Infant light adaptation shows Weber's law at photopic illuminances.

In the present study we investigate the dependence of photopic contrast thresholds on retinal illuminance in infants and adults. Contrast thresholds were measured at five retinal illuminances between about 6 and about 20,000 Td in subjects in both age groups. The forced-choice preferential looking technique was used in 3-month-old infants, and standard forced-choice techniques were used in adults. The stimulus was a 0.25 cy/deg squarewave grating phase alternated at 6 Hz. Infants' contrast thresholds were more than two log units higher than those of adults at all retinal illuminances. Contrast thresholds had a similar dependence on retinal illuminance in both infants and adults. For both age groups, contrast thresholds initially decreased with increasing retinal illuminance. However, at both ages, above a critical illuminance of about 200 Td, contrast thresholds remained constant, following Weber's law. Thus a vertical shift was sufficient to bring the two data sets into correspondence. In the context of a two-site model of light adaptation, our results imply that infants' elevated contrast thresholds cannot be explained solely on the basis of photoreceptoral immaturities. Later physiological immaturities must also limit infants' photopic contrast thresholds.

The transition from scotopic to photopic vision in 3-month-old infants and adults: an evaluation of the rod dominance hypothesis.

The scotopic to photopic transition was tested in adults and 12-week-old infants using a large field motion nulling technique at a series of luminances between -3.57 and 2.70 log cd m(-2). The stimuli were composed of 0.25 cyc deg(-1), 5.6 Hz blue/black and yellow/black sinusoidal grating components, superimposed and moving in opposite directions. The contrasts of the two components were traded off to determine motion nulls at each luminance level. An eye movement based response measure was used for infant subjects, whereas self-report was used in adults. In both age groups, the motion null values approached a scotopic asymptote consistent with V'(lambda) at the lowest luminance levels, and a photopic asymptote consistent with V(10)(lambda) at the highest luminance levels. The scotopic to photopic transition was gradual and occurred over about 3 log units between about -2 and 1 log cd m(-2) in both groups. The null values for infants and adults were highly similar at each luminance level, and the shapes of the transition curves were virtually identical at the two ages. These data suggest that at each different luminance level, the balance between rod-initiated and cone-initiated signals in the extrafoveal luminance channel is similar or identical in 12-week-old infants and adults.

Spatial frequency tuned covariance channels for red-green and luminance-modulated gratings: psychophysical data from human adults.

Both chromatic and luminance-modulated stimuli are served by multiple spatial-frequency-tuned channels. This experiment investigated the independence versus interdependence of spatial frequency channels that serve the detection of red-green chromatic versus yellow-black luminance-modulated stimuli at low spatial frequencies. Contrast thresholds for both chromatic and luminance-modulated gratings were measured within 12 individual subjects using a repeated-measures design. Spatial frequencies ranged from 0.27 to 2.16 c/deg. A covariance structure analysis of individual differences was applied to the data. We computed statistical sources of individual variability, used them to define covariance channels, and determined the number and frequency tuning of these channels. For luminance-modulated gratings, two covariance channels were found, including one above and one below 1 c/deg [cf. Peterzell, & Teller (1996). Individual differences in contrast sensitivity functions: the coarsest spatial pattern analyzer. Vision Research, 36, 3077-3085]. For chromatic gratings, correlations between thresholds for most spatial frequencies were uniformly high, yielding a single covariance channel covering all but the highest spatial frequency tested. A combined analysis of both data sets recovered the same three covariance channels, and showed that detection thresholds for low-frequency red-green chromatic and luminance-modulated stimuli are served by separate, statistically independent processes.

Spatial frequency tuned covariance channels for red-green and luminance-modulated gratings: psychophysical data from human infants.

This study concerns the spatial-frequency-tuned channels underlying infants' contrast sensitivity functions (CSFs) for red-green chromatic stimuli, and their relationship to the channels underlying infants' CSFs for luminance-modulated stimuli. Behavioral (forced-choice preferential-looking) techniques and stationary stimuli were used. In experiment 1. contrast thresholds were measured in 4- and 6-month-olds, using isoluminant red-green gratings with spatial frequencies ranging from 0.27 to 1.53 c deg. In experiment 2. contrast thresholds were measured in 4-month-olds. using both red-green and luminance-modulated gratings in the same low spatial frequency range. Covariance analyses of individual differences were performed. Experiment 1 revealed one dominant covariance channel for the detection of red-green gratings, with a second channel contributing to detection of the highest spatial frequencies used. Experiment 2 revealed two to three channels serving color and luminance: but surprisingly these channels were not statistically separable for luminance versus chromatic stimuli. Thus, covariance channels for color and luminance that are independent for adults [Peterzell & Teller (2000). Spatial frequency tuned covariance channels for red-green and luminance-modulated gratings: psychophysical data from human adults. Vision Research, 40, 417-430] are apparently interdependent in infants. These data suggest that for infants, detection thresholds for chromatic and luminance-modulated stimuli may be limited by common mechanisms.

Infant color vision: sharp chromatic edges are not required for chromatic discrimination in 4-month-olds.

In our previous demonstrations of chromatic discrimination in infants, we have used test and surround fields of different chromaticities that abutted each other at sharp chromatic edges. In order to see whether sharp chromatic edges are necessary for infants to make chromatic discriminations, 16-week-old infants were tested with three stimulus configurations in which sharp chromatic edges were eliminated. The three edge manipulations involved black borders, a dark surround, or blurred edges around the chromatic test field. In each case red, green, and violet test fields were used. Although performance decreased when sharp chromatic edges were eliminated, observers' percent correct scores remained clearly above chance for eight of the nine discriminations (three colors x three edge manipulations). We argue that all three edge manipulations reduce the likelihood of mediation of chromatic discrimination by M (magnocellular) cells. These data thus provide evidence that young infants have functional P (parvocellular) pathways, and use them for making chromatic discriminations.

Analysis of red/green color discrimination in subjects with a single X-linked photopigment gene.

Many subjects despite having only a single X-linked pigment gene (single-L/M-gene subjects) are able to make chromatic discriminations by Rayleigh matching, especially when large fields are used. We used a combination of psychophysics (Rayleigh match), electroretinograms (ERG), and molecular genetic techniques to rule out several possible explanations of this phenomenon. Use of rods for chromatic discrimination was unlikely since strong adapting fields were employed and the large-field match results were not consistent with rod participation. A putative mid- to long-wavelength photopigment that escapes detection by current molecular genetic analysis was ruled out by finding only a single L/M photopigment in flicker ERGs from 16 single-L/M-gene subjects. Large-field match results were not consistent with participation of S cones. Amino acid sequence polymorphisms in the S-pigment gene that might have shifted the S cone spectrum towards longer wavelengths were not found on sequencing. The mechanism of chromatic discrimination in the presence of a single photopigment therefore remains unknown. Further possible explanations such as variations in cone pigment density and retinal inhomogeneities are discussed.

Infants code the direction of chromatic quadrature motion.

The present experiment uses a quadrature motion paradigm to investigate the motion correspondence cues used by young infants for coding the direction of motion of red/green isoluminant gratings. Three-month-old infants and adults were tested with 0.25 c/d luminance-modulated or red/green isoluminant gratings, either moving continuously or shifted in spatial quadrature. Both direction-of-motion and detection thresholds were measured, and motion:detection (M:D) threshold ratios were examined. Infants, like adults, could code the direction of motion of red/green quadrature-shifted gratings. In adults, M:D ratios were similar for continuous and quadrature motion. In infants, M:D ratios were higher for quadrature than for continuous motion, but elevations of similar magnitude were seen for both luminance-modulated and red/green gratings. The results suggest that frequency-doubled signals, such as those often seen in the magnocellular (M-cell) pathway, are not necessary for coding the direction of motion of isoluminant gratings in infant subjects. Two other theoretical options--mediation by the scatter of isoluminance points in the M-cell population, and parvocellular (P-cell) mediation--are discussed.

Spatial and temporal aspects of infant color vision.

The present paper constitutes a review of the literature on young infants' chromatic discrimination capabilities. A series of early studies showed that infants as young as two months postnatal can make at least some chromatic discriminations between stationary, homogeneous fields of different wavelength compositions. Current studies of spatial and temporal contrast sensitivity functions (CSFs) for red/green isoluminant stimuli suggest that spatial chromatic CSFs show developmental changes in sensitivity and spatial scale, but not curve shape; while temporal chromatic CSFs (tCSFs) show developmental changes in sensitivity and curve shape, but not temporal scale. Infants can also code the direction of motion of moving isoluminant red/green gratings, for both continuous and quadrature motion. The possible mechanisms that underlie infants' chromatic discriminations are discussed.

Development of the spatio-chromatic visual evoked potential (VEP): a longitudinal study.

Most prior visual evoked potential (VEP) research on the development of color vision has employed pattern-reversing stimuli that are not optimal for producing chromatic responses. We measured infant VEPs using low spatial frequency, onset-offset stimuli, modulated along the three axes of a cone-based color space (Derrington et al. [J. Physiol 1984;357, 241-265.]). Three color-normal infants were tested in a longitudinal design over the first postnatal year. One red/green color-deficient infant was also tested at 197 days. We found that VEP responses to S-axis (tritan) stimuli have their initial onset later than responses to red/green (L-M) or achromatic stimuli, and that developmental changes in VEP waveforms are more complex and longer lasting for chromatic than for achromatic stimuli. Possible mechanisms underlying these changes are discussed.

Severity of color vision defects: electroretinographic (ERG), molecular and behavioral studies.

Earlier research on phenotype/genotype relationships in color vision has shown imperfect predictability of color matching from the photopigment spectral sensitivities inferred from molecular genetic analysis. We previously observed that not all of the genes of the X-chromosome linked photopigment gene locus are expressed in the retina. Since sequence analysis of DNA does not necessarily reveal which of the genes are expressed into photopigments, we used ERG spectral sensitivities and adaptation measurements to assess expressed photopigment complement. Many deuteranomalous subjects had L, M, and L-M hybrid genes. The ERG results showed that M pigment is not present in measurable quantities in deutan subjects. Using these results to determine gene expression improved the correlations between inferred pigment separation and color matching. Furthermore, we found a subject who had normal L and M genes and normal proximal promoter sequences, yet he had a single photopigment (M) by ERG and tested as a protanope. These results demonstrate the utility of ERG measurements in studies of molecular genetics of color vision deficiencies, and further support the conclusion that not all genes are expressed in color deficient subjects. In particular, deuteranomaly requires a presently unknown mechanism of selective expression which excludes normal M genes and allows expression of L-M hybrid genes in one cone type, and the normal L in another.

Development of pattern visual evoked potentials: longitudinal measurements in human infants.

PURPOSE: This experiment used longitudinal testing to trace the emergence of the major components of pattern visual evoked potentials (VEPs) in infants, using two paradigms: large-checkerboard pattern reversal and low spatial frequency pattern onset. METHODS: Testing with both pattern-reversal and pattern-onset stimuli was performed on the same infants. Testing was conducted at weekly intervals during the first three postnatal months, and at intervals of 2 weeks to 1 month thereafter. RESULTS: The pattern-reversal and early pattern-onset responses recorded within individual subjects showed remarkably systematic developmental sequences. The broad, positive component seen at 200 to 250 ms in infants could be traced readily through the developmental sequence, to become the more sharply tuned positive component seen at about 100 ms in adults. Responses to low spatial frequency pattern onsets in infants were larger and more reliable than those in adults. The late components of the pattern-onset response, generally attributed to pattern offset, emerged later and with more complex changes. In all cases, response amplitude was much more variable than response latency, both within and between subjects. CONCLUSIONS: Frequent VEP recording in a longitudinal design can reveal systematic and detailed transitions of wave-form during development.

Infant color vision: temporal contrast sensitivity functions for chromatic (red/green) stimuli in 3-month-olds.

In order to investigate the development of temporal contrast sensitivity functions (tCSFs) for chromatic (red/green) stimuli, we obtained chromatic contrast thresholds from 3-month-old infants and adults using behavioral techniques. Stimuli were moving or counterphase-reversing sinusoidal gratings of 0.25 c/deg. Five temporal frequencies were used: 0.7, 2.1, 5.6, 11 and 17 Hz (corresponding speeds = 2.8, 8.4, 22, 44 and 67 deg/sec). In order to compare chromatic results with those obtained under luminance-defined conditions, luminance tCSFs were also obtained from adults, and previously obtained infant luminance tCSFs were used (from Dobkins & Teller, 1996a). In accordance with previous studies, adults exhibited bandpass luminance tCSFs with peaks near 5 Hz and lowpass chromatic tCSFs that declined rapidly at temporal frequencies greater than 2 Hz, and the two curves crossed one another near 4 Hz. By contrast, infants exhibited bandpass rather than lowpass chromatic tCSFs with peaks near 5 Hz. These chromatic curves were quite similar in peak frequency and general shape to previously obtained infant tCSFs for luminance stimuli. Moreover, both chromatic and luminance tCSFs in infants were found to be quite similar in peak and shape to luminance tCSFs observed in adults. These findings point to the possibility that, for 3-month-old infants, both chromatic and luminance stimuli are detected by the same underlying mechanism under these conditions. We propose that such a mechanism is probably a physiological pathway dominated by magnocellular input. Earlier studies of infant color vision are discussed in this context.

The development of chromatic and achromatic contrast sensitivity in infancy as tested with the sweep VEP.

Swept-contrast visual evoked potential (VEP) techniques were used to measure the development of contrast sensitivity functions (CSFs) for achromatic and red/green isoluminant chromatic gratings. Subjects were infants of 8, 14, 20 and 32 weeks of age, and adults. Stimuli were 20 deg, 0.3-4 cyc/deg sinusoidal gratings, counterphased at 6 Hz and modulated through white. Achromatic and chromatic CSFs for all ages could be fit simultaneously with a double exponential equation of a common, lowpass shape. Both achromatic and chromatic CSFs exhibited developmental shifts in both sensitivity and spatial scale. From 8 weeks to adulthood, sensitivity increased by 0.64 log units for achromatic gratings and by 0.91 log units for chromatic gratings, yielding an 0.27 log unit larger sensitivity change for chromatic than for achromatic stimuli. Spatial scale shifts were closely similar across achromatic and chromatic CSFs, and were consistent with the factor of about four predicted on the basis of changes in foveal receptor packing density and eye size. The question of uniform vs differential loss of sensitivity for chromatic vs achromatic stimuli at fixed spatial frequencies is discussed.

Rayleigh match ranges of red/green color-deficient observers: psychophysical and molecular studies.

Large-field Rayleigh match ranges were measured in 27 red/green color-deficient male observers, using bright, temporally alternating, 3-9 deg annular test fields. The observers' X-linked opsin gene arrays were characterized by molecular genetic techniques, and used to infer the absorption maxima of each observer's L and/or M cone photopigment(s). Measured match ranges decreased rapidly as the inferred separation in pigment absorption maxima increased from 0 to 2-3 nm, and varied irregularly thereafter. Following He & Shevell [(1995) Vision Research, 35, 2579-2588] predicted match ranges were calculated for various pigment separations and assumed values of pigment optical density. The predicted variations in match range encompassed the measured match ranges of most (but not all) of the color-deficient observers. The calculations also showed that differences in pigment optical density, in two cone types containing the same pigment, are sufficient to allow a moderate degree of chromatic discrimination. Such models thus provide a possible account of the fact that some color-deficient observers, with only a single X-linked opsin gene, can make red/green chromatic discriminations.

Infant color vision: moving tritan stimuli do not elicit directionally appropriate eye movements in 2- and 4-month-olds.

The purpose of the present study was to investigate the capacity of infants to code the direction of motion of moving tritan-modulated gratings. Infant and adult subjects were tested with 0.2 c/d sinusoidal gratings moving at a speed of 20 deg/sec. Three conditions were tested: luminance-modulated gratings, tritan-modulated gratings, and luminance- vs tritan-modulated gratings superimposed and moving in opposite directions in a chromatic motion nulling paradigm. Two-month-old infants were tested in all three conditions, while 4-month-olds were tested in only the first two conditions. For infant subjects, an adult observer reported the direction of the slow phase of the infant's eye movements; adult subjects judged the perceived direction of motion of the stimuli. Luminance-modulated gratings produced directionally appropriate eye movements (DEM) in all age groups. Tritan gratings presented alone did not produce DEM in either 2- or 4-month-olds, but did so in adults. Mean equivalent luminance contrasts were near zero in 2-month-olds, and small but reliably above zero in adults. In sum, the present study provides no evidence that infants can code the direction of motion of moving tritan gratings.

Individual differences in contrast sensitivity functions: the lowest spatial frequency channels.

The number and nature of spatial channels tuned to low spatial frequencies in photopic vision was examined by measuring individual differences in the contrast sensitivity functions (CSFs) of seven visually normal adults. Stationary, 51 cd/m2, low spatial frequency sinusoidal gratings between 0.27 and 2.16 c/deg were used as stimuli. Correlational and factor analyses revealed that the set of CSFs contained only one statistical source of individual variability at spatial frequencies below 1 c/deg (tuned to a peak of about 0.8 c/deg), and a second source above 1 c/deg (tuned to about 1.4 c/deg). The sources ("factor-channels") mapped well onto the two coarsest spatial frequency channels from some existing computational models. The analysis was applied also to earlier data from 4-, 6- and 8-month-old infants, in which two sources of variability have been found below 1 c/deg [Peterzell, D. H., Werner, J. S. & Kaplan, P. S. (1995). Vision Research, 35, 961-980]. The combined results are consistent with the hypothesis that in photopic vision of the neonate, there are two channels with peak sensitivities below 1 c/deg, and that these channels shift their tuning from lower to higher spatial frequencies by about a factor of four during development.

Infant motion: detection (M:D) ratios for chromatically defined and luminance-defined moving stimuli.

In order to assess the relative contributions of chromatic vs luminance information to motion processing in infants, we employed a motion:detection (M:D) paradigm. Stimuli consisted of 27 deg by 40 deg, 0.25 c/deg sinusoidal gratings moving at 22 deg/sec (5.6 Hz), and were either chromatically defined or luminance-defined. Contrast thresholds for direction-of-motion (M) were obtained using a directional eye movement technique. Contrast thresholds for detection (D) were obtained using forced-choice preferential looking. M:D threshold ratios were obtained for individual infant subjects, and results were compared to those of adults. As expected, adult M:D threshold ratios were near 1:1 for luminance-defined stimuli, but greater than 1:1 for chromatically defined stimuli. This suggests that, for adults, luminance-defined, but not chromatically defined, stimuli are detected by mechanisms labeled for direction of motion. By contrast, infant M:D ratios for chromatically and luminance-defined stimuli were approximately equal and close to 1:1, suggesting that, for infants, luminance- as well as chromatically defined stimuli are detected by mechanisms that are labeled for direction of motion.

Infant color vision: motion nulls for red/green vs luminance-modulated stimuli in infants and adults.

Four-week-olds, 9-week-olds, and adult subjects were tested with low spatial frequency sinusoidal gratings moving at a speed of 25 deg/sec. Luminance-modulated and red/green gratings were presented either separately, or superimposed and moving in opposite directions in a chromatic motion nulling paradigm. An adult observer judged the direction of the slow phase of the subject's eye movements. Luminance-modulated gratings elicited directionally appropriate eye movements in all three age groups, with contrast thresholds decreasing markedly with age. For red/green gratings alone, 4-week-olds responded only marginally, but 9-week-olds and adults produced consistent directionally appropriate eye movements. In the motion nulling condition, 15% contrast luminance-modulated gratings were about equally effective in nulling the motion of the red/green gratings in all three age groups. A formal model of the motion nulling paradigm, separating threshold and equivalent luminance contrast parameters, was developed and applied to the data. Model fits showed that equivalent luminance contrast was constant or nearly constant across age groups. This outcome is consistent with the hypothesis that, with respect to adults, infants show a uniform rather than a differential loss of sensitivity to moving red/green vs luminance-modulated stimuli.

Infant contrast detectors are selective for direction of motion.

In order to investigate the presence of directionally selective mechanisms in 3-month-old infants, we employed a summation-near-threshold paradigm previously developed for studies of adult vision (Levinson & Sekuler, 1975 Journal of Physiology (London), 250, 347-366); Watson, Thompson, Murphy & Nachmias, 1980 Vision Research, 20, 341-347). The degree of contrast summation occurring between two sinusoidal gratings moving in opposite direction was determined by comparing the contrast threshold for a compound stimulus (a counterphase-reversing grating) with the contrast threshold for one of its components (a single moving grating). Using the forced-choice preferential looking (FPL) technique, contrast thresholds were obtained for both counterphase and single moving gratings within individual infant subjects. Data were collected at several speeds, ranging from 2.8 to 66.8 degrees/sec (temporal frequency range: 0.7-16.7 Hz). At slow speeds, infants' thresholds were approximately equal for counterphase and moving gratings, indicating that non-directional mechanisms were responsible for detection. At an intermediate speed (22.3 degrees/sec), thresholds were nearly twice as high for counterphase gratings as for single moving gratings, indicating the existence of directionally selective mechanisms at detection threshold for this speed. For faster speeds, relative thresholds for the two types of stimuli fell between the two extremes; a model incorporating probability summation between directionally selective mechanisms was sufficient to account for the data. These results demonstrate that, at speeds greater than or equal to 22.3 degrees/sec (t.f. = 5.6 Hz), 3-month-old infants possess directionally selective mechanisms at threshold.