A study of the Sherrington effect

In his studies of binocular fusion of flicker, Sherrington found a very small difference between CFFs when the eyes were synchronously stimulated and when they were alternately stimulated by two binocularly fused test patches. He concluded that the phenomena of alliance and antagonism which he had demonstrated in the motor system did not obtain in the sensory (visual) system. Later investigators found that this “Sherrington effect” was slightly larger than Sherrington reported, and it has been held as evidence of a definite, though perplexingly small, degree of binocular interaction. In the experiment reported here, the Sherrington effect was found to be about five times greater than it has previously been reported to be.     

The fusion frequency of flicker in the central and peripheral field with photopic levels of surround luminance

Flicker-fusion frequencies have been measured against a photopic level of surround luminance in the four quadrants of the visual field and the critical fusion frequences (c.f.f.s.) taken in direct vision compared with those determined at 15° and 35° in the periphery. The central c.f.f.s. of 47 subjects exceeded the peripheral values only in just over half of all the quadrants tested, although smaller differential thresholds were given almost without exception in the centre than in the periphery by the same observers and under similar conditions on a test of brightness discrimination. Likewise higher central than peripheral c.f.f.s. were recorded in the great majority of all quadrants when determinations were made against a dark surround. The contrasting types of flicker response observed against the photopic ground were shown to be relatively reproducible, and the same pattern was as a rule found in at least three quadrants of any one visual field. It was ascertained that the different patterns might result from a reciprocal shift of fusion frequencies either in central or peripheral vision.

Interpretations of the higher peripheral c.f.f.s which adduce the effect of accentuated pupillary constriction with central stimulation mere contra-indicated since such patterns of flicker perception survive after the photopupillary reflex has been abolished. Some evidence was, however, forthcoming for the view that peripheral fusion frequencies tend to exceed central values when the test object is larger than about 1·5° in angular size and more intense than the surround. It was suggested that the contrasting types of flicker response might be related to individual differences in the effect that stimulus size has on the flicker pattern.     

Varieties of Binocular Interaction in Human Vision

Binocular processing was investigated using a quantitative, process-oriented metatheory of response times. The analyses are not confined to particular distributional assumptions or specific models. Upper and lower performance boundaries for probability summation in parallel processing are defined and compared with observed distributions of reaction times using a variety of dichoptic stimuli. Performance that exceeds the upper bound strongly suggests faciliatory convergence between the two eyes (binocular channel summation). Performance below the lower bound suggests that inputs to the two eyes are processed serially. The results indicate that binocular channel summation in subjects with normal stereo vision requires targets of the same luminance polarity (paired increments or decrements) in corresponding retinal locations. When corresponding retinal locations are stimulated with opposing luminance polarities (increment to one eye, decrement to the other), performance is consistent with probability summation, indicating that parallel ON and OFF pathways remain segregated at least to the level of binocular fusion. Further analyses of data from a stereo-blind observer suggests serial processing of binocular inputs.     

On utrocular discrimination

Observers with good stereoacuity judged which eye received sine-wave grating patterns in a two-category forced-choice procedure. Large individual differences were found, but for most observers reliable discrimination was achieved at low spatial frequencies. No observer could perform the task above chance levels at high spatial frequencies. Discrimination was unaffected by retinal location, grating orientation, grating contrast, stimulus duration, or practice with feedback. Among observers who could perform the task, the following results were obtained: (1) Introduction of high spatial frequency components did not interfere with performance so long as a low spatial frequency component was present. (2) When gratings of low equal spatial frequency were presented to both eyes simultaneously at different contrast levels, observers could identify which eye received the higher contrast. (3) At low spatial frequencies, observers could distinguish monocular from binocular presentation. (4) Temporal frequency variations (counterphase flicker) influenced performance for some observers. Binocular summation and interocular transfer were unaffected by the spatial frequency variations which modulate utrocular discrimination. A new procedure for measuring stereopsis was developed which made possible comparison of utrocular discrimination with stereopsis at specific spatial frequencies. Stereopsis appeared mildly affected by spatial frequency.     

A simple device for producing synchronous and asynchronous binocular flicker

Many studies on binocular integration have compared in-phase and out-of-phase flicker stimulation. A new method of producing such stimulation is described. It differs from previous designs in that a combination of polarizers is used rather than mechanical light choppers or electronic flashers. The arrangement produces sinusoidal luminance pulses that can be presented with any degree of asynchrony between the two eyes.     

Transient increase of intact visual field size by high-frequency narrow-band stimulation

Three patients with visual field defects were stimulated with a square matrix pattern, either static, or flickering at frequencies that had been found to either promote or not promote blindsight performance. Comparison between pre- and post-stimulation perimetric maps revealed an increase in the size of the intact visual field but only for flicker frequencies previously found to promote blindsight. These changes were temporary but dramatic – in two instances the intact field was increased by an area of ∼30 deg² of visual angle. These results indicate that not only does specific high-frequency stimulus flicker promote blindsight, but that intact visual field size may be increased by stimulation at the same frequencies. Our findings inform speculation on both the brain mechanisms and the potency of temporal modulation for altering the functional visual field.     

Conscious perception of flickering stimuli in binocular rivalry and continuous flash suppression is not affected by tACS-induced SSR modulation

The content of conscious perception is known to correlate with steady-state responses (SSRs), yet their causal relationship remains unclear. Can we manipulate conscious perception by directly interfering with SSRs through transcranial alternating current stimulation (tACS)? Here, we directly addressed this question in three experiments involving binocular rivalry and continuous flash suppression (CFS). Specifically, while participants (N = 24) viewed either binocular rivalry or tried to detect stimuli masked by CFS, we applied sham or real tACS across parieto-occipital cortex at either the same or a different frequency and phase as an SSR eliciting flicker stimulus. We found that tACS did not differentially affect conscious perception in the forms of predominance, CFS detection accuracy, reaction time, or metacognitive sensitivity, confirmed by Bayesian statistics. We conclude that tACS application at frequencies of stimulus-induced SSRs does not have perceptual effects and that SSRs may be epiphenomenal to conscious perception.     

Frequency of fusion and of loss of fusion, and binocular depth perception with alternating stimulus presentation

Stereoscopic vision was investigated with an experimental design allowing dichoptic stimulus presentation at different frequencies of image alternation. For twenty subjects the frequency of binocular fusion and the frequency of loss of fusion to one stereoscopic image was measured as a function of the convergence angle. In thirteen subjects no dependence of the fusion frequency was found, while seven subjects showed a marked increase of the fusion frequency with increasing angle of convergence. In all cases the frequency of fusion was higher than the frequency of loss of fusion. Both frequencies, however, are lower than the flicker fusion frequency. Under conditions where no monocular cues and no references for stereoptic depth comparisons were presented, the apparent distance of the image from the observer could not be assessed, but perception of relative motion in depth was possible. All subjects assessed the direction of motion accurately down to changes of the convergence angle of 0.2 deg s-1.     

Binocular interactions in suprathreshold contrast perception

Suprathreshold binocular contrast interactions were studied psychophysically. A split-screen CRT display was used to present separate sine-wave gratings to the observer’s left and right eyes. The method of constant stimuli and a modified method of adjustment were used to find sets of binoculartest patterns that matched a given binocularstandard. Test patterns consisted of the simultaneous presentation of sine-wave gratings that differed in contrast to the left and right eyes. Standard patterns consisted of identical sine-wave gratings presented to the two eyes, and had the same spatial frequency as the test patterns. Binocular contrast matching functions were obtained for several standard contrasts at 1 and 8 c/deg. Binocular matching functions were obtained for luminance increments as well. The binocular contrast matching functions departed from a simple binocular averaging rule, and behaved as if the eye receiving the higher contrast disproportionately dominated the binocular contrast percept. Departures from the binocular averaging rule were slightly greater for higher standard contrasts. Spatial frequency had little effect, and the luminance increment matching functions also departed from the binocular averaging rule. There was evidence for a contrast version of Fechner’s paradox and for substantial individual differences in a form of ocular dominance. In a further experiment, additivity of suprathreshold binocular contrast summation was examined by testing the double-cancellation condition. We found no systematic violations of additivity at 1 and 8 c/deg. Models of suprathreshold binocular contrast summation were examined.     

Monocular-contingent and binocular-contingent aftereffects

Alternate monocular and binocular exposure to complementary stimulation can yield opposite but coexisting aftereffects that are contingent on whether the test display is viewed with one eye or two eyes. The motion aftereffect was studied by adapting each eye separately to a contracting spiral and both eyes together to an expanding spiral. The stationary test spiral subsequently appeared to be expanding when viewed monocularly, but to be contracting when it was seen with both eyes open. With respect to the McCollough effect, after monocular exposure to red-vertical and green-horizontal gratings and binocular exposure to red-horizontal and green-vertical gratings, the appearance of the color of the test gratings when viewed with one eye was different from that when viewed with both eyes. Opposite, coexisting aftereffects induced by complementary stimulation can be interpreted as evidence that there are unique binocular aspects to visual function.     

Binocular fixation in the newborn baby.

Three experiments are reported in which the newborn baby's ability to fixate binocularly was investigated, using the corneal reflection technique for measuring eye fixation position. Two criteria for consistent binocular fixation were assessed. These are (1) the two eyes will be optically more divergent when fixating more distant targets, and (2) each eye will be scored as being on-target when corrections for the expected deviations of the pupil center from the fixated stimulus are introduced.In the first experiment vertical arrays of lights were separately shown at distances of 10 and 20 in. from the subjects' eyes (with the retinal image size and luminance of the stimuli held constant). The 12 newborns who gave results at both viewing distances reliably converged to both stimuli, the optical divergence of the pupil centers of the eyes increasing with presentation of the more distant stimulus. In Expt 2 similar stimuli at 5 and 10 in. from the eyes were shown. It was again the case that the subjects reliably converged to the stimulus at 10 in. This was no so for the stimulus at 5 in., and many subjects fixated this stimulus with monocular vision. The failure to converge is probably due to an inability to accommodate to this near distance. In Expt 3 different stimuli (a vertical strip of light, an outline triangle and square, and an array of squares) were presented a constant distance (10 ± 1 in.) from the eyes. The majority of the 15 subjects binocularly fixated all three stimuli: for those subjects who failed to converge consistently to these stimuli the observed alternatives to binocular fixation were monocular fixation, divergent strabismus, and a third category of response that is most probably an indication of inattention to the stimulus. It can be concluded that the newborn baby possesses the ability to fixate binocularly an appropriately presented stimulus, and has the basic requirements for binocular vision.     

Suprathreshold binocular interactions for grating patterns

Binocular interactions of suprathreshold grating patterns have been investigated using a reaction time measure of contrast detection. Simple reaction times were determined for monocular and binocular viewing conditions over a contrast range from .63 to near threshold. The results from all subjects showed binocular summation for contrast levels near threshold, but there was considerable variation across subjects for contrast levels above threshold. Some subjects showed summation over the entire contrast range, but other subjects showed either binocular inhibition or binocular facilitation for some range of contrast levels. The pattern of binocular interaction for a given subject was consistent for several spatial frequencies. The differences in types of interaction between subjects, the variation in magnitude of binocular interaction with contrast level for each subject, and the data from experiments involving stimulation of noncorresponding retinal areas show that the binocular interactions found for suprathreshold stimuli cannot be accounted for on the basis of probability, and must, therefore, result from physiological interactions between the two eyes. These interactions have been investigated further under conditions of (1) induced fixation disparity, (2) horizontal gratings, and (3) orthogonally oriented gratings.     

Do background luminances interact during binocular fusion?

The question investigated in the experiments reported here was whether monocular background luminances sum during binocular fusion. Fusion was made explicit by using a random-dot stereogram (RDS) as a background stimulus. In the presence of the RDS, differential luminance thresholds were somewhat higher than in the uniform field: a full-field, binocular dot array acted as a mask for a full-field luminance change, but global depth had no effect at threshold. The amount of the binocular advantage at threshold was compared to the basic "threshold response," that is, the change in threshold resulting from raising the background luminance by a factor of 2. It was found that the amount of the binocular advantage was equivalent, on the average, to some 75% of the threshold response--significantly less than the 100% predicted by "simple summation." The amount of the binocular advantage varied substantially among observers and eyes, whereas the threshold response obeyed Weber's law in all cases: the variability was eye-, rather than threshold-dependent. Monocular thresholds did not decrease when taken with the nontest eye occluded rather than viewing a fused background. The proposition that the adaptation state of the visual system is increased during binocular fusion (Cogan, 1982) was not supported. Yet occluding the nontest eye, rather than presenting the test stimulus monocularly against a fused background, did change monocular thresholds in some eyes and observers. These findings are interpreted as evidence for a complex binocular background interaction involving both summation and inhibition.     

Temporal and spatial factors in the fading of diffuse visual images

Pattern vision breaks down after a period of inspecting a diffusely contoured dark figure upon a ground of uniform brightness and texture. When the field is illuminated with flickering light it is found that the persistence of pattern vision is inversely related to flicker frequency up to the region of fusion. At low flicker frequencies complex figures persist longer than less complex ones of equivalent area, but the reverse applies at frequencies above fusion. These findings are consistent with data obtained from experiments using stabilized retinal images     

An electrophysiological method for the study of binocular rivalry

A technique is described for recording cortical evoked responses from human Ss in binocular rivalry. Each target is illuminated with flickering light at different frequencies upon a steady background. While the S continuously indicates with a response key which target he is seeing, the cortical responses are tape recorded. The tape recording is later analyzed by average response computer separately for each eye, seeing and not seeing. This simultaneous record of the functioning of both eyes eliminates confounding variables found in earlier methods. Controls are included for rivalry changes during a stimulus cycle and for S motor response time.     

Critical flicker frequency under monocular and binocular conditions.

Critical flicker frequency of 40 undergraduate students, 20 men and 20 women whose mean age was 19.3 yr., was measured under monocular and binocular viewing conditions, using the Lafayette Visual Perception Control with Display Unit. Half of the subjects received monocular treatment first and binocular treatment second, and the other half received the treatments in the reversed order. A 2-min. rest was allowed between these treatments. Analysis shows that mean CFF under the monocular condition was significantly lower than that under binocular conditions. The findings were discussed in terms of visual constraint and eye strain and fatigue under the monocular viewing condition.     

Visual beats: Preliminary observations of perceived rate as a function of retinal locus stimulated

Visual beats produced by simultaneous presentation of two different flicker frequencies were used to determine the temporal processing of various retinal areas. The dichoptic presentation of two different flicker frequencies to the center and periphery of the retina resulted in a perceived beat rate that equalled the physical beat frequency. When the stimuli were presented to more discrete retinal areas (temporal vs temporal and nasal vs temporal), the perceived beat rate also equalled the physical beat frequency. The data indicate that different flicker frequencies stimulating divergent retinal loei can be represented accurately and simultaneously in the visual system: Differences in temporal resolution at the different retinal loei do not occur for this repetitive stimulation.     

Illusory colors promote interocular grouping during binocular rivalry

When dissimilar monocular images are presented separately to each of a person’s eyes, these images compete for visual dominance, with dominance of one image or the other alternating over time. While this phenomenon, called binocular rivalry, transpires, local image features distributed over space and between the eyes can become visually dominant at the same time; the resulting global figure implicates interocular grouping. Previous studies have suggested that color tends to influence the incidence of global dominance; in this study, we assess whether illusory color can also influence interocular grouping. To test this, we exploited the McCollough effect, an orientation-contingent color aftereffect induced by prolonged adaptation to different colors paired with different orientations. Results show that during binocular rivalry, illusory colors induced by the McCollough adaptation enhance strong interocular grouping relative to preadaptation testing, to an extent comparable in strength with the enhancement induced by real colors. Thus, illusory colors that are present only in an observer’s mind are sufficiently potent to influence low-level visual processes such as binocular rivalry.     

The psychophysical inquiry into binocular summation

Experiments that compare monocular and binocular visual performance of human psychophysical Os on a variety of visual tasks are reviewed. The review attempts to include all experiments published in English in this century, excluding work on stereopsis, rivalry, and evoked potentials. The concept of probability summation as a baseline for assessing the presence of neural summation is discussed, and the assumptions of several models for estimating probability summation are considered. Experiments are classified in terms of visual task, major categories being increment detection, flicker fusion, brightness magnitude, and contour resolution. A major conclusion is that binocular performance is superior for essentially all task categories and in most cases by a magnitude greater than that predicted by appropriate probability summation models.