Facts and reflections on thalamo cortical reverberation of impulses

Experiments on cats, either unanesthetized or anesthetized with various doses of pentobarbital, showed that the cortical rhythmic after-discharge ("slow after-activity"), which has been regarded as a manifestation of reverberation of impulses in thalamocortical circuits [17], consists of a burst of spontaneous "spindles" evoked by stimulation. This conclusion is supported by the following facts: Spontaneous "spindles" and the rhythmic after-discharge respond absolutely identically (disappear) to activation of the EEG and deepening of pentobarbital anesthesia. The absence of thalamocortical reverberation is also indicated by the preservation of a rhythmic after-discharge (to clicks), synchronous with the cortex, in the thalamic relay nucleus (the medial geniculate body) after cooling or after removal of its projection zone.     

Auditory evoked responses to rhythmic sound pulses in dolphins

The ability of auditory evoked potentials to follow sound pulse (click or pip) rate was studied in bottlenosed dolphins. Sound pulses were presented in 20-ms rhythmic trains separated by 80-ms pauses. Rhythmic click or pip trains evoked a quasi-sustained response consisting of a sequence of auditory brainstem responses. This was designated as the rate-following response. Rate following response peak-to-peak amplitude dependence on sound pulse rate was almost flat up to 200 s⁻¹, then displayed a few peaks and valleys superimposed on a low-pass filtering function with a cut-off frequency of 1700 s⁻¹ at a 0.1-amplitude level. Peaks and valleys of the function corresponded to the pattern of the single auditory brain stem response spectrum; the low-pass cut-off frequency was below the auditory brain stem response spectrum bandwidth. Rate-following response frequency composition (magnitudes of the fundamental and harmonics) corresponded to the auditory brain stem response frequency spectrum except for lower fundamental magnitudes at frequencies above 1700 Hz. These regularities were similar for both click and pip trains. The rate-following response to steady-state rhythmic stimulation was similar to the rate-following response evoked by short trains except for a slight amplitude decrease with the rate increase above 10 s⁻¹. The latter effect is attributed to a long-term rate-dependent adaptation in conditions of the steady-state pulse stimulation. Accepted: 18 June 1998     

KILLER WHALE (ORCINUS ORCA) AUDITORY EVOKED POTENTIALS TO RHYTHMIC CLICKS

Temporal auditory mechanisms were measured in killer whales ( Orcinus orca ) by recording auditory evoked potentials (AEPs) to clicks. Clicks were presented at rates from 10/sec to 1,600/sec. At low rates, clicks evoked an AEP similar to the auditory brainstem response (ABR) of other odontocetes; however, peak latencies of the main waves were 3–3.7 msec longer than in bottlenose dolphins. Fourier analysis of the ABR showed a prominent peak at 300–400 Hz and a smaller one at 800–1,200 Hz. High-rate click presentation (more than 100/sec) evoked a rate-following response (RFR). The RFR amplitude depended little on rate up to 400/sec, decreased at higher rates and became undetectable at 1,120/sec. Fourier analysis showed that RFR fundamental amplitude dependence on frequency closely resembled the ABR spectrum. The fundamental could follow clicks to around 1,000/sec, although higher harmonics of lower rates could arise at frequencies as high as 1,200 Hz. Both RFR fundamental phase dependence on frequency and the response lag after a click train indicated an RFR group delay of around 7.5 msec. This corresponds to the latency of ABR waves PIII-NIV, which indicates the RFR originates as a rhythmic, overlapping ABR sequence. The data suggest the killer whale auditory system can follow high click rates, an ability that may have been selected for as a function of high-frequency hearing and the use of rapid clicks in echolocation.     

Dynamics of brain rhythmic and evoked potentials

In this study, the simultaneously recorded and selectively averaged evoked potentials in some of the structures of the auditory pathway (acoustical cortex, medial geniculate nucleus, inferior colliculus), in the reticular formation and the hippocampus of the cat during sleep as well as the simultaneous amplitude frequency characteristics of these structures are given. The power spectral density functions computed from the simultaneously recorded spontaneous activities of these structures are also presented. Using these results, the following analyses are accomplished:(1) determination of the dynamics of potentials simultaneously obtained from various structures, in order to evaluate the common features of their system characteristics; (2) determination of the relationship (or interactions) between rhythmic activity and evoked potentials of the brain.     

Network dynamics and synchronous activity in cultured cortical neurons

Neurons extracted from specific areas of the Central Nervous System (CNS), such as the hippocampus, the cortex and the spinal cord, can be cultured in vitro and coupled with a micro-electrode array (MEA) for months. After a few days, neurons connect each other with functionally active synapses, forming a random network and displaying spontaneous electrophysiological activity. In spite of their simplified level of organization, they represent an useful framework to study general information processing properties and specific basic learning mechanisms in the nervous system. These experimental preparations show patterns of collective rhythmic activity characterized by burst and spike firing. The patterns of electrophysiological activity may change as a consequence of external stimulation (i.e., chemical and/or electrical inputs) and by partly modifying the "randomness" of the network architecture (i.e., confining neuronal sub-populations in clusters with micro-machined barriers). In particular we investigated how the spontaneous rhythmic and synchronous activity can be modulated or drastically changed by focal electrical stimulation, pharmacological manipulation and network segregation. Our results show that burst firing and global synchronization can be enhanced or reduced; and that the degree of synchronous activity in the network can be characterized by simple parameters such as cross-correlation on burst events.     

Dynamics of brain rhythmic and evoked potentials

In this study, the simultaneously recorded and selectively averaged evoked potentials in some of the structures of the auditory pathway (acoustical cortex, medial geniculate nucleus, inferior colliculus), in the reticular formation and the hippocampus of the awake cat as well as the simultaneous amplitude frequency characteristics of these structures are given. The power spectral density functions computed from the simultaneously recorded spontaneous activity of these structures are also presented. Using these results, the following analyses are accomplished: (1) determination of the dynamics of potentials simultaneously obtained from various brain structures, in order to evaluate the common features of their system characteristics; (2) determination of the relationship (or interactions) between rhythmic activity and evoked potentials of the brain, and (3) elaboration of a working hypothesis for the dynamics of potentials of the brain. Some suggestions and comments are also made for investigators working toward theories or dynamic models of signal transmission in the brain.     

Evoked potentials of the auditory cortex on paired stimuli

Changes in the responsiveness of the auditory cortex to an acoustic click and to direct stimulation of the medial geniculate body were studied by the method of evoked potentials in an extended experiment on cats with implanted electrodes. It is shown that the minimum interval between two stimuli for which a second click produces an EP in the auditory cortex is from 30 to 50 msec. The relative refractory period consists of two parts. The first (50–100 msec) is characterized by a rapid recovery, and the second (about 500 msec) by a slow recovery. In contrast with a click, direct stimulation of the geniculate body does not produce a refractory condition but one of facilitation. The effects of Nembutal and chloralose anesthesia and the state of alertness on the recovery of auditory cortex responsiveness were investigated. The reason for the absence and the reduction of an EP from the auditory cortex to a testing click during absolute and relative refractory periods is not a passive decrease of excitability of the usual refractory kind, but an active interplay of excitatory and inhibitory processes in the cerebral cortex, geniculate bodies, and reticular formation of the brain stem.A. A. Bogomolets' Institute of Physiology, Academy of Sciences, Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 1, pp. 54–64, July–August, 1969.     

Auditory Evoked Bursts in Mouse Visual Cortex during Isoflurane Anesthesia

General anesthesia is not a uniform state of the brain. Ongoing activity differs between light and deep anesthesia and cortical response properties are modulated in dependence of anesthetic dosage. We investigated how anesthesia level affects cross-modal interactions in primary sensory cortex. To examine this, we continuously measured the effects of visual and auditory stimulation during increasing and decreasing isoflurane level in the mouse visual cortex and the subiculum (from baseline at 0.7 to 2.5 vol % and reverse). Auditory evoked burst activity occurred in visual cortex after a transition during increase of anesthesia level. At the same time, auditory and visual evoked bursts occurred in the subiculum, even though the subiculum was unresponsive to both stimuli previous to the transition. This altered sensory excitability was linked to the presence of burst suppression activity in cortex, and to a regular slow burst suppression rhythm (∼0.2 Hz) in the subiculum. The effect disappeared during return to light anesthesia. The results show that pseudo-heteromodal sensory burst responses can appear in brain structures as an effect of an anesthesia induced state change.     

The excitability of neurons of the medial septal area in brain slices from hibernating and active Citellus undulatus susliks]

Spontaneous and evoked neuronal activity of the medical septum-diagonal band complex (MS-DB) has been investigated in slices from the brain of hibernating and active ground squirrels, as well as guinea pigs. In all experimental groups, the majority of the MS-DB neurones exhibited high regular of rhythmic burst spontaneous activity which persisted even after synaptic blockade in half of the neuronal population. Under the same conditions, the activity of the surrounding structures was completely suppressed. The density of the spontaneously active neurones in the slices, as well as the mean frequency of discharges in the MS-DB of hibernating ground squirrels, were significantly higher than in active ground squirrels and guinea pigs. Stimulation of the medial forebrain bundle evoked initial suppression of the activity in the majority of MS-DB units; in many of them, the suppression was followed by a burst discharge. Neurones with background rhythmic burst activity always reacted by resetting the spontaneous bursts. In total, 50-60% of the MS-DB neurones in active ground squirrels and guinea pigs reacted by post-inhibitory bursts, whereas in hibernating animals these responses were observed nearly in all neurones. Threshold values of the stimulating current were lower in hibernating animals; the intraburst density of spikes was increased.     

Effect of forward masking on evoked potentials of auditory cortex in guinea pigs

Thresholds of the event-related potentials (ERPs) appearance were measured for one stationary and four moving auditory images presented in silence or under forward masking conditions. The difference between thresholds in silence and after noise masker was considered as masking level. Under the forward masking, the amplitude of the ERP to the first click in the test series decreased in guinea pig auditory cortex. Masking level decreased with the time lag between signal and masker and didn't depend on the fused auditory image localization that corresponded to the first click in different test signals. This fact can support the hypothesis that for the long test signals the initial part can be masked more than the final one. The ERPs amplitude to next clicks in test series depended on interaction of two factors: forward masking in the "masker-signal" system and interaction of separate ERPs in the series evoked by the test signal.     

Responses of motor cortex neurons of the cat to auditory stimulation and their role in the instrumental food reflex

The responses of motor cortex neurons in the cat to the presentation of a single auditory click and a series of 10 clicks presented with 1,000/sec frequency were studied under conditions of chronic experiments before and after the development of an instrumental food reflex. After reflex development a single presentation of a positive conditioned stimulus (single click) markedly influenced for 7 sec the appearance of instrumental movements. At the same time, the immediate responses of motor cortex neurons to presentation of the conditioned auditory stimulus had no impact on the appearance in the motor cortex of discharges leading to the realization of instrumental movements. Consequently, motor cortex neurons do not require activation from afferent sensory inputs for the generation of such discharges. The immediate neuronal responses to conditioned stimulation did not inhibit the realization of the instrumental reflex. It is proposed that they are associated with the realization of motor function in the unconditioned defensive response evoked by the presentation of an auditory stimulus. The presence or absence of responses to auditory conditioned stimulation was dependent upon the signal meaning of the stimulus, its physical parameters, and the degree of excitability of the animal.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 539–550, July–August, 1985.     

Stimulation of the hippocampus and its effect on electrographic manifestations of the brain in unrestrained rats.

The dorsal hippocampus was electrically stimulated in unanaesthetized, unrestrained rats with a cobalt-gelatin rod in their cortex. The significance of the hippocampus in the elicitation of both physiological spontaneous rhythmic activity (episodic activity of 8--9/sec frequency bound, in rats, to a state of quiet wakefulness, and "sleep spindles") and pathological rhythmic activity of the self-sustained after-discharge (SSAD) type was determined from the aspect of the EEG and behavioural characteristics. 1. Single electrical pulses (0.1 msec, 1--10 V, 0.3/sec) elicited an evoked potential bilaterally in the somatosensory cortex. Elicitation of rhythmic after-activity (of the type of episodes or sleep spindles) was observed only in some cases in which an adequately strong stimulus was used. 2. Repeated series of rhythmic electrical stimuli following each other at short intervals (2--3 min) led to the formation of SSAD in about one third of the cases and at all stimulation frequencies (3-15/sec), although low frequencies (3--4/sec) were the least effective. The character of the SSAD and simultaneous behavioural phenomena differed fundamentally from those evoked by electrical stimulation of the thalamus (Chocholová et al. 1977). The development of paroxysmal after-activity was signalled by responses of a more or less distinct "recruiting" character during stimulation. On the basis of a comparison of electrographic and behavioural manifestations after electrical stimulation of the thalamus and hippocampus, the possibility of both thalamic and extrathalamic projection from the hippocampus to the cortical region is considered.     

Relation of unit spike discharges and evoked potentials in the auditory cortex of cats.

Extracellular microelectrode recordings were made from the auditory cortex of anaesthetized cats during acoustic click stimulation. The microelectrode of low resistance allowed to record evoked field potentials and unit discharges simultaneously. In distant extracellular leads the relation of unit discharges and field potentials was equivocal. Near extracellular leads revealed that the antidromic invasion of the somadendritic membrane by excitation is a frequency dependent process (just as evoked field potentials) while spike potentials can reliably be elicited from the initial segment at high frequencies. It is assumed that the excitation spreading from the initial segment to the soma-dendritic membrane represents an important component of the evoked potentials, and their frequency dependence may be traced back to inhibitions activated by afferent impulses.     

Magnetic stimulation of the human motor cortex evokes skin sympathetic nerve activity.

Single-pulse magnetic coil stimulation (Cadwell MES 10) over the cranium induces without pain an electric pulse in the underlying cerebral cortex. Stimulation over the motor cortex can elicit a muscle twitch. In 10 subjects, we tested whether motor cortical stimulation could also elicit skin sympathetic nerve activity (SSNA; n = 8) and muscle sympathetic nerve activity (MSNA; n = 5) in the peroneal nerve. Focal motor cortical stimulation predictably elicited bursts of SSNA but not MSNA; with successive stimuli, the SSNA responses did not readily extinguish (94% of discharges to the motor cortex evoked SSNA responses) and had predictable latencies [739 +/- 33 (SE) to 895 +/- 13 ms]. The SSNA responses were similar after stimulation of dominant and nondominant sides. Focal stimulation posterior to the motor cortex elicited extinguishable SSNA responses. In three of six subjects, anterior cortical stimulation evoked SSNA responses similar to those seen with motor cortex stimulation but without detectable movement; in the other subjects, anterior stimulation evoked less SSNA discharge than that seen with motor cortex stimulation. Contrasting with motor cortical stimulation, evoked SSNA responses were more readily extinguished with 1) peripheral stimulation that directly elicited forearm muscle activation accompanied by electromyograms similar to those with motor cortical stimulation; 2) auditory stimulation by the click of the energized coil when off the head; and 3) in preliminary experiments, finger afferent stimulation sufficient to cause tingling. Our findings are consistent with the hypothesis that motor cortex stimulation can cause activation of both alpha-motoneurons and SSNA.     

States of excitability in ovulation hormone producing neuroendocrine cells of Lymnaea stagnalis (gastropoda) and their relation to the egg-laying cycle

The electrotonically coupled network of about 100 neuroendocrine caudodorsal cells (CDC) of the freshwater snail Lymnaea stagnalis exhibits three states of excitability with distinct electrophysiological characteristics. Transitions between these states occur spontaneously or can be induced experimentally. The CDC produce an ovulation hormone, and the excitability states are clearly related to the egg-laying cycle of the snail. Two hours before egg laying, the cells enter an active state, which lasts one hour. This phase is characterized by a spontaneous firing pattern, which in preparations can be evoked as an afterdischarge, and during which the hormone is thought to be released. After this, the cells enter an inhibited state in which no other activity than directly stimulus-dependent ortho- and antidromic action potentials can be evoked. This phase lasts till about four hours after egg laying. The subsequent resting state is characterized by facilitation of the responses upon repetitive stimulation of the cells, leading to depolarization of the network and additional action potentials. In this phase, an afterdischarge can be evoked, which brings the cells into the active stage again.     

Postsynaptic neuronal response of a cat sensorimotor cortex during evoked and self-sustained rhythmic "spike and wave" activity

Intracellular correlates of complex sets of rhythmic cortical "spike and wave" potentials evoked in sensorimotor cortex and of self-sustained rhythmic "spike and wave" activity were examined during acute experiments on cats immobilized by myorelaxants. Rhythmic spike-wave activity was produced by stimulating the thalamic relay (ventroposterolateral) nucleus (VPLN) at the rate of 3 Hz; self-sustained afterdischarges were recorded following 8–14 Hz stimulation of the same nucleus. Components of the spike and wave afterdischarge mainly correspond to the paroxysmal depolarizing shifts of the membrane potential of cortical neurons in length. After cessation of self-sustained spike and wave activity, prolonged hyperpolarization accompanied by inhibition of spike discharges and subsequent reinstatement of background activity was observed in cortical neurons. It is postulated that the negative slow wave of induced spike and wave activity as well as slow negative potentials of direct cortical and primary response reflect IPSP in more deep-lying areas of the cell bodies, while the wave of self-sustained rhythmic activity is due to paroxysmal depolarizing shifts in the membrane potential of cortical neurons.I. S. Beritashvili Institute of Physiology, Academy of Sciences of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 18, No. 3, pp. 298–306, May–June, 1986.     

Dynamic characteristics of the auditory cortex of guinea pigs observed with multichannel optical recording

The spatiotemporal characteristics of neural activity in the guinea pig auditory cortex are investigated to determine their importance in neural processing and coding of the complex sounds. A multi-channel optical recording system has been developed for observing the cortical field of the mammalian brain in vivo. Using the voltage-sensitive dye: RH795, optical imaging was used to visualize neural activity in the guinea pig auditory cortex. Experimental results reveal a boomerang-shaped pattern of movement of activated neural cell regions for the evoked response to click as complex sounds. Parallel and sequential neural processing structure was observed. Although the exact frequency selectivities of single cells and tonotopical organization observed using microelectrode were not visible, the similar feature to the microelectrode evidences was imaged by extracting the strongly response field from the optical data.     

Fast detection of unexpected sound intensity decrements as revealed by human evoked potentials

The detection of deviant sounds is a crucial function of the auditory system and is reflected by the automatically elicited mismatch negativity (MMN), an auditory evoked potential at 100 to 250 ms from stimulus onset. It has recently been shown that rarely occurring frequency and location deviants in an oddball paradigm trigger a more negative response than standard sounds at very early latencies in the middle latency response of the human auditory evoked potential. This fast and early ability of the auditory system is corroborated by the finding of neurons in the animal auditory cortex and subcortical structures, which restore their adapted responsiveness to standard sounds, when a rare change in a sound feature occurs. In this study, we investigated whether the detection of intensity deviants is also reflected at shorter latencies than those of the MMN. Auditory evoked potentials in response to click sounds were analyzed regarding the auditory brain stem response, the middle latency response (MLR) and the MMN. Rare stimuli with a lower intensity level than standard stimuli elicited (in addition to an MMN) a more negative potential in the MLR at the transition from the Na to the Pa component at circa 24 ms from stimulus onset. This finding, together with the studies about frequency and location changes, suggests that the early automatic detection of deviant sounds in an oddball paradigm is a general property of the auditory system.     

Producing the cortical phenomenon of "kindling" during acute experiments on cats

Acute experiments on cats enabled spontaneous epileptogenic foci to be produced by means of electrical stimulation of the cerebral cortex. The central area of the suprasylvian gyrus was stimulated by 5-sec stimuli at 3-sec intervals. The strength of neuronal response gradually increased until spontaneous electrical activity set in as stimulation was set at a fixed intensity which had originally evoked local residual discharge at stimulation foci only, i.e., the phenomenon of "kindling" was observed. When "kindling" was produced in the cortical area under study, bursts of spindle activity were recorded on all ECoG with increasing frequency. Recordings of spindle activity then changed to a "spike-wave" pattern of activity. The results of these investigations, which were performed on an isolated strip of cortex, point to the involvement of subcortical structures in mediating "kindling" of the cortical focus.Institute of Clinical and Experimental Neurology, Ministry of Public Health of the Georgian SSR, Tbilisi. Translated from Neirofiziologiya, Vol. 17, No. 5, pp. 601–606, September–October, 1985.     

Neurotensin Regulation of Endogenous Acetylcholine Release from Rat Cerebral Cortex: Effect of Quinolinic Acid Lesions of the Basal Forebrain

The effects of neurotensin (NT) on endogenous acetylcholine (ACh) release from basal forebrain, frontal cortex, and parietal cortex slices were tested. The results show that NT differentially regulates evoked ACh release from frontal and parietal cortex slices without altering either spontaneous or evoked ACh release from basal forebrain slices. In the frontal cortex, NT significantly inhibited evoked ACh release by a tetrodotoxin (TTX)-insensitive mechanism, suggesting an action directly on cholinergic terminals. In the parietal cortex, NT enhanced evoked ACh release by a TTX-sensitive mechanism, suggesting an action of NT on the cholinergic neuron or in close proximity to the cholinergic neuron. The effects of NT on ACh release were confined to evoked ACh release; that is, spontaneous ACh release was not affected. NT did not affect spontaneous or potassium-evoked ACh release from occipital cortex slices. The second set of experiments tested the effects of quinolinic acid (QUIN) lesions of the basal forebrain cell bodies on the NT-induced regulation of evoked ACh release in the cerebral cortex. QUIN lesions of basal forebrain cell bodies caused decreases in choline acetyltransferase activity (27 and 28%), spontaneous ACh release (14 and 21%), and evoked ACh release (38 and 44%) in frontal and parietal cortex, respectively. In addition, 11 days following QUIN lesions of basal forebrain cell bodies, the action of NT to regulate evoked ACh release in frontal cortex or parietal cortex was no longer observed. The results suggest that in the rat frontal and parietal cortex, NT differentially regulates the activity of cholinergic neurons by decreasing and increasing evoked ACh release, respectively.(ABSTRACT TRUNCATED AT 250 WORDS)