OFF-center Ganglion Cells Research Articles

Selective Glycine Receptor α2 Subunit Control of Crossover Inhibition between the On and Off Retinal Pathways

In the retina, the receptive fields (RFs) of almost all ganglion cells (GCs) are comprised of an excitatory center and a suppressive surround. The RF center arises from local excitatory bipolar cell (BC) inputs and the surround from lateral inhibitory inputs. Selective antagonists have been used to define the roles of GABA(A) and GABA(C) receptor-mediated input in RF organization. In contrast, the role of glycine receptor (GlyR) subunit-specific inhibition is less clear because the only antagonist, strychnine, blocks all GlyR subunit combinations. We used mice lacking the GlyRα2 (Glra2(-/-)) and GlyRα3 (Glra3(-/-)) subunits, or both (Glra2/3(-/-)), to explore their roles in GC RF organization. By comparing spontaneous and visually evoked responses of WT with Glra2(-/-), Glra3(-/-) and Glra2/3(-/-) ON- and OFF-center GCs, we found that both GlyRα2 and GlyRα3 modulate local RF interactions. In the On pathway, both receptors enhance the excitatory center response; however, the underlying inhibitory mechanisms differ. GlyRα2 participates in crossover inhibition, whereas GlyRα3 mediates serial inhibition. In the Off pathway, GlyRα2 plays a similar role, again using crossover inhibition and enhancing excitatory responses within the RF center. Comparisons of single and double KOs indicate that GlyRα2 and GlyRα3 inhibition are independent and additive, consistent with the finding that they use different inhibitory circuitry. These findings are the first to define GlyR subunit-specific control of visual function and GlyRα2 subunit-specific control of crossover inhibition in the retina.

Life imitates op art

The beautiful, undulating orientation maps in visual cortex have motivated many developmental models. A new study finds that this functional organization could be seeded in the retina by moire interference between mosaics of ON-center and OFF-center retinal ganglion cells.

Difference in regeneration and survival between OFF-center ganglion cells and ON-center cells in cat retina

Difference in regeneration and survival between OFF-center ganglion cells and ON-center cells in cat retina

Visual Response Properties of Retinal Ganglion Cells in the Royal College of Surgeons Dystrophic Rat

Alterations in retinal ganglion cell response patterns were profiled in dystrophic Royal College of Surgeons (hereafter RCS) rats over the first 100 postnatal days as a baseline for retinal rescue and vision restoration strategies. This method enabled the evaluation of the extent to which postreceptoral neuronal attributes in degenerating retinas mirror inferred declines in photoreceptor function. Single-unit responses from large retinal ganglion cells were recorded from age-matched dystrophic RCS (RCS-rdy(-)) and congenic RCS-p(+) (hereafter wild-type or wt) rats were recorded in vitro under visual control. Cells were profiled with conventional spatial and flux stimulus modulations. Ganglion cell single unit and population attributes alter slowly over the course of photoreceptor degeneration in dystrophic RCS rats, with significant decreases in apparent receptive field size, contrast sensitivity, and threshold sensitivity detected by the first month of life. Spatial frequency tuning and contrast responses were extremely weak by postnatal day (P)76, paralleled by a progressive decline in signal-to-noise (S-N) ratio to roughly unity by postnatal day (P)107. This decline was only a simple loss of responsivity, as background firing rates increased substantially over time. Whereas wt retinas were dominated by ON-center cells (15/23 cells), dystrophic animals were dominated by OFF-center cells by P47 (24/27 cells). The first definitive signs of degeneration in dystrophic RCS rats are parallel decreases in ganglion cell threshold sensitivity and receptive field size, followed by deterioration in spatial summation. Arguably, these changes can be qualitatively explained as photoreceptor signaling losses. However, the apparent shift in population profile from ON- to OFF-center ganglion cells long before loss of the b-wave at P90 implies that a reactive mechanism such as bipolar cell rewiring and/or transformation of neuronal phenotypes occur during the early phase of photoreceptor stress, before rod and cone death.

Convergence and segregation of the multiple rod pathways in mammalian retina.

Using a multidisciplinary approach, we demonstrate that three different pathways are responsible for the transmission of rod signals across the mouse retina. Each pathway serves a primarily nonoverlapping range of stimulus intensities, with ganglion cells receiving either segregated or convergent inputs. For both on-center (ON) and off-center (OFF) ganglion cells, the primary rod pathway carries signals with the lowest threshold, whereas the secondary rod pathway is less sensitive by approximately 1 log unit. In addition, OFF signaling uses a tertiary rod pathway that is approximately 1 log unit less sensitive than the secondary. Although some ganglion cells received rod inputs exclusively from one of the pathways, others showed convergent inputs. Using pharmacological and genetic approaches, we defined classes of ON and OFF ganglion cells for which the scotopic inputs derive only from the primary pathway or from both primary and secondary pathways. In addition, we observed a class of OFF ganglion cell receiving mixed input from primary and tertiary pathways. Interestingly, OFF ganglion cells receiving convergent inputs from all three rod pathways or from the secondary and tertiary pathways together were never observed. Overall, our data show a complex arrangement of convergence and segregation of rod inputs to ganglion cells in the mammalian retina.

Function and plasticity of homologous coupling between AII amacrine cells

The AII amacrine cells are critical elements in the primary rod pathway of the mammalian retina, acting as an obligatory conduit of rod signals to both on- and off-center ganglion cells. In addition to the chemical synaptic circuitry they subserve, AII cells form two types of electrical synapses corresponding to gap junctions formed between neighboring AII cells as well as junctions formed between AII cells and on-center cone bipolar cells. Our recent results indicate that coupling between AII cells and cone bipolar cells forms an obligatory synapse for transmission of scotopic visual signals to on-center ganglion cells. In contrast, AII–AII cell coupling acts to maintain the sensitivity of the primary rod pathway by allowing for summation of synchronous activity and the attenuation of asynchronous background noise. Further, the conductance of AII–AII cell gap junctions is highly dynamic, regulated by ambient light conditions, thereby preserving the fidelity of rod signaling over the scotopic operating range from starlight to twilight.

The midget pathways of the primate retina.

Midget ganglion cells in the foveal slope, parafovea, near periphery and far periphery of human and monkey retinas have been studied by electron microscopy (EM). Five human foveal ganglion cells were reconstructed and found to share input from seven midget bipolar cells. The OFF center ganglion cells were in a one to one relationship with their midget bipolar cells. But the ON center cells received input from two to three midget bipolar cells, of which one was dominant in terms of numbers of ribbon synapses directed at the midget ganglion cell dendrites. In the human parafovea every midget ganglion cell received input from only one midget bipolar cell (previously published, Kolb and DeKorver, 1991). At 4 mm of eccentricity, the near peripheral ON midget ganglion cell received input from three midget bipolar cells and thus from three cones. In far peripheral retina (12 mm) the ON midget ganglion cell received input from three to four midget bipolar cells. The peripheral midget bipolar cells probably contacted three cones each: therefore between nine and 12 cones could have input to such midget ganglion cell. The relationship of the increasing dendritic field size and increasing convergence of cones to the midget ganglion cells with eccentricity from the fovea is discussed in terms of color processing and resolution.

The network-selective actions of quinoxalines on the neurocircuitry operations of the rabbit retina

We examined the contribution of N-methyl- d-aspartate (NMDA) and alpha-amino-3-hydroxy-5-methyl-4-isoxalole-4-propionic acid (AMPA)/kainate (KA) receptors to the light-responses of rabbit retinal neurons. In the outer retina, bath application of the AMPA/KA receptor antagonists 6,7-dinitro-quinoxaline-2,3-dione (DNQX) and 2,3,dihydroxy-6-nitro-7-sulfamoyl-benzo- f-quinoxaline (NBQX) blocked the light-responses of horizontal cells. Application of quinoxalines enhanced ON-bipolar cell light-responses, and was associated with a hyperpolarization of their resting potentials. In the inner retina, application of both AMPA/KA and NMDA antagonists to AII amacrine-like cells only partially blocked their light-responses. Their residual responses may reflect electrical coupling to neighboring ON-center cone bipolar cells, and can inhibit OFF-center ganglion cells. ON-sustained ganglion cells were highly sensitive to the quinoxalines, which reduced their light-evoked firing, while the firing of ON-transient cells remained as NMDA-mediated light-responses. Quinoxalines had differential effects on the firing rates of ON- and OFF-center ganglion cells: ON-cells were reduced, while OFF-cells were increased. In contrast, firing rates of ON–OFF ganglion cells were not excited by NBQX, and showed a recovered light-response mediated by NMDA receptors. The receptive field surround was lost in ganglion cells. For comparison, NMDA antagonists had only moderate effects on all ganglion cell light-responses. Our results indicate that NMDA and AMPA/KA receptors both contribute to ganglion cell light-responses. However, AMPA/KA receptors also significantly effect the light-response of neurons presynaptic to retinal ganglion cells, altering the observed pharmacology at the ganglion cell level.

Axonal conduction velocities of functionally characterized retinal ganglion cells in goldfish.

1. Visual response properties and conduction velocities of retinal ganglion cells were studied by extracellular recordings in the intact goldfish eye. Visually responsive single units were confirmed as ganglion cells by collision testing, and their receptive fields were mapped. 2. From compound action potentials, we identified groups I-V in the optic nerve, with overall conduction velocities of 11.5 +/- 1.17, 7.1 +/- 0.79, 4.4 +/- 0.56, 3.1 +/- 0.31 and 2.3 +/- 0.18 m s-1 (mean +/- S.D.) at 23 degrees C. 3. Ganglion cells were classified by their receptive fields as off-, on-off- or on-centre. Nearly all confirmed ganglion cells had axonal conduction velocities in groups II, III and IV; none fell in the fastest group, I. 4. Off-centre ganglion cells had conduction velocities only in the fast group, II. On-off-centre cells fell mainly in group III, with some in group, II. On-centre cells fell in groups II-V, but mainly in groups III and IV. 5. Receptive field centre diameters were 5-30 deg measured with a photopic background. The mean diameters for off-, on-off- and on-centres were 24, 15 and 18 deg, respectively. The relatively larger diameter and higher rate of spontaneous firing of the off-centre cells were maintained under different adaptation conditions. 6. The off-centre cells can be identified with an anatomical class of large, alpha-like ganglion cells in the goldfish retina.

Neural Mechanisms of Visual Perception

It is more than half a century ago since Keffer H Hartline published his classical receptive fields studies of single optic nerve fibres in the frog. World War II intervened and the full impact of his work did not become apparent until the early fifties, when Horace Barlow extended Hartline's analysis in the frog and Stephen W Kuffler showed the on-centre and off-centre type ganglion cells in the cat retina. The next advances were made in the late fifties when Jerome Lettvin and Humberto Maturana described cells in the frog tectum with very complex response properties and when David Hubel and I discovered that cells in the cat striate cortex were sensitive to orientation of contours and binocular stimulation. Vision research has gone a long distance since that time—nonetheless we have just begun the long journey towards a detailed mechanistic understanding of the neural basis of visual perception. In this lecture I discuss the processing of visual information at the level of the striate cortex in the cat and monkey, and describe technical advances that have greatly facilitated the analysis of the neural mechanisms of visual perception.

A differential effect of APB on ON- and OFF-center ganglion cells in the dark adapted rabbit retina

The glutamate analog, 2-amino-4-phosphonobutyric acid (APB) is a proven tool in exploring the retinal circuit; it has been shown to interfere specifically with the transmission from photoreceptor to depolarizing bipolar cell. Consequently, in photopic retinae, the application of APB disrupts the ON-channel leaving the OFF-channel undisturbed; on the other hand, in the scotopic state, APB application blocks all ganglion cell responses. In this paper, we will show that the ON- and OFF-channels have a differential sensitivity to application of APB. That is to say, APB blocks center responses in ON-ganglion cells at mean concentration of 22 ± 5.1 μM (mean ± standard error of the mean; n = 15) and in OFF-ganglion cells at mean concentration of 91 ± 15.5 μM ( n = 16. Since considerable data rule out direct effects of APB on ganglion cells, we hypothesize that this effect is due to a difference in the synaptic gain of ON and OFF pathways in the inner retina.

An alternative pathway for signal flow from rod photoreceptors to ganglion cells in mammalian retina.

Rod signals in the mammalian retina are thought to reach ganglion cells over the circuit rod-->rod depolarizing bipolar cell-->AII amacrine cell-->cone bipolar cells-->ganglion cells. A possible alternative pathway involves gap junctions linking the rods and cones, the circuit being rod-->cone-->cone bipolar cells-->ganglion cells. It is not clear whether this second pathway indeed relays rod signals to ganglion cells. We studied signal flow in the isolated rabbit retina with a multielectrode array, which allows the activity of many identified ganglion cells to be observed simultaneously while the preparation is stimulated with light and/or exposed to drugs. When transmission between rods and rod depolarizing bipolar cells was blocked by the glutamate agonist 2-amino-4-phosphonobutyric acid (APB), rod input to all On-center and briskly responding Off-center ganglion cells was dramatically reduced as expected. Off responses persisted, however, in Off-center sluggish and On-Off direction-selective ganglion cells. Presumably these responses were generated by the alternative pathway involving rod-cone junctions. This APB-resistant pathway may carry the major rod input to Off-center sluggish and On-Off direction-selective ganglion cells.

The role of NMDA and non-NMDA excitatory amino acid receptors in the functional organization of primate retinal ganglion cells.

The role of excitatory amino acid (EAA) receptors in primate retinal ganglion cell function was analyzed in a superfused retina-eyecup preparation using single-unit, extracellular recording techniques. The effects of bath applied L-2-amino-4-phosphonobutyrate (APB), N-methyl-D-aspartate (NMDA), and non-NMDA EAA receptor agonists and antagonists were examined on the light-evoked responses and resting firing rates of ganglion cells. APB (30-100 microM) reduced or blocked the light-evoked responses and resting firing rates of all ON-center ganglion cells; higher doses of APB (100 microM) were required to block the light-evoked responses of ON-transient cells. In contrast, an increase in resting firing rates was observed when L-APB was applied to some OFF-center ganglion cells. The EAA agonists kainate (KA) (10-20 microM) and NMDA (200-350 microM) increased the firing rate of virtually all ganglion cells examined. Quisqualate (10-20 microM) increased firing in most cells, but occasionally (4/13 cases) produced inhibition. The NMDA antagonist D-amino-phosphono-heptanoic acid (D-AP7) (200-250 microM) reduced the light-evoked responses of ganglion cells by an average of 12% from control levels, while resting firing rates declined 37%. In the presence of D-AP7, the basic receptive-field characteristics of cells were not significantly altered. In contrast, two non-NMDA receptor antagonists, NBQX (2,3-Dihydroxy-6-nitro-7-sulfamoyl-benzo-(F)-quinoxalinedione) and DNQX (6,7-dinitro-quinoxaline-2,3-dione), produced substantial reductions in the light-evoked responses (82%) and resting firing rates (87%) of all ganglion cell classes. A striking observation in some neurons was the recovery of a persistent transient light-evoked response in the presence of NBQX. This NBQX-insensitive, light-evoked response was always blocked by adding D-AP7. Thus, neurotransmission from bipolar to ganglion cells in primates is mediated predominantly by non-NMDA EAA receptors, with NMDA receptors forming a minor component of the light-evoked response.

Simulation of visual cortex development under lid-suture conditions: enhancement of response specificity by a reverse-Hebb rule in the absence of spatially patterned input.

In this report, I show that a reverse-Hebb synaptic modification rule leads to the enhancement of response specificity of simulated visual cortex neurons in the absence of spatial patterning of the afferent activity. Although it is clear that receptive fields in the visual cortex can be modified by experience, many studies have shown a substantial increase of response specificity in cats deprived of pattern vision by lid suture, leading some to conclude that receptive field properties are essentially hard-wired. The hard-wired vs. experience-dependent controversy can be resolved by assuming that while Hebb-type plasticity is responsible for developmental synaptic changes, the organization of presynaptic activity which exists under conditions of visual deprivation is sufficient to drive the neurons towards greater specificity (Linsker 1986a-c; Miller 1989, 1992; Miller et al. 1989). As a reverse-Hebb rule enhances response specificity by balancing the push-pull system of ON- and OFF-center afferents, the sufficient condition is that the activity of ON- and OFF-center retinal ganglion cells be negatively correlated, a condition which will be met by diffuse illumination as seen through sutured eyelids. Unlike the models of Linsker and Miller and colleagues, which are based on a standard-Hebb rule, the model presented here does not require the presence of a "Mexican hat" spatial patterning of the afferent correlations, which has not been observed experimentally.

Effects of vasoactive intestinal peptide on ganglion cells in the rabbit retina.

The effects of vasoactive intestinal peptide (VIP) on the extracellularly recorded activity of ganglion cells were studied in superfused rabbit retinas. VIP, applied to the bathing solution, significantly increased the maintained activity of both OFF-center and ON-center brisk ganglion cells. The transient, excitatory responses of these cells to flashes of light (spots or annuli centered over the receptive field) were either unaffected or moderately reduced. VIP did not affect the responses of most ON/OFF directionally selective ganglion cells to a moving light stimulus. Furthermore, maintained activity of most of these cells remained absent. Effects of VIP on the light responses of ganglion cells were pronounced in retinas that were bathed with the dopamine D1 antagonist (+)-SCH 23390. For OFF-center brisk ganglion cells, VIP brought out both the center and surround excitatory responses that were selectively reduced by (+)-SCH 23390. Similarly, VIP brought out the leading edge responses that were reduced by (+)-SCH 23390 in ON/OFF directionally selective ganglion cells to a moving light stimulus. VIP did not however reverse the effects of (+)-SCH 23390 on ON-center brisk ganglion cells. It is argued that VIP counteracts the effects of (+)-SCH 23390 on OFF-center brisk ganglion cells and ON/OFF directionally selective ganglion cells by stimulating adenylate cyclase activity in dopamine-receptive cells of the retina.

Effects of serotonergic agonists and antagonists on ganglion cells in the goldfish retina.

Extracellular recordings were made from the isolated goldfish retina during superfusion with various serotonergic agonists and antagonists to determine the effects of these drugs on the maintained activity and response properties of the ganglion cells. Superfusion of the retina with serotonin (25-500 microM) increased the maintained activity of OFF-center ganglion cells and decreased the maintained activity of ON-center ganglion cells. In addition, serotonin also attenuated the excitatory responses to annular stimuli, suggesting a decrease in the strength of surround input to the ganglion cells. The effects of serotonin on OFF-center ganglion cells were mimicked by the nonselective 5-HT1 agonist 5-MeOT and the 5-HT1A receptor agonist 8-OH-DPAT, while only 5-MeOT mimicked the action of serotonin on ON-center ganglion cells. The effects of exogenously applied serotonin on the ganglion cells could be blocked by the mixed 5-HT1/5-HT2 receptor antagonist methysergide but not by the 5-HT2 receptor antagonist mianserin or the dopamine receptor antagonist haloperidol. These results support previous anatomical and biochemical evidence that serotonin functions in a neurotransmitter or neuromodulatory role in the teleost retina and suggest that serotonin may be involved in modulating the maintained activity and surround input to the ganglion cells. The results also indicate that two different types of receptors may mediate the actions of serotonin in the ON and OFF pathways, respectively.

Push-pull modulation of ganglion cell responses of carp retina by amacrine cells

The responses of a ganglion and an amacrine cell were recorded simultaneously in the carp retina. Sinusoidal current injected into amacrine cells modulated ganglion cell discharges either in phase (excitation) or in opposite phase (inhibition). ON-center ganglion cells received excitatory inputs and OFF-center ganglion cells received inhibitory inputs from ON-center amacrine cells. They received inputs of opposite polarity from OFF-center amacrine cells. Namely, inputs from ON-center and OFF-center amacrine cells augment the responses of ON-center and OFF-center ganglion cells in a push-pull manner.

Analysis of the horizontal cell contribution to the receptive field surround of ganglion cells in the rabbit retina.

1. The influence of horizontal cells on ganglion cells, the output neuron of the retina, was examined in an in vitro rabbit eyecup preparation. The extracellular spike activity of ganglion cells was monitored while pulsatile DC or sinusoidally modulated current was injected intracellularly into nearby horizontal cells. Interactions between the effects of light stimulation and horizontal cell current injections on ganglion cell responses were also examined. 2. Horizontal cells were found to contribute to the receptive field surround of ganglion cells. In particular, horizontal cells contributed to surround excitability and to surround antagonism of the centre light response. 3. Brisk, sluggish and direction-selective ganglion cells were all affected by current injections into horizontal cells. However, brisk ganglion cells responded to lower amplitude currents than did sluggish or direction-selective cells. 4. Horizontal cells with receptive fields that overlap those of ganglion cells were able to affect ganglion cell discharge. Moreover, the closer a horizontal cell was to the receptive field centre of a ganglion cell, the more effective were current injections in modulating ganglion cell discharge rate. The length constant of the horizontal cell contribution to the ganglion cell receptive field was approximately 200 microns. These results indicate that horizontal cells which are located within or outside of a ganglion cell's receptive field centre can influence that ganglion cell's activity. 5. The influence of horizontal cells on ganglion cell discharges was relatively weak at low temporal frequencies of sinusoidally modulated current. 6. Application of 2-amino-4-phosphonobutyrate (APB), a glutamate analogue, blocked the modulation of spike activity of on-centre ganglion cells that was induced by sinusoidally modulated current injected into nearby horizontal cells. The spike activity of off-centre ganglion cells was not blocked. 7. These findings suggest that horizontal cells contribute to the surround of ganglion cells and bipolar cells primarily through a feedback pathway onto cone photoreceptor cells.

Involvement of glycinergic neurons in the diminished surround activity of ganglion cells in the dark-adapted rabbit retina

Previous studies have reported that the surround responses of retinal ganglion cells weaken or disappear upon dark adaptation. The mechanism(s) by which this occurs is largely unknown, although changes in activity of retinal dopaminergic neurons have been implicated. In the light-adapted rabbit retina, the surround ON responses of OFF-center ganglion cells have been shown to be markedly reduced or abolished by a dopamine antagonist. This effect of a dopamine antagonist was recently shown to be reversed by the glycine antagonist strychnine and by compounds that elevate intracellular cAMP levels. The present study was conducted to determine whether strychnine and cAMP-elevating compounds could bring out the surround ON responses in OFF-center ganglion cells that are diminished upon dark adaptation. Extracellular recordings of OFF-center brisk ganglion cells were made from isolated, superfused retinal preparations. During the course of dark adaptation, the surround ON responses of many cells decreased markedly. Application of low micromolar concentrations of strychnine to the bathing solution brought out the surround ON responses in both brisk-transient and brisk-sustained OFF-center ganglion cells. The center OFF responses of these cells, on the other hand, were not enhanced by strychnine. Of the cAMP-elevating compounds tested, 8-(4-chlorophenylthio) cyclic AMP was the most effective in bringing out the surround ON responses in dark-adapted OFF-center ganglion cells. The effects of bath application of this cAMP analog were very similar to those of strychnine. The findings from this study suggest that under dark-adapted conditions glycinergic neurons inhibit the surround component of OFF-center ganglion cells. The release of glycine from these neurons is suggested to be regulated by a cAMP-dependent mechanism.

GABA‐ergic and glycinergic pathways in the inner plexiform layer of the goldfish retina

GABA-ergic and glycinergic circuitry in the inner plexiform layer of the goldfish retina was evaluated by electron microscopic autoradiography of 3H-GABA and 3H-glycine uptake, combined with retrograde horseradish peroxidase (HRP) labeling of ganglion cells. GABA-ergic and glycinergic synapses were found on labeled ganglion cells throughout the inner plexiform layer. This reinforces the idea that physiological evidence of GABA-ergic and glycinergic influence on a variety of ganglion cells in goldfish and carp often reflects direct inputs. Double-labeled synapses are presented as evidence of direct type Ab amacrine cell input to on-center ganglion cells. At least one population of type Aa sustained-off GABA-ergic amacrine cell is proposed, on the basis of profuse GABA-ergic inputs onto bipolar cells in sublamina a. Similar GABA-labeled profiles are shown to synapse onto HRP-labeled probable off-center ganglion cells. Thus GABA-ergic amacrine cells not only provide the predominant feedback to depolarizing (on-center) and hyperpolarizing (off-center) bipolar cells but also provide feed-forward inputs to on- and off-center ganglion cells. Large-caliber GABA-ergic dendrites present in both sublaminae a and b resemble those expected of a previously described bistratified, transient amacrine cell. These processes synapse onto HRP-labeled ganglion cell profiles in both sublaminae. Two morphologies of glycinergic amacrine cell are proposed on the basis of light microscopic autoradiography, 1) the previously described small pyriform cell and 2) a multipolar cell. The differential connectivity of the glycinergic neurons described, however, remains indistinguishable. Whereas abundant glycinergic inputs to ganglion cells occur throughout the inner plexiform layer, contacts between glycinergic profiles and bipolar cells are extremely rare. Therefore, interpreting the meaning of glycinergic input to ganglion cells will require further study of amacrine cell circuitry.