Synchronization of visual responses between the cortex, lateral geniculate nucleus, and retina in the anesthetized cat

Synchronization of spatially distributed responses in the cortex is often associated with periodic activity. Recently, synchronous oscillatory patterning was described for visual responses in retinal ganglion cells that is reliably transmitted by the lateral geniculate nucleus (LGN), raising the question of whether oscillatory inputs contribute to synchronous oscillatory responses in the cortex. We have made simultaneous multi-unit recordings from visual areas 17 and 18 as well as the LGN and the retina to examine the interactions between subcortical and cortical synchronization mechanisms. Strong correlations of oscillatory responses were observed between retina, LGN, and cortex, indicating that cortical neurons can become synchronized by oscillatory activity relayed through the LGN. This feedforward synchronization occurred with oscillation frequencies in the range of 60-120 Hz and was most pronounced for responses to stationary flashed stimuli and more frequent for cells in area 18 than in area 17. In response to moving stimuli, by contrast, subcortical and cortical oscillations dissociated, proving the existence of independent subcortical and cortical mechanisms. Subcortical oscillations maintained their high frequencies but became transient. Cortical oscillations were now dominated by a cortical synchronizing mechanism operating in the 30-60 Hz frequency range. When the cortical mechanism dominated, LGN responses could become phase-locked to the cortical oscillations via corticothalamic feedback. In summary, synchronization of cortical responses can result from two independent but interacting mechanisms. First, a transient feedforward synchronization to high-frequency retinal oscillations, and second, an intracortical mechanism, which operates in a lower frequency range and induces more sustained synchronization.     

The influence of the visual cortex on the spatiotemporal response properties of lateral geniculate nucleus cells

Previous studies of the cortical input to the mammalian dorsal lateral geniculate nucleus (LGN) have identified a number of possible functions for the corticogeniculate pathway, including alteration of LGN spatial frequency selectivity and facilitation of both binocular interactions and orientation selectivity. These changes may be due to either a tonic or a phasic cortical facilitation or both. The temporal differences between each of these inputs suggests that their impact on LGN cell temporal tuning should be unique. To test this hypothesis, we reversibly blocked the visual cortex (VI) and measured the effects on several indices of the temporal properties of LGN cells, including peak frequency, bandwidth, and response phase. Macaque monkeys were anesthetized and paralyzed during single cell recording from the LGN while area VI was cryogenically deactivated. Single-cell responses were visually evoked with drifting, luminance-modulated, sine-wave gratings and discrete-Fourier analyzed. Cortical cooling produced statistically significant increases or decreases in response amplitude in 64% of cells recorded. In most cases, alterations in response amplitude occurred for stimuli that varied in spatial as well as temporal frequency. For those cells influenced by changes in stimulus temporal frequency, the majority showed changes over a broad range of frequencies. A minority of cells showed changes in either peak temporal tuning or temporal frequency bandwidth. Response phase angles for all temporal frequencies tested were unaffected by cortical cooling. Overall, these results suggest that the cortical input may alter the temporal response properties of LGN cells, perhaps by tonic, but not exclusively excitatory, corticofugal influences.     

Human primary visual cortex and lateral geniculate nucleus activation during visual imagery

The functional magnetic resonance (fMRI) technique can be robustly used to map functional activation of the visual pathway including the primary visual cortex (V1), the lateral geniculate nucleus (LGN), and other nuclei of humans during visual perception stimulation. One of the major controversies in visual neuroscience is whether lower-order visual areas involve the visual imagery process. This issue was examined using fMRI at high magnetic field. It was demonstrated for the first time that the LGN was activated during visual imagery process in the human brain together with V1 and other activation. There was a tight coupling of the activation between V1 and the LGN during visual imagery.     

Functional organization of owl monkey lateral geniculate nucleus and visual cortex

The nocturnal, New World owl monkey (Aotus trivirgatus) has a rod-dominated retina containing only a single cone type, supporting only the most rudimentary color vision. However, it does have well-developed magnocellular (M) and parvocellular (P) retinostriate pathways and striate cortical architecture [as defined by the pattern of staining for the activity-dependent marker cytochrome oxidase (CO)] similar to that seen in diurnal primates. We recorded from single neurons in anesthetized, paralyzed owl monkeys using drifting, luminance-modulated sinusoidal gratings, comparing receptive field properties of M and P neurons in the lateral geniculate nucleus and in V1 neurons assigned to CO "blob," "edge," and "interblob" regions and across layers. Tested with achromatic stimuli, the receptive field properties of M and P neurons resembled those reported for other primates. The contrast sensitivity of P cells in the owl monkey was similar to that of P cells in the macaque, but the contrast sensitivities of M cells in the owl monkey were markedly lower than those in the macaque. We found no differences in eye dominance, orientation, or spatial frequency tuning, temporal frequency tuning, or contrast response for V1 neurons assigned to different CO compartments; we did find fewer direction-selective cells in blobs than in other compartments. We noticed laminar differences in some receptive field properties. Cells in the supragranular layers preferred higher spatial and lower temporal frequencies and had lower contrast sensitivity than did cells in the granular and infragranular layers. Our data suggest that the receptive field properties across functional compartments in V1 are quite homogeneous, inconsistent with the notion that CO blobs anatomically segregate signals from different functional "streams."     

Synaptic excitation of principal cells in the cat''s lateral geniculate nucleus during focal epileptic seizures in the visual cortex

Psoriasis and its related arthritis are chronic inflammatory disorders affecting predominantly the skin and synovium. Although their etiology remains to be established, multiple factors seem to play important roles in their pathogenesis. These environmental (eg, infectious agents and trauma), genetic, and immunologic factors are reviewed in this article.     

Correlated size variations in human visual cortex, lateral geniculate nucleus, and optic tract

We have examined several components of the human visual system to determine how the dimensions of the optic tract, lateral geniculate nucleus (LGN), and primary visual cortex (V1) vary within the same brain. Measurements were made of the cross-sectional area of the optic tract, the volumes of the magnocellular and parvocellular layers of the LGN, and the surface area and volume of V1 in one or both cerebral hemispheres of 15 neurologically normal human brains obtained at autopsy. Consistent with previous observations, there was a two- to threefold variation in the size of each of these visual components among the individuals studied. Importantly, this variation was coordinated within the visual system of any one individual. That is, a relatively large V1 was associated with a commensurately large LGN and optic tract, whereas a relatively small V1 was associated with a commensurately smaller LGN and optic tract. This relationship among the components of the human visual system indicates that the development of its different parts is interdependent. Such coordinated variation should generate substantial differences in visual ability among humans.     

The ventral lateral geniculate nucleus and the intergeniculate leaflet: interrelated structures in the visual and circadian systems

The ventral lateral geniculate nucleus (vLGN) and the intergeniculate leaflet (IGL) are retinorecipient subcortical nuclei. This paper attempts a comprehensive summary of research on these thalamic areas, drawing on anatomical, electrophysiological, and behavioral studies. From the current perspective, the vLGN and IGL appear closely linked, in that they share many neurochemicals, projections, and physiological properties. Neurochemicals commonly reported in the vLGN and IGL are neuropeptide Y, GABA, enkephalin, and nitric oxide synthase (localized in cells) and serotonin, acetylcholine, histamine, dopamine and noradrenalin (localized in fibers). Afferent and efferent connections are also similar, with both areas commonly receiving input from the retina, locus coreuleus, and raphe, having reciprocal connections with superior colliculus, pretectum and hypothalamus, and also showing connections to zona incerta, accessory optic system, pons, the contralateral vLGN/IGL, and other thalamic nuclei. Physiological studies indicate species differences, with spectral-sensitive responses common in some species, and varying populations of motion-sensitive units or units linked to optokinetic stimulation. A high percentage of IGL neurons show light intensity-coding responses. Behavioral studies suggest that the vLGN and IGL play a major role in mediating non-photic phase shifts of circadian rhythms, largely via neuropeptide Y, but may also play a role in photic phase shifts and in photoperiodic responses. The vLGN and IGL may participate in two major functional systems, those controlling visuomotor responses and those controlling circadian rhythms. Future research should be directed toward further integration of these diverse findings.     

Spatiotemporal receptive field organization in the lateral geniculate nucleus of cats and kittens

We have studied the spatiotemporal receptive-field organization of 144 neurons recorded from the dorsal lateral geniculate nucleus (dLGN) of adult cats and kittens at 4 and 8 wk postnatal. Receptive-field profiles were obtained with the use of a reverse correlation technique, in which we compute the cross-correlation between the action potential train of a neuron and a randomized sequence of long bright and dark bar stimuli that are flashed throughout the receptive field. Spatiotemporal receptive-field profiles of LGN neurons generally exhibit a biphasic temporal response, as well as the classical center-surround spatial organization. For nonlagged cells, the first temporal phase of the response dominates, whereas for lagged neurons, the second temporal phase of the response is typically the largest. This temporal phase difference between lagged and nonlagged cells accounts for their divergent behavior in response to flashed stimuli. Most LGN cells exhibit some degree of space-time inseparability, which means that the receptive field cannot simply be viewed as the product of a spatial waveform and a temporal waveform. In these cases, the response of the surround is typically delayed relative to that of the center, and there is some blending of center and surround during the time course of the response. We demonstrate that a simple extension of the traditional difference-of-Gaussians (DOG) model, in which the surround response is delayed relative to that of the center, accounts nicely for these findings. With regard to development, our analysis shows that spatial and temporal aspects of receptive field structure mature with markedly different time courses. After 4 wk postnatal, there is little change in the spatial organization of LGN receptive fields, with the exception of a weak, but significant, trend for the surround to become smaller and stronger with age. In contrast, there are substantial changes in temporal receptive-field structure after 4 wk postnatal. From 4 to 8 wk postnatal, the shape of the temporal response profile changes, becoming more biphasic, but the latency and duration of the response remain unchanged. From 8 wk postnatal to adulthood, the shape of the temporal profile remains approximately constant, but there is a dramatic decline in both the latency and duration of the response. Comparison of our results with recent data from cortical (area 17) simple cells reveals that the temporal development of LGN cells accounts for a substantial portion of the temporal maturation of simple cells.     

Reversal of the morphological effects of monocular deprivation in the kittens's lateral geniculate nucleus

1. Eleven kittens were deprived of vision in one eye until the age of between 5 and 14 weeks. Their eyes were then reverse-sutured, they were allowed to survive for a further 3-63 days, and their brains were then examined histologically. 2. Measurement of the cross-sectional area of cells in the lateral geniculate nucleus (LGN) showed that when the reversal of lid suture was performed at the age of 8 or 14 weeks, the mean cell size was smaller in laminae connected to the initially closed right eye than it was in other laminae. 3. When the reversal of lid suture took place at 5 or 6 weeks of age there was a reversal of interlaminar size differences: the initially deprived eye was then connected to laminae containing larger cells. Even within 3 days after the reversal of lid suture, most of the morphological effects of the initial suture had been abolished, and they were fully reversed within 12 days. 4. These results are compared with physiological changes in the visual cortex of these and similarly reared animals.     

Effects of saccades on the activity of neurons in the cat lateral geniculate nucleus

Effects of saccades on individual neurons in the cat lateral geniculate nucleus (LGN) were examined under two conditions: during spontaneous saccades in the dark and during stimulation by large, uniform flashes delivered at various times during and after rewarded saccades made to small visual targets. In the dark condition, a suppression of activity began 200-300 ms before saccade start, peaked approximately 100 ms before saccade start, and smoothly reversed to a facilitation of activity by saccade end. The facilitation peaked 70-130 ms after saccade end and decayed during the next several hundred milliseconds. The latency of the facilitation was related inversely to saccade velocity, reaching a minimum for saccades with peak velocity >70-80 degrees /s. Effects of saccades on visually evoked activity were remarkably similar: a facilitation began at saccade end and peaked 50-100 ms later. When matched for saccade velocity, the time courses and magnitudes of postsaccadic facilitation for activity in the dark and during visual stimulation were identical. The presaccadic suppression observed in the dark condition was similar for X and Y cells, whereas the postsaccadic facilitation was substantially stronger for X cells, both in the dark and for visually evoked responses. This saccade-related regulation of geniculate transmission appears to be independent of the conditions under which the saccade is evoked or the state of retinal input to the LGN. The change in activity from presaccadic suppression to postsaccadic facilitation amounted to an increase in gain of geniculate transmission of approximately 30%. This may promote rapid central registration of visual inputs by increasing the temporal contrast between activity evoked by an image near the end of a fixation and that evoked by the image immediately after a saccade.     

Development of contrast sensitivity and temporal-frequency selectivity in primate lateral geniculate nucleus

We studied the development of spatial contrast-sensitivity and temporal-frequency selectivity for neurons in the monkey lateral geniculate nucleus. During postnatal week 1, the spatial properties of P-cells and M-cells are hardly distinguishable, with low contrast-sensitivity, sluggish responses, and poor spatial resolution. The acuity of P-cells improves progressively until at least 8 months, but there is no obvious increase in their maximum contrast-sensitivity with age. The contrast sensitivity of M-cells is already clearly higher than that of P-cells by 2 months, and at 8 months of age this characteristic difference between M- and P-cells approaches the adult pattern. There is a major increase in responsiveness during the first 2 postnatal months, especially for M-cells, the peak firing rate of which rises fivefold, on average, between birth and 2 months. Many P-cells in the neonatal and 2-month-old animals did not give statistically reliable responses to achromatic gratings, even at the highest contrasts: this unresponsiveness of P-cells might result from low gain and/or chromatic opponency. The upper limit of temporal resolution in the neonate is low--about one-third of that in the adult. Among M-cells, the improvement in temporal resolution, like that in contrast sensitivity, is rapid over the first 2 months, followed by a slower change approaching the adult value by 8 months of age. The development of contrast sensitivity, responsiveness and temporal tuning are little affected, if at all, by binocular deprivation of pattern vision from birth for even a prolonged period.     

Visual receptive-field properties of single neurons in cat's ventral lateral geniculate nucleus

1. Visual receptive-field characteristics were determined for 154 cells in the ventral lateral geniculate nucleus (VLG) of cats anesthetized with nitrous oxide. All cells were verified histologically to be within the VLG. Responses of 182 cells from laminae A and A1 of the dorsal lateral geniculate nucleus (DLG) were tested for comparison. 2. The VLG cells could be grouped into one of seven classes according to their responses to light stimulation. Twenty-seven percent of the cells had uniform receptive fields. They responded maximally to stationary stimuli flashed on or off anywhere within the receptive field and showed no evidence for antagonistic surround mechanisms. About 19.5% of the VLG cells had concentric receptive fields. They were similar to the uniform type, with the addition of a concentric inhibitory surround. Eight percent of the VLG cells had ambient receptive fields. These cells were characterized by an unusually regular maintained discharge which varied in rate in relation to the level of receptive-field illumination or of full-field ambient illumination. About 4% of the VLG cells were movement sensitive. They gave little or no response to stationary stimuli flashed on or off in the receptive field, and responded best to a contour moving across the receptive field in any direction. An additional 2.5% of the VLG cells were direction sensitive. Their response was dependent on the direction of stimulus movement through the receptive field. Sixteen percent of the VLG cells had indefinite receptive fields. They responded to whole-eye illumination or to localized visual-field stimulation; however, specific receptive-field properties could not be adequately defined. Approximately 23% of the VLG cells studied gave no convincing response to visual stimulation. 3. Responses of DLG cells agreed with those reported in previous studies. Almost all (97%) had concentric receptive fields, and a few (3%) had uniform receptive fields with no apparent antagonistic surround. None of the DLG cells had receptive fields like those in the other classes found for VLG cells. 4. The VLG cells tended to have large receptive fields; mean diameter was 10.6 degrees of visual arc. This was substantially larger than the diameter of receptive fields for DLG cells. In addition, VLG cells generally required larger stimuli than DLG cells to respond. There was no consistent relationship between receptive-field size and visual-field eccentricity for VLG cells, in contrast to the DLG. Most (57%) VLG cells were driven only by the contralateral eye, 30% were binocularly driven, and 13% were driven only by the ipsilateral eye. 5. A systematic visuotopic organization was present in the VLG. The lower visual field was represented anteriorly in the nucleus and the upper visual field posteriorly. The vertical meridian was represented along the dorsomedial border of the VLG where it abuts the DLG, and the temporal periphery was represented ventrolaterally. 6. Responses to electrical stimulation of the optic chiasm were studied for 55 VLG cells...     

Activity-evoked extracellular pH shifts in slices of rat dorsal lateral geniculate nucleus

Activity-dependent extracellular pH shifts were studied in slices of the rat dorsal lateral geniculate nucleus (dLGN) using double-barreled pH-sensitive microelectrodes. In 26 mM HCO3--buffered media, afferent activation (10 Hz, 5 s) elicited an early alkaline shift of 0.04+/-0.02 pH units associated with a later, slow acid shift of 0.05+/-0.03 pH units. Extracellular pH shifts in the ventral lateral geniculate nucleus were rare, and limited to acidifications of approximately 0.02 pH units. The alkaline shift in the dLGN increased in the presence of benzolamide (1-2 microM), an extracellular carbonic anhydrase inhibitor. The mean alkaline shift in benzolamide was 0.10+/-0.05 pH units. In 26 mM HEPES-buffered saline, the alkaline response averaged 0.09+/-0.03 pH units. The alkaline shifts persisted in 100 microM picrotoxin (PiTX) but were blocked by 25 microM CNQX/50 microM APV. If stimulation intensity was raised in the presence of CNQX/APV, a second alkalinization arose, presumably due to direct activation of dLGN neurons. The direct responses were amplified by benzolamide, and blocked by either 0 Ca2+/EGTA, Cd2+ or TTX. In 0 Ca2+, addition of 500 microM-5 mM Ba2+ restored the alkalosis. Alkaline shifts evoked with extracellular Ba2+ were larger and faster than those elicited by equimolar Ca2+. In summary, synchronous activation in the dLGN results in an extracellular H+ sink, via a Ca2+-dependent mechanism, similar to activity-dependent alkaline shifts in hippocampus.     

A dynamic and quantitative study of pattern visual evoked potentials and gamma-aminobutyric acid neurones in the lateral geniculate nucleus and the visual cortex of monocular deprivation cats

PURPOSE: To assess the effects of monocular lid closure during critical period on cortical activity. METHOD: Pattern visual evoked potentials (PVEP) of the normal and the monocular deprivation (MD) cats were dynamically measured and the number of gammaaminobutyric acid immunopositive (GABA-IP) neurones of the area 17 of the visual cortex and the lateral geniculate nucleus (LGN) was quantitatively compared by using immunohistochemical method (ABC). RESULTS: The amplitude of the N1-P1 attenuated in deprived eyes (DE), NE/DE at postnatal week (PNW) 7-8 (P < 0.05), NE/DE at PNW 15-16 (P < 0.01); while P1 latency delayed, NE/DE at PNW 7-8 (P > 0.05), NE/DE at PNW 15-16 (P< 0.05). The numbers of GABA-IP neurones in layer A1 of the ipsilateral LGN and in layer A of the contralateral LGN, compared to those in the corresponding normal laminae, were not significant at PNW 7-8 and PNW 11-12 (P > 0.05), while in the same cats a reduction in the number of GABA-IP neurones was found in layer IV of area 17 at PNW 11-12 (P < 0.05). However, with longer survival of 3-4 weeks in duration, the numbers of GABA-IP neurones in the deprived laminae of LGN were remarkably reduced (P < 0.05). CONCLUSIONS: The amplitude of N1-P1 components is sensitive to the effects of monocular deprivation. Monocular deprivation in cats during critical period leads to dramatic changes of the number of GABA-IP neurones in the LGN and cortical layer IV receiving inputs from the deprived eye in cats. The deprivation-induced reduction in GABA-IP neurones is delayed in the LGN compared with the visual cortex. PVEP of the MD cats is consistent with the damage of its GABA system in visual cortex.     

Properties of relay cells in cat's lateral geniculate nucleus: a comparison of W-cells with X- and Y-cells

1. Observations are presented on the physiological properties of W-, X-, and Y-type relay cells in the cat's lateral geniculate nucleus (LGN). Emphasis is placed on the most recently recognized type, W-cells; data are presented on X- and Y-cells by way of comparison. 2. Seventy-seven W-cells were recognized on 70 microelectrode penetrations through the LGN. They resembled W-type retinal ganglion cells in their responses to visual stimuli. Tonic (on-center and off-center) W-cells, phasic (on-, off- and on-off center) W-cells, suppressed-by-contrast, and color-coded cells were recognized. 3. W-type relay cells also resembled retinal W-cells in their maintained activity and receptive field-center diameters. 4. W-type relay cells comprised 11.5% X-cells 48.4%, and Y-cells 22.3% of all LGN cells encountered on a reference sample of 62 electrode tracks. W-cells were found in laminae C, C1, and C2, comprising 36.5% of the sample in these laminae, but were not encountered in laminae A or A1. X- and Y-cells were found in laminae A, A1, and C. Within lamina C there was a tendency for X- and Y-cells to be located dorsal to W-cells. There was thus a substantial dorsoventral segregation of W-cells from X- and Y-cells. W-cells being found in the ventral parvocellular component of the dorsal LGN. 5. Cells considered to be W-type relay cells were shown to respond to electrical stimulation of the optic nerve and chiasm at latencies which were longer than those of X- and Y-cells, and were consistent with their receiving monosynaptic input from retinal W-cells. Geniculate W-cells of all subtypes were activated antidromically from the visual cortex. Their antidromic latencies were, on the average, longer than for Y- or X-cells, indicating that W-type relay cells had slower axons as well as slower retinal afferents, than X- or Y-cells. 6. The visual cortex thus appears to receive input from all three major types of retinal ganlion cells (W-, X-, and Y-cells) relayed separately, in parallel, by different groups of relay cells.     

The distribution of calcium-binding proteins in the lateral geniculate nucleus and visual cortex of a New World monkey, the marmoset, Callithrix jacchus

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

On the significance of temporally structured activity in the dorsal lateral geniculate nucleus (LGN)

Higher organisms perceive information about external or internal physical or chemical stimuli with specialized sensors that encode characteristics of that stimulus by a train of action potentials. Usually, the location and modality of the stimulus is represented by the location and specificity of the receptor and the intensity of the stimulus and its temporal modulation is thought to be encoded by the instantaneous firing rate. Recent studies have shown that, primarily in cortical structures, special features of a stimulus also are represented in the temporal pattern of spike activity. Typical attributes of this time structure are oscillatory patterns of activity and synchronous discharges in spatially distributed neurons that respond to inputs evoked by a coherent object. The origin and functional significance of this kind of activity is less clear. Cortical, subcortical and even very peripheral sources seem to be involved. Most of the relevant studies were devoted to the mammalian visual system and cortical findings on temporally structured activity were reviewed recently (Eckhorn, 1994, Progr. Brain Res., Vol. 102, pp. 405-426; Singer and Gray, 1995, Annu. Rev. Neurosci., Vol. 18, pp. 555-586). Therefore, this article is designed to give an overview, especially of those studies concerned with the temporal structure of visual activity in subcortical centers of the primary visual pathway, which are the retina and the dorsal lateral geniculate nucleus (LGN). We discuss the mechanisms that possibly contribute to the generation and modulation of the subcortical activity time structure and we try to relate to each other the subcortical and cortical patterns of sensory activity.