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J I Nelson

Publications and source records attributed to J I Nelson.

At least 19 recordsLinked to original sources

Structural basis of cortical synchronization. I. Three types of interhemispheric coupling.

1. Single-unit and multiunit activities were recorded at the area 17-18 border of each cortical hemisphere in paralyzed cats anesthetized with nitrous oxide supplemented with halothane. Cross-correlation histograms (CCHs) were computed between 86 pairs of single units and 99 pairs of multiunit activities. Visually evoked peaks in the CCHs were removed by subtracting the shift predictor. 2. Three types of CCH peaks were observed: T peaks with narrow widths (4-28 ms), C peaks with intermediate widths (30-100 ms), and H peaks with large widths (100-1,000 ms). Oscillatory coupling was observed rarely. This tripartite distribution of CCH peaks is similar to that reported in an earlier study on the temporal coupling between areas 17 and 18. Different types of peaks occurred in isolation or in combination. Combination of different peak types was more often observed in multiunit recordings. 3. CCH peaks of all types were usually centered, meaning that units in opposite hemispheres tend to synchronize their discharges. 4. T peaks were observed almost exclusively for units with overlapping receptive fields and preferentially for units with similar optimal orientations. No dependence on receptive field position or optimal orientation was observed for the encounter rate of C and H peaks. 5. A new method, called the peristimulus CCH, was developed to study the time course of the temporal coupling. This showed that H peaks can occur during visual stimulation and that their time course follows that of the visual responses of the coupled neurons. 6. Using one single bar or two simultaneously presented light bars as stimuli, we studied the effect of visual stimulation on the strength of H coupling. This showed that H coupling observed under stimulation with a single moving light bar can be completely abolished, with little change in visual responses, when the stimulus is changed to two noncoherently moving bars. This was related to a strong decrease of the H peaks in the autocorrelograms. 7. These results demonstrate that T, C, and H peaks constitute, together with high-frequency oscillations, universal forms of temporal coupling between neurons located in different cortical areas. The following paper reports on the effects of cortical lesions on the encounter rate and strength of these different types of coupling.

Anesthesia↗

Structural basis of cortical synchronization. II. Effects of cortical lesions.

1. To understand the structural basis of the different types of interhemispheric synchronizations described in the preceding paper, we made sections of the corpus callosum and lesions of extrastriate cortex. We measured the effects of such operations on the frequency of encounter, width and strength of T, C, and H peaks in cross-correlation histograms computed from single-unit and multiunit recordings from areas 17-18 of opposite cortical hemispheres in the cat. 2. Sectioning of the corpus callosum led to a complete abolition of T and C couplings and a strong reduction of encounter rate and strength of H coupling. A section limited to the posterior half of the corpus callosum abolished T and C couplings completely. This suggests that T and C couplings are mediated by the direct reciprocal connections between visual cortical areas circulating through the posterior part of the corpus callosum. 3. The encounter rate of H peaks depended on the extent of the callosal cut. Larger lesions gave a more pronounced reduction of the number of H peaks. From this observation we conclude that H peaks are at least partially mediated by polysynaptic pathways involving widely distributed cortical regions. 4. Extensive lesions of extrastriate cortex were made by aspiration of the gray matter or injections of ibotenic acid. These lesions removed the direct inputs from cortical areas sending ipsilateral as well as contralateral inputs to the area 17-18 border region. Encounter rate and coupling strength of C and H peaks were decreased, whereas little effect was observed on T peaks. 5. These results demonstrate that all types of interhemispheric synchronization are mediated by corticocortical connections and that T and C peaks are generated by reciprocal connections between areas 17 and 18 of each hemisphere. Feedback connections play a role in mediating or facilitating the C and H types of interhemispheric synchronization.

Animals↗

The RF-cinematogram. A cross-correlation technique for mapping several visual receptive fields at once.

We present a spike-triggered averaging method capable of mapping the visual receptive fields of several neurons simultaneously. The stimulation is general and the mapping proceeds automatically without the need to match the stimulation to the cells' preference for position, orientation, direction, etc. The maps are spatiotemporal; receptive field (RF) structures are quantitatively determined in three dimensions: the two dimensions of visuotopic space, and time. The method presented is one of a family of "reverse correlation" or "spike-triggered averaging" techniques (DeBoer and Kuyper 1968) capable of revealing linear aspects of stimulus-response coupling. The formal relationship of these methods to stimulus-response cross-correlation is shown. The analysis is extended to provide some second-order axis-of-motion information ("direction marks"). The stimulus is a constantly illuminated, randomly jumping bright or dark spot, not an elongated bar. Spot diameters between one-third to 1 x RF width are effective. The method ascertains for each recorded action potential or "spike" the prior visual field position of the spot. The average or most probable spot positions define the receptive field spatially. Repeating the process for a succession of times prior to observed spikes defines the field temporally, presented here as a succession of spatial maps. We term this portrayal a receptive field cinematogram, RFc or ciné. The RFc reveals and economically portrays the spread of excitability and suppression across the receptive field, culminating in the generation of a spike. RFcs for LGN neurons and for simple cells recorded in cat cortical areas 17 and 18 are presented and interpreted in terms of classic ON/OFF regions. The availability of temporal information permits the separation of an excitatory exit response, generated when a moving bright spot leaves an OFF region, from an excitatory entrance response occurring when a bright spot enters an ON region, because these responses occur at different times (exit responses earlier). Spike emission remains coupled to (cross-correlated with) stimulus events over time periods as long as 96 ms, implying that some stimulus drive or afferent visual input is delayed by as much as 96 ms more than other input. This is a striking instance of temporal dispersion in the visual system. In some cells, said to be "spatiotemporally inseparable", the delay (latency) varies systematically across the visual field; i.e., the place for optimal stimulation varies with the time prior to spike emission. In these cells, the RFc shows receptive field structures which move across the visual field over trajectories equal to approximately twice the total conventional RF width.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Spatial and temporal coherence in cortico-cortical connections: a cross-correlation study in areas 17 and 18 in the cat.

Visual cortical areas are richly but selectively connected by "patchy" projections. We characterized these connections physiologically with cross-correlograms (CCHs), calculated for neuron pairs or small groups located one each in visual areas 17 and 18 of the cat. The CCHs were then compared to the visuotopic and orientation match of the neurons' receptive fields (RFs). For both spontaneous and visually driven activity, most non-flat correlograms were centered; i.e. the most likely temporal relationship between spikes in the two areas is a synchronous one. Although spikes are most likely to occur simultaneously, area 17 spikes may occur before area 18 or vice versa, giving the cross-correlogram peak a finite width (temporal dispersion). Cross-correlograms fell into one of three groups according to their full-width at half peak height: 1-8 ms (modal width, 3 ms), 15-65 ms (modal width 30 ms), or 100-1000 ms (modal width 400 ms). These classificatory groups are nonoverlapping; the three types of coupling appeared singly and in combination. Neurons whose receptive fields (RFs) are nonoverlapping or cross-oriented may yet be coupled, but the coupling is more likely to be the broadest type of coupling than the medium-dispersed type. The sharpest type of coupling is found exclusively between neurons with at least partially overlapping RFs and mostly between neurons whose stimulus orientation preferences matched to within 22.5 deg. The maximum spatial dispersion observed in the RFs of coupled neurons compares well with the maximum divergence seen anatomically in the A18/A17 projection system. We suggest three different mechanisms to produce each of the three different degrees of observed spatial and temporal coherence. All mechanisms use common input of cortical origin. For medium and broad coupling, this common input arises from cell assemblies split between both sides of the 17/18 projection system, but acting synchronously. Such distributed common-input cell assemblies are a means of overcoming sparse connectivity and achieving synaptic transmission in the pyramidal network.

Animals↗

Intracortical facilitation among co-oriented, co-axially aligned simple cells in cat striate cortex.

Most neurons in cat striate visual cortex show inhibitory effects when moving contours are presented beyond the limits of classic receptive field regions. Facilitatory effects are also present in about 40% of simple cells. Here, we report a highly specific form of this facilitation, mediated only by neurons possessing both an orientation tuning matched to the test unit, and a receptive field position aligned with its long axis. This finding illustrates one of the intracortical interconnection schemes hypothesized by Mitchison and Crick (1982). Periodic clustering in long, intrinsic axons may signify a neuron seeking specific functional interactions like these across columnar systems in both the spatial and orientation domains.

Animals↗

Comparisons among Snellen, psychophysical, and evoked potential visual acuity determinations.

Acuity limits were determined for gratings of progressively finer spatial frequency directly from the visually evoked potential. The evoked potential was retrieved in real time, while spatial frequency was electronically changed or swept. The results of the swept evoked potential technique are compared to acuity thresholds determined psychophysically with similar gratings, and with Snellen acuity determinations over the range 6/60 (20/200) to 6/3.6 (20/12), obtained by defocusing with positive spherical lenses. One line of Snellen acuity is easily discriminated; the absolute Snellen level can often be identified to within two lines with 95% certainty. Correlations between visually evoked response (VER) grating and Snellen optotype acuity levels are poorer than correlations between VER- and psychophysically-determined grating acuity limits, perhaps because of variability inherent in the Snellen task itself. The swept evoked potential affords a rapid, objective, and potentially useful measure of acuity in young or minimally cooperative subjects.

Adult↗

Spatiotemporal conditions which elicit or abolish the oblique effect in man: direct measurement with swept evoked potential.

Reversing sine wave gratings were electronically swept in spatial frequency and contrast. The acuity limits and contrast thresholds of 4 observers were inferred from evoked potential stimulus-response functions elicited by these stimuli and retrieved with a quadrature lock-in amplifier. The evoked potential functions, linearized in the case of contrast by increasing contrast logarithmically with time, were extrapolated to the point of zero response. This point provides an electrophysiologically defined threshold value for acuity and for contrast. An oblique effect (superior sensitivity for HV-oriented gratings) could reliably be demonstrated in both acuity and contrast threshold performance. This oblique effect could readily be abolished under low spatial/high temporal frequency conditions. The findings are discussed in terms of shifting relative strengths of X and Y contributions to the steady-state evoked potential.

Adult↗

A rapid evoked potential index of cortical adaptation.

Contrast thresholds and acuity limits were measured in 4 observers with the swept visual evoked potential (VEP) technique. In this technique, grating contrast or grating spatial frequency is electronically varied while the subject's evoked response is retrieved in real time (without averaging). Contrast or spatial frequency variation make the stimulus vary in intensity; zero VEP response amplitude indicates the threshold intensity. Large shifts occur in the indicated threshold when stimulus sweep direction is reversed. Thresholds are always relatively elevated when the run begins with the strongest stimulus value. These shifts do not have a technical origin in the delay of the instrument (Nelson et al. 1984b). Here, it is shown that the shifts are due to orientation and spatial frequency selective adaptation, probably of cortical origin. Measureable adaptation is produced by momentary exposure to contrasts as low as 1.25%; nearly maximum adaptation (0.6 log units) is reached with 20% contrast. These findings support the concept of a contrast gain control mechanism in visual cortex, and pose practical problems for visual assessment with the evoked potential.

Adaptation, Physiological↗

Lock-in techniques for the swept stimulus evoked potential.

We discuss the use of synchronous-demodulation (lock-in) techniques for evoked potential retrieval. Application to electronically swept visual displays is emphasized. These techniques permit a visual threshold to be specified in 20 s, but their application to visual assessment requires careful consideration of several problems, notably alleged delay in the instrumentation, specification of the baseline response level, and the nature of EEG interference. In addition, since stimulus waveform information is lost in all lock-in methods, questions concerning what activity is contributing to the measured response must be answered. A technique addressing these issues and combining phase-sensitive detection and vector computation is presented.

Computers↗

The assessment of evoked potential contrast thresholds using real-time retrieval.

Electrophysiologic contrast sensitivity functions (CSF) have been estimated using lock-in amplifier signal retrieval of the visually evoked response (VER). These CSFs were compared with CSFs obtained psychophysically using the same stimulus conditions. The two measures of contrast sensitivity behave similarly in response to variations of temporal and spatial frequency. The major advantage of using real-time retrieval is speed. Threshold for a single spatiotemporal condition can be estimated in as little as 20 sec, making the application of electrophysiologic contrast sensitivity testing feasible for clinical populations.

Adult↗

Contrast sensitivity loss in multiple sclerosis. Selectivity by eye, orientation, and spatial frequency measured with the evoked potential.

Multiple sclerosis can produce highly selective losses in visual function. Psychophysical studies have demonstrated contrast sensitivity deficits for spatial frequencies or for stimulus orientations. Using real-time lock-in retrieval of the visual evoked potential, the authors measured contrast sensitivity in 15 cases with probable or definite multiple sclerosis and acuities of 20/40 or better. Sine-wave grating contrast threshold determinations for three spatial frequencies (1, 4, and 8 cycles/deg) and four orientations (0, 45, 90, and 135 deg) revealed contrast deficits in at least one spatial frequency and orientation in every case. In most cases the visual losses were spotty or multifocal, and not the same in both eyes. Some cases with highly selective patterns of orientation or spatial frequency losses were observed and are discussed in terms of involvement of cortical functional architecture in the disease.

Adult↗

The 20/20 eye in multiple sclerosis.

Using clinical and electrophysiologic measures, we evaluated the visual pathway of patients who had multiple sclerosis, 20/20 Snellen acuity, and no history of optic neuritis. Delayed latencies were found in the transient visual evoked potentials (VEPs) of 38% of the patients, and interocular latency differences were abnormal in 67%. Contrast VEPs were abnormal in 46%. Psychophysical determinations of contrast sensitivity were abnormal in 78%. Only 17% of the patients had dyschromatopsia, 36% had afferent pupillary abnormalities, and 59% had optic nerve pallor or nerve fiber layer loss. Psychophysical contrast evaluations and VEP studies were superior to other clinical evaluations in demonstrating visual dysfunction in these patients.

Evoked Potentials, Visual↗

A neurophysiological model for anomalous correspondence based on mechanisms of sensory fusion.

Normal retinal correspondence is not stable. The arguments for the plasticity of correspondence in normal binocular vision have been given into previous papers (Nelson, 1975, 1977). In this paper, both laboratory research and the clinical strabismus literature are reviewed to show similarities between normal and abnormal binocular vision. In particular, it is argued that sensory fusion (Panum's areas) and anomalous retinal correspondence (AC) obey similar principles, and so a sensory fusional model of AC may be developed. Recent advances in the neurophysiology of binocular vision are reviewed, but current laboratory knowledge cannot account for many phenomena known clinically unless certain postulates are made. Two hypothesized intracortical interactions among binocular disparity detectors, termed disparity domain inhibition and spatial domain facilitation, play key roles in extending the neurophysiology of binocular vision to an account of both normally - and clinically - observed plasticities of correspondence. The fusional model of retinal correspondence developed here from postulated domain interactions contrasts with the older concept of fixed corresponding points, an approach which has failed to provide a unified foundation for the treatment of normal and abnormal binocular vision.

Animals↗

Discrimination of orientation and position disparities by binocularly activated neurons in cat straite cortex.

1. We have examined and compared the ability of binocularly activated striate neurons to make both position disparity and orientation disparity discrimination in the anesthetized (N2O/O2) and paralyzed cat preparation. 2. Accurate knowledge of eye position is essential for disparity studies. Using a retinal projection technique able to detect eye drifts of less than 3' arc per retinal landmark and less than 18' arc cyclorotation disparity, we determined eye drift during the course of 2- to 4-day experiments. After the initial eye rotation due to the anesthesia and the onset of paralysis (see below), rotational drift thereafter was mainly excyclorotatory and, from all causes, rarely totaled more than 4 degrees disparity. All our data have been corrected for this residual cyclorotatory drift. 3. Optimal stimulus orientation disparities were determined from quantitative monocular orientation tuning curves for 74 binocularly activated striate cells (37 simple, 3 hypercomplex I, 31 complex, 3 hypercomplex II) from nine cats. Without exception, the mean optimal stimulus orientation disparity in each of our animals showed a departure from zero disparity equivalent to an incyclorotation of the eyes (mean, 9.2 degrees; range, 2.7 degrees-15.9 degrees). 4. We attribute this mean optimal stimulus orientation disparity shift to ocular cyclorotation as a result of the initial anesthesia and paralysis. Assuming equal intortion, incyclorotation for each eye averages 4.6 degrees. On the assumption that the mean optimal stimulus orientation disparity is zero in normal life, we pooled results from the nine animals about their individual means. For the 74 cells the resulting distribution of the optimal stimulus orientation disparities had a range of about +/-15 degrees (simple cells: SD 4.9 degrees; complex cells: SD 7.4 degrees). 5. We examined the relationship of the sharpness of the orientation tuning curves to ocular dominance, to absolute orientation preference, and to other unit properties. The striking observation was the high correlation between the sharpness of orientation tuning curves for the two eyes of a binocular neuron. For simple cells the mean difference for the half-widths of half-height was only 2.54 degrees, with sharpness showing a high correlation between the two eyes (r=0.915) over half-width at half-heights ranging from 8.5 degrees to 41.8 degrees. Complex cells showed a similar, albeit weaker, correlation. 6. Having shown that, assessed monocularly binocular units show different orientation tunings in the two eyes, we undertook binocular experiments to ascertain if these differences were the optimal disparities of sharply tuned stimulus orientation disparity channels. Using a matrix stimulation paradigm to minimize the effects of spontaneous changes in responsiveness, we have simultaneously extracted bionocular stimulus orientation disparity and position disparity tuning curves from single striate neurons...

Animals↗