Auditory evoked transient and sustained potentials in the human EGG: II. Effects of small doses of ethanol

The effect of small doses of ethanol (0.4 ) on auditory evoked transient and sustained potentials was studied. Tones of 1-second duration were presented in trains of four stimuli (interstimulus interval = 1 second; intertrain interval = 1 minute). The electroencephalogram was recorded from derivation Cz-Al. Ethanol depressed the transient responses both at the first stimulus of the train and during repeated stimuli. The sustained potentials elicited by the first stimuli of the train were not affected by ethanol, whereas the sustained potentials elicited by repeated stimuli were larger in amplitude under the influence of ethanol than during control experiments. It is suggested that the decrease of the transient responses under the influence of ethanol is mainly due to depression of the reticular formation, whereas the increase of sustained potentials reflects ethanol-induced release of intracortical inhibition.     

Auditory evoked transient and sustained potentials in the human EEG: I. Effects of expectation of stimuli

The characteristics of auditory evoked transient and sustained potentials were recorded using trains of four-tone stimuli of 1-second duration (interstimulus interval = 1 second) presented once every minute. The subject either attentively expected the stimuli or ignored them while reading. The electroencephalogram was recorded from derivations Cz-Al and Fz-Al. Expectation of the stimuli was associated with increased amplitudes of the transient responses both at the first stimulus of the train and during stimulus repetition. In contrast, the sustained potential at the first stimulus of the train was unchanged or smaller when the subject expected the stimuli. During stimulus repetition, however, the amplitude of the potential was enhanced by expectation of the stimuli. The results support the hypothesis of two sustained potential components and stress the importance of stimulus repetition rate when sustained potentials are studied.     

The development of memory trace depending on the number of the standard stimuli.

The mismatch negativity (MMN) component of the event-related potentials reflects the automatic detection mechanism of sound change. MMN is elicited by a neuronal mismatch process between deviant (infrequent) auditory input and the sensory memory trace of the standard (frequent) stimuli. Although many previous studies have investigated MMN to reveal the sensory memory mechanism, the development of memory representation still remains unclear, in particular, the topographical aspect of the trace-development in sensory memory has not been clarified. We measured the frontal and the temporal MMN components, respectively, when the sound trace was developed as the number of standard stimuli was changed to 1, 3, 5 or 7. In this experiment, the inter-train interval was 15 sec. The stimulus train with the different frequency of 800 Hz, 900 Hz, or 1000 Hz was repeatedly presented. Thus, we reduced the influence of the previous train. For the first time, we found not only the enhanced amplitude but also the shortened latency for both MMN components when the number of standard stimuli was increased. These findings indicate that both frontal and temporal MMN components reflect the development of memory trace depending on the number of standard stimuli.     

Cortical responsiveness during inner speech in schizophrenia: an event-related potential study.

OBJECTIVE: The study assessed the effects of inner speech on auditory cortical responsiveness in schizophrenia. METHOD: Comparison subjects (N=15) and patients with schizophrenia (N=15) were presented with acoustic and visual stimuli during three conditions: while subjects were silent, when spontaneous inner speech might occur; during directed inner speech, while subjects repeated a statement silently to themselves; and while subjects listened to recorded speech. N1 event-related potentials were recorded during the three conditions. RESULTS: N1 event-related potentials elicited by acoustic stimuli, but not by visual stimuli, were lower during directed inner speech than during the silent baseline condition in the comparison subjects but not in the patients. CONCLUSIONS: Abnormal auditory cortical responsiveness to inner speech in patients with schizophrenia may be a sign of corollary discharge dysfunction, which may potentially cause misattribution of inner speech to external voices.     

Auditory event-related potentials in the squirrel monkey: Parallels to human late wave responses

Event-related potentials (ERPs) were recorded from the brain surface in squirrel monkeys during the presentation of two auditory stimulus paradigms which have previously been utilized to elicit scalp-recorded ERPs in humans. In the first paradigm, inter-stimulus interval (ISI) was systematically varied during the presentation of a series of tone pips. The tones produced a negative (70 ms)-positive (130 ms) sequence of components similar in morphology to the human scalp-recorded N1-P2 'vertex' potential. The amplitude of the N70 and P130 components recorded from midline electrodes decreased with decreasing ISI, as previously shown for the human vertex potential. However, this amplitude change with ISI was not observed in ERPs recorded from lateral frontal and temporal electrodes. These results agree with previous studies of monkeys and humans which suggest at least two different sources contribute to N1-P2 components recorded in response to tones. The effects of stimulus probability and novelty on ERP morphology and amplitude were studied in the second paradigm. ERPs elicited by frequent (P = 0.92) and infrequent (P = 0.08) tone pips presented in an unpredictable order were compared. N70 - P130 components were produced by both stimuli, and the infrequent stimuli also elicited a broad, long latency (300 ms) positive complex that decreased in amplitude with repeated presentations. In humans the same infrequent auditory stimuli produce a frontally distributed late positive component that has been interpreted as indicating the activation of orientation mechanisms or of a 'mismatch detector'. These data suggest that in these paradigms squirrel monkeys exhibit ERPs which are similar in several respects to ERPs recorded to identical stimuli in humans.     

Auditory and visual event-related perturbations in the 40 Hz auditory steady-state response.

The effects of salient auditory and visual 'foreground' stimuli on responses to 'background' probe stimuli were investigated. The foreground stimuli were given at long and aperiodic intervals and required a discriminative judgment. Simultaneously, evoked potentials were obtained in response to background probe auditory stimuli presented in a continuous train at about 40/sec. The 40 Hz steady-state rhythm (SSR) evoked under such conditions was extracted using digital averaging and filtering techniques and examined continuously for evidence of change in latency or amplitude during the period surrounding the foreground stimulus. Within the first 200-300 msec after the onset of an acoustic foreground stimulus the latencies of individual peaks in the rhythm were momentarily reduced by a mean of 5.5 msec. A shift in the 40 Hz rhythm was also seen following visual foreground stimuli, although the shift was about one-third that following acoustic stimuli. A latency shift of comparable magnitude was not produced by deliberate manipulation of intensity or signal-to-noise ratio of the stimuli used to evoke the rhythm. The latency shift response is discussed in terms of a transient period of sensory facilitation during orienting or alerting associated with the foreground stimuli.     

Recovery and refractoriness of auditory evoked fields after gaps in click trains

When clicks are presented in a train at a rate above approximately 5 Hz, they evoke a sustained field in human auditory cortex that can be recorded by magnetoencephalography. In this study we evaluated how this sustained field continues when a click train is interrupted by a silent gap. The stimuli were click trains with interclick intervals of either 12 or 24 ms, which produce pitches of 83.3 or 41.7 Hz, respectively. The click trains were 996 ms in duration with a gap of 12, 24, 48, 96, or 192 ms beginning 504 ms post-stimulus onset. The sustained field for click trains with short gaps was similar to the one evoked by a continuous click train. Subtraction of the response evoked by a solitary click train of 504 ms enabled estimation of the sustained field in the interval after the gap. The comparison revealed that the sustained field amplitude after the gap was larger than that at the onset of the initial click train in the interval from 150 to 350 ms after onset, and the difference decreased with gap duration. In contrast, the transient P1m was refractory for gaps up to 48 ms, but had nearly recovered its initial amplitude for gaps of 192 ms. We discuss how these results might relate to the perception, i.e. if an interrupted click train is perceived as one continuous sound with a transient gap or as two successive events.     

Reduced suppression or labile memory? Mechanisms of inefficient filtering of irrelevant information in older adults

Cognitive aging theories emphasize the decrease in efficiency of inhibitory processes and attention control in normal aging, which, in turn, may result in reduction of working memory function. Accordingly, some of these age-related changes may be due to faster sensory memory decay or to inefficient filtering of irrelevant sensory information (sensory gating). Here, event-related brain potentials and the event-related optical signal were recorded in younger and older adults passively listening to tone trains. To determine whether age differentially affects decay of sensory memory templates over short intervals, trains were separated by delays of either 1 or 5 sec. To determine whether age affects the suppression of responses to unattended repeated stimuli, we evaluated the brain activity elicited by successive train stimuli. Some trains started with a shorter-duration stimulus (deviant trains). Results showed that both electrical and optical responses to tones were more persistent with repeated stimulation in older adults than in younger adults, whereas the effects of delay were similar in the two groups. A mismatch negativity (MMN) was elicited by the first stimulus in deviant trains. This MMN was larger for 1- than 5-sec delay, but did not differ across groups. These data suggest that age-related changes in sensory processing are likely due to inefficient filtering of repeated information, rather than to faster sensory memory decay. This inefficient filtering may be due to, or interact with, reduced attention control. Furthermore, it may increase the noise levels in the information processing system and thus contribute to problems with working memory and speed of processing.     

Distractible children show abnormal orienting to non-attended auditory stimuli.

Event-related potentials were recorded in response to intermittently presented, non-attended trains of identical auditory stimuli in healthy 9-year-old children. In abnormally distractible children (n =24), the first tone in each train elicited a significantly larger N1 vertex response than in the non-distractible children (n 24), suggesting that increased distractibility may be associated with an abnormally strong cerebral orienting towards non-attended stimuli. A later negativity at around 300 ms, which increases in amplitude with stimulus repetition and may thus reflect the building up of a functional neuronal representation of the stimulus properties, was significantly smaller in the distractible than in the non-distractible children. These findings demonstrate that event-related potential measures may be useful in helping to understand the information processing found in distractible children.     

The habituation of event-related potentials to speech sounds and tones

We examined the short- and long-term habituation of auditory event-related potentials (ERPs) elicited by tones, complex tones and digitized speech sounds (vowels and consonant-vowel-consonant syllables). Twelve different stimuli equated in loudness and duration (300 msec) were studied. To examine short-term habituation stimuli were presented in trains of 6 with interstimulus intervals of 0.5 or 1.0 sec. The first 4 stimuli in a train were identical standards. On 50% of the trains the standard in the 5th position was replaced by a deviant probe stimulus, and on 20% of the trains the standard in the 6th position was replaced by a target, a truncated standard that required a speeded button press response. Short-term habituation (STH) was complete by the third stimulus in the train and resulted in amplitude decrements of 50-75% for the N1 component. STH was partially stimulus specific in that amplitudes were larger following deviant stimuli in the 5th position than following standards. STH of the N1 was more marked for speech sounds than for loudness-matched tones or complex tones at short ISI. In addition, standard and deviant stimuli that differed in phonetic structure showed more cross-habituation than did tones or complex tones that differed in frequency. This pattern of results suggests that STH is a function of the acoustic resemblance of successive stimuli. The long-term habituation (LTH) of the ERP was studied by comparing amplitudes across balanced 5.25 m stimulus blocks over the course of the experiment. Two types of LTH were observed. The N1 showed stimulus-specific LTH in that N1 amplitudes declined during the presentation of a stimulus, but returned to control levels when a different stimulus was presented in the subsequent condition. In contrast, the P3 elicited by the deviant stimuli showed non-specific LTH, being reduced across successive blocks containing different stimuli. P3s elicited by target stimuli remained stable in amplitude.     

Interstimulus interval effect on event-related potential N270 in a color matching task.

Event-related brain potentials were recorded in a matching task, in which subjects were asked to discriminate if the color of the second stimulus (S2) was the same as the first stimulus (S1). The interstimulus interval between the two stimuli of a pair was of three levels (150 ms, 500 ms and 1000 ms). A negative component about 270 ms after the presentation of S2 was elicited when the color of the two stimuli was not identical for the interstimulus interval of 500 ms and 1000 ms, but not for the interval of 150 ms. This may suggest that N270 represented the response of the brain to conflicting information between different cortical levels.     

Identification of afferent C units in intact human skin nerves

Single unit potentials were recorded with microelectrodes from intact human skin nerves, and the unitary responses to electrical and natural skin stimuli were studied. The unitary discharges were derived from afferent C fibres since the impulses were conducted at C velocity and persisted after preferential blocking of activity in myelinated fibres by nerve compression, whereas they were abolished before the A fibre discharges by lidocaine.The afferent C units adapted slowly to mechanical stimuli and some of them exhibited afterdischarges following withdrawal of the stimuli. The excitability decreased as a consequence of repeated mechanical stimulation. The most vigorous responses were elicited by various intense stimuli such as pinpricks or heat stimulation. Although no definite classification was made, the responses of the human C units to different stimuli reminded of responses in ‘polymodal’ C receptors identified in the cat and the monkey.Painful stimuli elicited the most intense discharges in the units, but considerable activity was also elicited by non-painful stimuli. Some evidence suggested that single or a few repeated impulses in an afferent C unit need not reach consciousness.     

Responses of medullary neurons to stimulation of locomotor and inhibitory loci in the brain stem of the cat

The medullary locomotor point (L) and the pontine inhibitory point (I) were found in mesencephalic decerebellate cats. Repetitive (60/s) microstimulation of L elicited stepping of forelimbs which terminated during repetitive microstimulation of I. Responses of neurons were evoked applying 1.5 s-1 single or paired stimuli to L or a train of 2-4 stimuli to I, the interstimulus interval being 2 ms. Medial neurons (N-301) gave PSPs or action potentials of 1 stimulation three times as often as to L stimulation. Contrary, lateral neurons (N-166) responded two times as often as to L stimulation. IPSPs were recorded in both groups of neurons two times as seldom as EPSPs. In medial neurons IPSPs were produced mainly by I stimulation. I stimulation did not evoke usually IPSPs in neurons excited from L. Possible mechanisms of termination of stepping due to I stimulation are discussed.     

Event-related brain potentials dissociate visual working memory processes under categorial and identical comparison conditions

Event-related potentials (ERPs) have been successfully employed to examine the functional and neuronal characteristics of working memory processes. In the present study, we examined the ERP waveforms in a delayed matching task to examine the cognitive processes underlying category and identity comparison and the effects of stimulus complexity. Subjects had to decide whether two visual stimuli are (a) physically identical (identical comparison condition, IC) or (b) identical, irrespective of their orientation (categorial comparison condition, CC). The stimuli were structured five-point patterns, which varied in complexity. For the ERPs elicited during the 1500 ms retention interval, the following pattern of results was obtained: Stimuli in the CC-condition elicited larger P300 components than in the IC-condition. In the IC-condition, the P300 was followed by a broadly distributed negative slow wave. Moreover, complex patterns elicited a posteriorily distributed negativity at 350 ms (N350), whereas the less complex patterns gave rise to a fronto-centrally distributed slow wave that started around 500 ms. These results suggest that S1 was more elaborately processed in the CC-condition, while the more complex figures were associated with an early classification process during the retention interval.     

Effects of musical training on the early auditory cortical representation of pitch transitions as indexed by change‐N1

The effects of musical training on the early auditory cortical response to pitch transitions in music were investigated by use of the change‐N1 component of auditory event‐related potentials. Musicians and non‐musicians were presented with music stimuli comprising a melody and a harmony under various listening conditions. First, when the subjects played a video game and were instructed to ignore the auditory stimuli, the onset of stimuli elicited a typical, fronto‐central onset‐N1, whereas melodic and harmonic pitch transitions within the stimuli elicited so‐called change‐N1s that were more posterior in scalp distribution. The pitch transition change‐N1s, but not onset‐N1, were enhanced in musicians. Second, when the listeners attended to the same stimuli as above to detect infrequently occurring target stimuli, the change‐N1 elicited by pitch changes (in non‐target stimuli) was augmented, in non‐musicians only when the target was easily detectable, and in both musicians and non‐musicians when it was difficult to detect. Thus, the early, obligatory cortical response to pitch transitions during passive listening was chronically enhanced by training in musicians, and, reflecting this training‐induced enhancement, the task‐related modulation of this response was also different between musicians and non‐musicians. These results are the first to demonstrate the long‐term effects of training, short‐term effects of task and the effects of their interaction on the early (~100‐ms) cortical processing of pitch transitions in music. The scalp distributions of these enhancement effects were generally right dominant at temporal electrode sites, suggesting contributions from the radially oriented subcomponent of change‐N1, namely, the Tb (N1c) wave of the T‐complex.     

Synchronized clusters of action potentials can increase or decrease the excitability of the axons of magnocellular hypothalamic neurosecretory cells

Extracellular recordings were made from supraoptic nucleus (SON) cells in urethane anaesthetized male rats in vivo. Two stimulating electrodes were positioned to activate the cells antidromically, one in the mid axon region of the cells and the other at the axon terminals. Trains of 5-20 just-subthreshold stimuli at 5 s intervals decreased the threshold for antidromic activation from both sites. Whereas neither single stimuli, nor the stimuli at the beginning of a train of 20 stimuli evoked antidromic action potentials, later action potentials did so. Paradoxically, trains of 20 just-suprathreshold stimuli increased the threshold for activation of both axons and terminals. In recordings from the same cells, stimuli were applied singly at 5 s intervals at an intensity which almost invariably evoked an antidromic action potential. Identical stimuli were then applied in trains of 20 stimuli at 50 Hz. After the first train, the initial stimulus pulses of the trains frequently fell below threshold. Following a conditioning train of five stimuli applied to one electrode, the period of decreased threshold (increased excitability) at the other electrode lasted less than 100 ms and the period of increased threshold (decreased excitability) after 12 trains of 20 stimuli lasted between 5 and 10 s. Both decreased and increased excitability were seen at axons and terminals of both putative oxytocin and vasopressin cells. Since the excitability changes were shown in vivo at frequencies encountered during recordings, it is likely that they influence the probability of spike propagation and hormone secretion under physiological conditions.     

Electrophysiological investigation of β, β′-iminodipropionitrile neuropathy: Intracellular recordings in spinal cord

β, β′-Iminodipropionitrile (IDPN) was given to cats (50 mg/kg/week for 5 weeks) to induce giant axonal swellings in the proximal internodes of motor axons. Conventional intracellular recording techniques were used to investigate the influence of the axon swellings on axonal impulse conduction and generation of action potentials in unidentified lumbosacral motoneurons (MN).Action potentials recorded from axon swellings, verified by lack of orthodromically or antidromically elicited EPSPs or IPSPs, afterhyperpolarization potentials or initial segment-somaldendritic (IS-SD) inflections, were variable in shape. Some were indistinguishable from recordings in normal axons. Component or extra potentials occurred in 45% of recordings from axon swellings; their position on the action potential depended on the direction of impulse invasion into the swelling. Many action potentials were broad, with amplitudes less than 30 mV. Impulse conduction was estimated to be blocked in 19% of motor axons tested.Action potentials recorded in MN of IDPN treated cats resembled in many aspects those recorded in chromatolytic MN, with increased latencies upon antidromic stimulation and decreased IS conduction times and SD thresholds; other parameters did not differ significantly. The minimal interval between two stimuli which each evoked action potentials increased from3.3 ± 0.1to5.8 ± 0.5ms. IS-SD portions of the action potentials could not be fractionated in 49% of cells regardless of interpulse interval. Many MN failed to follow frequencies as low as 10 Hz. Delayed depolarizations were observed in 14% of MN recordings. Repetitive action potentials were elicited by single stimuli in 14% of MN and more frequently by orthodromic than antidromic stimulation. Action potentials could often be elicited in the same MN by stimulation of more than one ventral root filament. The incidence of this ephaptic transmission or crosstalk was estimated to be 12%. The findings are discussed in terms of the influence of proximal axon swellings on action potential generation in MN, propagation along non-homogeneous regions of axons and functional chromatolysis.     

Temporal correlates of information processing during visual short-term memory.

The question is raised whether the sequence of spikes of a cortical neuron, i.e. its spike train, is related to cognitive functions. Neuronal patterns of firing in the inferotemporal cortex of monkeys performing visual delayed-matching tasks showed that short-term memory was accompanied by decreased bursting and changes in the incidence of recurrent spike-interval patterns. The temporal structure of the spike train suggests an inverse relationship between the incidence of repeated patterns and the degree of selectivity of sustained firing frequency elicited by the memorandum (sample stimulus).     

Increased transneuronal excitation of the cat lateral geniculate nucleus after acute deafferentation

Single neurons and sum potentials were recorded from the cat optic tract (OT), the lateral geniculate nucleus (LGN) and the optic radiation (OR) before and after visual deafferentation obtained by locally restricted photocoagulation of the retina or by 'total' photocoagulation of the optic disc of 'monocular' cats. Immediately after deafferentation, the spontaneous activity of single LGN-neurons fell to a very low level (is much less than 0.1 impulses/sec). This nearly total depression of neuronal activity was followed by a slow but incomplete recovery of the spontaneous impulse rate which reached about 1-2 impulses/sec, 2-4 days after deafferentation. The 'normal' LGN neuron impulse rate was about 8-30 impulses/sec. After deafferentation the impulse pattern showed an increased occurrence of double discharges separated by very long intermittent intervals. The excitability of LGN principal cells activated by OT electrical stimuli increased immediately after deafferentation. Double and multiple discharges were then elicited by single optic tract stimuli. The deafferentation hyperexcitability was abolished temporarily by OT stimulus trains with a frequency range > 10 stimuli/sec, and returned to the prestimulus level within 0.5-1sec after the stimulus train was terminated. Antidromic conditioning stimuli did not influence the deafferentation hyperexcitability. The time course of the postdeafferentation hyperexcitability was separated into two parts: an immediate rise in excitability after deafferentation was followed by a further slow increase in excitability within the first 2 h after deafferentation. The hyperexcitability measurable at a single unit level was already maximal after locally restricted deafferentation and did not increase with the extension of the retinal lesion beyond 1-2 mm diameter, corresponding to 4-8 degrees in the visual field. Intracellular recordings after deafferentation displayed no significant increase in the amplitude of subthreshold EPSPs, but additional slow, aperiodic, depolarizing waves were found to modify the resting membrane potential. From the multiple discharge pattern elicited by single OT electrical stimuli, one can conjecture that the suprathreshold EPSPs, however, increased considerably in their amplitude. The evoked potentials (OR-waves r1, r2) elicited by electrical OT stimuli increased in amplitude and the small waves r3-r6 became more prominent after deafferentation. All postsynaptic r-waves reached their maximum values about 1.5 h after interruption of the optic nerve signal flow and did not change further within the following 28 h.