Walter Makous

Enhancing the contrast sensitivity function through action video game training

The contrast sensitivity function (CSF) is routinely assessed in clinical evaluation of vision and is the primary limiting factor in how well one sees. CSF improvements are typically brought about by correction of the optics of the eye with eyeglasses, contact lenses or surgery. We found that the very act of action video game playing also enhanced contrast sensitivity, providing a complementary route to eyesight improvement.

Spatial frequency, phase, and the contrast of natural images

We examined contrast sensitivity and suprathreshold apparent contrast with natural images. The spatial-frequency components within single octaves of the images were removed (notch filtered), their phases were randomized, or the polarity of the images was inverted. Of Michelson contrast, root-mean-square (RMS) contrast, and band-limited contrast, RMS contrast was the best index of detectability. Negative images had lower apparent contrast than their positives. Contrast detection thresholds showed spatial-frequency-dependent elevation following both notch filtering and phase randomization. The peak of the spatial-frequency tuning function was approximately 0.5-2 cycles per degree (c/deg). Suprathreshold contrast matching functions also showed spatial-frequency-dependent contrast loss for both notch-filtered and phase-randomized images. The peak of the spatial-frequency tuning function was approximately 1-3 c/deg. There was no detectable difference between the effects of phase randomization and notch filtering on contrast sensitivity. We argue that these observations are consistent with changes in the activity within spatial-frequency channels caused by the higher-order phase structure of natural images that is responsible for the presence of edges and specularities.

A visual nonlinearity fed by single cones

An intensive nonlinearity in the visual system can produce distortion products, or difference frequency gratings, when observers view two high contrast, high spatial frequency interference fringes of slightly different frequency or orientation added together at the retina. These distortion products are visible even when the two fringes imaged on the retina are above the resolution limit. Our experiments take advantage of this nonlinearity to measure the spatial filtering in the visual system following the formation of the retinal image, but preceding the site of the nonlinearity. The point spread function corresponding to this spatial filter is so small that it can be entirely explained by light integration within the apertures of foveal and parafoveal cones. The small size of this point spread function implies that (1) laser interferometry avoids contrast losses inherent in the eyes optics at spatial frequencies as high as 130 c/deg, (2) retinal scatter causes negligible image degradation in the fovea and parafoveal retina, (3) eye movements have little or no effect on contrast sensitivity to the distortion product and (4) that there is no neural spatial summation in the visual system prior to the site of the nonlinearity. Distortion products could also be observed when a bright interference fringe was briefly flashed on the fovea and a test interference fringe was viewed through the resulting afterimage. Measurements of the point spread function at stages in the visual system that precede the generation of this distortion product were similar to those obtained with simultaneous presentation of the two fringes, implying that the aftereffect of light adaptation is extremely local, no larger than the dimensions of single cones.

A three channel model of temporal frequency perception.

This paper derives the constraints on a set of channels that would be consistent with the results of several experiments on the temporal properties of the visual system, and it describes a specific set of channels that meet these constraints. Data on simultaneous detection and discrimination require a minimum of three channels. Temporal frequency discrimination at and above threshold constrain the bandwidths and locations of the channels. The shape of the temporal modulation sensitivity function constrains their sensitivities. The functions that meet these constraints are similar to those derived from masking data, and they can account for data on flicker matching, notch losses in modulation sensitivity, and changes of perceived temporal frequency with changes of modulation depth.

Electrical Coupling between Mammalian Cones

BACKGROUND Cone photoreceptors are noisy because of random fluctuations of photon absorption, signaling molecules, and ion channels. However, each cones noise is independent of the others, whereas their signals are partially shared. Therefore, electrically coupling the synaptic terminals prior to forward transmission and subsequent nonlinear processing can appreciably reduce noise relative to the signal. This signal-processing strategy has been demonstrated in lower vertebrates with rather coarse vision, but its occurrence in mammals with fine acuity has been doubted (even though gap junctions are present) because coupling would blur the neural image. RESULTS In ground squirrel retina, whose triangular cone lattice resembles the human fovea, paired electrical recordings from adjacent cones demonstrated electrical coupling with an average conductance of approximately 320 pS. Blur caused by this degree of coupling had a space constant of approximately 0.5 cone diameters. Psychophysical measurements employing laser interferometry to bypass the eyes optics suggest that human foveal cones experience a similar degree of neural blur and that it is invariant with light intensity. This neural blur is narrower than the eyes optical blur, and we calculate that it should improve the signal-to-noise ratio at the cone terminal by about 77%. CONCLUSIONS We conclude that the gap junctions observed between mammalian cones, including those in the human fovea, represent genuine electrical coupling. Because the space constant of the resulting neural blur is less than that of the optical blur, the signal-to-noise ratio can be markedly improved before the nonlinear stages with little compromise to visual acuity.

Cones block signals from rods

Abstract Incremental thresholds were measured against four kinds of background light that had identical effects on rods: blue or red light entering the pupil near its edge or near its center. The match of scotopic luminance for each subject was based on his own absolute rod threshold for the two colors. Data from dark adaptation curves and the Stiles-Crawford effect show that only the rod system could detect the test flash. Yet, the threshold was either 0.3 or 0.5 log units higher against the red background than against the scotopically matched blue background, depending upon whether the red background light entered the pupil near its edge or near its center, respectively. The rods were shown to be indifferent to the loci at which the background light entered the pupil. The incremental threshold rose by more than a log unit when the test flash was presented at the same time as the change of locus at which the red background light entered the pupil. It follows that excitation of cones can prevent detection of a test flash by the rod system.

Effects of ethanol on turnover and function of striatal dopamine

Acute oral administration of ethanol increased the rate of depletion of dopamine in the striata of rats injected with α-methyl-p-tyrosine. This effect was eliminated by pretreatment with atropine or by lesioning of the striato-nigral tract. Ethanol also attenuated the inhibitory effect of apomorphine on turnover of striatal dopamine. Unilateral injection of ethanol into the neostriatum of rats followed by intraperitoneal injection of either apomorphine or amphetamine elicited marked ipsilateral head-to-tail body turning. This turning was blocked by pretreatment with haloperidol. Chronic intubation of ethanol to rats enhanced contralateral body turning elicited by unilateral intrastriatal injection of dopamine. Injection of 6-hydroxydopamine into the substantia nigra led to denervation supersensitivity of dopaminergic functions in the neostriatum. This effect was not seen in rats that were given ethanol postinjection of 6-hydroxydopamine. These results suggested that ethanol has an inhibitory effect on the nigrostriatal dopaminergic system.

Enhanced motion aftereffect for complex motions.

We measured the magnitude of the motion after effect (MAE) elicited by gratings viewed through four spatial apertures symmetrically positioned around fixation. The gratings were identical except for their orientations, which were varied to form patterns of global motion corresponding to radiation, rotation or translation. MAE magnitude was estimated by three methods: the duration of the MAE; the contrast required to null the MAE and the threshold elevation for detecting an abrupt jump. All three techniques showed that MAEs for radiation and rotation were greater than those for translation. The greater adaptability of radiation and rotation over translation also was observed in areas of the display where no adapting stimulus had been presented. We also found that adaptation to motion in one direction had equal effects on sensitivity to motion in the same and opposite directions.

Conditioned Insulin Secretion in the Albino Rat

Summary Although the phenomenon of conditioned hypoglycemia is now well established, little is known about its mechanism. The present experiments lead to the conclusion that the mechanism involves a release of insulin: Experiment 1 showed that rats given an injection of streptozotocin, a drug which destroys the beta cells of the islets of Langerhans, did not show a conditioned hypoglycemia, whereas rats given only the vehicle for the streptozotocin did; and in experiment 2, blood drawn from conditioned rats just before the conditioned hypoglycemia would normally occur showed greater insulinlike activity than blood drawn from control rats. These results demonstrate a conditioned release of some hypoglycemic agent that depends upon the integrity of the beta cells. A neural control over release of insulin must be inferred to explain them. This study was supported by the USPHS Training Grant No. 5TO1 GM 00666. Thanks are due to Dr. Clara A. Muehlbaecher and to Dr. Daniel Porte, Jr. for their helpful comments; to Jane Hajdu for the histological work; and to Dr. Raymond Pictet for supplying the streptozotocin.

Radial motion looks faster.

Current models of motion perception depend on unidirectional motion-sensitive mechanisms that provide local inputs for complex pattern motion, such as optic flow. To test the generality of such models, we asked observers to compare the speed of radial gratings with the translational speed of vertical gratings. The speed of the radial gratings was consistently overestimated by 20-60% relative to that of translating gratings that were identical in all other respects. The speed bias was not associated with a general spatial or temporal processing bias, nor with the high relative speed of points about the center of expansion/contraction. The bias increased non-linearly with the size of sectors of the radiating pattern exposed. As the motion of the two patterns was locally identical but judged differently, the apparent speed of both kinds of motion cannot be served by any mechanism, nor described by any model, that is based entirely on local motion signals. We speculate that the greater apparent speed of the radial motion has to do with apparent motion in depth.