Sensory thresholds obtained from MEG data: cortical psychometric functions.

Sensory sensitivity is typically measured using behavioural techniques (psychophysics), which rely on observers responding to very large numbers of stimulus presentations. Psychophysics can be problematic when working with special populations, such as children or clinical patients who may lack the compliance or cognitive skills to perform the behavioural tasks. We used an auditory gap-detection paradigm to develop an accurate measure of sensory threshold derived from passively-recorded magnetoencephalographic (MEG) data. Auditory evoked responses were elicited by silent gaps of varying durations in an on-going noise stimulus. Source modelling was used to spatially filter the MEG data and sigmoidal 'cortical psychometric functions' relating response amplitude to gap duration were obtained for each individual participant. Fitting the functions with a curve and estimating the gap duration at which the amplitude of the evoked response exceeded one standard deviation of the prestimulus brain activity provided an excellent prediction of psychophysical threshold. Accurate sensory thresholds can therefore be reliably extracted from MEG data recorded while participants listen passively to a stimulus. Because our paradigm required no behavioural task, the method is suitable for studies of populations where variations in cognitive skills or vigilance make traditional psychophysics unsuitable.

Masking effect produced by Mach bands on the detection of narrow bars of random polarity.

Difficulties arise in measuring masking by Mach bands because very-low-contrast signals distort the bands. [J. Opt. Soc. Am. A 17, 1147 (2000).] Adding narrow luminance increments (bright bars) in the dark Mach band widens the dark band; adding decrements (dark bars) narrows the dark band, and conversely in the bright bands. Randomizing signal polarity prevents observers from using the distortion of the Mach bands as a cue to the presence of the signal. We measured (two-alternative-forced-choice) Mach bands' masking of randomly selected bright (incremental) or dark (decremental) bars. Detection was worse in both dark and bright Mach bands than on the neighboring plateaus. Separate analysis of trials containing only one polarity signal revealed 9-cycle/deg oscillations in performance as a function of location. Oscillations in the two polarities were approximately 180 degrees out of phase.

Contrast discrimination with pulse trains in pink noise.

Detection performance was measured with sinusoidal and pulse-train gratings. Although the 2.09-cycles-per-degree pulse-train, or line, grating contained at least eight harmonics all at equal contrast, it was no more detectable than its most detectable component. The addition of broadband pink noise designed to equalize the detectability of the components of the pulse train made the pulse train approximately a factor of 4 more detectable than any of its components. However, in contrast-discrimination experiments, with a pedestal or masking grating of the same form and phase as the signal and with 15% contrast, the noise did not affect the discrimination performance of the pulse train relative to that obtained with its sinusoidal components. We discuss the implications of these observations for models of early vision, in particular the implications for possible sources of internal noise.

Contrast discrimination with sinusoidal gratings of different spatial frequency.

The detectability of contrast increments was measured as a function of the contrast of a masking or "pedestal" grating at a number of different spatial frequencies ranging from 2 to 16 cycles per degree of visual angle. The pedestal grating always had the same orientation, spatial frequency, and phase as the signal. The shape of the contrast-increment threshold versus pedestal contrast (TvC) functions depends on the performance level used to define the "threshold," but when both axes are normalized by the contrast corresponding to 75% correct detection at each frequency, the TvC functions at a given performance level are identical. Confidence intervals on the slope of the rising part of the TvC functions are so wide that it is not possible with our data to reject Weber's law.

Monaural and binaural detection of sinusoidal phase modulation of a 500-Hz tone.

The detectability of phase modulation was measured for three subjects in two-alternative temporal forced-choice experiments. In experiment 1, the detectability of sinusoidal phase modulation in a 1500-ms burst of an 80-dB (SPL), 500-Hz sinusoidal carrier presented to the left ear (monaural condition) was measured. The experiment was repeated with an 80-dB, 500-Hz static (unmodulated) tone at the right ear (dichotic condition). At a modulation rate of 1 Hz, subjects were an order of magnitude more sensitive to phase modulation in the dichotic condition than in the monaural condition. The dichotic advantage decreased monotonically with increasing modulation rate. Subjects ceased to detect movement in the dichotic stimulus above 10 Hz, but a dichotic advantage remained up to a modulation rate of 40 Hz. Thus, although sound movement detection is sluggish, detection of internal phase modulation is not. In experiment 2, thresholds for detecting 2-Hz phase modulation were measured in the dichotic condition as a function of the level of the pure tone in the right ear. The dichotic advantage persisted even when the level of the pure tone was reduced by 50 dB or more. The findings demonstrate a large dichotic advantage which persists to high modulation rates and which depends very little on interaural level differences.

Detection of incremental and decremental bars at different locations across Mach bands and related stimuli.

Two-alternative forced-choice procedures were used to measure the detectability of bright and dark bars at various locations across luminance patterns that produced Mach bands. Detection performance was significantly affected by both dark and bright Mach bands: poor detection performance was observed at locations near, but not in, the Mach bands; relatively good detection performance at locations within the Mach bands was caused by reliable changes in the width, depth, or symmetry of the bands produced by the signal bars. The changes were apparent with signals of lower luminance than that needed for detection in the plateau regions far from the bands, but, because the cues were not sufficiently reliable to allow errorless performance, unusually shaped psychometric functions were obtained.

Muscle performance in an urban population sample of 40- to 79-year-old men and women.

An urban population sample of 40 to 79-year-old men and women was investigated to evaluate the influence of age and activity level on muscle strength and endurance and to establish a reference material. During the investigation 144 persons were tested bilaterally, except for ankle strength, when only the right side was examined. Isometric muscle strength was determined in the knee extensors and flexors. Isokinetic (at 60 degrees/s and at 180 degrees/s) muscle strength was determined concentrically and eccentrically in the knee extensors and flexors. The dynamic and static endurance of the extensors was measured. Isometric strength was determined in the ankle plantar and dorsiflexor muscles. Isokinetic ankle plantar flexion strength was determined concentrically at 60 degrees/s with and without prior eccentric muscle contraction. Hand-grip strength was evaluated with a dynamometer. Walking velocity and the number of heel-rises were recorded. Physical activity level was assessed by questionnaire. Muscle biopsies were taken from the vastus lateralis muscle for histochemical and enzymatic analyses. Walking and the different muscle tests declined with age, and with a slight gender difference. Muscle biopsies showed a trend toward smaller muscle fibers with age. The results of our study can be used as reference material for clinical studies in different age groups.

Forward and backward masking with brief impulsive stimuli.

The forward and backward masking effect of a 20-micros pulse was measured for delays ranging from 0 to -/+4 ms. Masking is not a monotonic function of delay in either forward or backward masking. For two of the three observers, the asymmetry in which forward masking exceeds that of backward masking is small for delays less than 500 micros. The implications of the data for the contribution of masking to the precedence effect are considered.

Some aspects of the lateralization of echoed sound in man. II. The role of the stimulus spectrum.

The lateralization of clicks and their "echoes" was investigated with a view to determining the role of spectral characteristics in lateralization. Lateralization-discrimination performance was measured in a number of two-interval, two-alternative forced-choice experiments using three pairs of binaural clicks designed to elucidate how spectral cues are used in lateralization. The stimulus in one observation interval comprised a diotic click followed, after the interclick interval (ICI), by a dichotic click with either (1) an interaural time delay or (2) an interaural amplitude difference. The dichotic click was in turn followed, after an ICI of the same size, by another diotic click. In the second observation interval, the signals to the two ears were interchanged. The stimulus has the property that the signals delivered to the two ears had either (1) identical energy-density spectra but nonzero interaural-phase differences (IPDs) or (2) zero IPDs but nonidentical energy-density spectra. Under certain circumstances, observers perceived these stimuli as arising from the side of the head opposite that which would be predicted from the direction of the interaural cue in the temporal waveform. Joint consideration of the psychophysical data and the spectral characteristics of the stimuli strongly suggest a spectral "dominance region" for lateralization near 750 Hz, observers' lateralization performance was determined predominantly by the IPD cues from this region. In general, the results demonstrate that echoes of transients that arrive within about 2-3 ms of an initial transient are not suppressed, but have a substantial effect on lateralization through their contribution to the resultant spectral characteristics. The results contradict models that represent the precedence effect as the temporary suppression or inhibition of directional information in echoes over 2-3 ms after an initial transient.

Some aspects of the lateralization of echoed sound in man. I. The classical interaural-delay based precedence effect.

The precedence effect in two-click stimuli was investigated by measuring observers' sensitivity to interaural time delays (ITDs) as a function of interclick interval (ICI). A two-interval two-alternative forced-choice discrimination paradigm was used in two stimulus configurations: type I, a dichotic click with a given ITD preceded a diotic click; and type II, a dichotic click followed a diotic click. Threshold ITDs were measured in each configuration for a finely sampled distribution of ICIs that ranged from 0.1 to 25.6 ms. Performance was characterized by the "threshold elevation factor" (TEF) which normalized each of the observers' type I and type II ITD thresholds relative to their ITD threshold for a single dichotic click. The finer sampling of ICIs revealed two novel results: First, for two observers, sensitivity to ITD in the later arriving ITD (type II) oscillated in a consistent and systematic way with changes in ICI. Second, when the ICI reached 12.8 ms, ITD thresholds in the type I and type II configurations were equal but nearly a factor of 2 greater than for a single dichotic click. Some aspects of the data are consistent with the phenomenon of binaural adaptation.

No role for motion blur in either motion detection or motion-based image segmentation.

The human contrast sensitivity function is bandpass in form for stimuli of low temporal frequency but low pass for flickering or moving stimuli. Because the loss in sensitivity to moving stimuli is large, images moving on the retina have little perceptible high-spatial-frequency content. The loss of high-spatial-frequency content--often referred to as motion blur--provides a potential cue to motion. The amount of motion blur is a function of stimulus velocity but is significant at velocities encountered by the visual system in everyday situations. Our experiments determined the influence of high-spatial-frequency losses induced by motion of this order on motion detection and on motion-based image segmentation. Motion detection and motion-based segmentation tasks were performed with either spectrally low-pass or spectrally broadband stimuli. Performance on these tasks was compared with a condition having no motion but in which form differences mimicked the perceptual loss of high spatial frequencies produced by motion. This allowed the relative salience of motion and motion-induced blur to be determined. Neither image segmentation nor motion detection was sensitive to the high-spatial-frequency content of the stimuli. Thus the change in perceptual form produced in moving stimuli is not normally used as a cue either for motion detection or for motion-based image segmentation in ordinary situations.

Evidence for a sound movement area in the human cerebral cortex.

Human listeners can localize sounds by the difference in both arrival time (phase) and loudness between the two ears. Movement of the sound source modulates these cues, and responses to moving sounds have been detected in animals in primary auditory cortex and in humans in other cortical areas. Here we show that detection of changes in the interaural phase or amplitude difference occurs through a mechanism distinct from that used to detect changes in one ear alone. Moreover, a patient with a right hemisphere stroke is unable to detect sound movement, regardless of whether it is defined by phase or by loudness cues. We propose that this deficit reflects damage to a distinct cortical area, outside the classical auditory areas, that is specialized for the detection of sound motion. The deficit is analagous to cerebral akinotopsia (motion blindness) in the visual system, and so the auditory system may, like the visual system, show localization of specialized functions to different cortical regions.

Muscle fiber characteristics, capillaries and enzymes in patients with fibromyalgia and controls.

In 11 female patients with Fibromyalgia Syndrome (FS), biopsies from the m. vastus lateralis were analyzed, in order to reveal any possible changes which might explain muscular weakness and fatigue. Nineteen healthy subjects served as a control group. Light microscopy did not show any gross histopathological findings. Fiber composition and fiber areas did not differ between the two groups, except for a greater coefficient of variation of the area of type II A fibers and of the mean fiber area in the FS group. The number of capillaries per square millimeter and also the fiber area in relation to the capillaries, was lower in the FS patients. Analyses of enzymes showed decreased levels of 3-hydroxy-CoA-dehydrogenase and citrate synthase in the patient group. The reduced oxidative enzyme levels and capillarization indicate reduced physical activity, although this does not associate with muscle fiber hypotrophy.

Changes in muscle morphology, strength and enzymes in a 4-5-year follow-up of subjects with poliomyelitis sequelae.

Twenty subjects with polio sequelae were studied on two occasions 4-5 years apart by means of dynamometer measurements of knee-extension and flexion strength and muscle biopsy for histochemical and enzymatic analyses. The subjects were divided into those who reported (unstable, n = 12) and did not report (stable, n = 8) new or increased muscle weakness in the tested leg between the two examinations. Muscle strength decreased significantly in the unstable subjects, but only for knee-flexion in the stable subjects. However, the endurance test comprising 50 consecutive knee-extensions at 180 degrees/sec showed increased fatigability at the second examination only in the stable subjects. Most subjects had markedly increased muscle fiber areas, which in some subjects increased further, but in those already with very extreme hypertrophy the fiber size decreased. Capillarization and activity of citrate-synthase were decreased at the initial examination, but no significant further reduction was observed at the second examination. The results demonstrate individual patterns in the compensatory process for the presumed loss of motor units.

Speed, spatial-frequency, and temporal-frequency comparisons in luminance and colour gratings.

The perceived speed, temporal frequency, and spatial frequency of matched colour and luminance gratings were compared in separate experiments. The large factor by which colour gratings are perceived to be slower moving than matched luminance gratings cannot be explained by systematic differences in the perceived spatial frequency or in the perceived temporal frequency of the two types of grating.

Implications of motion detection for early non-linearities.

Analysis of motion may be accomplished using the spatiotemporal variations produced when a spatially varying luminance waveform moves across linear receptive fields. Moving contrast-modulated patterns which consist of coarse-scale spatial variations in the contrast of fine-scale luminance patterns cannot be analysed in this way. The human visual system can analyse the motion of contrast-modulated patterns and this suggests it may contain mechanisms that use non-linear transformations. Non-linear transformation of contrast-modulated patterns would give rise to a component (a distortion product) that varies on the same spatial scale as the contrast variation; this can be analysed to extract motion. Is the non-linearity simply an inherent part of the transduction process or is it a characteristic of a mechanism specialized for the analysis of the motion of such patterns? Comparisons of the spatial and temporal limitations of motion discrimination using luminance and contrast-modulated patterns suggest that the mechanisms which analyse the two types of patterns are different, although recent physiological evidence suggests that they may have common elements.

Discriminating the direction of second-order motion at short stimulus durations.

We measured the ability of human observers to discriminate the direction of motion of different spatial patterns presented for durations ranging from 0.021 to 0.67 sec. The patterns were: (1) a vertical grating (spatial frequency 0.93 c/deg at 5% contrast); (2) a "beat" pattern made by adding vertical gratings of 6.3 and 5.4 c/deg both at 5% contrast moving in opposite directions (this pattern appears as a horizontally moving, 0.93 c/deg "beat"; i.e. spatial variation in the contrast of a stationary vertical grating of 5.8 c/deg); and (3) a "plaid" pattern made by adding gratings of 5.9 c/deg orientated +/- 81 deg from vertical (this pattern can also be expressed as a horizontally moving 1.9 c/deg beat in a horizontal grating of 5.8 c/deg). The direction of motion of the grating and the plaid pattern were discriminable at all durations tested. The direction of motion of the beat could only be discriminated at durations above approx. 200 msec. We suggest that this is a consequence of the fact that the moving beat is only visible to second-order mechanisms, and that second-order mechanisms for the analysis of motion operate more slowly than first-order mechanisms.

Errors in direction-of-motion discrimination with dichoptically viewed stimuli.

At durations shorter than about 150 msec, a complex grating comprising a static 1-c/deg grating and a moving 3-c/deg grating is perceived as moving in the direction opposite that of the physical direction of motion. Here the phenomenon is further examined by measuring the perceived direction of motion of the fused images of a 1-c/deg grating presented to one eye and a moving 3-c/deg grating presented to the other. The strength of the illusion is almost unaffected by dichoptic presentation. This observation is consistent with the hypothesis that perceived motion is a consequence of the way the visual system integrates signals arising from different detectors tuned to the two component gratings.

Detecting and discriminating the direction of motion of luminance and colour gratings.

Human observers were required to discriminate the direction of motion of vertically moving, 1 c/deg luminance and colour gratings. The gratings had different contrasts and moved at temporal frequencies between 0.5 and 32 Hz. Sensitivity [the reciprocal of the contrast at which performance reached 75% correct in a temporal two-alternative forced-choice (2 AFC) discrimination task] was a band-pass function of temporal frequency for luminance gratings, and a low-pass function of temporal frequency for colour gratings. Further, when colour contrast was expressed in terms of the modulation in cone excitation produced by the stimulus, sensitivity to colour gratings was greater than sensitivity to luminance gratings at frequencies below 2 Hz. On the other hand, at temporal frequencies above 4 Hz, sensitivity to colour gratings was comparable with sensitivity to luminance gratings of double the temporal frequency. Detection sensitivity was measured for luminance and colour gratings of 1, 4 and 16 Hz. With either colour or luminance gratings, detection thresholds were very similar to those for direction-of-motion discrimination. This result confirms findings of Mullen and Boulton [(1992) Vision Research, 32, 483-488] and Cavanagh and Anstis [(1991) Vision Research, 31, 2109-2148], but is different from that reported by Lindsey and Teller [(1990) Vision Research, 30, 1751-1761] who used a smaller stimulus seen in a parafoveal region and found that motion discrimination thresholds were higher than detection threshold for colour gratings. We repeated our threshold measurements using parafoveal viewing conditions similar to those used by Lindsey and Teller (1990). We found that, although for luminance gratings detection thresholds were very close to direction-discrimination thresholds, for colour gratings, they were lower. The result is in qualitative agreement with Lindsey and Teller (1990). Our results suggest that low-level, or "first-order" motion mechanisms are not as sensitive to chromatic gratings as are colour-detection mechanisms.