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The effects of the experience of forming visual images on the spatial organization of the EEG.

The spatial organization of biopotentials in the cerebral cortex of 23 subjects who were students at the Faculty of Graphic Arts ("professionals") as well as 39 subjects lacking systematic experience of visual images ("non-professionals") was compared with the aim of identifying EEG correlates of the experience of visual images (image formation) in humans. Changes in measures of the spatial organization of biopotentials (spatial synchronization, spatial disordering, coherence, and spectral power) were analyzed as subjects mentally composed visual images consisting of two simple graphic elements - right angles and oblique lines. The total number of image elements increased in each of four sequential tasks, from a number which could be analyzed at the conscious level (4-7 elements) to a number exceeding analysis at the conscious level (8-16). Intergroup differences, particularly increases in the spatial disordering of biopotentials (non-linear processes), were detected when large numbers of elements were used (tasks 3 and 4). This measure increased more markedly in professionals than in non-professionals. Changes were significant in the anterior areas of the right hemisphere. Spatial synchronization of biopotentials (linear processes) increased in non-professionals in the posterior areas of the right hemisphere. Coherence and spectral power increased in professionals in a larger number of narrow-band EEG frequency subranges than in non-professionals. These data show that experience of visual imagery results in a more complex neurodynamic process during the activity, with non-linear dynamics and a multitude of EEG resonance systems at different frequencies.

Adolescent↗

Spatial correlates of hippocampal unit activity are altered by lesions of the fornix and endorhinal cortex.

Behavioral and electrophysiological evidence supports the role of the hippocampus in the processing of spatial information. In the present study, neuronal activity recorded from chronically implanted hippocampal microelectrodes was correlated with a rat's spatial orientation while traversing a radial maze for food reward. Place units were found in all fields of the dorsal hippocampus and dentate gyrus. Rotation of the maze relative to extramaze cues failed to disrupt the intact animal's spatial task performance of the spatial correlates of the unit activity. Lesions of the fornix or entorhinal cortex disrupted performance of the task. Unit activity correlated to the animal's spatial orientation was also disrupted by either lesion. There was no correlation between the disruption of the unit activity and location of the unit within hippocampal fields. Unit activity from lesioned animals showed correlation to the physical properties of the maze rather than to the orientation of the maze in space. These results further support the role of the hippocampus in the processing of spatial information.

Animals↗

Temporal and spatial response characteristics of the cat superior colliculus.

We have examined the responses of 72 cells of the cat superior colliculus to drifting gratings of sinusoidal luminance profile as a function of spatial frequency velocity and contrast. Of 72 cells, 66 responded to gratings either by change in mean firing rate only (58/72) or in a temporally modulated pattern in addition to the change in mean firing rate (8/72). The remaining 6 showed no change in discharge rate in response to any of the gratings tested. Many cells (24/72) were inhibited or excited by particular combinations of spatial and temporal frequencies. Some (8/72) demonstrated selective inhibition or excitation to a particular temporal frequency independent of spatial frequency and velocity and could therefore be said to be tuned specifically to temporal frequency. No cells were tuned only to a constant spatial frequency or a constant velocity. (24/72) cells displayed maximum inhibition or excitation only at a particular combination of spatial and temporal frequencies. Some cells (8/72) demonstrated a temporal modulation synchronous with the drifting grating in addition to an elevated mean discharge rate. The change in discharge rates evoked by gratings are generally less than those evoked by presentation of moving small slits or spots of light. Collicular cells often demonstrate a center-surround organization in their response to gratings. The center and surround often differ in their spatial frequency and velocity preferences. Compared to cortical and retinal ganglion cells, individual collicular cells are extremely non-linear. On a cell population basis, however, a linear Fourier analysis on grating response predicts the collicular cells' preference for movement of small objects.

Animals↗

Amelioration of spatial memory impairment by intrahippocampal grafts of mixed septal and raphe tissue in rats with combined cholinergic and serotonergic denervation of the forebrain.

Previous studies in the rat have shown that a serotonergic depletion greatly potentiates the learning and memory impairments produced by pharmacological or lesion-induced cholinergic blockade in the forebrain. The impairment produced by combined serotonergic-cholinergic lesions is reminiscent of that seen in memory-impaired aged rats. In the present experiment, we investigated whether grafts of cholinergic septal tissue and serotonergic mesencephalic raphe tissue, placed in the hippocampus, could reverse the severe memory impairment produced by combined cholinergic-serotonergic lesions. Adult rats were given an intraventricular injection of 5,7-dihydroxytryptamine followed by a radiofrequency lesion of the septum 1-2 weeks later. Three weeks after lesion surgery, the rats were given bilateral intrahippocampal cell suspension grafts of either fetal septal or mesencephalic raphe tissue, or both. The rats were tested for spatial learning and memory in the Morris water maze task at 4 and 10 months after grafting. At 4 months, lesioned and grafted groups were all impaired compared to the normal controls in their swim time and distance swum to find the platform, and they did not show any spatially focussed search strategy in the spatial probe trial when the platform was removed from the tank. At 10 months, the rats with mixed cholinergic and serotonergic grafts were no longer impaired compared to normals in their swim time and distance to find the platform, and they were significantly improved compared to the other grafted groups. Moreover, in the spatial probe trial, the rats with mixed cholinergic and serotonergic grafts displayed a spatially focussed search behaviour over the previous platform site, which was not seen in the lesioned control rats or in the other graft groups. Morphological analysis of the hippocampus revealed that the septal grafts produced an acetylcholinesterase-positive innervation but were totally devoid of serotonin innervation. The raphe grafts produced mainly a serotonin innervation, of both acetylcholinesterase- and serotonin-positive fibres. The results suggest that a mixture of septal and raphe tissue is required when grafted to the hippocampal formation in order to ameliorate the severe spatial learning and memory impairments produced by a combined cholinergic and serotonergic denervation, and that each of these graft types separately are not sufficient to ameliorate such deficits.

Animals↗

Physical activity enhances spatial learning performance with an associated alteration in hippocampal protein kinase C activity in C57BL/6 and DBA/2 mice.

The effects of physical activity on spatial learning performance and associated hippocampal functioning were examined in C57BL/6Ibg (C57) and DBA/2Ibg (DBA) mice. C57 and DBA mice, 3 months of age, were subjected to 8 weeks of a physical activity regime (consisting of moderate-pace treadmill running 5 days/week, 60 min/day, 0% grade, 12 m/min) or remained sedentary in their cages. Mice were then tested on the Morris water maze task for 6 days followed by 12 days of testing on the place learning-set task (8 trials/day with each task). Both C57 and DBA run mice showed no difference in swim speed compared to controls. Hippocampal protein kinase C (PKC) activity was measured in cytosolic, loosely bound, and membrane-bound homogenate fractions. Mice subjected to the physical activity protocol were compared to sedentary controls from the same set of litters. Physical activity produced a 2- to 12-fold enhancement in spatial learning performance on both the Morris (P < 0.0001) and place learning-set (P < 0.02) probe trials in both C57 and DBA mice. DBA mice, which characteristically perform poorly in comparison to C57 mice, were enhanced to perform similarly to C57 control mice. This physical activity-induced enhancement in spatial learning performance was accompanied by alterations in hippocampal bound PKC activity (P < 0.05). These data provide further support for our previous hypotheses of a PKC activity involvement in spatial learning and enhancement of spatial learning performance in rodents by physical activity. In addition, these data indicate that hippocampal PKC activity may be involved in the physical activity-induced enhancement of spatial learning performance.

Analysis of Variance↗

Spatial organization of extracellular matrix and fibroblast activity: effects of serum, transforming growth factor beta, and fibronectin.

The goal of our research is to understand reciprocal relationships between cell function and tissue organization. We studied the regulation of fibroblast activity in an in vitro culture model that recapitulates in continuous fashion the cycle of events occurring during connective tissue repair. We present evidence that concomitant with spatial reorganization of the extracellular matrix, there was a dramatic decline in extracellular matrix synthesis and cell proliferation. Therefore, spatial reorganization was a crucial turning point for fibroblast activity. Factors that regulated the timing of spatial reorganization included serum, transforming growth factor beta, and fibronectin. By accelerating spatial reorganization of the cultures, transforming growth factor beta led to a relative decrease in cell proliferation and extracellular matrix synthesis. By retarding spatial reorganization of the cultures, fibronectin led to a relative increase in cell proliferation and extracellular matrix synthesis. The results indicate that spatial information in the three-dimensional cell-matrix interaction permits higher order, tissue-level regulation of fibroblast function.

Blood Proteins↗

The development of basic mechanisms of pattern vision: spatial frequency channels.

The mature visual system possesses mechanisms that analyze visual inputs into bands of spatial frequency. This analysis appears to be important to several visual capabilities. We have investigated the development of these spatial-frequency channels in young infants. Experiment 1 used a masking paradigm to test 6-week-olds, 12-week-olds, and adults. The detectability of sine wave gratings of different spatial frequencies was measured in the presence and the absence of a narrowband noise masker. The 12-week data showed that at least two spatial-frequency channels with adultlike specificity are present at 12 weeks. The 6-week data did not reveal the presence of narrowband spatial-frequency channels. Experiment 2 used a different paradigm to investigate the same issue. The detectability of gratings composed of two sine wave components was measured in 6-week-olds and adults. The results were entirely consistent with those of experiment 1. The 12-week and adult data indicated the presence of narrowband spatial-frequency channels. The 6-week data did not. The results of these experiments suggest that the manner in which pattern information is processed changes fundamentally between 6 and 12 weeks of age.

Adult↗

Visuo-spatial working memory: structures and variables affecting a capacity measure.

The present paper examines the issue of the capacity of visuo-spatial working memory. A series of experiments test the hypothesis that two different components are critical in visuo-spatial working memory (passive store and active imagery operations), and, thereafter, attempt to specify the variables that affect the capacity of the passive store component. In the experiments, congenitally blind and sighted participants were asked to remember the spatial positions of target objects in two-dimensional matrices, with or without simultaneously performing a sequence of spatially-based imagery operations. We considered both the positions recall performance (the passive storage component) and the sequential imagery processing performance (the active processing component). We suggest that the two components of visuo-spatial working memory are independent. We also propose that both the number of relevant matrices and the number of target objects within each matrix affect the capacity of visuo-spatial working memory, with the latter factor possibly playing a greater role than the former one.

Adult↗

Deficits in spatial working memory after unilateral temporal lobectomy in man.

Forty neurosurgical patients and 20 controls were tested on a series of computerized tasks (the executive golf, structured golf and rotate tasks) designed to investigate spatial working memory. As defined by Olton [Spatial Abilities, Academic Press, New York, 1982], spatial working memory involves the encoding of specific and contextual information within the spatial domain. Right temporal lobectomy patients were significantly impaired on all three tasks, while the left temporal lobectomy patients showed a less significant overall impairment only on the structured golf task. Although there was no statistically significant differences between the two patient groups on the three tasks, the results point towards a robust deficit in spatial memory associated with right temporal lobectomy. The results provide further evidence for the role of the mesial temporal lobe structures in the processing and encoding of spatial information.

Adult↗

Discrimination thresholds in the two-dimensional spatial frequency domain.

In two experiments we have determined the discriminability between two sinusoidal gratings as a function of orientation and spatial frequency differences. Twelve orientation (15 degrees steps) and four spatial frequencies (2, 4, 8, 12 c/deg) were considered and corresponding discrimination thresholds were determined. Results indicated that: (a) spatial frequency discrimination thresholds did not significantly vary over all frequency and orientation positions, and averaged at +/- 1/8 octave limits. (b) Orientation thresholds only slightly increased with spatial frequency (from +/- 5 degrees at 2 c/deg to +/- 6 degrees at 12 c/deg). (c) A large and consistent anisotropy occurred with orientation thresholds showing most sensitivity in the horizontal and vertical directions, with largest thresholds at the oblique angles. No oblique effect was observed with spatial frequency thresholds. These results confirm past observations, providing an extensive set of threshold measurements in the two-dimensional spatial domain.

Discrimination, Psychological↗

Spatial frequency and the pattern onset-offset response.

Visually evoked responses (VERs) were recorded with vertical sinusoidal gratings presented in the on-off mode at rates of 0.5 and 1 Hz. Stimulus spatial frequency ranged from 0.5 to 16 c/deg and its contrast varied from near-threshold to the value of 0.3. At low spatial frequencies, 0.5 and 1 c/deg, the onset and offset VERs were of similar shape, magnitude and contrast dependence. At spatial frequencies higher than 2 c/deg the onset VERs were usually larger and persisted at lower contrast levels than the offset VERs. The results are consistent with the hypothesis that a gradual transition from the operation of transient channels to the operation of sustained channels takes place on increasing stimulus spatial frequency and that within a wide spatial frequency range (at least from 2 to 8 c/deg) suprathreshold gratings effectively stimulate both types of channels. It is confirmed that both onset and offset VERs are delayed at high spatial frequencies. In addition, a longer delay of the late onset waves was found in comparison with the delay of the early onset waves.

Adolescent↗

Facilitatory and inhibitory after-effect of spatially localized grating adaptation.

Aftereffects of spatially localized grating adaptation were measured for different locations of the adaptation grating relative to test grating. When the adaptation grating was located on or near the retinal area occupied by the test grating, contrast sensitivity was markedly reduced. When the adaptation grating was spatially separated from the test grating, contrast sensitivity was significantly increased. This aftereffect of spatially localized grating adaptation suggests that spatial-frequency-selective detectors are not spatially independent, but tonically inhibited by spatially contiguous mechanisms. Thus the adaptation of these mechanisms might cause an increase in contrast sensitivity of detectors subserving the test grating.

Adaptation, Ocular↗

Spatial frequency interference on grating-induction.

Spatial frequency interference and facilitation in suprathreshold vision were studied using the grating-induction effect [McCourt, Vision Res. 22, 119-134 (1982)] as a sensitive probe. The effects on grating-induction magnitude produced by variations in "interfering" and "inducing" grating spatial frequency, contrast and phase were examined in four experiments. A limited range of high spatial frequency interfering gratings reduced the contrast of gratings induced by spatially coextensive lower frequency inducing gratings. Both phase-dependent and phase-independent interference was observed. Facilitation of grating-induction was produced by interfering gratings of lower frequency than the inducing grating. It is hypothesized that the grating-induction interference effect is due to inhibition of the low spatial frequency selective mechanisms responsible for induction, by channels tuned to higher frequencies. The functional significance of induction and spatial frequency inhibition is discussed, and a mathematical description of the results is presented.

Form Perception↗

The effect of various anaesthetics on the spatial tuning of two major wave peaks in the transient pattern electroretinogram of the cat: evidence for pattern and luminance components.

The main PERG component of the transient contrast reversal pattern electroretinogram (PERG) in cats was a negative wave (3.5 microV average, SD 1.7 microV) peaking at about 130 msec (N130) with a spatial resolution above 5.5 c/deg, close to behavioural estimates. The early positivity (P35) was more variable, smaller and had lower spatial resolution. Different anaesthetic protocols affected both the waveform and the amplitude by spatial frequency functions. Responses of urethane anaesthetised cats were like those reported previously for decerebrate cats or cats paralysed and ventilated with N2O/O2/CO2 (75%/24%/1%). P35 was evoked only by coarse stimuli and N130 amplitude decreased linearly as spatial frequency increased. When the luminance response amplitude, predicted from the optical transfer function of the eye, was subtracted, spatial tuning appeared. An anaesthetic mixture of ketamine hydrochloride and xylazine depressed both P35 and N130 at low spatial frequencies while enhancing them at high frequencies. In paralysed animals ventilated with N2O/O2 (67%/33%) P35 was larger and recordable to 1.6 c/deg. Peak times were reduced and the inter-peak time halved. Other anaesthetics depressed the ERGs. These effects suggest that cats are a good model for studying N130 in isolation or its interaction with P35 and that both PERG peaks include luminance and pattern components.

Anesthetics↗

Spatial frequency tuning of human stereopsis.

A masking paradigm was employed to measure the spatial frequency selectivity of channels underlying human stereopsis. Observers viewed spatially filtered (0.4 octave bandwidth) random-dot stereograms in which a disparate bar appeared in either the top or bottom half of the display; superimposed on one RDS half-image was a noise target whose spatial frequency content was varied relative to that of the RDS. A staircase procedure was used to measure the monocular noise energy (and hence the signal-to-noise ratio) at which observers could judge the location of the disparate bar on 71% of trials. Statistical analyses showed that the resulting stereoscopic masking functions could be grouped into two sets, one with peak sensitivity at 3 c/deg and the other with peak sensitivity at 5 c/deg. These two channels were observed for both crossed and uncrossed disparities ranging from coarse to fine. Essentially the same results were obtained with binocular noise and with stereo displays flashed too briefly to be affected by eye movements. Our results are inconsistent with models of stereopsis in which the disparity range to which a channel is sensitive varies with that channel's peak spatial frequency. These data imply that the spatial frequency selectivity of stereopsis differs from the tuning of spatial channels underlying the detection and discrimination of form.

Depth Perception↗

Inhibitory refinement of spatial frequency selectivity in single cells of the cat striate cortex.

Single cells in the cat striate cortex are more selective for the spatial frequency of sinewave grating stimuli than are cells of the retina or lateral geniculate nucleus. We have explored the possibility that this enhancement of selectivity results from spatial-frequency-selective inhibition. Stimulation with two superimposed gratings, one to excite the cell and one to prove for inhibition, revealed spatial frequency-dependent response suppression in 74% of the total population studied. Suppression was slightly more prevalent in simple cells (80%) than in complex cells (68%). In 93% of the cases where suppression was found, its tuning was complementary to excitatory spatial frequency tuning, and the strongest suppression was usually found where the excitatory tuning function approached zero imp./sec. Characteristics of the phenomenon were independent of cortical layers. We conclude that organized inhibitory mechanisms serve to refine the spatial frequency bandpass of striate cortical cells. This provides evidence for another degree of nonlinearity in the organization of cortical receptive fields and supports the hypothesis that a fundamental function of the visual cortex is image dissection in the domain of spatial frequency.

Adaptation, Physiological↗

Orientation bandwidth: the effect of spatial and temporal frequency.

The orientation bandwidths of psychophysically defined channels of human vision were estimated by two techniques for a wide range of spatial and temporal frequencies. The first technique was an adaptation paradigm, where the subjects' ability to see patterns of various orientations was measured before and after adapting to a high contrast pattern. The second technique evaluated subjects' ability to discriminate between two gratings of different orientations in relation to their ability to detect the patterns. Bandwidths were unaffected by temporal frequency at high spatial frequencies but increased with temporal frequency at low spatial frequencies. Bandwidths increased modestly with decreasing spatial frequency at low temporal frequencies but more drastically at high temporal frequencies. Both techniques gave similar results except for patterns with very low spatial and high temporal frequencies. In this region the stimulus appears "spatial-frequency doubled" which may be used as a cue for the orientation discrimination task.

Adaptation, Ocular↗

Spatial frequency selective mechanisms underlying the motion aftereffect.

The motion aftereffect (MAE) was used to study the spatial frequency selectivity of suprathreshold motion perception. Observers were adapted to drifting sine-wave gratings confined to a retinal eccentricity of approx. 4 deg. The magnitude of the subsequent MAE was measured while viewing a stationary sine-wave grating test surface of one of a number of spatial frequencies. The largest MAE was found when the spatial frequency of the test stimulus was the same as that of the adapting stimulus. This phenomenon held for spatial frequencies between 0.5 and 4 c/deg, and was robust with changes in contrast of either adapting or test gratings. However, at an adapting spatial frequency of 0.25 c/deg, the peak MAE was observed at 0.5 c/deg. Control experiments indicated that this peak shift was not the result of the reduced number of cycles in the stimulus, nor the temporal frequency. There was no measurable MAE at spatial frequencies lower than 0.25 c/deg. These results suggest the existence of a "lowest adaptable channel" for the motion aftereffect.

Adaptation, Ocular↗