Axon-bearing amacrine cells in mammalian retinae are encountered relatively infrequently during electrophysiological investigations, and thus very little is known about their physiological properties. Patch-clamp electrodes were used to record light responses from two axon-bearing amacrine cells in flat-mounted, dark-adapted rabbit retina. The recorded cells were stained, and the morphology visualized. Both cells were capable of generating action potentials. In one case, a linear relationship between mean depolarization and action-potential frequency was demonstrated. The cells had a proximal dendritic arbor and a morphologically distinct, much larger axon terminal system. The receptive field of the center response was coextensive with the dendritic arbor, and thus also much smaller than the axon terminal system. The center response was suppressed by activation of an inhibitory surround. Both cells responded to center illumination with an inward current which became more transient as the size of the illuminating spot was increased. It is suggested that axon-bearing amacrine cells receive input over a receptive field defined by the dendritic arbor, and distribute their output over a much more extensive axon terminal system, most probably via action potentials.

Adaptation of visual cortical cells' responses is observed following repeated presentation of grating stimuli. Grating adaptation is believed to exist only at the cortical level. The purpose of this study was to see if grating adaptation also occurs in the lateral geniculate nucleus. We studied the responses of 164 relay cells in layer A and A1 of the dorsal lateral geniculate nucleus (LGNd) to grating stimuli. Normal cats, as well as cats in which visual cortex was ablated, were studied. Adaptation was investigated using repeated presentation of gratings of different contrasts and orientations. The results showed the following: (1) Grating adaptation reduced the responses of 46% of the LGNd cells recorded. The responses normally decreased within 30 s and then stabilized. However, there was heterogeneity in the effects observed. About 38% of the cells studied were not affected by the adapting gratings. Some cells (16%) showed facilitation rather than habituation of their responses to test stimuli. (2) There was no significant difference between X and Y cells in their susceptibility to adaptation. This suggests that grating adaptation is a general property, independent of cell type. (3) The contrast-response curves of 57% of the LGNd cells studied shifted down after exposure to high-contrast adapting gratings. (4) Adapting gratings of the cells' preferred orientation decreased the orientation sensitivity of 56% of the orientation-sensitive cells. Adapting gratings at the nonpreferred orientation did not affect orientation sensitivity. (5) Prolonged grating adaptation also reduced the responses of 49% of the LGNd cells after inactivation of cortical inputs to the LGNd.

Although the neural location of the plaid motion coherence process is not precisely known, the middle temporal (MT) cortical area has been proposed as a likely candidate. This claim rests largely on the neurophysiological findings showing that in response to plaid stimuli, a subgroup of cells in area MT responds to the pattern direction, whereas cells in area V1 respond only to the directions of the component gratings. In Experiment 1, we report that the coherent motion of a plaid pattern can be completely abolished following adaptation to a grating which moves in the plaid direction and has the same spatial period as the plaid features (the so-called “blobs”). Interestingly, we find this phenomenon is monocular: monocular adaptation destroys plaid coherence in the exposed eye but leaves it unaffected in the other eye. Experiment 2 demonstrates that adaptation to a purely binocular (dichoptic) grating does not affect perceived plaid coherence. These data suggest several conclusions: (1) that the mechanism determining plaid coherence responds to the motion of plaid features, (2) that the coherence mechanism is monocular, and thus (3), that it is probably located at a relatively low level in the visual system and peripherally to the binocular mechanisms commonly presumed to underlie two-dimensional (2-D) motion perception. Experiment 3 examines the spatial tuning of the monocular coherence mechanism and our results suggest it is broadly tuned with a preference for lower spatial frequencies. In Experiment 4, we examine whether perceived plaid direction is determined by the motion of the grating components or the features. Our data strongly support a feature-based model.

We have examined the retinotopy in the superior colliculus of pigmented and albino ferrets using both anatomical and electrophysiological methods. While the distribution of contralaterally projecting retinotectal ganglion cells is characterized by the presence of an area centralis superimposed on a visual streak in both strains, the ipsilateral projection from temporal hemiretina is strongly reduced in albinos. In spite of the significantly altered retinotectal projection pattern, the collicular visual field map in the albino ferret reveals the same characteristics as in the pigmented animal with a strongly enlarged representation of the center of visual space. An areal comparison between retinotectal ganglion cell distribution and collicular areal magnification shows that the increase in areal magnification factor between the periphery and the representation of the central visual hemifield exceeds the corresponding increase in retinal ganglion cell density between peripheral retina and area centralis by a factor of three in pigmented and a factor of four in albino ferrets. The areal magnification factor of the representation of the retinal visual streak does not exceed the increase in retinotectal ganglion cell density. Thus, our results suggest that the representation of visual space in the superior colliculus of albino and pigmented ferrets does not simply follow the retinotectal ganglion cell density, but that there is an enhanced representation of the frontal central visual field. The possibility is discussed that the collicular visual field map may be determined either by both retinotectal and corticotectal projections or by the colliculus' intrinsic structure.

Occipital visual evoked potentials (VEPs) were recorded in response to low-contrast, low spatial-frequency chromatic, and achromatic gratings. Fast Fourier Transform (FFT) and time-domain analysis were used to reveal differences in harmonic content and amplitude of chromatic and achromatic response components over a wide range of temporal frequencies. The chromatic ON/OFF VHP is dominated by the fundamental component indicating that onset and offset responses are different. This type of response is typical of neurons with sustained type response characteristics. Conversely, the achromatic onset VEP contains a predominant second harmonic component in addition to the fundamental. This similarity between onset and offset responses suggests that transient mechanisms are responsible for the generation of achromatic components. Frequency analysis of VEPs elicited by phase-reversing stimuli reveals that all of the response energy is concentrated at the second harmonic of the stimulating frequency. The magnitude of the second harmonic component is maximal for achromatic stimuli and undergoes a distinct minimum for isoluminant, chromatic stimuli. This behavior indicates that under the stimulus conditions used, magnocellular neurons with transient characteristics dominate the reversal VEP.

Electrical stimulation in pigeons of the input from the medial subdivision of the nucleus of Edinger-Westphal (EWM) to the choroidal neurons of the ipsilateral ciliary ganglion, which themselves have input to the choroidal blood vessels of the ipsilateral eye, increases choroidal blood flow (ChBF). Since the EWM receives input from the contralateral suprachiasmatic nucleus (SCN), which in turn receives contralateral retinal input, the present study sought to determine if activation of the SCN by microstimulation or by retinal illumination of the contralateral eye would also yield increases in ChBF in that same eye. Using laser Doppler flowmetry (LDF) to measure ChBF, we found that electrical activation of the contralateral SCN by 100-Hz anodal pulse trains yielded increases in ChBF that were stimulus related and proportional to the stimulating current. These increases in ChBF elicited by the SCN stimulation were accompanied by increases in choroidal volume (vasodilation), but not by increases in systemic blood pressure. Furthermore, the increases could be blocked reversibly by lidocaine injection into the EWM. These results suggest that the increases in ChBF in the eye contralateral to the SCN stimulation were specifically mediated by the SCN-EWM pathway. Retinal illumination with a fiber optic light source was also found to increase ChBF in the illuminated eye, and these effects too could be blocked reversibly with lidocaine injection into the EWM or permanently by the EWM lesion. Control studies confirmed that the light-elicited increases were mediated by increases in choroidal volume (i.e. vasodilation), were not accompanied by systemic blood pressure increases, and were not artifactually generated by transocular illumination of the LDF probe. Thus, the SCN-EWM circuit may be involved in regulating ChBF in response to the level of retinal illumination and/or the visual patterns falling on the retina.

This paper describes changes in the appearance and distribution of microglia in postnatal cat retina as demonstrated by a new antibody, H386F. This fractionated IgM antibody was created via an intrasplenic immunization of a single BALB/C mouse with about 2–3 × 105 large, whole cells isolated from 46 minced cat retinae. To confirm that the labeled cells are microglia, the staining properties of H386F were compared with those of four commercially available antibodies, OX-33, OX-41, OX-42, and ED-1, that have been used by others to distinguish between microglia and other cells in rat brain. These experiments show that H386F is the only antibody of the five to label only microglia in both the cat retina and hippocampus.

The role of N-methyl-D-aspartate (NMDA) receptors in the mediation of cortical inputs to visual neurones in the superficial layers of the superior colliculus (SSC) has been investigated. Extracellular recording with iontophoresis in the SSC of cortically intact cats has demonstrated that visual responses of most neurones were reduced by iontophoretic application of the NMDA receptor antagonist D-2-amino-5-phosphonopentanoate (APS). Following inactivation of areas 17 and 18 of the visual cortex with topical lignocaine, the visual responses of 11, previously AP5-sensitive, neurones were no longer reduced by APS ejection. The cortical input is generally assumed to influence the directional responses of visual neurones in SSC. However, detailed study of the directional bias showed that the degree of directional tuning in SSC neurones was similar to that of retinal ganglion cells, as previously described by others. Moreover, inactivation of the visual cortex with topical lignocaine did not alter the directional bias of SSC neurones. Likewise, the directional bias of SSC neurones was not reduced by iontophoretic ejection of APS in the SSC. These data imply that NMDA receptors have an important role in mediating the cortical input to the SSC. However, cortical input does not appear to be responsible for conferring directional bias upon SSC neurones' visual responsiveness.

Endogenous opioid systems (i.e. opioid peptides and opioid receptors) modulate developmental events in the neonatal mammalian retina. In the present study, the mRNA encoding preproenkephalin A (PPE), the prohormone for the opioid growth factor (OGF), [Met5]-enkephalin, was studied in the developing and the adult retinas of rats. Northern analysis indicated the presence of a 1.4-kb message in the developing and adult retinas corresponding to rat PPE mRNA. Quantitation showed that PPE message was present on postnatal day 1 at 5% of the adult level, and increased during development until the adult quantity was reached by postnatal day 27. In situ hybridization experiments first detected the presence of PPE mRNA in retinal tissues during late gestation. In late prenatal and neonatal retinas, PPE message was associated with areas of the developing retina containing proliferating neuroblasts and postmitotic cells. Later in development, message appeared to be located primarily within the inner retina, with abundant PPE mRNA associated with putative horizontal cells of the inner nuclear layer (INL). The adult retina showed a similar pattern of PPE gene expression in the cells of the INL. These findings document that the gene expression in the retina for PPE begins in the fetus, continues during retinal development, and coincides with the presence of a PPE mRNA derivative ([Met5]-enkephalin) that regulates DNA synthesis during retinal ontogeny. Our results are also the first to show the presence of PPE message in the adult mammalian retina, suggesting transcription of an opioid gene in the mature visual system.

Previous studies have shown that directionally selective (DS) retinal ganglion cells cannot only discriminate the direction of a moving object but they can also discriminate the sequence of two flashes of light at neighboring locations in the visual field: that is, the cells elicit a DS response to both real and apparent motion. This study examines whether a DS response can be elicited in DS ganglion cells by simply stimulating two neighboring areas of the retina with high external K+. Extracellular recordings were made from ON-OFF DS ganglion cells in superfused rabbit retinas, and the responses of these cells to focal applications of 100 mM KCl to the vitreal surface of the retina were measured. All cells produced a burst of spikes (typically lasting 50–200 ms) when a short pulse (10–50 ms duration) of KCl was ejected from the tip of a micropipette that was placed within the cell's receptive field. When KCl was ejected successively from the tips of two micropipettes that were aligned along the preferred-null axis of a cell, sequence-dependent responses were observed. The response to the second micropipette was suppressed when mimicking motion in the cell's null direction, whereas an enhancement during apparent motion in the opposite direction frequently occurred. Sequence discrimination in these cells was eliminated by the GABA antagonist picrotoxin and by the Ca2+-channel blocker ω-conotoxin MVIIC, two drugs that are known to abolish directional selectivity in these ganglion cells. The spatiotemporal properties of the K+-evoked sequence-dependent responses are described and compared with previous findings on apparent motion responses of ON-OFF DS ganglion cells.

The effects of changing retinal ganglion cell (RGC) density and availability of presynaptic sites on the development of RGC dendritic arbor in the developing chick retina were contrasted. Visual form deprivation was used to induce ocular enlargement and expanded retinal area resulting in a 20–30% decrease in RGC density. In these retinas, RGC dendritic arbors increased in a compensatory manner to maintain the inner nuclear layer to RGC convergence ratio in a way that is consistent with simple stretching; RGC dendritic arbors become larger with increased branch lengths, but without change in the total number of branches. In the second manipulation, partial optic nerve section was used to produce areas of RGC depletion of approximately 60% in the central retina. This reduction in density is comparable to the density of locations in the normal peripheral retina. In RGC-depleted retinas, dendritic arbor areas of RGCs in the central retina grow to match the size of normal peripheral arbors. In contrast to the expanded case, two measures of intrinsic arbor structure are changed in RGC-depleted retinas; the branch density of RGC dendrites is greater, and the relative areas of the two arbors of bistratified cells are altered. We discuss the potential roles of retinal growth, local RGC density, and availability of presynaptic terminals in the developmental control of RGC dendritic arbor.

The morphology of horizontal cells in ox, sheep, and pig retinae as observed after Lucifer Yellow injections are described and compared with the descriptions of Golgi-stained cells by Ramón y Cajal (1893). Horizontal cells in the retinae of less domesticated species, wild pig, fallow and sika deer, mouflon, and aurochs were also examined. All these retinae have two types of horizontal cell; their morphologies are in common, although with some familial differences. Their basic appearance is as Cajal described; except in one important respect, a single axon-like process could not be identified on the external horizontal cells. It is concluded that external horizontal cells of artiodactyls correspond to the axonless (A-type) cells of other mammals. Cajal's internal horizontal cells have a single axon which contacts rods. This type corresponds to the B-type cells of other mammalian retinae. Artiodactyl A- and B-type horizontal cells differ from those of many other mammals in that the B-type dendritic tree is robust and the A-type dendritic tree is delicate. Historically, this morphological difference between orders of mammals has led to some confusion. The comparisons presented here suggest that the morphological types of primate horizontal cells can be integrated into a general mammalian classification.

We have developed two brain slice preparations for studying tectofugal visual pathways in the chick: conventional, 400-μm slices (“thin slices”), and “thick slices” which encompass the rostral pole of the optic tectum and the contralateral optic nerve. Stimulation was delivered with a bipolar electrode positioned in stratum opticum in thin slices and in the contralateral optic nerve in thick slices. While the latter preparation provided a means of exclusively and unambiguously activating retinal afferents, several lines of evidence also indicated that the evoked field potentials in thin slices were chiefly consequent to retinal afferent excitation: (1) the similarity of evoked field potentials in thin slices to those in thick slice preparations; (2) their precise localization in retinorecipient layers as shown by prelabeling from retina with FITC-coupled cholera toxin; (3) transmission delays appropriate for retinal afferents as established with the thick slice preparation; (4) patterns of labeled afferents resulting from applications of Dil crystals to slices fixed after recording; and (5) the similarity in transmitter pharmacology between thin and thick slice preparations. Pharmacological manipulations carried out with bath-applied antagonists indicated that glutamate is the principal retinotectal transmitter. The broadly active glutamate receptor blocker, kynurenic acid, reversibly eliminated the postsynaptic component of the field potential as confirmed with 0 Ca2+ saline. A complete block was also effected by the non-NMDA antagonists CNQX and DNQX. The specific NMDA antagonist, APS, caused a smaller and variable reduction in response amplitude. The GABA antagonist, bicuculline, caused a prolongation of the monosynaptic field epsp in retinorecipient layers and an enhancement of the long-latency, negative wave in cellular layers below, supporting a late, excitation-limiting role for this inhibitory transmitter.

Vertical Slices of postnatal day 6 (P6) rat retina were cut and cultured using the roller-tube technique. The organotypic differentiation during a culture period of up to 30 days has been described in a previous study (Feigenspan et al., 1993a). Here we concentrated on the synaptic organization in the retinal slice culture. Electron microscopy revealed the presence of ribbon synapses in the outer plexiform layer and conventional and ribbon syanpses in the inner plexiform layer. Immunofluroscence with antibodies that recognize specific subunits of GABAA or glycine receptors revealed a punctuate distribution of the receptors. They were aggregated in “hot spots” that correspond to a concentration of receptors at postsynaptic sites. Different isoforms of GABAA and glycine receptors occured in the slice cultures. The experiments show that there is a differentiation of synapses and a diversity of transmitter receptors in the slice cultures that is comparable to the in vivo retina.

Using a combination of retrograde horseradish peroxidase (HRP) labelling, silver staining, and electron microscopy, we have assessed the relationship between retinal ganglion cell soma size and axon diameter in the adult ferret (Mustela putorius furo). Retinal ganglion cells were labelled following injections of HRP into the lateral geniculate nucleus (LGN), superior colliculus (SC), or LGN+SC. The soma size distributions following LGN, SC, or LGN+SC injections were all unimodal showing considerable overlap between different cell classes. This was confirmed for alpha cells, identified on the basis of dendritic filling or from neurofibrillar-stained retinae. Analysis of the soma size and axon diameters of a population of heavily labelled retinal ganglion cells showed a significant correlation between the two. However, the overall distribution of intraretinal axon diameter was bimodal with an extended tail. Analysis of the ganglion cell distributions in the adult ferret indicates that beta cells comprise about 50.5–55%, gamma 42.5–47%, and alpha 2.5% of the ganglion cell population. This implies that the proportion of gamma, beta, alpha cells in both cat and ferret retina is highly conserved despite differences in visual specialization in the two species.

Previous experiments in animals have shown that early unilateral eyelid suture, a model of amblyopia induced by cataract, causes shrinkage of ocular dominance columns serving the deprived eye in the striate cortex. It is unknown whether the ocular dominance columns are affected in amblyopia produced by strabismus. We examined specimens of striate cortex obtained postmortem from a 79-year-old woman with a history of amblyopia in her left eye (20/800) since age 2 from accommodative esotropia. Four years prior to her death, she suffered an ischemic infarct of the left optic disc. This injury to the left optic disc made it possible to label the ocular dominance columns using cytochrome oxidase histochemistry. The pattern of ocular dominance columns was reconstructed throughout most of the right striate cortex. No shrinkage of columns was found. In the left cortex only half the column mosaic was labelled, because the patient had some residual vision in the temporal retina of her left eye. The columns within the labelled portion of the overall mosaic appeared normal. These findings indicate that shrinkage of ocular dominance columns does not occur in humans with amblyopia caused by accommodative esotropia. The ocular dominance columns are probably no longer susceptible to shrinkage at the age when most children with this condition begin to develop amblyopia.

Visual motion, i.e. the pattern of changes on the retinae caused by the motion of objects or the observer through the environment, contains important cues for the accurate perception of the three-dimensional layout of the visual scene. In this study, we investigate if neurons in the visual system, specifically in area MT of the macaque monkey, are able to differentiate between various velocity gradients. Our stimuli were random dot patterns designed to eliminate stimulus variables other than the orientation of a velocity gradient. We develop a stimulus space (“deformation space”) that allows us to easily parameterize our stimuli. We demonstrate that a substantial proportion of MT cells show tuned responses to our various velocity gradients, often exceeding the response evoked by an optimized flat velocity profile. This suggests that MT cells are able to represent complex aspects of the visual environment and that their properties make them well suited as building blocks for the complex receptive-field properties encountered in higher areas, such as area MST to which many cells in area MT project.