Timo Järvilehto

Short-term habituation of the auditory evoked response in man ☆

Abstract 1. 1. In six healthy subjects tha changes in the auditory evoked vertex response occurring in the course of a short train of clicks were studied in order to determine quantitatively the dependence of the response amplitude on the number of stimulus presentations and on the stimulus interval. Care was taken to control the subjects vigilance and attention. 2. 2. The peak-topeak amplitude of the individual response to the first stimulus of a train varied considerably and was on the average larger when the train was attentively expected. In most cases the amplitude distributions differed from normal. The peak-to-peak amplitude of the computed average response was always smaller than the calculated mean amplitude of the corresponding single responses. 3. 3. With the presentation of a stimulus train all components of the average response except P2a decreased rapidly in amplitude. With a stimulus interval of 1 sec the response had reached the asymptotic level at the third stimulus. At a stimulus interval of 3 sec the decrease was less rapid and less pronounced. In all cases the amplitude decrease followed approximately a negative exponential function of the number of stimulus presentations. In part of the components the amplitude decrease coincided with a latency reduction. 4. 4. The results show that human vertex response satisfies several main characteristics of habituation.

Effect of stimulus repetition on negative sustained potentials elicited by auditory and visual stimuli in the human EEG.

Effect of repetition of auditory and visual stimuli on sensory-evoked sustained potentials was studied. The stimuli were tones and flashes of 1 sec duration presented in trains of six stimuli with an inter-stimulus-interval of 1 sec. The repetition rate of the trains was 1 train/min. The EEG was recorded from electrodes located at Cz, Pz and Oz. Both auditory and visual stimuli elicited negative sustained potentials which during the first stimuli of the trains were maximal in amplitude and shortest in latency at Cz. Repetition of the stimulus resulted in almost complete disappearance of the sustained potentials; for auditory stimulation a small negative shift was recorded only at Cz, whereas for visual stimulation a small sustained potential was seen only at Pz and Oz. The results indicate that the slow potentials evoked by short tones or lights are composed of at least two components. One of these components may reflect neural processing associated with orienting reflex. The other component seems to be related to specific neural processing of the stimulus.

Peripheral neural basis of tactile sensations in man: II. Characteristics of human mechanoreceptors in the hairy skin and correlations of their activity with tactile sensations

Properties of the human mechanoreceptors in the hairy skin of the back of the hand were studied by microelectrode measurements from the radial nerve. Correlations of unit activity with sensations elicited by tactile pulses (single cycle sinusoids of 20, 60 and 150 HZ) were examined with simultaneous measurements of unit activity and sensation thresholds and magnitude. A total of 264 mechanoreceptive units were identified. Of all units 66% were classified as slowly adapting (SA) and 34% as rapidly adapting (RA) units. Mechanical thresholds of the units as well as simultaneously measured sensation thresholds decreased with increasing frequency of the pulse. The thresholds of several SA units were identical with the subjective thresholds. The responses of the units to supraliminal pulses consisted maximally of 7 impulses. Most SAI and RA units were able to code to some extent the stimulus amplitude on the basis of number of impulses, but only RA units had stimulus-response functions indicating velocity coding. Comparisons of the estimates of sensation magnitude with the number of impulses in the response indicated that the estimate may be based mainly on activity in a population of RA units. The comparison of the present results with earlier reports on properties of receptors in the glabrous skin of the human hand indicates that there are some differences between the characteristics of receptors in the hairy and glabrous skin. However, human receptors in the hairy skin do not seem to differ from the corresponding receptors in the animals.

Paradoxical heat sensations during moderate cooling of the skin

Paradoxical heat sensations during cooling of the skin were examined in two experiments. In Expt. I the number of occurrences of sensation was studied in 19 naive test subjects (Ss) when cooling from thermal indifference both without and with preceding heating. Without preceding heating 13 Ss reported sensations of paradoxical heat (9.8% of all stimulations). Preheating markedly facilitated the occurrence of the sensations (35% of all stimulations). In Expt. II the effects of cooling velocity (velocities 0.4, 0.7 and 2.0 degrees C/s) and the type of skin area stimulated (hairy or glabrous skin of the hand) on the thresholds of paradoxical sensations were studied in 4 Ss without and with preheating. Cooling velocity, type of skin area and preheating had significant effects on the sensation thresholds, the thresholds being the higher (i.e. the sensation appearing at lower stimulation temperatures) the higher the cooling velocity, if the stimuli were applied to the glabrous skin, or if no preheating was used. The results confirm the existence of paradoxical heat sensations during cooling of the skin and suggest that the sensation is mediated by polymodal units supplied by C-fibers.

AUDITORY and VISUAL EVOKED POTENTIALS OF SCHOOLBOYS WITH SPELLING DISABILITIES

EEG evoked potentials were studied in 23 young adolescent poor spellers and 21 controls. Findings typical of poor spellers were long auditory evoked potential (AEP) latencies and low amplitudes of early AEP déflections. à high frequency of spelling errors concerning the word as à whole was associated with low AEP P50 and P300 amplitudes. Poor reading was associated with high AEP sustained potential amplitudes and à long AEP P185 latency. Generally, poor spellers seem to have problems with the early filtering processes of of attention, whereas spelling errors concerning the word as à whole seem to be associated with problems of late attentional processes. The abnormal findings in the poor spellers are interprétéd in terms of à brain maturational delay that presents as an attentional disorder.

Peripheral neural basis of tactile sensations in man: I. Effect of frequency and probe area on sensations elicited by single mechanical pulses on hairy and glabrous skin of the hand

Tactile thresholds and sensation magnitudes for single mechanical pulses of varying frequency and probe area were studied in order to provide psychophysical data for correlations of tactile sensations with functional properties of different types of human peripheral nerve fibers. Single mechanical pulses were delivered to the hairy or glabrous skin of the hand by means of a perspex probe (area 0.07, 0.7, 3.1, or 12.5 sq.-mm) fixed to the moving coil of an electromechanical vibrator. The frequency of the single pulse was 20, 60 or 150 HZ. Absolute and touch thresholds were measured with a method of production. Suprathreshold sensations were measured with the method of magnitude estimation (6 predetermined displacement amplitudes ranging from 50 to 950 micrometers). Both absolute and touch thresholds were significantly higher on the hairy than on the glabrous skin. The absolute thresholds varied between 44 and 140 micrometers on the hairy and between 9 and 45 micrometers on the glabrous skin; the touch threshold varied between 75 and 300 micrometers and 24 and 153 micrometers, respectively. Lower threshold values were obtained on both skin areas with increasing frequency of the pulse and probe area, the latter effect being significant only for the hairy skin. Standardized magnitude estimation functions could be described with power functions on both skin areas, the exponents of the functions varying between 0.70 and 1.41 on the hairy and between 0.55 and 0.86 on the glabrous skin. The equi-sensation contours showed the difference of sensitivity between the skin areas. Larger sensation magnitudes were obtained with increasing frequency of the pulse and probe area on both skin areas. These psychophysical findings indicate that there is a functional difference at absolute sensation thresholds between hairy and glabrous skin. This difference, however, disappears at higher sensation levels.

Vibrotactile atonal interval correlated with activity in peripheral mechanoreceptive units innervating the human hand

The quality of sensations and the vibrotactile atonal interval-the gap between detection and vibration thresholds-were studied with vibratory stimuli of varying frequency (20, 80 and 160 Hz) and duration (100, 400 and 800 ms) applied to the hairy and glabrous skin of the hand. Detection and vibration thresholds were also determined while simultaneously recording single unit activity from the radial nerve innervating the hairy skin of the hand. Both thresholds were lower on the glabrous than the hairy skin, and the thresholds decreased on both skin areas with increasing vibration frequency. A sensation of short duration was elicited at detection threshold only with a 20-Hz stimulus of 100-ms duration; with other frequency-duration combinations sensations of longer duration were reported. Considerably larger vibration amplitudes were needed on both skin areas for the sensations to be unequivocal with respect to duration and pitch (vibration threshold). There was no significant effect of stimulus duration on vibrotactile thresholds. The width of the average atonal intervals was above 10 dB on both skin areas, and with increasing vibration frequency, decreasing values of atonal intervals were obtained on the hairy skin, whereas considerably increasing values were obtained on the glabrous skin. Recording of single unit activity indicates that on the hairy skin detection of the stimulus at 20 Hz is correlated with activation of slowly adapting (SA) type II and the most sensitive rapidly adapting (RA) units, while distinct vibratory sensations involve entrainment of RA units. Also at 80 Hz, non-pacinian units could contribute to the mechanism of vibrotactile thresholds, whereas at 160 Hz only pacinian (PC) units are involved.

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.

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 g/kg) 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.

Somatosensory evoked potentials associated with tactile stimulation at detection threshold in man

Somatosensory evoked potentials elicited by sub- and supraliminal tactile pulses were studied during continuous threshold measurement. Two threshold methods were used: a modified tracking method and a detection method. With both methods threshold estimates of the same order of magnitude were obtained. Stimuli just above the threshold elicited a distinct somatosensory evoked potential with components P50, P100, N190 and P400. No such SEP was associated with the subliminal stimuli. However, in some cases some potential oscillations were obtained, time-locked to the subliminal stimuli; these are suggested to be due to errors in responding. In control experiments detected stimuli elicited a distinct SEP, but no SEP was associated with undetected stimuli of identical amplitude. The results indicate that in the psychophysical threshold determination the neural processing reflected in the SEP is associated only with supraliminal tactile stimulation. The lack of evoked brain activity associated with subliminal tactile stimuli supports the hypothesis derived from human microneurographic studies, stating that the tactile detection threshold may be based on an extremely small peripheral input, perhaps only on a single impulse in a single peripheral fiber.