower frequencies were more sensitive to variations in azimuth, whereas higher frequencies were more sensitive to variations in elevation. 5. The loci of spatial coordinates that form iso-IID contours and iso-ITD contours form a non-orthogonal grid that relates binaural disparity cues to sound location.     

Postural interneurons in the abdominal nervous system of lobster

Using intracellular recording and dye-filling techniques, a survey of postural interneurons was undertaken by impaling their somata in the 2nd abdominal ganglion of lobster. During the course of study approximately fourty different intersegmental interneurons in this ganglion were sampled. Of these, 8 evoked unique, patterned responses in the postural (superficial) motoneurons; each could be identified morphologically. Five of the 8 interneurons had caudally directed axons; 4 of these projected beyond the 4th abdominal ganglion. The remainder projected rostrally, beyond the 1st abdominal ganglion. The postural interneurons were classified according to the motor program they elicited. Five were flexion producing interneurons (FPIs), one was extension producing (EPI), and two generated only inhibitory motor outputs. All motor responses were bilateral and occurred in several segments, including A2. Two neurons, FPIs 201 and 301, produced the full motor reciprocity that typically is observed when flexion command fibers are stimulated. However, three of the FPIs and the single EPI did not express complete reciprocity in synergistic and antagonistic motoneurons. The results indicate that some interneurons displaying all of the properties of command neurons are located entirely within the abdominal nervous system. The overall organization of posture-evoking interneurons appears to be similar to that found in crayfish, suggesting an even more fundamental homology in the neuronal connectivities of these two species than has been established previously.     

Diversity of intersegmental auditory neurons in a bush cricket

Various auditory interneurons of the duetting bush cricket Ancistrura nigrovittata with axons ascending to the brain are presented. In this species, more intersegmental sound-activated neurons have been identified than in any other bush cricket so far, among them a new type of ascending neuron with posterior soma in the prothoracic ganglion (AN4). These interneurons show not only morphological differences in the prothoracic ganglion and the brain, but also respond differently to carrier frequencies, intensity and direction. As a set of neurons, they show graded differences for all of these parameters. A response type not described among intersegmental neurons of crickets and other bush crickets so far is found in the AN3 neuron with a tonic response, broad frequency tuning and little directional dependence. All neurons, with the exception of AN3, respond in a relatively similar manner to the temporal patterns of the male song: phasically to high syllable repetitions and rhythmically to low syllable repetitions. The strongest coupling to the temporal pattern is found in TN1. In contrast to behavior the neuronal responses depend little on syllable duration. AN4, AN5 and TN1 respond well to the female song. AN4 (at higher intensities) and TN1 respond well to a complete duet.     

A gain-control mechanism for processing of chorus sounds in the afferent auditory pathway of the bushcricket Tettigonia viridissima (Orthoptera; Tettigoniidae)

The representation of alternative conspecific acoustic signals in the responses of a pair of local interneurons of the bushcricket Tettigonia viridissima was studied with variation in intensity and the direction of sound signals. The results suggest that the auditory world of the bushcricket is rather sharply divided into two azimuthal hemispheres, with signals arriving from any direction within one hemisphere being predominantly represented in the discharge of neurons of this side of the auditory pathway. In addition, each pathway also selects for the most intense of several alternative sounds. A low-intensity signal at 45 dB sound pressure level is quite effective when presented alone, but completely suppressed when given simultaneously with another signal at 60 dB sound pressure level. In a series of intracellular experiments the synaptic nature of the intensity-dependent suppression of competitive signals was investigated in a number of interneurons. The underlying synaptic mechanism is based on a membrane hyperpolarisation with a time-constant in the order of 5–10 s. The significance of this mechanism for hearing in choruses, and for the evolution of acoustic signals and signalling behaviour is discussed. Accepted: 20 November 1999     

Characterization of tactile-sensitive interneurons in the abdominal ganglia of the cockroach,Periplaneta americana

Tactile stimulation of an insect's abdomen evokes various behaviors including grooming and vigorous escape responses. We tested a sample of 37 tactile-sensitive abdominal interneurons for various morphological and physiological characteristics, including their ability to excite thoracic interneurons that are known to integrate wind information conducted by giant interneurons in the classical escape response. The results suggest that abdominal tactile-sensitive interneurons are heterogeneous both in anatomical and physiological properties. In general, these cells are very small interganglionic interneurons that respond to tactile stimulation at more than one abdominal segment. However, the larger population contained virtually all types of cells. Some projected anteriorly, others posteriorly, and still others projected in both directions. For most cells, the soma was on the side opposite to their axons, but in 24% of the cells it was on the same side. Patterns of dendritic arbors also varied among cells. However, tactile sensitivity was in general consistent with the morphological bias noted in dendritic branch patterns. We were able to document the existence of tactile abdominal interneurons that connect directly to thoracic interneurons involved in escape (TI_As). However, instances of demonstrated connectivity were rare. One cell that did show connectivity (AI652) was characterized in detail, and its properties were appropriate for conducting tactile signals in a directional escape system. The dendritic arbors were biased to the side that was ipsilateral to the cell's soma and axon. As a result, this cell's abdominal inputs and thoracic outputs are on the same side. This pattern is appropriate for generating the sensory fields recorded previously in TI_As. Its axon was located in the ventral median tract, which should bring it close to the integrating region of the TI_As. © 1998 John Wiley & Sons, Inc. J Neurobiol 34: 227–241, 1998     

Network activity of "interneurons" and large cells of amygdala in active and passive rabbits

By plotting cross-correlation histograms differences were found in interaction of conjectural small "interneurons" and large principal cells of the central and basal amygdalar nuclei in negative emotional situations. The network activity of "nterneurons" was higher than in principal cells. "Interneurons" more frequently had excitatory and inhibitory input or output connections with neighbouring cells, latency of their connections with other cells was smaller than in principal neurons. Interaction of "interneurons" and principal cells differed in animals with active and passive behavioural strategy in negative emotional situations. As compared to active animals, in passive rabbits inhibitory connections to "interneurons" from other cells occurred more frequently, excitatory or inhibitory connections from "interneurons" to principal cells appeared more rare.     

Microfluidics for in vivo imaging of neuronal and behavioral activity in Caenorhabditis elegans

The nematode C. elegans is an excellent model organism for studying behavior at the neuronal level. Because of the organism's small size, it is challenging to deliver stimuli to C. elegans and monitor neuronal activity in a controlled environment. To address this problem, we developed two microfluidic chips, the 'behavior' chip and the 'olfactory' chip for imaging of neuronal and behavioral responses in C. elegans. We used the behavior chip to correlate the activity of AVA command interneurons with the worm locomotion pattern. We used the olfactory chip to record responses from ASH sensory neurons exposed to high-osmotic-strength stimulus. Observation of neuronal responses in these devices revealed previously unknown properties of AVA and ASH neurons. The use of these chips can be extended to correlate the activity of sensory neurons, interneurons and motor neurons with the worm's behavior.     

Processing of calling songs by an L-shaped neuron in the prothoracic ganglion of the female cricket, Acheta domesticus

ABSTRACT. An L-shaped auditory intemeuron (LI) has been recorded from extracellularly and intracellularly, and identified morphologically (by Lucifer yellow or cobalt injection) in the prothoracic ganglion of mature female Acheta domesticus. The morphology of the LI is very similar to ascending, prothoracic acoustic interneurons that are most sensitive to higher carrier frequencies in both A. domesticus and other gryllid species. Its terminations in the brain are similar to ascending acoustic interneurons found in other gryllids. The LI neuron is most sensitive to 4–5 kHz model calling songs (CSs), the main carrier frequency of the natural call. Thresholds to high frequencies (8–15 kHz) are 15–20 dB higher. Increasing CS intensities of up to 15 dB above threshold at 4–5 kHz result in increased firing rates by the LI. More than 15 dB increase in intensity causes saturation with little increase in spiking rate until the intensity surpasses 80 dB. In response to 70 dB or higher stimulus intensities, the LI responds to the second and third CS syllables with one or two spikes, pauses, and then produces a burst of nerve impulses with the same or greater latency than for lower intensity stimuli. In response to CS syllables of changing duration (10–30 ms) this neuron responds with a rather constant duration burst of impulses. Syllable periods of the CS stimuli were accurately encoded by the LI. Progressively stronger injection of hyperpolarizing current reduces, and ultimately stops spiking of the LI in response to CS stimuli. More intense stimulation with reduced hyperpolarization shows an initial spike, pause and burst of spikes. Intracellular recording from axonal regions of the neuron shows large spikes, small EPSPs and a developing hyperpolarization through the response to a CS chirp. Inhibitory input to the LI is demonstrated at 4.5, 8 and 16 kHz. This probably explains the specialized response characteristics of the LI which enhanced its encoding of CS syllable period.