An important mechanism for the discrimination of direction of motion in the retina is a spatially asymmetric inhibition. This inhibition has been postulated to operate either as a subtraction, like in difference-of-Gaussians' models, or as a division, like in shunting-inhibition models of directional selectivity. The latter, but not the former, is nonlinear. This raises the question of whether the inhibitory mechanism involved in directional selectivity is nonlinear. To investigate this issue, we studied the linearity of the contrast dependence of the extracellularly recorded responses to apparent motions in ON-OFF directionally selective ganglion cells of the rabbit retina. The results show that the inhibition underlying directional selectivity is nonlinear and fits shunting-inhibition models well. Other biophysical mechanisms that might account for the type of nonlinearity observed in the data are also considered.

The echidna, Tachyglossus aculeatus, a monotreme mammal, is thought to possess an all-rod retina (O' Day, 1952). This study provides anatomical evidence for the presence of cone-like photoreceptors in the retina of the echidna. The cones, which constitute 10–15% of the photoreceptors, have all of the ultrastructural characteristics previously shown in the cones of placental mammals, and, like the cones of other animals (Blanks & Johnson, 1984), they bind peanut agglutinin. Unlike the cones of another monotreme, the platypus, the cones of the echidna retina do not possess oil droplets. Twin cones, pairs of cones in which there is no obvious difference in the size, shape, or ultrastructural features of the members of a pair, are common. The density of cones varies from 9000 cells/mm2 in the superior periphery to 22,000 cells/mm2 in the central retina. Nearest-neighbor analysis suggests that the cone mosaic in the echidna retina results from the presence of single and twin cones in a relatively regular array.

The light-adaptation properties of goldfish photoreceptor mechanisms were examined using Stiles' two-color threshold technique. Threshold vs. background intensity (TVI) curves were determined for isolated cone and rod mechanisms using the heart-rate conditioning technique. The principal aim of this study was to compare the light-adaptation properties of the ultraviolet (UV)-sensitive cone mechanism to the other receptor mechanisms of goldfish. This examination revealed several striking functional differences: (1) The UV-sensitive cone mechanism threshold vs. background intensity (TVI) exhibited a slope of 0.65 (compared to the approximate 1.0 for the other cone mechanisms on a log/log plot) and thus was not in accordance with Weber's law. This may in part be related to the intrusion of the blue-sensitive mechanism at the upper radiance range. (2) The operation of the UV-sensitive cone mechanism was limited to intermediate intensities (i.e. not very dim or bright). (3) The UV-sensitive cone mechanism exhibited a Weber fraction or luminance contrast threshold of 0.316 that was approximately six times larger than the other cone mechanisms but comparable to the rod mechanism. This indicates that the UV-sensitive cone mechanism performs relatively poorly in terms of brightness contrast detection.

In the mammalian retina, the dendritic arbors of cholinergic amacrine neurons have a unique starburst shape; these arbors lie in narrow sublaminae within the inner plexiform layer, where they provide input to a wide variety of ganglion cell types. Immunocytochemistry has been used to identify cholinergic cells in one poikilotherm, the goldfish (Tumosa et al., 1984), but there has been no description of the detailed dendritic morphology of these cells in the lower vertebrates. In the present study, cholinergic neurons have been characterized, by immunocytochemistry and dye filling, in the retina of the Pacific Coast dogfish, Squalus acanthias.

The inner nuclear layer contained two populations of choline acetyltransferase-immunoreactive amacrine cells, of different sizes (average soma diameters 12.2 vs. 16.3 μm); 70% of the immunoreactive cells were of the smaller type. Cholinergic dendrites from these two cell populations formed two narrow strata within the inner plexiform layer, at depths of 14% and 31%. In the ganglion cell layer, 40% of the cells were immunoreactive for choline acetyltransferase (ChAT); these cells were very homogeneous in size, had an average diameter of 12.6 μm, and appeared to represent a single class of cholinergic amacrine. The dendrites of these cells formed a single, narrow stratum within the inner plexiform layer, at a depth of 59%.

In living preparations, the smallest cell bodies in the ganglion cell layer were filled iontophoretically with Lucifer Yellow, under microscopic control. Such cells invariably had a stellate morphology; in many cases, they appeared quite similar to the starburst cholinergic amacrine cells described in rabbit and rat (Vaney, 1984; Voigt, 1986). Although double-label experiments failed to demonstrate ChAT immunoreactivity in specific dye-filled cells, the dendritic arbors of individual dye-filled stellate dogfish amacrines did co-stratify precisely with the proximal ChAT-immunoreactive sublamina of the inner plexiform layer. In addition, dye injection and ChAT immunocytochemistry appeared to label the same population of dogfish neurons, as suggested by the close structural similarity, and similar numerical proportion, of the cells identified with these two techniques.

Similarities between the displaced cholinergic amacrine neurons of the dogfish retina, and the cholinergic, “starburst” amacrine neurons of the rabbit retina, are discussed.

The density recovery profile is a plot of the spatial density of a set of points as a function of the distance of each of those points from all the others. It is based upon a two-dimensional point autocorrelogram. If the points are randomly distributed, then the profile is flat, with a value equal to the mean spatial density. Thus, any deviation from this value indicates that the presence of the object represented by the point alters the probability of encountering nearby objects of the same set. Increased value near an object indicates clustering, decreased value near an object indicates anticlustering. The method appears to be unique in its ability to provide quantitative measures of the anticlustered state. Two examples are presented. The first is based upon a sample of the distribution of the somata of starburst amacrine cells in the macaque retina; the second is based upon the distribution of the terminal enlargements on the dendrites of a single macaque ganglion cell that projects to the superior colliculus. In both cases, the density recovery profile is initially lower than the mean density, and increases up to the plateau at the value of the mean density. Two useful measures can be derived from this profile: an intensive parameter termed the effective radius, which quantifies the extent of the region of decreased probability and is insensitive to random undersampling of the underlying distribution, and an extensive parameter termed the packing factor, which quantifies the degree of packing possible for a given effective radius, and is insensitive to scaling. An extension of this method, applicable to correlations between two superimposed distributions, and based upon a two-dimensional point cross-correlogram, is also described.

The cholinergic amacrine cells of the rabbit retina form two mosaics placed symmetrically on either side of the inner plexiform layer. Recently, these cells have been reported to contain immunocytochemical markers for GABA. In this paper, we labeled the cholinergic cells with DAPI, then incubated the retina in [3H]-muscimol, a neuronal marker for GABA. Subsequently, we converted the DAPI fluorescence of the displaced cholinergic matrix to an opaque product by photooxidation in the presence of DAB. Autoradiography showed that all of the displaced cholinergic amacrine cells were labeled with ]3H]-muscimol, thus confirming the immunocytochemical results. The cholinergic cells account for approximately 80% of the cells in the ganglion cell layer which take up ]3H]-muscimol.

Using patch-clamp techniques, properties of cGMP-activated channel were studied at a single-channel level in order to examine (1) whether any differences are recognized between the cGMP-activated channels of rods and cones in the same animal species, and (2) whether the channel properties of the same photoreceptor class differ in different animal species. Experiments were performed on inside-out membrane patches excised from outer segments of rods and morphological subtypes of cones in the carp retina. Single-channel activities could be recorded when the patches were perfused with low concentrations of cGMP (<10 μM). Throughout five morphological subtypes of cones and rod, single-channel currents showed no significant rectification at membrane hyperpolarization in a low divalent cation solution, and single-channel conductances were almost the same: 13.8 ± 0.2 pS (mean ± s.e.m., n = 23) in cones and 12.7 ± 0.8 pS (n = 3) in rods. These values were significantly smaller than that reported in catfish cones (about 50 pS), and that in rods of the toad and the tiger salamander (about 25 pS). In rods and all subtypes of cones of the carp, open durations of cGMP activated channels were brief. In addition, kinetic parameters of channel openings and closings showed no differences throughout all subtypes of cones and rod.

The vertebrate retina processes visual information in parallel neural pathways known as the ON and OFF pathways. These pathways encode increments and decrements of light independently as excitatory responses. We examined the photopic spectral response of ON and OFF mechanisms in goldfish by measuring the sensitivity of optic nerve responses to the onset and termination of stimuli of various wavelengths. Using various adapting backgrounds, we found that the ON and OFF responses have different spectral sensitivities. The weighting of the cone inputs to the responses was estimated by an algebraic summation model. This model suggests that for the ON response, input from S-cones is stronger and more independent than for the OFF response, and M- and L-cones show stronger antagonism in the ON response than in the OFF response. The OFF response probably receives input from all cone types, but spectral antagonism is weak and its dominant input is from L-cones.

A subpopulation of cells developing in dissociated neuronal cultures prepared from 8-day-old embryonic chick retinae can be identified as putative in vitro counterparts of GABAergic amacrine cells by immunocytochemical and autoradiographic markers and by their electrophysiological responses to transmitter agonists. In the present study, transmitter-gated conductances expressed by these neurons were examined using the whole-cell patch-clamp technique. At negative holding potentials, the excitatory amino acid agonists N-methyl-D-aspartate (NMDA), kainate quisqualate, and glutamate induced inward currents with reversal potentials close to 0 mV in most of the cells selected for recording. NMDA-evoked responses were selectively blocked by the noncompetitive inhibitor MK 801 and by Mg2+ (in a voltage-dependent manner) and were potentiated in the presence of submicromolar concentrations of glycine. Glutamate apparently interacted with both NMDA and non-NMDA type receptors. All cells tested responded to the inhibitory transmitters GABA and glycine. Both inhibitory agonists could be shown to activate chloride conductances. Responses to GABA and glycine were specifically inhibited in the presence of bicuculline and strychnine, respectively. Thus, GABAergic neurons in retinal cultures express at least two different excitatory amino acid receptors–NMDA and non-NMDA–and two different inhibitory amino acid receptors–the GABAA and the glycine receptor. The results demonstrate the ability of the cultured neurons to develop an apparently mature phenotype and contribute to the understanding of the functional properties of GABAergic amacrine cells in the vertebrate retina.

We have compared the responses of simple cells to laterally moving sinusoidal gratings and to stationary temporally-modulated gratings. From the amplitudes and temporal phases of the responses to stationary gratings of different spatial phases, it should be possible to predict the preferred direction of movement, the amplitudes of the responses to gratings moving in the preferred and nonpreferred directions and, thence, the degree of directional preference (Reid et al., 1987). The preferred direction can be predicted reliably. However, the magnitude of the directional preference cannot be predicted, since the measured amplitude of the response in the nonpreferred direction of movement is very much less than that predicted by a linear theory. Nonlinearities in the relationship between response amplitude and contrast may contribute to the failure of the predictions, but this contribution is small. We conclude that the magnitude of the directional preference seems to be determined predominantly by nonlinear suppression of the response in the nonpreferred direction of movement.

This paper reviews evidence that the superior colliculus (SC) of the midbrain represents visual direction and certain aspects of saccadic eye movements in the distribution of activity across a population of cells. Accurate and precise eye movements appear to be mediated, in part at least, by cells of the SC that have large sensory receptive fields and/or discharge in association with a range of saccades. This implies that visual points or saccade targets are represented by patches rather than points of activity in the SC. Perturbation of the pattern of collicular discharge by focal inactivation modifies saccade amplitude and direction in a way consistent with distributed coding. Several models have been advanced to explain how such a code might be implemented in the colliculus. Evidence related to these hypotheses is examined and continuing uncertainties are identified.

We propose a network model that accounts for the kinetic depth in structure from motion phenomena. Using plausible neural mechanisms, the model accounts for (1) fluctuations in perception when viewing a simple kinetic depth stimulus, (2) disambiguation of this stimulus with stereoscopic information, and (3) subsequent bias of the percept of this stimulus following stereoscopic adaptation. The model comprises two levels: a layer of monocular directionally selective motion detectors that provide input to a second layer of disparity- selective and direction-selective binocular mechanisms. The network of facilitatory and inhibitory connections between binocular mechanisms gives rise to fluctuations in network activity that mimic the fluctuations in perception of kinetic depth in the absence of disparity information. The results of a psychophysical experiment are consistent with the nature of the proposed interactions.

In the retina of a variety of vertebrate species, a monoclonal antibody against protein kinase C (PKC) has been shown to label preferentially bipolar cells. Although the functional consequences of PKC activation in these cells is yet to be revealed, the present study was motivated in part by the possibility that the antibody might be used as a selective marker for examining the development of bipolar cells in the rat retina. Here, the developmental pattern and the dynamic changes of retinal cells expressing PKC-like immunoreactivity (PKC-LI) were studied and analyzed throughout postnatal life until adulthood. Upon its initial detection by immunohistochemistry on postnatal day (PD)-10, faint PKC-LI was limited to the central region of the retina, labeling cell bodies located at the scleral margin of the inner nuclear layer (INL) adjacent to the outer plexiform layer (OPL). On subsequent days, PKC-LI spread progressively to the peripheral retina and axon terminal bulbs at the vitreal margin of theinner plexiform layer (IPL) began showing the first signs of immunoreactive labeling. Not until PD-15, the time of eye opening, did PKC-LI in these cells increase to the extent such that their thin axons were immunoreactive. Each of these axons traversed the entire thickness of the IPL and divided into two or three short branches before ending as enlarged terminal bulbs. The morphology and the location of PKC-LI cells in both the developing and adult retina observed in our study are consistent with them being rod bipolar cells. By the end of the fourth postnatal week, the rod bipolar cells appeared mature, resembling those found in the adult. Overall, more dynamic changes occurred at the axon terminal bulbs than at the cell bodies during the maturational process of rod bipolar cells. Interestingly, PKIC-LI was expressed precociusly in these cells when rat pups were reared in complete darkness starting from the day of birth.

Selective labeling of intravitreal Procion yellow dye by presumed blue-sensitive cone photoreceptors has been demonstrated in primate retina. To determine whether Procion yellow is selective for this cone type in an unrelated vertebrate species, labeling by this dye was studied in goldfish retina, where cone pigment type can be directly inferred from photoreceptor morphology. At low vitreal concentrations of the dye (<0.4%), only cone outer segments were labeled. At vitreal concentrations of 0.4–0.5%, the inner segments of short-double cones and a subset of long single cones (presumed green-sensitive cones) were selectively stained. At still higher vitreal concentrations (0.6–0.7%), the inner segments of short-single cones and miniature short-single cones (presumed blue-sensitive cones) showed evidence of Procion label, but were not as heavily labeled. The inner segments of long-double cones and a subset of long-single cones (presumed red-sensitive cones) did not label at any of these concentrations. These results show that Procion yellow is not a selective marker for blue-sensitive cones in the goldfish retina. In addition, stained rod and cone nuclei were observed at each dye concentration, including those concentrations at which no inner segments were labeled.

We have explored the visuotopic organization of the territory surrounding the middle suprasylvian sulcus (MSS) of cat cerebral cortex by electrophysiological mapping, and by tracing the topography of its cortical and subcortical connections using wheatgerm-agglutinin horseradish peroxidase (WGA-HRP). Observations from the two approaches were concordant, and confirmed the presence of two separate visual areas in the MSS that approximate, but do not exactly correspond, to the location and internal organization of the posterior medial and posterior lateral lateral suprasylvian (PMLS, PLLS) areas of Palmer et al. (1978).

We define as part of the lateral suprasylvian (LS) area the territory on the medial bank and caudal end of the lateral bank of the MSS that receives a topographically organized projection from the region of area 17 representing the lower visual quadrant. This territory is connected with other structures that are themselves striate-recipient (cortical areas 18 and 19, and the lateral division of the lateral posterior (LPI) nucleus), and with a variety of nuclei that receive direct retinal input, such as the C-laminae of the LGd, the medial interlaminar nucleus (MIN), and the superficial layers of the superior colliculus (SC). Its connections with the LP1, LGd, MIN, and SC correspond topographically with the input from area 17. Revised maps of area LS were produced from the physiological and connectional data: its rostral border is formed by a representation of lower visual elevations with the horizontal meridian represented caudally, and its lateral border is formed by the vertical meridian; area LS shares a representation of the center of gaze with the visual area of the lateral bank at its caudal end.

The adjacent lateral bank area has larger receptive fields than area LS, and very different connectivity. It receives no input from area 17 and little input from striate-recipient structures, including area LS, but instead is connected to more remote extrastriate visual areas, such as the anterior ectosylvian visual (AEV) area in insular cortex, and to zones of the thalamus in receipt of tectal input (LPm and the lateromedial-suprageniculate nuclear complex). According to both mapping approaches, the lateral bank area contains representations of both the upper and lower visual quadrants but a rather limited degree of visuotopic order. We refer to it as the posterior ectosylvian visual (PEV) area, because it appears to be functionally and connectionally dissociated from area LS, but is possibly a functional antecedent of area AEV.

We investigated whether regenerating mature axons recapitulate embryonic features essential to successful reconnectivity within the injured nervous system. Strips from embryonic and adult chick retinae were cultured, and outgrowing axons were examined morphometrically and immunohistochemically. In addition, the target-recognition properties of adult neurites were analyzed. Regenerating adult axons elongate on a poly-L-lysine/laminin substratum with a speed about one order of magnitude slower than that of embryonic axons. Morphologically, adult axonal tips differ dramatically from embryonic growth cones in that they possess only filopodial extensions whereas embryonic growth cones possess both lamellipodial and filopodial processes. Both embryonic and adult neurites express the growth-associated protein GAP-43. When cultured on alternating stripes of anterior and posterior embryonic tectal membranes, both adult and embryonic retinal axons distinguish between the two membrane preparations. Our results demonstrate that during axonal regeneration the mature neurons express embryonic properties that are involved in the recognition of tectal positional cues.

The ultrastructure of the pigeon and quail ventral lateral geniculate nucleus was analyzed with standard electron microscopy and horseradish peroxidase tracing of its retinal and tectal afferents. Six types of neurons were distinguished: two large, two medium-sized, and two small types.

Using the whole-cell patch clamp technique, we have examined the voltage-gated currents present in adult chicken cone cells. When calcium and calcium-gated currents are blocked, hyperpolarizing voltage steps elicit slowly increasing inward currents as has been shown for photoreceptors in other species. Unlike the case for other species, chicken cones appear to lack the inward-rectifying cationic current Ih that contributes to the shaping of the photovoltage. Instead of Ih, these cones possess an anionic inward-rectifying current that in kinetics, activation range and probably function is remarkably similar to Ih. This anion channel is unusual in that both nitrate and acetate are more permeant than chloride ions.

Both kainic acid (KA) and N-methyl-d-aspartatic acid (NMDA) depolarize luminosity-type horizontal cells (L-type H cells) in normal turtle retina. The presence of both NMDA and non-NMDA receptors for excitatory amino acids (EAAs) on these cells was highlighted by an unusual effect of the noncompetitive NMDA-antagonist, MK-801. In retinas that had been exposed to MK-801, the action of NMDA was irreversibly altered to one of hyperpolarization, while the depolarizing effect of KA was unaltered. The aim of the present study was to further characterize these receptors on L-type H cells and to extend the investigation to color-opponent H cells (C-type H cells). Intracellular recording was used to study the effects of KA, NMDA, MK-801, the competitive NMDA antagonists, 2-amino-5-phosphonopentanoic acid (AP5) and 2-amino-7-phosphonoheptanoic acid (AP7), and the nonspecific EAA antagonist, kynurenic acid (KYN) on the light responses of L-type and C-type H cells in turtle retina. The effects of combinations of these drugs were also studied. In L-type H cells the agonists caused depolarization and loss of light response, KYN caused hyperpolarization and loss of light response, and MK-801, AP5 or AP7 had no direct effect. However, application of NMDA following MK-801, AP5 or AP7, but not KYN, caused hyperpolarization and loss of light response. The depolarizing effect of KA was unaltered by these antagonists. These data confirm the presence of an unusual NMDA receptor on L-type H cells. In the case of red\green C-type H cells, application of KA caused loss of responses to both red and green light, with loss of green responses preceding loss of red responses. NMDA initially removed responses to both red and green light. The most striking effect of NMDA was seen during early washout where the responses to red were reversed (hyperpolarizing). These responses eventually recovered their normal polarity. These results suggest that the depolarizing response of C-type H cells to red light is mediated by L-type H cells, but not via inhibition of the excitatory input from green cones to C-type H cells.