The contribution of interocular orientation differences to depth perception, at either the neuronal or the psychophysical level, is unclear. To understand the responses of binocular neurons to orientation disparity, we extended the energy model of Ohzawa et al. (1990) to incorporate binocular differences in receptive-field orientation. The responses of the model to grating stimuli with interocular orientation differences were examined, along with the responses to random dot stereograms (RDS) depicting slanted surfaces. The responses to combinations of stimulus orientations in the two eyes were left–right separable, which means there was no consistent response to the binocular orientation difference. All existing neuronal data concerning orientation disparity can be well described by this type of model (even a version with no disparity selectivity). The disparity sensitive model is nonetheless sensitive to changes in RDS slant, although it requires narrow orientation bandwidth to produce substantial modulation. The disparity-insensitive model shows no selectivity to slant in this stimulus. Several modifications to the model were attempted to improve its selectivity for orientation disparity and/or slant. A model built by summing several disparity-sensitive models showed left–right inseparable responses, responding maximally to a consistent orientation difference. Despite this property, the selectivity for slant in RDS stimuli was no better than the simple disparity-selective model. The range of models evaluated here demonstrate that interocular orientation differences are neither necessary nor sufficient for signaling slant. In contrast, within the framework of the energy model, positional disparity sensitivity appears to be both necessary and sufficient.

Nitric oxide (NO) is a neuronal messenger molecule that mediates pathway refinement in some brain regions. We used nicotinamide adenine dinucleotide phosphate diaphorase (NADPHd) histochemistry to examine the development of NO expression in the superior colliculus (SC) of kittens aged E28–E58 and P2–P57 and adults in order to determine if NO expression is correlated with pathway refinement. At E28, labeled cells were seen only within the subventricular zone (SVZ). At E36–E41, labeled cells were also found within the deep gray layer (DGL) of SC. At E51 and E58, a few labeled neurons were also present in the intermediate gray layer (IGL). These neurons already had extensive dendritic fields and well-developed morphologies at the time that they first expressed nitric oxide synthase (NOS). The number of neurons labeled in the DGL and IGL increased postnatally, reaching a peak density between P14 and P35. Neurons within the optic (OL) and superficial gray layers (SGL) were first visible at P7 and increased slightly in number until adulthood. However, SGL-labeled neurons were relatively limited in number and lightly labeled at all ages examined. We conclude that (1) NADPHd expression occurs in SC beginning in the second trimester in kittens and progresses in a ventral to dorsal pattern between E36–P35; (2) few neurons in kitten SGL are labeled by NADPHd and these appear relatively late in postnatal development; and (3) there is no correlation between NOS expression and retinocollicular pathway refinement in kittens, a result different from that seen in rodents.

In binocular vision, the lateral displacement of the eyes gives rise to both horizontal and vertical disparities between the images projected onto the left and right retinae. While it is well known that horizontal disparity is exploited by the binocular visual system of birds and mammals to enable depth perception, the role of vertical disparity is still largely unclear. In this study, neuronal activity in the visual forebrain (visual Wulst) of behaving barn owls to vertical disparity was investigated. Single-unit responses to global random-dot stereograms (RDS) were recorded with chronically implanted electrodes and transmitted via radiotelemetry. Nearly half of the cells investigated (44%, 16/36) varied the discharge as a function of vertical disparity. Like horizontal-disparity tuning profiles, vertical-disparity tuning curves typically exhibited periodic modulation with side peaks flanking a prominent main peak, and thus, could be fitted well with a Gabor function. This indicates that tuning to vertical disparity was not caused by disrupting horizontal-disparity tuning via vertical stimulus offset, but by classical disparity detectors whose orientation tuning was tilted. When tested with horizontal in addition to vertical disparity, almost all cells investigated (92%, 12/13) were tuned to both kinds of disparity. The emergence of disparity detectors sensitive in two dimensions (horizontal and vertical) is discussed within the framework of the disparity energy model.

Visual stimulation of a region outside the receptive field of single cells in visual cortex often results in the modulation of their responses. The modulatory effects are thought to be mediated through lateral connections within visual cortex. Research on lateral interactions commonly shows suppression. There has been no systematic study of the optimal conditions for facilitation. Here we have studied the nature of the modulation using a new type of compound stimulus: contrast reversal of pattern stimuli made of three discrete grating patches. The middle patch, optimally fitted to the receptive field in orientation, size, and spatial as well as temporal frequencies, was flanked by two similar patches presented well outside the receptive field. We found that (1) both facilitation and suppression occurred often in the same cells, when orientations of the target and flankers matched the receptive-field's optimal orientation; (2) facilitation with collinear flankers occurred most frequently at target contrasts just above the cell's firing threshold and suppression prevailed at high contrasts; (3) facilitative or suppressive modulation was obtained with target-flankers separation of up to 12 deg or more; (4) collinear facilitation was lost when flankers' orientation was rotated by 90 deg, while keeping all other parameters the same; and (5) neither the modulation mode nor the proportion of modulated cells was related to the cell types (simple vs. complex cells) and cells' laminar locations. Here we have provided physiological evidence for contrast-dependent, collinear facilitation probably underlying perceptual grouping in humans.

The visual pigment from the ultraviolet (UV) cone photoreceptor of the tiger salamander has been cloned, expressed, and characterized. The cDNA contains a full-length open reading frame encoding 347 amino acids. The phylogenetic analysis indicates that the highest sequence homology is to the visual pigments in the S group. The UV opsin was tagged at the carboxy-terminus with the sequence for the 1D4 epitope. This fusion opsin was expressed in COS-1 cells, regenerated with 11-cis retinal (A1) and immuno-purified, yielding a pigment with an absorbance maximum (λmax) of 356 nm which is blue shifted from the absorption of retinal itself. The transducin activation assay demonstrated that this pigment is able to activate rod transducin in a light-dependent manner. Regeneration with 11-cis 3,4-dehydroretinal (A2) yielded a pigment with a λmax of 360 nm, only 4 nm red shifted from that of the A1 pigment, while bovine rhodopsin generated with A2 showed a 16-nm red shift from the corresponding A1 pigment. These results demonstrate that the trend for a shorter wavelength pigment to have a smaller shift of λmax between the A1 and A2 pigments also fits UV pigments. We hypothesize that the small red shift with A2 could be due to a twist in the chromophore that essentially isolates the ring double bond(s) from conjugation with the rest of the polyene chain.

At reduced luminance levels, the visual system integrates light over extended periods of time. Although the general effects of this process are known, specific changes in the visual cortex have not been identified. We have studied the physiological changes that occur during a transition from high to low luminance by measurements of single neurons in the cat's primary visual cortex. Under low-luminance conditions, we find increased latencies, expanded temporal responses, and a loss of temporal structure. This results in temporal-frequency tuning curves that are peaked at relatively low frequencies. To examine parallel perceptual changes, we compared perceived temporal frequency in human subjects under high- and low-luminance conditions. Low-luminance flickering patterns are perceived to modulate at relatively high rates. This occurs even though peak sensitivity is shifted to relatively low temporal frequencies. To explore further the perceptual component, we measured perceived temporal frequency in human subjects with unilateral optic neuritis for whom optic nerve transmission is known to be relatively slow and generally similar to the normal physiological state under low luminance. These subjects also perceive relatively high modulation rates through their affected eye. Considered together, these results demonstrate an inverse relationship between the physiological and the perceptual consequences of reduced stimulus luminance. This relationship may be accounted for by shifts of neuronal population responses between high- and low-luminance levels.

An investigation of retinal specializations was carried out in larval and juvenile dhufish, Glaucosoma hebraicum (Glaucosomidae, Teleostei). The development of photoreceptors and formation of the retinal mosaic was followed by light and electron microscopy. At hatching the eye was undifferentiated. Cone photoreceptors were present by day 3 posthatch (dph), when exogenous feeding began. Single and multiple cones were present in a row arrangement from 3 dph to 20 dph, when the first rod nuclei were observed. Between 20 dph and approximately 3 months posthatch (mph), the row arrangement was replaced by a square mosaic of four double cones surrounding a single cone, and the cones increased in size, with the outer segments reaching up to 30 μm in length. During the period of spatial rearrangement, triple cones were often observed. From their first appearance, rod photoreceptors were added rapidly. Investigation of ganglion cell topography in 3-mph fish that had attained the adult-like square photoreceptor mosaic was carried out using retinal wholemounts. The highest densities of neurones in the ganglion cell layer were in temporal retina but no well-defined area centralis was observed. Microspectrophotometric measurements of the visual pigments within the outer segments of the photoreceptors of 3-mph fish revealed double cones with identical absorption spectra in each member of the outer segment, and the wavelength of maximum absorption (λmax) located at 522 nm. Single cones were found to possess a visual pigment with λmax at 460 nm and rods with a λmax of 498 nm. The results imply that the larvae and juveniles are adapted for survival in coastal waters and may be active in relatively low light levels from early stages of development.

Nicotinic acetylcholine receptors are localized on retinotectal axons' terminals in lower vertebrates. The effects of activation of these receptors by endogenous acetylcholine were observed under stimulation of mass optic fibers. This study was designed to determine whether endogenous acetylcholine facilitates frog retinotectal transmission, provided only the synapses of an individual optic axon are activated, and to evaluate the feasible extent of nicotinic facilitation in these synapses by applied agonist. To this end, the effects of cholinergic drugs on the extracellular action and synaptic potentials recorded from the terminal arborization of a separate retinotectal fiber (in layer F of the tectum) were investigated in vivo. Glutamatergic nature of retinotectal synapses was reexamined by treatment with kynurenic acid. Both kynurenic acid (0.25–1 mM) and d-tubocurarine chloride (10–15 μM) significantly depressed the synaptic potentials. Carbamylcholine chloride (50–150 μM) evoked a large augmentation of the synaptic potentials and a slight but statistically significant decrease of the action potentials. D-tubocurarine reduced the effect of carbamylcholine. Pilocarpine hydrochloride (50 μM) had only a weak effect. The paired-pulse facilitation of the synaptic potentials changed significantly under the action of carbamylcholine and d-tubocurarine. The obtained results suggest that the glutamate release from activated synapses of individual retinotectal axons is facilitated by endogenous acetylcholine via presynaptic nicotinic receptors. Under used stimulation conditions, this modulation mechanism was employed only partially since its activation by applied carbamylcholine could enhance synaptic transmission up to 2.8 times.

The retina possesses subpopulations of amacrine cells, which utilize different transmitters, including acetylcholine (ACh), GABA, and dopamine. We have examined interactions between these neurones by studying the effects of nicotinic agonists on GABA and dopamine release. Isolated rabbit retinas were incubated with [3H]dopamine and then superfused. Fractions of the superfusate (2 min) were collected and the [3H]dopamine in each sample was measured. Endogenous GABA release was examined by incubating retinas in a small chamber. At 5-min intervals, the medium was changed and the GABA measured by high-pressure liquid chromatography (HPLC). Exposure of the retina to nicotine, epibatidine, and other nicotinic agonists increased the release of both GABA and dopamine. The effects of nicotine and epibatidine were blocked by mecamylamine, confirming an action on nicotinic receptors. The action of epibatidine on dopamine release was unaffected by glutamate antagonists but was blocked by picrotoxin and gabazine. These results suggested that nicotine might increase dopamine release indirectly by stimulating the release of GABA, which in turn inhibited the release of an inhibitory transmitter acting tonically on the dopaminergic amacrines. Exposure of the retina to GABA caused a small increase in dopamine release. This hypothetical inhibitory transmitter was not GABA, an opioid, adenosine, glycine, nociceptin, a cannabinoid, or nitric oxide because appropriate antagonists did not affect the resting release of dopamine. However, metergoline, a 5HT1/5HT2 receptor antagonist, and ketanserin, a 5HT2A receptor antagonist, but not the 5HT1A antagonist WAY100635, increased the resting release of dopamine and blocked the effects of nicotine. The 5HT1A/5HT7 agonist 8-hydroxy DPAT inhibited both the nicotine and GABA-evoked release of dopamine. We conclude that nicotinic agonists directly stimulate the release of GABA, but the evoked release of dopamine is indirect, and arises from GABA inhibiting the input of an inhibitory transmitter, which we tentatively identify as serotonin.

Chromaticity (C-type) horizontal cells in the retina of cold-blooded vertebrates receive antagonistic inputs from cone photoreceptors of different spectral types leading to color opponency. The relative contribution of each spectral type of cones can be selectively altered by chromatic background illumination. Therefore, the spectral properties of C-type horizontal cells are expected to change when the intensity and color of ambient illumination are altered. In this study, we investigated the effects of chromatic background lights upon color opponency in Red/Green (RGH) and Yellow/Blue (YBH) C-type horizontal cells in the everted eyecup preparation of the turtle Mauremys caspica. Photoresponses were elicited by long-wavelength and short-wavelength light stimuli in the dark-adapted state and under conditions of chromatic background illumination. We found that the total voltage range, within which graded depolarizing and the hyperpolarizing photoresponses could be elicited, either increased or decreased depending upon the color of the background light. However, the maximal and minimal potential levels determined respectively by long-wavelength and short-wavelength light stimuli of supersaturating intensity remained unchanged, regardless of the wavelength and intensity of the background. These findings indicate that turtle C-type horizontal cells operate as push–pull devices. A sufficiently bright short-wavelength stimulus can push them all the way to the maximal hyperpolarizing level while a very bright long-wavelength stimulus can pull them towards the most depolarizing potential.

The mammalian visual cortex harbors a number of functional maps that represent distinct attributes of stimuli in the visual environment. How different functional maps are accommodated within the same cortical space, especially in species that show marked irregularities in one or more functional maps, remains poorly understood. We used optical imaging of intrinsic signals and electrophysiological techniques to investigate the organization of the maps of orientation preference, ocular dominance, and visual space in ferret. This species shows striking nonuniformity in the arrangement of ocular dominance domains and disruption of the mapping of visual space along the V1/V2 border. We asked whether these irregularities would be reflected in the organization of the map of orientation preference. The results show that orientation preference is mapped consistently within both V1 and V2, and across the interareal boundary, with no reflection of the irregularities in the other maps. These observations demonstrate the accommodation of multiple functional maps within the same cortical space without systematic geometrical relationships that necessarily constrain the organization of each representation. Furthermore, they imply that the structure of the map of orientation preference reflects the architecture and activity patterns of cortical circuits that are independent of other columnar systems established in layer 4.

Anatomical findings are presented that identify cortical and subcortical sources of afferents to the nucleus reticularis tegmenti pontis (NRTP) and basal pontine nuclei. Projections from the middle temporal visual area (MT), medial superior temporal visual area (MST), lateral intraparietal area (LIP), and areas 7a and 7b to the basal pontine nuclei were studied using 3H-leucine autoradiography. The results complemented a parallel study of retrograde neuronal labeling attributable to injecting WGA-HRP into NRTP and neighboring pontine nuclei. Small 3H-leucine injections confined to MT, MST, LIP, area 7a, or area 7b, produced multiple patches of pontine terminal label distributed as follows: (1) An injection within MT produced terminal label limited to the dorsolateral and lateral pontine nuclei. (2) Injections restricted to MST or LIP showed patches of terminal label in the dorsal, dorsolateral, lateral, and peduncular pontine nuclei. (3) Area 7a targets the dorsal, dorsolateral, lateral, peduncular, and ventral pontine nuclei, whereas area 7b projects, additionally, to the dorsomedial and paramedian pontine nuclei. Notably, no projections were seen to NRTP from any of these cortical areas. In contrast, injections made by other investigators into cortical areas anterior to the central sulcus revealed cerebrocortical afferents to NRTP, in addition to nuclei of the basal pontine gray. With our pontine WGA-HRP injections, retrograde neuronal labeling was observed over a large extent of the frontal cortex continuing onto the medial surface which included the lining of the cingulate sulcus and cingulate gyrus. Significant subcortical sources for afferents to the NRTP and basal pontine nuclei were the zona incerta, ventral mesencephalic tegmentum, dorsomedial hypothalamic area, rostral interstitial nucleus of the medial longitudinal fasciculus, red nucleus, and subthalamic nucleus. The combined anterograde and retrograde labeling data indicated that visuo-motor cortico-pontine pathways arising from parietal cortices target only the basal pontine gray, whereas the NRTP, together with select pontine nuclei, is a recipient of afferents from frontal cortical areas. The present findings implicate the existence of parallel direct and indirect cortico-pontine pathways from frontal motor-related cortices to NRTP and neighboring pontine nuclei.

Monkey electrophysiological and human neuroimaging studies indicate the existence of specialized neural systems for the perception and execution of actions. To date, the dynamics of these neural systems in humans have not been well studied. Here, we investigated the spatial and temporal behavior of human neural responses elicited to viewing motion of the face, hand, and body. Scalp event-related potentials (ERPs) were recorded in 20 participants viewing videotaped mouth (opening, closing), hand (closing, opening), and body stepping (forward, backward) movements. ERP peak differences within the movements of each body part were compared using topographical maps of voltage, voltage difference, and Student's t-test at ERP peak latencies. Predominantly temporoparietal negative ERPs occurred to motion of all body parts within 200 ms postmovement onset. Hand closure elicited a significantly greater negativity than opening, particularly in the left hemisphere. Vertex positive ERPs within 300 ms postmovement onset were elicited to hand and body motion. A significantly greater positivity occurred for the body stepping forward relative to stepping backward. The ERP topography was consistent with observed activation foci in human neuroimaging studies. Our data indicate that the neural activity of a system dedicated to the perception of high-level motion stimuli can rapidly differentiate between movements across and within body parts.

The present study has examined the emergence of cholinergic stratification within the developing inner plexiform layer (IPL), and the effect of ablating the cholinergic amacrine cells on the formation of other stratifications within the IPL. The population of cholinergic amacrine cells in the ferret's retina was identified as early as the day of birth, but their processes did not form discrete strata until the end of the first postnatal week. As development proceeded over the next five postnatal weeks, so the positioning of the cholinergic strata shifted within the IPL toward the outer border, indicative of the greater ingrowth and elaboration of processes within the innermost parts of the IPL. To examine whether these cholinergic strata play an instructive role upon the development of other stratifications which form within the IPL, one-week-old ferrets were treated with l-glutamate in an attempt to ablate the population of cholinergic amacrine cells. Such treatment was shown to be successful, eliminating all of the cholinergic amacrine cells as well as the alpha retinal ganglion cells in the central retina. The remaining ganglion cell classes as well as a few other retinal cell types were partially reduced, while other cell types were not affected, and neither retinal histology nor areal growth was compromised in these ferrets. Despite this early loss of the cholinergic amacrine cells, which are eliminated within 24 h, other stratifications within the IPL formed normally, as they do following early elimination of the entire ganglion cell population. While these cholinergic amacrine cells are present well before other cell types have differentiated, apparently neither they, nor the ganglion cells, play a role in determining the depth of stratification for other retinal cell types.

Visual stimulation of zones extending beyond the classical receptive field can modulate the contrast gain of neurons in the lateral geniculate nucleus (LGN) of cats, but little is known about the effect of extra-classical visual stimulation on the LGN of primates. Hence, we compare the effect of long-range interactions in parvocellular and magnocellular LGN layers of the marmoset monkey Callithrix jacchus using optimal, incremental spots flashed on the classical receptive field either alone or simultaneously with the shift of a grating (98% contrast; 0.1 cycles/deg) confined to a peripheral annulus (radii: 5–15 deg). The contrast required to drive the response halfway to saturation (c50) of most LGN neurons was raised by remote pattern shifts. The c50 ratio [(shift+spot)/spot] in OFF-center magnocellular neurons was significantly higher than in OFF-center parvocellular neurons. OFF-center magnocellular neurons closer to the fovea (<10 deg eccentricity) tended to have a higher c50 ratio than in more peripheral neurons. A significant drop in visual sensitivity to 25% contrast spots was observed during remote motion: d′ fell from 1.8 to 1.4 in parvocellular neurons and from 2.2 to 1.7 in magnocellular neurons. Such long-range interactions produce a reduction in visual sensitivity by changing the gain of the geniculate relay and point to an inhibitory, motion-sensitive extra-classical receptive field in both parvocellular and magnocellular pathways, which may be involved in saccadic suppression and attentional mechanisms in early vision.

Observers performed red–green and yellow–blue hue cancellation tasks for a 0.8-deg circular test field on a dark surround, by manipulating the excitation level of one cone class while the other two classes were held constant. The results of the red–green judgments conformed to classical opponent color theory in that both L- and S-cone excitation levels were antagonistic to M-cone signals. The yellow–blue judgments revealed a nonmonotonic nonlinearity in which the S-cone signal could act either antagonistically or synergistically with M- and L-cone signals. These results demonstrate that fixed hue sensations should not be associated with a given class of cone, even at the level of opponent neural coding.

Retinal neurotransmitters are known to play a role in postnatal ocular development and eye growth. The success of muscarinic antagonists in blocking form-deprivation myopia has implicated retinal acetylcholine in the control of ocular growth. The present study investigated whether steady-state content of acetylcholine (ACh) and its metabolite choline (Ch) are altered in the retina of eyes developing axial myopia, in both tree shrews and chicks. Retinal ACh and Ch content were measured using reverse-phase high-performance liquid chromatography. Posterior and anterior retinal samples were analyzed from myopic (form deprived) and control eyes as well as age-matched normal eyes. Normative data on retinal neurotransmitter content demonstrated that chick retinas contained less than half the ACh and Ch neurotransmitter content of tree shrews when normalized to retinal protein (ACh: 61 ± 3 vs. 130 ± 6 ng, Ch: 131 ± 5 vs. 347 ± 25 ng). There was no significant difference in either ACh or Ch content between myopic and contralateral control eyes in either tree shrews or chicks, irrespective of the degree of myopia. This finding was consistent for both posterior, anterior, and consequently whole retinal samples. In contrast, dopamine and DOPAC contents were found to be reduced in myopic compared to control eyes of the same tree shrews (dopamine −6.9% and DOPAC −15.5%) and chicks (dopamine −12.3% and DOPAC −28.2%). These findings demonstrate that, contrary to dopamine and DOPAC content, steady-state retinal acetylcholine and choline content is not significantly altered during myopia development.

In the marmoset Callithrix jacchus, ocular dominance columns (ODC) have been reported to be present in young animals, but absent in adults (Spatz, 1989). We have studied in juvenile and adult animals the postnatal organization of the retino-geniculo-cortical afferents by means of transneuronal labeling. We show in the present work that ODC are present in the primary visual cortex of Callithrix jacchus, both in the adult and in the juvenile animal. The present work confirms the presence of ODC in the visual cortex of juvenile marmoset before the end of the first postnatal month. In 2-month-old animals, ODC are well demarcated in IVcα and IVcβ. In the adult marmosets, the present data clearly show that the primary visual cortex is also organized with ODC. In horizontal sections, they form a mosaic through the ventral and dorsal calcarine cortex and through the dorso-lateral occipital part of the striate cortex. In frontal sections, their presence is manifest in IVcβ within the calcarine cortex and they only faintly appear in IVcα. These new findings are important since they underline the usefulness of the adult New World Monkeys as a model in visual research.

We studied the visuomotor activity of corticotectal (CT) cells in two visual cortical areas [area 17 and the posteromedial lateral suprasylvian cortex (PMLS)] of the cat. The cats were trained in simple oculomotor tasks, and head position was fixed. Most CT cells in both cortical areas gave a vigorous discharge to a small stimulus used to control gaze when it fell within the retinotopically defined visual field. However, the vigor of the visual response did not predict latency to initiate a saccade, saccade velocity, amplitude, or even if a saccade would be made, minimizing any potential role these cells might have in premotor or attentional processes. Most CT cells in both areas were selective for direction of stimulus motion, and cells in PMLS showed a direction preference favoring motion away from points of central gaze. CT cells did not discharge with eye movements in the dark. During eye movements in the light, many CT cells in area 17 increased their activity. In contrast, cells in PMLS, including CT cells, were generally unresponsive during saccades. Paradoxically, cells in PMLS responded vigorously to stimuli moving at saccadic velocities, indicating that the oculomotor system suppresses visual activity elicited by moving the retina across an illuminated scene. Nearly all CT cells showed oscillatory activity in the frequency range of 20–90 Hz, especially in response to visual stimuli. However, this activity was capricious; strong oscillations in one trial could disappear in the next despite identical stimulus conditions. Although the CT cells in both of these regions share many characteristics, the direction anisotropy and the suppression of activity during eye movements which characterize the neurons in PMLS suggests that these two areas have different roles in facilitating perceptual/motor processes at the level of the superior colliculus.

Simultaneous recordings from relay cells in the lateral geniculate nucleus (LGN) and their retinal afferents were used to examine the rules governing the transmission of spikes across the retino-geniculate synapse. Retinal spikes that terminate short retinal interspike intervals are much more likely to be transmitted across the synapse than spikes terminating longer intervals. This facilitation can be observed for interspike intervals as long as 50 ms when retinal firing rates are low, but the range of effective intervals decreases exponentially as retinal firing rate increases. Contribution, the fraction of LGN spikes triggered by an individual retinal afferent, is typically much higher during visual stimulation than during maintained activity, and these differences are unrelated to presynaptic or postsynaptic firing rate. It is suggested that this effect is a manifestation of increased synchronization of spikes among retinal afferents to the geniculate cell during structured visual stimulation, and that this synchronization offers a means of enhancing signal-to-noise ratio at the retino-geniculate synapse. Cross-correlograms between geniculate burst spikes and retinal afferents often contain two distinct peaks; a short latency peak that results from direct coupling between burst spikes and retinal input spikes, and a longer latency peak that represents indirect coupling in which retinal spikes trigger the calcium spike underlying the burst. Direct coupling is most likely to occur for the later spikes in the burst, and is present regardless of whether the calcium spike underlying the burst is triggered by the same or a different retinal afferent. These results further illuminate the relationship between tonic and burst modes of retino-geniculate transmission and indicate that bursts in LGN relay cells can be viewed as a mechanism of signal amplification, producing signals whose timing is potentially related to the temporal structure of a stimulus, independent of presynaptic and postsynaptic firing rate. This mechanism also appears to capitalize on the synchronization that is present among parallel retinal afferents to a geniculate cell.