A rapid evoked potential index of cortical adaptation

Contrast thresholds and acuity limits were measured in 4 observers with the swept visual evoked potential (VEP) technique. In this technique, grating contrast or grating spatial frequency is electronically varied while the subject's evoked response is retrieved in real time (without averaging). Contrast or spatial frequency variation make the stimulus vary in intensity; zero VEP response amplitude indicates the threshold intensity.Large shifts occur in the indicated threshold when stimulus sweep direction is reversed. Thresholds are always relatively elevated when the run begins with the strongest stimulus value. These shifts do not have a technical origin in the delay of the instrument (Nelson et al. 1984b). Here, it is shown that the shifts are due to orientation and spatial frequency selective adaptation, probably of cortical origin.Measurable adaptation is produced by momentary exposure to contrasts as low as 1.25%; nearly maximum adaptation (0.6 log units) is reached with 20% contrast. These findings support the concept of a contrast gain control mechanism in visual cortex, and pose practical problems for visual assessment with the evoked potential.     

Adaptation to warming but not cooling at slow rates of stimulus change in thermal threshold measurements

The relationship between thermal detection threshold and rate of temperature change of the thermal stimulus when slow (<1 degrees C s(-1)) rates of change are employed was investigated. Using both the reaction time (RT) inclusive Method of Limits and RT exclusive Method of Levels healthy volunteers had warming (WDT) and cooling detection thresholds (CDT) measured at four different rates of temperature change (0.3, 0.5, 0.7 and 1.0 degrees C s(-1)) from the thenar and/or mental regions using a contact thermode. With the Method of Limits, CDT increased linearly with rate of temperature change suggesting increments were due to RT artefacts. This was further supported by threshold assessment with the Method of Levels which showed CDT were unaffected by the rate of change in the RT exclusive method (P > 0.1). In contrast, WDT did not increase linearly with rate of stimulus temperature change when the Method of Limits was used and threshold assessment with the Method of Levels showed WDT assessed using a 0.3 degrees C s(-1) ramp rate were significantly higher than those measured with a 1 degrees C s(-1) rate of change (P < 0.05). This study indicates that adaptation to a warming stimulus can occur at faster rates of stimulus change than previously anticipated and identifies differences in warming and cooling pathways in sensitivity to adaptation.     

Energy model for contrast detection: spatiotemporal characteristics of threshold vision

A model for contrast detection of spatiotemporal stimuli is proposed which consists of a spatiotemporal linear filter, an energy device and a threshold device. Assuming the existence of independent intrinsic noise, the probability of stimulus detection was approximated by a Weibull function of the response energy. With this assumption, the stimulus energy is a constant at fixed detection probability. This energy model for contrast detection satisfactorily accounted for the elliptical threshold contours of line pairs at stimulus separations within the range 2–30 min and at stimulus onset asynchronies within the range 20–140 ms. The threshold contour at a large stimulus onset asynchrony (300 ms) was in the form of a rounded square. This finding was explained by assuming that the probability of seeing the line pair was determined by the joint probability that at least one stimulus had been detected. With the energy model, the temporal and spatial autocorrelation functions of the response to a flashed line were evaluated. The autocorrelation functions thus determined were used to predict the temporal contrast sensitivity function to a flickering line stimulus and the spatial contrast sensitivity function to flashed gratings, which were in agreement with the experimental data. The data obtained were fitted adequately by an impulse response approximated by a spatiotemporal Gabor-like function. Received: 08 December 1997 / Accepted in revised form: 26 January 1999     

The tactile perception of stimulus orientation

Studies of the visual system suggest that, at an early stage of form processing, a stimulus is represented as a set of contours and that a critical feature of these local contours is their orientation. Here, we characterize the ability of human observers to identify or discriminate the orientation of bars and edges presented to the distal fingerpad. The experiments were performed using a 400-probe stimulator that allowed us to flexibly deliver stimuli across a wide range of conditions. Orientation thresholds, approximately 20 degrees on average, varied only slightly across modes of stimulus presentation (scanned or indented), stimulus amplitudes, scanning speeds, and different stimulus types (bars or edges). The tactile orientation acuity was found to be poorer than its visual counterpart for stimuli of similar aspect ratio, contrast, and size. This result stands in contrast to the equivalent spatial acuity of the two systems (at the limit set by peripheral innervation density) and to the results of studies of tactile and visual letter recognition, which show that the two modalities yield comparable performance when stimuli are scaled appropriately.     

Auditory steady-state responses to multiple simultaneous stimuli

Steady-state responses can follow multiple simultaneous auditory stimuli. If the stimuli are modulated at different rates, responses specific to each stimulus can be assessed by measuring in the frequency domain response the spectral component corresponding to the rate of modulation. When each stimulus has a different carrier frequency or different ear of presentation, the responses when 8 stimuli are presented simultaneously are not significantly different than when each stimulus is presented alone. Since significant responses can be recognized down to intensities that average 14 dB above behavioral threshold, this technique may be useful in objective audiometry. It is also possible to record steady-state responses to multiple modulations of the same carrier frequency. In this case, the amplitude of the responses when the stimuli are combined is smaller than when the stimuli are presented alone. The decrease in amplitude depends upon the number of concomitant stimuli and their relative intensities. These effects are probably due to the compressive rectification occurring during cochlear transduction, and the data may be used to model cochlear processing of auditory stimuli.     

Effect of stimulus (postsynaptic current) shape on fibre excitation.

Effects of variation of the stimulus pulse shape on the excitation of a nonmyelinated nerve fibre were studied using a mathematical model based on the Hodgkin-Huxley equations. Efficiency of smoothly changing pulses was compared with that of rectangular pulses. For pulses shorter than the time to excitation, the rate of the stimulus rise did not determine the ability of a smoothly changing pulse to excite the fibre. For a given stimulus duration, the main factor was the pulse area or the charge delivered by the pulse. The strength-duration curve for smoothly changing pulses was a nonmonotonic function, in contrast to the curve for rectangular pulses. The dependence of latency on changes in the pulse area was non-linear. It would be nonmonotonic when the pulse area variation were due to the stimulus duration or the stimulus rise duration. More that one propagating intracellular action potential (IAP) could arise upon fibre activation by a long smoothly changing threshold stimulus. Upon activation of relatively short fibres the IAP could arise not at the site of the smoothly changing stimulus injection. The rectangular pulses of long duration were more efficient than the corresponding smoothly changing ones. Irrespective of the shape, the pulses whose duration at the foot is 1-2 ms, are more suitable for a prolonged threshold fibre activation.     

Time course and action spectrum of vibrotactile adaptation

In a series of experiments designed to explore the processes underlying adaptation of the sense of flutter-vibration, vibrotactile threshold was measured on the pad of the index finger, using Békésy tracking. Unadapted thresholds were first measured, for a number of frequencies (4-90 Hz) and contactor sizes (1-8 mm diameter). As expected, these measurements indicated the presence of (1) a Pacinian system possessing spatial summation and increasing in sensitivity, as frequency was raised, at the rate of 12 dB/octave; and (2) a non-Pacinian system showing little spatial summation, and with a frequency characteristic matching that of the NP I mechanism of Bolanowski et al. (1988). These baseline data of Experiment 1 guided the selection of stimulus parameters for subsequent experiments, in which threshold for a test stimulus was measured before, during, and after periods of vibrotactile adaptation. In Experiment 2, test stimuli of 10 Hz and 50 Hz were combined factorially with 30-dB SL adapting stimuli of the same two frequencies. When the test stimulus was 10 Hz, the two adapting frequencies were equally effective in raising threshold; however, when the 50-Hz test stimulus was used, the 50-Hz adapting stimulus raised threshold by a greater amount than did the 10-Hz adapter. These results confirm on the finger the independence of adaptation in Pacinian and non-Pacinian channels, a result previously established on the thenar by other workers. For all four frequency combinations, threshold rose exponentially with a time constant of 1.5-2 min. In Experiment 3, an action spectrum was determined, showing the adapting amplitude needed at each of a series of frequencies to raise the threshold of a 10-Hz stimulus by 10 dB; this spectrum was essentially flat from 30 to 90 Hz. The results, taken in conjunction with what is known about rapidly adapting cutaneous mechanoreceptors, imply that the effectiveness of an adapting stimulus is not determined solely by the amount of activity it generates in first-order afferents.     

Time Course and Action Spectrum of Vibrotactile Adaptation

In a series of experiments designed to explore the processes underlying adaptation of the sense of flutter-vibration, vibrotactile threshold was measured on the pad of the index finger, using Békésy tracking. Unadapted thresholds were first measured, for a number of frequencies (4-90 Hz) and contactor sizes (1-8 mm diameter). As expected, these measurements indicated the presence of (1) a Pacinian system possessing spatial summation and increasing in sensitivity, as frequency was raised, at the rate of 12 dB/octave; and (2) a non-Pacinian system showing little spatial summation, and with a frequency characteristic matching that of the NP I mechanism of Bolanowski et al. (1988). These baseline data of Experiment 1 guided the selection of stimulus parameters for subsequent experiments, in which threshold for a test stimulus was measured before, during, and after periods of vibrotactile adaptation.

In Experiment 2, test stimuli of 10 H_z and 50 H_z were combined factorially with 30-dB SL adapting stimuli of the same two frequencies. When the test stimulus was 10 Hz, the two adapting frequencies were equally effective in raising threshold; however, when the 50-Hz test stimulus was used, the 50-Hz adapting stimulus raised threshold by a greater amount than did the 10-Hz adapter. These results confirm on the finger the independence of adaptation in Pacinian and non-Pacinian channels, a result previously established on the thenar by other workers. For all four frequency combinations, threshold rose exponentially with a time constant of 1.5-2 min.

In Experiment 3, an action spectrum was determined, showing the adapting amplitude needed at each of a series of frequencies to raise the threshold of a 10-Hz stimulus by 10 dB; this spectrum was essentially flat from 30 to 90 Hz. The results, taken in conjunction with what is known about rapidly adapting cutaneous mechanoreceptors, imply that the effectiveness of an adapting stimulus is not determined solely by the amount of activity it generates in first-order afferents.     

How Long Depends on How Fast—Perceived Flicker Dilates Subjective Duration

How do humans perceive the passage of time and the duration of events without a dedicated sensory system for timing? Previous studies have demonstrated that when a stimulus changes over time, its duration is subjectively dilated, indicating that duration judgments are based on the number of changes within an interval. In this study, we tested predictions derived from three different accounts describing the relation between a changing stimulus and its subjective duration as either based on (1) the objective rate of changes of the stimulus, (2) the perceived saliency of the changes, or (3) the neural energy expended in processing the stimulus. We used visual stimuli flickering at different frequencies (4–166 Hz) to study how the number of changes affects subjective duration. To this end, we assessed the subjective duration of these stimuli and measured participants'' behavioral flicker fusion threshold (the highest frequency perceived as flicker), as well as their threshold for a frequency-specific neural response to the flicker using EEG. We found that only consciously perceived flicker dilated perceived duration, such that a 2 s long stimulus flickering at 4 Hz was perceived as lasting as long as a 2.7 s steady stimulus. This effect was most pronounced at the slowest flicker frequencies, at which participants reported the most consistent flicker perception. Flicker frequencies higher than the flicker fusion threshold did not affect perceived duration at all, even if they evoked a significant frequency-specific neural response. In sum, our findings indicate that time perception in the peri-second range is driven by the subjective saliency of the stimulus'' temporal features rather than the objective rate of stimulus changes or the neural response to the changes.     

Contrast sensitivity and appearance in briefly presented illusory figures.

We examined the contributions of brightness enhancement, illusory figure formation and figural completion to changes in contrast sensitivity in contour gaps. The brightness on the border of a Kanizsa-square and an outline square was measured as the point of subjective equality with the background (PSE) for small line targets. Increment and decrement thresholds were measured at the same location. We found that contrast thresholds were lower than in a control condition without inducers, and that the threshold reduction was independent of the contrast polarity of the inducers. This reduction cannot be explained by a simple summation of stimulus contrast and induced brightness. In a second experiment the inducers that define the contour of the Kanizsa and the outline square were changed so that the figure was no longer closed, keeping the local stimulus surround constant. Thresholds were equally reduced for all conditions, independently of whether the figure was completed or not, or whether an illusory contour was perceived or not. The results suggest that the reduction of contrast threshold in contour gaps is independent of the brightness perceived in these gaps and of the formation of an illusory figure. Processes that cause contrast threshold reduction in contour gaps also seem to operate independently of figural completion.     

Adaptation to warming but not cooling at slow rates of stimulus change in thermal threshold measurements

The relationship between thermal detection threshold and rate of temperature change of the thermal stimulus when slow (<1°C?s^?1) rates of change are employed was investigated. Using both the reaction time (RT) inclusive Method of Limits and RT exclusive Method of Levels healthy volunteers had warming (WDT) and cooling detection thresholds (CDT) measured at four different rates of temperature change (0.3, 0.5, 0.7 and 1.0°C?s^?1) from the thenar and/or mental regions using a contact thermode. With the Method of Limits, CDT increased linearly with rate of temperature change suggesting increments were due to RT artefacts. This was further supported by threshold assessment with the Method of Levels which showed CDT were unaffected by the rate of change in the RT exclusive method (P?>?0.1). In contrast, WDT did not increase linearly with rate of stimulus temperature change when the Method of Limits was used and threshold assessment with the Method of Levels showed WDT assessed using a 0.3°C?s^?1 ramp rate were significantly higher than those measured with a 1°C?s^?1 rate of change (P?<?0.05). This study indicates that adaptation to a warming stimulus can occur at faster rates of stimulus change than previously anticipated and identifies differences in warming and cooling pathways in sensitivity to adaptation.     

Visual discrimination of objects differing in spatial depth by goldfish

Training experiments were performed to investigate the ability of goldfish to discriminate objects differing in spatial depth. Tests on size constancy should give insight into the mechanisms of distance estimation. Goldfish were successfully trained to discriminate between two black disk stimuli of equal size but different distance from the tank wall. Each stimulus was presented in a white tube so that the fish could see only one stimulus at a time. For each of eight training stimulus distances, the just noticeable difference in distance was determined at a threshold criterion of 70% choice frequency. The ratio of the retinal image sizes between training stimulus and comparison stimulus at threshold was about constant. However, in contrast to Douglas et al. (Behav Brain Res 30:37–42, 1988), goldfish did not show size constancy in tests with stimuli of the same visual angle. This indicates that they did not estimate distance, but simply compared the retinal images under our experimental conditions. We did not find any indication for the use of accommodation as a depth cue. A patterned background at the rear end of the tubes did not have any effect, which, however, does not exclude the possibility that motion parallax is used as a depth cue under natural conditions.     

Behavioral and neurophysiological assessment of lateral line sensitivity in the mottled sculpin,Cottus bairdi

1. The unconditioned feeding response of the mottled sculpin, Cottus bairdi, was used to measure threshold sensitivity of the lateral line system to a vibrating sphere as a function of stimulus position (i.e., sphere near head, trunk or tail) and vibration frequency. In addition, extracellular recording techniques were used to measure threshold sensitivity curves for posterior lateral line nerve fibers for the same stimulus positions used for measuring trunk sensitivity in behavioral measurements. 2. For all stimulus positions, behaviorally-measured threshold sensitivity was relatively independent of vibration frequency from 10 to 100 Hz when defined in terms of water acceleration, rather than velocity or displacement. Best thresholds for stimuli placed 15 mm away from the head were around -75 dB re: 1m/s(2), approximately 20 dB less than that for stimuli placed at the same distance near the tail. Trunk sensitivity was intermediate. 3. Physiologically-measured threshold sensitivity, in terms of acceleration, was also relatively independent of of frequency from 10 to 100 Hz in most fibers. A smaller number of fibers showed a decline in acceleration sensitivity after 10-30 Hz, with the rate of decline being equivalent to equal velocity sensitivity. Best sensitivity of all fibers fell between -40 and -70 dB re: 1m/s (2). 4. These results indicate that (a) behavioral thresholds are based on acceleration-sensitive endorgans--most likely lateral line canal (rather than superficial) neuromasts, (b) behavioral performance can be accounted for on the basis of information from a single population of fibers, and (c) sensitivity varies along the fish's body in a manner that corresponds to the size and distribution of neuromasts.     

Receptive field types in the lateral geniculate body and their frequency characteristics]

Time amplitude -- frequency characteristics of the I and II types of receptive fields (RF) of lateral geniculate and their dependence on the contrast and spatial parameters of the light stimulus were studied. It is shown that the frequency characteristics of the RF I type depends on the contrast and area of the light stimulus, the higher being the contrast at a small area the smaller are the low frequencies. However at a large area of the stimulus the inhibition of low frequencies is greater at a small contrast. The transmitting band of frequency characteristics of RF II type does not depend on the contrast at a small area of the stimulus, at a large area a fall of low frequencies takes place at high contrasts of the stimulus. Such different behaviour of the receptive fields is explained by the models, which take into account RF spatial characteristics.     

Perceived contrast following adaptation: the role of adapting stimulus visibility

The issue of whether contrast adaptation can reduce the perceived contrast of gratings oriented orthogonal to the adapting stimulus to a greater extent than parallel gratings has been the subject of considerable debate (Snowden and Hammett, 1992; Ross and Speed, 1996). We compared the reductions in perceived contrast of various test gratings oriented parallel and orthogonal to the adapting stimulus across a range of spatial frequencies (2.25-9 c/deg) and adaptation contrasts (0.19-1.0). Our results show that when the adapting stimulus is low in contrast, parallel adaptation effects are always greater than the effects of orthogonal adaptation. When the adapting contrast is increased, however, the difference between parallel and orthogonal effects is reduced. Further increases in adapting contrast can produce a situation where cross-orientation adaptation effects exceed iso-orientation effects. This was observed at low spatial frequencies (2.25 and 4.5 c/deg) only. The difference in the pattern of results obtained at low and high spatial frequencies can be explained in terms of the adapting stimulus visibility. We conclude that cross-orientation adaptation effects can be greater than iso-orientation effects, but only when the adapting stimulus is highly suprathreshold.     

正常和卡那霉素中毒豚鼠听觉脑干诱发反应及频率跟随反应的比较研究

通过测定阈值和频谱分析,比较了正常和卡那霉素中毒后豚鼠听觉脑干反应(ABR)及频率跟随反应(FFR)的特性.正常FFR的非线性随刺激频率的减低而增加,表现为高次谐波的出现.从频谱中基频成分的峰很容易识别FFR阈值.卡那霉素作用后,高频FFR首先受到影响,波幅下降,阈值显著升高.低频FFR受到影响较小.同一动物ABR阈值尚未变化时高频FFR的基频峰值已下降,乃至消失.实验结果表明FFR对耳毒性药物中毒反应由于其包含有频率特性比ABR更灵敏,更便于定量分析.     

Behavioural examination of the infrared sensitivity of ball pythons

The ability to detect infrared (IR) radiation is a characteristic trait in boids and pitvipers. These snakes possess highly sensitive IR receptors, which enable them to perceive IR sources and assess their direction and distance independently of visual cues. Electrophysiological studies have been conducted to determine IR detection thresholds in boids and pitvipers. This, however, is the first behavioural study that focuses on the detection threshold of a boid snake to IR stimuli. Blindfolded ball pythons Python regius were exposed to a moving IR stimulus of constant size and temperature at various distances (10–100 cm). Distinct behavioural changes during stimulus presentation (S-form posture, freeze and fix, follow and fix) allowed quantification of the behavioural responses. The threshold to elicit behavioural responses was used to assess the IR detection threshold. The results revealed that P. regius can detect a moving IR stimulus resembling a mouse in temperature and size up to a distance of 30 cm, which corresponds to an irradiance contrast of 38.83 × 10⁻⁶ W cm⁻². This irradiance contrast detection threshold value is about one-third lower (reveals a 1.5 times higher sensitivity) than the results from earlier electrophysiological studies.     

Sweetness intensity enhancement by pulsatile stimulation: effects of magnitude and quality of taste contrast

Upon stimulation with continuously alternating (pulsatile) taste concentrations, humans report higher average taste intensities than for continuous stimulation with the same average tastant concentration. We investigated the effect of the magnitude of concentration changes (concentration contrast) and the effect of taste quality changes (quality contrast) between alternating tastants on sweet taste enhancement. The perceived sweetness intensity increased with the magnitude of the sucrose concentration contrast: The pulsatile stimulus with the highest concentration difference (average sucrose concentration: 60 g/L) was rated as the sweetest in spite of the fact that the gross sucrose concentrations were identical over stimuli. Moreover, this stimulus was rated equally sweet as a continuous reference of 70 g/L sucrose. On alternation of sucrose with the qualitatively different citric acid, sweet taste enhancement remained at the level observed for alternation with water at citric acid concentration levels up to 3 times its detection threshold. Alternation of a sucrose solution with a citric acid solution at 9 times its threshold concentration, resulted in an attenuation of the pulsation-induced enhancement effect. Upon alternation of citric acid pulses at concentrations around the threshold with water intervals only, no taste enhancement was observed compared with continuous citric acid stimuli of the same net concentration. We propose that the magnitude of pulsation-induced taste enhancement is determined by the absolute rather than relative change of tastant concentration. This explains why 1) pulsation-induced sweet taste enhancement is determined by the magnitude of the sucrose pulse-interval contrast and 2) the alteration of citric acid with water does not enhance taste intensity at detection threshold level.     

Modulation rate transfer functions in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>) with normal hearing and high-frequency hearing loss

Envelope following responses were measured in two bottlenose dolphins in response to sinusoidal amplitude modulated tones with carrier frequencies from 20 to 60 kHz and modulation rates from 100 to 5,000 Hz. One subject had elevated hearing thresholds at higher frequencies, with threshold differences between subjects varying from ±4 dB at 20 and 30 kHz to +40 dB at 50 and 60 kHz. At each carrier frequency, evoked response amplitudes and phase angles were plotted with respect to modulation frequency to construct modulation rate transfer functions. Results showed that both subjects could follow the stimulus envelope components up to at least 2,000 Hz, regardless of carrier frequency. There were no substantial differences in modulation rate transfer functions for the two subjects suggesting that reductions in hearing sensitivity did not result in reduced temporal processing ability. In contrast to earlier studies, phase data showed group delays of approximately 3.5 ms across the tested frequency range, implying generation site(s) within the brainstem rather than the periphery at modulation rates from 100 to 1,600 Hz. This discrepancy is believed to be the result of undersampling of the modulation rate during previous phase measurements.     

Electrophysiological and psychophysical quantification of temporal summation in the human nociceptive system

Animal experiments have shown that the nociceptive reflex can be used as an indicator of central temporal integration in the nociceptive system. The aim of the present study on humans was to investigate whether the nociceptive reflex, evoked by repetitive strong electrical sural nerve stimuli, increased when summation was reported by the volunteers. The reflexes were recorded from the biceps femoris and rectus femoris muscles in eight volunteers following a series of stimulations at 0.1, 1, 2, and 3 Hz. Each series consisted of five consecutive stimuli. Using 0.1- and 1-Hz stimulation, the reflex was not facilitated in the course of the five consecutive stimuli. Following 2- and 3-Hz stimulation, the reflex size (root mean square amplitude) increased significantly during the course of the fifth stimulus. This reflex facilitation was followed by a significant increase (summation) in the pain magnitude when compared with 1- and 0.1-Hz stimulation. Furthermore, the threshold for psychophysical summation could be determined. This threshold (stimulus intensity) decreased when the stimulus frequency (1–5 Hz) of the five consecutive stimuli was increased. The nociceptive reflex and the psychophysical summation threshold might be used to clarify and quantify aspects of temporal summation within the human nociceptive system.