A psychophysical analog to cortical receptive-field end-stopping has been demonstrated previously in spatial filters tuned to a wide range of spatial frequencies (Yu & Levi, 1997a). The current study investigated tuning characteristics in psychophysical spatial filter end-stopping. When a D6 (the sixth derivative of a Gaussian) target is masked by a center mask (placed in the putative spatial filter center), two end-zone masks (placed in the filter end-zones) reduce thresholds. This “end-stopping” effect (the reduction of masking induced by end-zone masks) was measured at various spatial frequencies and orientations of end-zone masks. End-stopping reached its maximal strength when the spatial frequency and/or orientation of the end-zone masks matched the spatial frequency and/or orientation of the target and center mask, showing spatial-frequency tuning and orientation tuning. The bandwidths of spatial-frequency and orientation tuning functions decreased with increasing target spatial frequency. At larger orientation differences, however, end-zone masks induced a secondary facilitation effect, which was maximal when the spatial frequency of end-zone masks equated the target spatial frequency. This facilitation effect might be related to certain types of contour and texture perception, such as perceptual pop-out.

Many rodents do not see well. For a target to be resolved by a rat or a mouse, it must subtend a visual angle of a degree or more. It is commonly assumed that this poor spatial resolving capacity is due to neural rather than optical limitations, but the quality of the retinal image has not been well characterized in these animals. We have modified a double-pass apparatus, initially designed for the human eye, so it could be used with rodents to measure the modulation transfer function (MTF) of the eye's optics. That is, the double-pass retinal image of a monochromatic (λ = 632.8 nm) point source was digitized with a CCD camera. From these double-pass measurements, the single-pass MTF was computed under a variety of conditions of focus and with different pupil sizes. Even with the eye in best focus, the image quality in both rats and mice is exceedingly poor. With a 1-mm pupil, for example, the MTF in the rat had an upper limit of about 2.5 cycles/deg, rather than the 28 cycles/deg one would obtain if the eye were a diffraction-limited system. These images are about 10 times worse than the comparable retinal images in the human eye. Using our measurements of the optics and the published behavioral and electrophysiological contrast sensitivity functions (CSFs) of rats, we have calculated the CSF that the rat would have if it had perfect rather than poor optics. We find, interestingly, that diffraction-limited optics would produce only slight improvement overall. That is, in spite of retinal images which are of very low quality, the upper limit of visual resolution in rodents is neurally determined. Rats and mice seem to have eyes in which the optics and retina/brain are well matched.

We report VEP studies which delineate interactions between chromatic and luminance contrast signals. We examined responses to sinusoidal luminance gratings undergoing 4-Hz square-wave contrast reversal, upon which standing gratings with various admixtures of luminance and chromatic contrast were alternately superimposed and withdrawn. The presence of the standing grating induced a VEP component at the fundamental frequency of the contrast-reversal grating. This VEP component appeared without any appreciable lag, and did not vary in amplitude over the 4 s during which the standing grating was present. The observed fundamental response differed from the fundamental component that would be expected from the known interaction between the luminance component of the standing grating with the modulated grating (Bodis-Wollner et al., 1972; Bobak et al., 1988), in three ways: (1) The fundamental response was not nulled for standing gratings that were isoluminant or near-isoluminant. (2) The chromatic dependence of the fundamental response implied an S-cone input to the interaction. (3) No single mechanism (driven by a linear combination of cone signals) could account quantitatively for the size of this response, particularly when the standing grating strongly modulated two cones in phase.

Antibodies directed against the calcium-binding proteins, parvalbumin and calbindin, can be used to label distinct neuronal subgroups in the primate visual pathway. We analyzed parvalbumin immunoreactivity (P-IR) and calbindin immunoreactivity (C-IR) in the lateral geniculate nucleus (LGN) and visual cortex of the marmoset, Callithrix jacchus. We compared marmosets which were identified as having dichromatic or trichromatic color vision. Within the LGN, the density of P-IR neurones is highest in the parvocellular and magnocellular laminae, but C-IR neurones are found mainly in the koniocellular division of the LGN, that is, the interlaminar zones and S laminae. Not all interlaminar zone cells are C-IR. In the visual cortex, P-IR neurones are present in all laminae except lamina 1, in areas V1 and V2. Neurones which are strongly C-IR are mainly located in laminae 2 and 3 in V1 and V2. Lightly C-IR neurones are concentrated in lamina 4, and are more numerous in V1 than in V2. Quantitative analysis showed no differences in the density or distribution of IR neurones in either LGN or visual cortex when dichromat and trichromat animals were compared. We conclude that this functional difference is not associated with differences in the neurochemistry of calcium-binding proteins in the primary visual pathways.

To assess the dolphin's capacity for color vision and determine the absorption maxima of the dolphin visual pigments, we have cloned and expressed the dolphin opsin genes. On the basis of sequence homology with other mammalian opsins, a dolphin rod and long-wavelength sensitive (LWS) cone opsin cDNAs were identified. Both dolphin opsin cDNAs were expressed in mammalian COS-7 cells. The resulting proteins were reconstituted with the chromophore 11-cis-retinal resulting in functional pigments with absorption maxima (λmax) of 488 and 524 nm for the rod and cone pigments respectively. These λmax values are considerably blue shifted compared to those of many terrestrial mammals. Although the dolphin possesses a gene homologous to other mammalian short-wavelength sensitive (SWS) opsins, it is not expressed in vivo and has accumulated a number of deletions, including a frame-shift mutation at nucleotide position 31. The dolphin therefore lacks the common dichromatic form of color vision typical of most terrestrial mammals.

Reverse correlation was used in conjunction with ternary white noise to estimate the first-order spatiotemporal receptive-field structure of LGN cells in the anesthetized, paralyzed cat. Based on a singular-value decomposition of these data, we conclude that most LGN cells are approximately space–time separable. An analysis of the timecourses of the first singular values revealed a strongly bimodal but continuous distribution of rise times and waveforms. The two modes represented cells generally associated with the lagged and nonlagged classes of Mastronarde (1987a,b), and this was confirmed by their responses to step and sine-modulated spots in their field centers. The intermediate cells, rather than appearing to constitute a separate group, smoothly filled the region between the obviously lagged and nonlagged cells in every respect. These conclusions are limited to X-cells although the data from a much smaller population of Y-cells conform to the same scheme. We conclude that lagged and nonlagged cells represent the modes of a continuous and very broad distribution of temporal responses in the cat LGN.

The population of retinal ganglion cells which project ipsilaterally in the brain was examined in the fat-tailed dunnart, Sminthopsis crassicaudata, following injection of horseradish peroxidase into one optic tract. Retinae were examined as wholemounts and optic nerves as serial sections. In addition, visual fields were measured ophthalmoscopically. Ipsilaterally projecting ganglion cells were located temporal to a line which ran vertically through the middle of the area centralis and extended medially to define a ventrolateral crescent. Temporal to the naso-temporal division, a mean of 77% of ganglion cells projected ipsilaterally; these cells represented 20% of the total ganglion cell population. The magnitude and retinal location of the ipsilateral projection correlated with the extensive binocular field which measured 180 deg in the vertical (from 20 deg below the horizontal axis to 70 deg beyond the zenith) and 140 deg in horizontal meridian. Ipsilaterally projecting axons were restricted to the lateral third of the optic nerve along its length, sharing territory with contralaterally projecting axons.

The distributions of rod and cone photoreceptors have been determined in the retina of the California ground squirrel, Spermophilus beecheyi. Retinas were fixed by perfusion and the rods and cones were detected with indirect immunofluorescence using opsin antibodies. Local densities were determined at 2-mm intervals across the entire retina, from which total numbers of each receptor type were estimated and isodensity distributions were constructed. The ground squirrel retina contains 7.5 million cones and 1.27 million rods. The peak density for the cones (49,550/mm2) is found in a horizontal strip of central retina 2 mm ventral to the elongated optic nerve head, falling gradually to half this value in the dorsal and ventral retinal periphery. Of the cones, there are 14 M cones for every S cone. S cone density is relatively flat across most of the retina, reaching a peak (4500/mm2) at the temporal end of the visual streak. There is one exception to this, however: S cone density climbs dramatically at the extreme dorso-nasal retinal margin (20,000/mm2), where the local ratio of S to M cones equals 1. Rod density is lowest in the visual streak, where the rods comprise less than 5% of the local photoreceptor population, increasing conspicuously in the ventral retina, where the rods achieve 30% of the local photoreceptor population (13,000/mm2). The functional importance of the change in S to M cone ratio at the dorsal circumference of the retina is compromised by the extremely limited portion of the visual field subserved by this retinal region. The significance for vision, if any, remains to be determined. By contrast, the change in rod/cone ratio between the dorsal and ventral halves of the retina indicates a conspicuous asymmetry in the ground squirrel's visual system, suggesting a specialization for maximizing visual sensitivity under dim levels of illumination in the superior visual field.

ninaED1, a dominant allele of the major Drosophila rhodopsin gene, expresses a rhodopsin that is predominantly recovered in a 80-kD complex that likely represents rhodopsin dimers. By driving either ninaED1 or ninaE+ expression from a heat-shock promoter, we show that the 80-kD rhodopsin complex forms immediately after gene activation. In wild type, but not ninaED1, rhodopsin monomeric forms are detected at later times. The generation of monomeric forms of wild-type rhodopsin is suppressed in vitamin A-deprived flies or in flies heterozygous for the dominant rhodopsin mutation. We also show that ninaED1 expression does not affect the maturation of another Drosophila visual pigment, Rh3. These results are consistent with the view that the ninaED1 rhodopsin antagonizes an early posttranslation process that is specific for maturation of the ninaE-encoded rhodopsin.

The common brush-tailed possum Trichosurus vulpecula is a small diprotodont marsupial common to both urban and natural environments. This is the first analysis of the neurotransmitter content of its retinal cells and, as the possum is a nocturnal forager, it was appropriate to begin with the dopaminergic amacrine cells that form an essential link in the modulation of the rod pathways subserving nocturnal vision. These results were compared with those from another diprotodont, the marsupial wallaby or quokka (Dann, 1996) to establish whether the dopaminergic systems were similar between these two diprotodont marsupials and also to compare these findings with those of other mammals. This study describes a population of amacrine cells in the possum retina that were immunolabelled with an antibody raised against tyrosine hydroxylase (TH). These TH-immunoreactive (IR) cells were located within the inner nuclear layer (INL) and their dendrites predominantly ramified within the most sclerad layers of the inner plexiform layer (IPL). The TH-IR amacrines formed a sparse cell population, of around 2400 cells, distributed over the entire retina. There was little evidence of a concentration gradient except for a slight elevation in density in the naso-temporal axis in dorsal retina. The formation of rings within the dendritic plexus, a feature common to TH-IR cells in other species, was also present in the possum and these appeared relatively frequently. This latter finding was rather unexpected since, in the marsupial quokka (Dann, 1996), the TH-IR dendrites formed fewer rings despite having the same density of TH-IR amacrines as the possum. This suggests that there may be subtle differences in the way the rod pathways are interconnected even within the same marsupial group and may also be a reflection of relative rod dominance across species.

Action potentials of single perigeniculate (PGN) cells and relay cells of the dorsal lateral geniculate nucleus (dLGN) with topographically matched or at least partially overlapping receptive fields (RF) were simultaneously recorded in the anesthetized and paralyzed cat during visual stimulation with moving gratings or flashing light spots of different size. In many cases, PGN cells showed an activity pattern which appeared like a mirror image of distinct periods of dLGN activity. Flashing spots evoked transient volleys of activity in PGN cells which increased in strength and shortened in latency with increasing size of the stimulus. These responses were temporally matched with inhibitory phases in the early part of visual responses in the dLGN. The spatio-temporal properties of the RFs were established by reverse correlation of the spike activity with the spatially random presentation of bright and dark spots within an array of 20 × 20 positions. Anticorrelated firing patterns of such kind could also be elicited as interocular inhibition with stimulation of the perigeniculate RF in the nondominant eye. Inversely correlated changes in spontaneous and visually induced activity were also visible during spontaneous changes in EEG pattern. With increasing synchronization of the EEG (predominance of delta-waves) the strength of geniculate visual responses declined while maintained perigeniculate activity increased. A weakened interocular and monocular inhibition of dLGN relay cells during visual stimulation of PGN RFs could be achieved with local reversible inactivation of PGN areas topographically matched with the dLGN recording sites. The results indicate that the PGN contributes to the state-dependent control of retino-geniculate transmission and to the monocular and interocular inhibitory processes that shape the visual responses in the dLGN.

Noise recorded in visual neurons, or variability in psychophysical experiments, may be quantified in terms of quantal fluctuations from an “equivalent” steady illumination. The conversion requires assumptions concerning how photon signals are pooled in space and time, i.e. how to pass from light fluxes to numbers of photon events relevant to the Poisson statistics describing signal/noise. It is usual to approximate real weighting profiles for the integration of photon events in space and time (the sensitivity distribution of the receptive field [RF] and the waveform of the impulse response [IR]) by sharp-bordered apertures of “complete,” equal-weight summation of events. Apertures based on signal-equivalence cannot provide noise-equivalence, however, because greater numbers of events summed with smaller weights (as in reality) have lower variances than smaller numbers summed with full weight. Thus sharp-bordered apertures are necessarily smaller if defined for noise- than for signal-equivalence. We here consider the difference for some commonly encountered RF and IR profiles. Summation areas, expressed as numbers of photoreceptors (cones or rods) contributing with equal weight, are denoted NS for signal and NN for noise; sharply delimited summation times are correspondingly denoted tS and tN. We show that the relation in space is NN = 0.5NS for the Gaussian distribution (e.g. for the RF center mechanism of retinal ganglion cells). For a photoreceptor in an electrically coupled network the difference is even larger, e.g., for rods in the toad retina NN = 0.2NS (NS = 13.7 rods and NN = 2.8 rods). In time, the relation is tN ≈ 0.7tS for realistic quantal response waveforms of photoreceptors. The surround input in a difference-of-Gaussians RF may either decrease or increase total noise, depending on the degree of correlation of center and surround noise. We introduce a third useful definition of sharp-bordered summation apertures: one that provides the same signal-to-noise ratio (SNR) for large-long stimuli as the real integration profiles. The SNR-equivalent summation area is N* = NS 2/NN and summation time t* = tS2/tN.

Mammalian horizontal cells are believed to be GABAergic because, in most species, they contain both GABA and glutamic acid decarboxylase (GAD), and their terminals are presynaptic to GABA receptors. In adult rabbit, however, GABA and GAD immunoreactivity have not been consistently demonstrated in horizontal cells, casting doubts on the assumption that they too are GABAergic. Here we demonstrate that all rabbit horizontal cell terminals—dendritic terminals of type A, and both dendritic and axonal terminals of type B—immunostain for one isoform of GAD, GAD67. In addition, we show that type A horizontal cell somas and primary dendrites in the visual streak (identified by their immunoreactivity to calbindin) are immunoreactive for the other GAD isoform, GAD65. Double-labeling experiments for GAD65 and GABA reveal that every cell that stains for GAD65 also stains for GABA. The presence of GAD67 in horizontal cell terminals suggests that rabbit horizontal cells are GABAergic. The segregation of the two GAD isoforms to different cell compartments suggests that GABA is released at different sites, possibly by two different mechanisms.

Photoreceptors in the isolated turtle retina of two species of turtle, Chelydra serpentina and Pseudemus scripta elegans, were penetrated with double-barrel electrodes. Physiological responses were recorded through one barrel and Neurobiotin tracer was injected from the other. Intracellular injection of Neurobiotin revealed patterns of tracer-coupled photoreceptors. Both the patterns of tracer coupling and the electrophysiology suggest a high degree of specificity of connections. Rods seem to be coupled only to rods and green and red cones seem to be coupled to cones of the same spectral type. Receptive-field profiles, measured with a thin, sharply focused slit of light, often had well-defined peaks and troughs in sensitivity. We have taken advantage of this observation and used the position of a peak in sensitivity to locate the position on the retina of a coupled cell. In one rod, it was possible to correlate physiological and morphological data and to show that the peaks in the physiological receptive field occurred at positions on the retina where there were dye-coupled cells. This provides direct evidence that gap junctions produce the physiological coupling between rods.

Observation of the spread of biotinylated or fluorescent tracers following injection into a single cell has become one of the most common methods of demonstrating the presence of gap junctions. Nevertheless, many of the fundamental features of tracer movement through gap junctions are still poorly understood. These include the relative roles of diffusion and iontophoretic current, and under what conditions the size of the stained mosaic will increase, asymptote, or decline. Additionally, the effect of variations in amount of tracer introduced, as produced by variation in electrode resistance following cell penetration, is not obvious. To examine these questions, Neurobiotin was microinjected into the two types of horizontal cell of the rabbit retina and visualized with streptavidin-Cy3. Images were digitally captured using a confocal microscope. The spatial distribution of Neurobiotin across the patches of coupled cells was measured. Adequate fits to the data were obtained by fitting to a model with terms for diffusion and amount of tracer injected. Results indicated that passive diffusion is the major source of tracer movement through gap junctions, whereas iontophoretic current played no role over the range tested. Fluorescent visualization, although slightly less sensitive than peroxidase reactions, produced staining intensities with a more useful dynamic range. The rate constants for movement of Neurobiotin between A-type horizontal cells was about ten times greater than that for B-type horizontal cells. Although direct extrapolation to ion conductances cannot be assumed, tracer movement can be used to give an estimate of relative coupling rates across cell types, retinal location, or modulation conditions in intact tissue.

We analyzed the relationship between eye movements and neuronal responses recorded from area MT in alert monkeys trained to maintain visual fixation during the presentation of moving patterns. The monkeys made small saccades which moved the eyes with velocities that spanned the sensitivity range of MT neurons. The saccades evoked changes in the neuronal response that depended upon (1) the level of stimulus-evoked activity amidst which the saccade occurred and (2) the direction of the saccade relative to the preferred direction of the neuron. Most notably, saccades were able to suppress stimulus-evoked activity when they caused retinal image flow that opposed the neuron's preference and were able to elicit a response or enhance weak activity when they caused flow in the neuron's preferred direction. On average, the disturbance lasted 40 ms beginning about 40 ms following saccade onset. Using these parameters, we simulated synthetic spike trains from an imaginary pair of similarly tuned neurons and determined that the interneuronal correlation due to saccades should be negligible at all but the lowest ongoing firing rates. This conclusion was supported from our data by the observation that response variance for single MT spike trains was not measurably reduced during periods of stable gaze compared to periods when eye movement exceeded a stability criterion (0.1 deg during 0.5 s). While the intrusions caused by saccades are too short-lived and infrequent to account for the variability of MT neuronal response (counter to the finding in V1 of Gur et al., 1997), the clear directional signal that they carry in area MT suggests that motion perception is not blocked during saccades by suppression at early stages in the visual pathway.