Psychophysical measurements using achromatic grating resolution acuity in peripheral vision show a prominent retinal asymmetry in acuity which is consistent with predicted values based on available estimates of midget ganglion cell density. Recent studies have shown that peripheral grating resolution acuity values for short-wavelength-sensitive (SWS) isolating gratings in normal observers are closely related to predicted values based on the underlying small bistratified ganglion cell density. By measuring SWS resolution acuity at different locations across the visual field, we wished to see if any significant acuity asymmetry exists for the short-wavelength system. In addition to this, we wanted to compare SWS and achromatic resolution acuity at different retinal locations of equal eccentricity. SWS and achromatic grating resolution acuity was measured in two observers at a number of different retinal meridians of 10- and 25-deg eccentricity from the fovea, and out to 35-deg eccentricity along the horizontal meridian. Achromatic resolution acuity was higher than SWS resolution acuity at all locations. At 10-deg eccentricity there was slight radial asymmetry in SWS and achromatic acuity, both displaying highest acuity along the horizontal meridian. At 25-deg eccentricity, SWS and achromatic acuity showed significant asymmetry with acuity being higher in the nasal retina compared to the temporal retina and with higher acuity in the superior retina compared to the inferior retina. At 35-deg eccentricity, the acuity asymmetry along the horizontal meridian was maintained with acuity for both significantly higher in the nasal retina. The SWS acuity changes with eccentricity and meridian were qualitatively similar to that found for achromatic acuity at the majority of retinal locations. Like achromatic acuity, SWS acuity shows significant asymmetry at different retinal locations of equal eccentricity. This suggests that both the midget and small bistratified ganglion cell population density changes significantly with retinal location and eccentricity. SWS acuity appears to change in parallel with achromatic acuity for the majority of retinal locations measured, although the amount of nasotemporal asymmetry appears to be slightly less for the SWS system at 25- and 35-deg eccentricity.

The two-dimensional organization of receptive fields (RFs) of 44 cells in the cat visual cortex and four cells from the cat LGN was measured by stimulation with either dots or bars of light. The light bars were presented in different positions and orientations centered on the RFs. The RFs found were arbitrarily divided into four general types: Punctate, resembling DOG filters (11%); those resembling Gabor filters (9%); elongate (36%); and multipeaked-type (44%). Elongate RFs, usually found in simple cells, could show more than one excitatory band or bifurcation of excitatory regions. Although regions inhibitory to a given stimulus transition (e.g. ON) often coincided with regions excitatory to the opposite transition (e.g. OFF), this was by no means the rule. Measurements were highly repeatable and stable over periods of at least 1 h. A comparison between measurements made with dots and with bars showed reasonable matches in about 40% of the cases. In general, bar-based measurements revealed larger RFs with more structure, especially with respect to inhibitory regions. Inactivation of lower cortical layers (V-VI) by local GABA injection was found to reduce sharpness of detail and to increase both receptive-field size and noise in upper layer cells, suggesting vertically organized RF mechanisms. Across the population, some cells bore close resemblance to theoretically proposed filters, while others had a complexity that was clearly not generalizable, to the extent that they seemed more suited to detection of specific structures. We would speculate that the broadly varying forms of cat cortical receptive fields result from developmental processes akin to those that form ocular-dominance columns, but on a smaller scale.

Tritiated proline and leucine were injected into areas 17 and 18 of intact cats and into the medial bank of the lateral suprasylvian (LS) cortex of intact cats and cats from which areas 17 and 18 had been removed on postnatal day 1 (P1), P28, or in adulthood (A). The density of label transported to the superior colliculus was quantified using image-analysis equipment. The results from the intact cats confirmed previous reports that areas 17 and 18 project most heavily to stratum zonale (SZ) and stratum griseum superficiale (SGS) and LS cortex projects most heavily to stratum opticum (SO) of the superior colliculus. However, in cats with lesions of areas 17 and 18, the projections from LS cortex showed an age-dependent reorganization. LS projections to SGS and SZ were enhanced following ablation of areas 17 and 18 on P1, and projections to SGS were enhanced following an ablation on P28. The pattern of LS-collicular projection following ablations incurred in adulthood was indistinguishable from the pattern presented by intact cats. This study demonstrates that the LS corticocollicular projection expands in SGS and possibly substitutes for inputs eliminated by the removal of areas 17 and 18 from the immature brain. This enhanced pathway may contribute to compensatory neuronal changes and to spared behaviors that accompany damage of immature cortex.

Effects of light adaptation on contrast processing in the outer retina were investigated over nearly four decades of background illumination by analyzing the intracellular responses of 111 bipolar cells, 66 horizontal cells, and 22 cone photoreceptors in the superfused eyecup of the tiger salamander (Ambystoma tigrinum). Light adaptation had striking and similar effects on the average contrast responses of the hyperpolarizing (Bh) and depolarizing (Bd) classes of bipolar cells: Over the lower two decades of background illumination, the contrast gain increased 7-fold to reach values as high as 20–30, the dynamic range and the half-maximum contrast decreased by about 60%, the total voltage range increased some 40%, and contrast dominance changed from highly positive to more balanced. At higher levels of background, most aspects of the contrast response stabilized and Weber's Law then held closely. In this background range, the contrast gain of bipolar cells was amplified some 20× relative to that of cones whereas the corresponding amplification in horizontal cells was about 6×. Differences in the growth of contrast gain with the intensity of the background illumination for cones versus bipolar cells suggest that there are at least two adaptation-dependent mechanisms regulating contrast gain. One is evident in the cone photoresponse such that an approximately linear relation holds between the steady-state hyperpolarization and contrast gain. The other arises between the voltage responses of the cones and bipolar cells. It could be presynaptic (modulation of cone transmitter release by horizontal cell feedback or other mechanisms) and/or postsynaptic, that is, intrinsic to bipolar cells. Contrast gain grew with the background intensity by a larger factor in horizontal than in bipolar cells. This provides a basis for the widely held view that light adaptation increases the strength of surround antagonism in bipolar cells. On average, the effects of light adaptation and most quantitative indices of contrast processing were remarkably similar for Bd and Bh cells, implying that both classes of bipolar cells, despite possible differences in underlying mechanisms, are about equally capable of encoding all primary aspects of contrast at all levels of light adaptation.

Thresholds for psychophysically opposite stimuli—light and dark, or equiluminous red and green, or equiluminous blue and yellow—are usually nearly equal. This color threshold symmetry is sometimes violated in subjects who have optic nerve hypoplasia, a congenital loss of retinal ganglion cells. We describe a subject who has optic nerve hypoplasia, who exhibits large red-green and blue-yellow detection threshold asymmetries for equiluminous spots. Temporal and spatial integration for equiluminous red and green test spots also differed from normal; static perimetric thresholds for equiluminous green, blue, and yellow (but not red) spots lacked the normal “V” shaped minimum at the fovea. These asymmetries may relate to a developmental paucity of some ganglion cell subtypes. Optic nerve hypoplasia may allow the contributions to detection made by individual ganglion cell subtypes to be isolated psychophysically, in analogy to the study of cone spectral sensitivity in dichromats.

This study examined the influence of the pontomesencephalic peribrachial region (PBR) on the visual response properties of cells in the dorsal lateral geniculate nucleus (LGN). The response of single cells to a stationary flashing light spot was recorded with accompanying electrical stimulation of the PBR. The major objectives were to compare the effects of PBR stimulation on lagged and nonlagged cells, to examine how the visual response pattern of lagged cells could be modified by PBR stimulation and to examine whether the physiological criteria used to classify lagged and nonlagged cells are applicable during increased PBR input to the LGN. During PBR stimulation, the visual response was enhanced to a similar degree for lagged and nonlagged cells and the latency to half-rise of the visual response was reduced, particularly for the lagged X cells. The latency to half-fall of the visual response of lagged cells was not changed by PBR stimulation. Accordingly, the division of LGN cells into lagged and nonlagged cells based on visual response latencies was maintained during PBR stimulation. The initial suppression that a visual stimulus evokes in lagged cells was resistant to the effects of PBR stimulation. For the lagged cells, the largest response increase occurred for the initial part of the visual response. For the nonlagged cells, the largest increase occurred for the tonic part of the response. The results support the hypothesis that the differences in temporal response properties between lagged and nonlagged cells belong to the basic distinctions between these cell classes.

Changes of color perception in the peripheral field are measured using an asymmetric simultaneous matching paradigm. The data confirm previous observations in that saturation changes can be neutralized if the test target is increased in size. However, this compensation does not apply to hue shifts. We show that some hues remain unchanged with eccentricity whereas others exhibit substantial changes. Here the color shifts are plotted in terms of a second-stage cone opponent model. The data suggest that the S-L+M channel is more robust to increasing eccentricity than the L-M channel. Observations are interpreted in terms of the known underlying morphological and physiological differences in these channels.

Neurons in the superficial layers of the midbrain superior colliculus (SC) exhibit distinct tuning properties for visual stimuli, but, unlike neurons in the geniculocortical visual pathway, most respond best to visual stimuli that are smaller than the classical receptive field (RF). The mechanism underlying this size selectivity may depend on the number and pattern of feedforward retinal inputs and/or the balance between inhibition and excitation within the RF. We have previously shown that chronic blockade of NMDA receptors (NMDA-R), which increases the convergence of retinal afferents onto SC neurons, does not alter size selectivity in the SC. This suggests that the number of retinal inputs does not determine size selectivity. Here we show, using single unit extracellular recordings from the SC of normal hamsters, that size selectivity in neurons selective for small stimulus size is correlated with the strength of inhibition within the RF. We also show that dark rearing causes concomitant reductions in both inhibition and size selectivity. In addition, dark rearing increases the percentage of neurons non-selective for stimulus size. Finally, we show that chronic blockade of NMDA-R, a procedure that does not alter size tuning, also does not change the strength of inhibition within the RF. Taken together, these results argue that inhibition within the RF underlies selectivity for small stimulus size and that inhibition must be intact for size tuning to be preserved after developmental manipulations of activity. In addition, these results suggest that regulation of the balance between excitation and inhibition within the RF does not require NMDA-R activity but does depend on visual experience. These results suggest that developmental experience influences neural response properties through an alteration of inhibitory circuitry.

The N-methyl-D-aspartate receptor (NMDAR) is an ionotropic glutamate receptor that is important in neurotransmission as well as in processes of synaptic plasticity in the mammalian superior colliculus (SC). Despite the importance of this receptor in synaptic transmission, there is as yet no evidence that demonstrates directly the synaptic localization of the NMDAR receptor in SC. We have used electron-microscope (EM) immunocytochemistry to localize the NMDAR1 subunit of this receptor protein and its association with sensory afferents in the cat SC. Retinal synaptic terminals were identified by normal morphology and cortical synaptic terminals by degeneration after lesions of areas 17–18 of the visual cortex. At the light-microscope level, label was densest within the superficial gray and upper optic layers, but also present in all other layers. Label was contained within cell bodies, dendrites, and a few putative axons. At the EM level, antibody labeling was found along postsynaptic densifications and internalized within the cytoplasm of a variety of dendrites and some cell bodies. Postsynaptic profiles labeled by NMDAR1 included conventional dendrites and presynaptic dendrites which contained pleomorphic synaptic vesicles and are known to be GABAergic. Many of the labeled postsynaptic densifications of both of these profile types received synaptic inputs from retinal or cortical terminals. Virtually no NMDAR1 immunoreactivity was found on thin dendritic thorns or putative spines, even when these were postsynaptic to retinal or cortical terminals. In summary, these results show that the NMDAR1 subunit is postsynaptic to both retinal and cortical afferents, which are known to be glutamatergic, and are consistent with physiological evidence showing that stimulation of either pathway can activate the NMDA receptor.

The eyes of adult barn owls (Tyto alba) are virtually fixed in the head in positions that are highly consistent from one individual to the next. However, early in development the eyes are exodeviated; the eyes achieve their adult positions during the owl's second month of life. Disruption of binocular vision in baby owls leads to permanent, highly abnormal eye positions and interocular alignment. Of three owls raised with both eyelids sutured closed, two developed exotropic strabismus and one developed esotropic strabismus. Two owls reared with monocular vision developed esotropic strabismus, whereas three owls reared with fused, but optically deviated binocular vision developed normal eye positions. Thus, the alignment of the eyes in adults results from an active process that depends on fused binocular vision during early life.

Extracellular microelectrode recordings from the optic tecta of strabismic owls reveal that many units retain binocular inputs from corresponding points of the two eyes: the left-eye and right-eye receptive fields of individual units are misaligned by an amount predicted by the direction and magnitude of the strabismus. These results indicate that an innately determined pattern of connections in the brain anticipates the eye positions necessary to achieve binocular fusion. The hypothesis is put forth that the powerful activation of such binocular neurons by strong, synchronous inputs from the two eyes is the signal required by the optimotor system that proper eye alignment has been attained.

The purpose of this study was to specify isoluminance at different retinal eccentricities and to characterize macular pigment optical density (MPOD) and distribution using the steady-state visual evoked potential (VEP). Red–green (R/G) and blue–green (B/G) gratings were generated within two circular stimulus fields (radius = 0.55 or 1.1 deg) and within four annular fields (maximum mean radius = 6.0 deg) on a color monitor. Temporal frequency was 15 Hz. Isoluminance was determined for each stimulus using minimum flicker photometry. Steady-state onset–offset VEPs were recorded to the same annular stimuli as the luminance ratio between adjacent chromatic components was changed from 0.25 to 0.85 in 11 automated steps (0.5 representing photometric isoluminance). Fourier analysis showed that the power of the first harmonic was minimized at the isoluminant ratio specific to each subject. Relative OD was computed by comparing the isoluminant ratio at any location with that for the most eccentric annulus. To compensate for the broadband characteristics of the monitor, OD values were corrected according to minimum flicker measurements made through known concentrations of carotenoid solution. MPOD was additionally measured using minimum motion photometry.

There was high correlation between the isoluminant ratios determined by minimum flicker and VEPs for both R/G and B/G stimulation (r = 0.91, P < 0.005, slope = 1). Calibrated OD values computed from VEP estimates of B/G isoluminance correlated with those derived from minimum flicker (r = 0.96, P < 0.0005, slope = 0.85) and motion photometry (r = 0.94, P < 0.0005, slope = 0.88). OD values derived from B/G VEPs increased toward the fovea and corresponded closely with minimum flicker and minimum motion assessment of MP distribution profiles. The steady-state VEP can be used to determine isoluminance at different retinal eccentricities. MPOD and distribution can be measured by steady-state VEPs to B/G stimuli.

Horizontal cells and cone photoreceptors in the vertebrate retina are interconnected by a complex network of synapses leading to the generation of color-coded responses in chromaticity horizontal cells. A simple cascade model of excitatory feedforward and inhibitory feedback synapses had been suggested to underlie these observations. In this study, the photoresponses of cones and horizontal cells were recorded intracellularly from the turtle eyecup. Three different approaches were adopted in order to test quantitatively the cascade model. Comparing linearity functions between these neurons indicated multiple excitatory inputs to each type of horizontal cells. The depolarizing photoresponses of R/G C-type horizontal cells were considerably faster than those of L-type horizontal cells but slower than those recorded from L-cones. This observation disagrees with the basic assumption of the cascade model that assign the depolarizing photoresponses of R/G C-type horizontal cells to a negative feedback pathway from L-type horizontal cells onto M-cones. Finally, the action spectra of each of the three types of horizontal cells could not be solely accounted for by input from one spectral type of cones. Only by assuming excitatory and inhibitory inputs from all spectral types of cones, the action spectra of all types of horizontal cells could be reconstructed. These findings suggest that the negative feedback pathways from horizontal cells onto cones in the turtle retina cannot solely account for the chromatic properties of the horizontal cells and support a direct inhibitory inputs from cones to turtle horizontal cells.

In the frog optic tectum, the spatiotemporal pattern of neuronal activity evoked by electrical stimulation of the optic tract was examined by means of a current source density (CSD) analysis. The CSD depth profile was highly reproducible in different experiments. In all seven CSD profiles, three current sinks A, B, and D were observed in the retinorecipient layers. Four out of the seven profiles show additional two sinks C and E below the retinorecipient layers. Very small and short lasting sinks related to afferent fiber activities precede sinks A and B by about 1 ms, which could be accounted for by monosynaptic delay, in the corresponding depth region. The earliest prominent sink A at the bottom of the retinorecipient layers reflects only excitatory monosynaptic activities derived from R3 and/or R4 retinal ganglion cells. The second prominent sink B in the superficial retinorecipient layer is composed partly of excitatory monosynaptic activity from medium-sized myelinated optic fibers. It may involve excitatory monosynaptic activity from unmyelinated optic fibers and further polysynaptic activity. The fourth prominent sink D in the intermediate retinorecipient layer partially reflects excitatory monosynaptic activity derived from unmyelinated optic fibers. It may also involve further polysynaptic activity. In contrast with these three sinks, the third prominent sink C and fifth sink E exclusively reflect intratectal polysynaptic activity that has not been reported in any previous CSD studies in the frog optic tectum. These sinks almost overlap spatially in the tectal layer. We also measured the intratectal resistance changes and computed inhomogeneous CSD depth profiles to show that the results from homogeneous CSD computation assuming constant conductivity are valid for our present study. Finally, we compared the present results with previously reported CSD studies on the frog optic tectum and discuss consistencies and discrepancies among these experiments.

The uses of light for cephalopods living at various depths are described. Aphakic apertures are shown in the eyes of Amphitretus and bolitaenids. In cirrate octopods, the eye is an open cup without lens and the retinal rhabdoms are disorganized. The photosensitive vesicles of cephalopods are extraocular receptors present either in the mantle or on the head. In some mesopelagic forms, they serve to compare the downwelling light with that emitted by the animal's own photophores, thus allowing regulation of counterillumination. In bathypelagic species, the photosensitive vesicles are very large and may serve to ensure reproduction at great depths.

Some of the uses of the paired eyes in shallow water species are discussed. The mechanism for visual learning consists of a system for allowing many possible combinations of the output from numerous feature detectors. This begins with a set of columns in the optic lobes, followed by a tangential system. Outputs from the optic lobe lead to either attack or retreat: a third output leads to a memory system of four matrices allowing for interaction among the visual signals and between them and signals of taste or pain. These matrices allow conjunctive interaction between particular sets of signals and the setting up of memories ensuring appropriate responses. The matrices may be considered as analogous with those of the mammalian hippocampus. They include re-excitation among themselves and with the optic lobes. The tactile memory apparatus of the octopus has four similar lobes and also makes use of the four lobes of the visual system. These are therefore striking examples of adaptive networks allowing learned reactions by statistical selection among numerous channels. The anatomy, function, and generalizing powers of these networks emerged from Boycotts's early work, whose significance for computation can now be appreciated.

The influence of postnatal neuronal activity on the magnitude of retinal ganglion cell death has been studied in cats. A constant blockade of activity in one eye starting just after birth does not change the severity of naturally occurring ganglion cell death, and as in normal animals, the ganglion cell population declines from 250,000 to 160,000 over a 4- to 6-week period. However, the population of retinal ganglion cells in the active untreated eye of monocularly deprived cats is increased 12% above normal (180,000 vs. 160,000 in each of four cases). This increase of 20,000 cells is permanent, and presumably reflects the competitive advantage in their target nuclei that the still active axons have over their silenced companions from the treated eye. Surprisingly, in one animal treated successfully for long duration with TTX in both eyes, the population of ganglion cells was elevated in both eyes (200,000 and 208,000 ganglion cells). This increase matches that achieved by early unilateral enucleation (Williams et al., 1983). Our results demonstrate that the complete blockade of activity reduces the severity of naturally occurring cell death in a population of CNS sensory neurons. The effects of unilateral blockade emphasize that the activity-dependent modulation of neuron death only occurs under conditions that do not place the inactive population of neurons at a competitive disadvantage.

The laminar and tangential organization of visual callosal projections of areas 17 and 18 were investigated in the adult ferret, using histochemical methods to visualize axonally transported horseradish peroxidase (HRP). Normal adult ferrets were given injections of HRP throughout one visual cortex or had gelfoam soaked in HRP applied to the transected corpus callosum. The ferret callosal cell distribution has a greater tangential extent in area 18 than in area 17. In addition, the radial organization of callosal cells in areas 17 and 18 differs: three times as many infragranular cells are present in area 18 than in area 17, although the number of supragranular cells is similar for both areas 17 and 18. Since the projections of alpha retinal ganglion cells are reported to be exclusively contralateral in the ferret (Vitek et al., 1985), callosal projections may make a major contribution to the binocularity of neurons in area 18.

Some visual stimuli produce a strong percept of motion, even though they fail to excite motion detectors based on Fourier energy or cross correlation. Models which suffice to explain the motion percept in these non-Fourier motion (NFM) stimuli include linear spatiotemporal filtering, followed by rectification, followed by standard motion analysis (Chubb & Sperling 1988). We used the human “motion-onset” evoked potential, which has been assigned to area 17 on the basis of work in the macaque (van Dijk et al., 1986; van Dijk & Spekreijse, 1989), to investigate the neural substrate of the processing stages postulated in the above models. Motion-onset VEPs elicited by FM and NFM matched for spatial and temporal characteristics were indistinguishable in temporal characteristics and scalp topography at a transverse chain of electrodes. Addition of textural cues (granularity and higher-order form) did not influence the response dynamics or scalp topography of NFM responses. However, comparison of responses to NFM stimuli and related stimuli without coherent motion but similar spatial and temporal properties showed that the motion-onset responses were distinct from responses to the onset of fixed flicker-defined contours not undergoing coherent motion. We discuss the implications of these results for computational models of motion analysis.

The corticogeniculate circuit is an evolutionarily conserved pathway linking the primary visual cortex with the visual thalamus in the feedback direction. While the corticogeniculate circuit is anatomically robust, the impact of corticogeniculate feedback on the visual response properties of visual thalamic neurons is subtle. Accordingly, discovering the function of corticogeniculate feedback in vision has been a particularly challenging task. In this review, the morphology, organization, physiology, and function of corticogeniculate feedback is compared across mammals commonly studied in visual neuroscience: primates, carnivores, rabbits, and rodents. Common structural and organizational motifs are present across species, including the organization of corticogeniculate feedback into parallel processing streams in highly visual mammals.

Previous studies on fish retina have suggested that a curved, non-fused embryonic fissure is associated with, and perhaps caused by, asymmetric growth along the retina's marginal germinal zone (where neurons and Miiller glia are added appositionally throughout life). In this report retinal growth was measured directly in adult green sunfish (Lepomis cyanellus), which has a curved, non-fused embryonic fissure. Growth was asymmetric in both small and large fish: ventral and nasal retina grew more than temporal and dorsal retina. This asymmetry was due to different net rates of cellular addition, rather than differential passive expansion. The absolute rates of retinal growth in the centroperipheral direction were roughly exponential functions of fish size—smaller fish grow faster than large fish—but the area of new retina added per unit time did not vary with fish size. Visual implications of asymmetric retinal growth are evaluated.

A class 1 aldehyde dehydrogenase (ALDH) catalyzes oxidation of retinaldehyde to retinoic acid in bovine retina. We used immunocytochemistry and in situ hybridization to localize this enzyme in adult and fetal bovine retinas. Specific ALDH immunoreactivity was present in the cytoplasm of wide-field amacrine cells restricted in distribution to the dorsal part of the adult retina. The somata diameters ranged from ∼8 μ to ∼15 μ, and the cells increased in density from ∼125 cells/mm2 near the horizontal meridian to ∼425 cells/mm2 in the superior far periphery. The ALDH-positive cells had somata on both sides of the inner plexiform layer (IPL) and processes in two IPL strata. The majority of ALDH-positive cells were unreactive with antibodies against known amacrine cell enzymes and neurotransmitters, including GABA and glycine. The ALDH-positive amacrine cells also did not react with anti-cellular retinoic acid-binding protein, which was present in a subset of GABA-positive amacrine cells. In flat-mounted retinas processed by in situ hybridization, the larger ALDH-positive amacrine cells tended to be more heavily labeled. In addition to amacrine cells, Müller cell processes in the inner retina were weakly immunoreactive for ALDH; however, these glial cells did not contain ALDH mRNA. The pattern of ALDH expression in fetal bovine retinas was documented by immunocytochemistry. No ALDH reactivity was found before 5.5 months; for the remainder of the fetal period, ALDH immunoreactivity was present in amacrine cells similar to those in adult retina. The ALDH-positive amacrine cells in bovine retina are novel, being limited in distribution to the dorsal retina and unlabeled with other amacrine cell-specific markers. Identification of ALDH in amacrine cells provides additional evidence that cells of the inner retina are involved in retinoid metabolism.