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M S Cynader

Publications and source records attributed to M S Cynader.

At least 91 records · Page 5Linked to original sources

Surface organization of orientation and direction selectivity in cat area 18.

Two-dimensional maps of orientation and direction preference were made in area 18 of the cat's visual cortex using multiple electrode penetrations 150-300 micron apart. The maps were then analyzed by autocorrelation and Fourier transformation. The power spectrum of the orientation map was sharply peaked below the theoretical cutoff frequency for the sampling function (the pattern of electrode penetrations) used to obtain the map. This suggested that it would be possible to interpolate orientation values between the sample points in the cortex. This was done and the resulting fine-grain maps of orientation preference were studied. Empirical testing showed that the interpolated orientation values were accurate to within +/- 30 degrees. Study of the fine-grain maps showed that iso-orientation domains for broad orientation ranges (0 degrees-90 degrees) were branching bands running from an anterolateral to a posteromedial direction across the surface of the cortex with a periodicity of 1.25 +/- 0.13 mm. Domains for smaller orientation ranges (0 degrees-30 degrees) were periodically spaced, but somewhat irregularly shaped patches. The orientation maps contained numerous point singularities where orientation changed discontinuously. These were spaced about 750 micron apart and most were surrounded by a single 180 degrees cycle of orientations. Autocorrelation analysis of the maps of preferred direction revealed local clustering that extended over a distance of 250-300 micron, but spectral analysis failed to reveal any evidence of periodicity. The absence of periodicity was probably due to a relatively large number of 180 degrees reversals in the map of direction preference, which do not affect the continuity of the orientation map and are not sufficiently numerous to destroy local continuity in the direction map. It is shown that the direction map, if it is to be as continuous as possible, must nevertheless contain lines across which direction preference reverses by 180 degrees. These lines run from one orientation singularity to another. Evidence that the direction map approaches this degree of continuity is presented.

Action Potentials↗

Physiological segregation of geniculo-cortical afferents in the visual cortex of dark-reared cats.

Transneuronal autoradiography in cats dark-reared from birth shows that geniculate terminals from left and right eyes are spread throughout layer IV in area 17 of the visual cortex. In area 18 however, segregation into alternating left and right eye ocular dominance patches appears to be normal. We attempted to correlate this difference between the two areas with the ocular dominance distribution of physiological activity of the geniculate afferents. To do this, microelectrode recordings of multi-unit activity, believed to represent the activity of a number of geniculate terminal axons in the vicinity of the electrode tip, were made in layer IV of areas 17 and 18 in dark-reared cats. The ocular dominance distribution of this activity was predominantly monocular in area 18, as the anatomical results predicted, but surprisingly there were also many regions in area 17 where activity could only be elicited by stimulation of one eye. This has to be reconciled with the anatomical results showing that inputs from both eyes are present throughout layer IV in area 17 of dark-reared cats. Reasons for the discrepancy are discussed.

Afferent Pathways↗

Response properties of single units in the accessory optic system of the dark-reared cat.

The visual responses of single units in the lateral and dorsal terminal nuclei (LTN and DTN) of the accessory optic system (AOS) were studied in adult cats reared in total darkness. In the LTN of the normal cat equal numbers of cells prefer upward and downward vertical stimulus motion (previous results). While direction selectively continued to be a characteristic property of LTN and DTN units in dark-reared animals, the distribution of preferred and non-preferred directions of LTN cells was radically altered such that almost every LTN cell examined in the dark-reared cat preferred downward stimulus motion. In contrast, the distribution of preferred directions among DTN cells was largely unaffected by dark rearing. Both normal and dark-reared cat DTN cells responded best to stimuli moving horizontally toward the recorded hemisphere. The velocity preferences of DTN units of the dark-reared cat were, however, much slower than those of normal DTN units. LTN units responding to downward motion in dark-reared cats showed similar velocity preferences to those downward direction-selective LTN units in normal animals. Unlike the highly binocular responses of AOS cells encountered in the normal cat, the ocular dominance distribution obtained from units in the LTN and DTN of the dark-reared cat is completely monocular, favoring the contralateral eye. Thus, dark rearing renders the distribution of preferred directions most affected in the LTN, velocity preference most affected in the DTN and ocular dominance strongly affected in both nuclei. The physiological response properties of the dark-reared cat presented in this report bear a close resemblance to those we have described in the AOS of acutely decorticated animals (previous results). Data obtained from the dark-reared cat support our earlier suggestion that the visual cortex is a major source of upward direction selectively, high-velocity tuning and ipsilateral eye input for AOS cells. Some of the functional consequences of these findings are discussed in relation to frontal eye placement and optokinetic nystagmus.

Animals↗

Spatial receptive-field properties of direction-selective neurons in cat striate cortex.

Responses of direction-selective neurons in cat striate cortex (area 17) were studied with flashed-bar stimuli. Spatial parameters of interactions within the receptive field giving rise to direction selectivity and of receptive-field subunits were quantitatively determined for the same cells and correlated. A bar stimulus flashed sequentially at two nearby locations in the receptive field produced direction-selective behavior comparable with that elicited by continuously moving stimuli. Each cell exhibited a characteristic optimal spatial displacement, Dopt, for which responses in the presumed preferred and null directions were maximally distinct. In all cases, Dopt was much smaller than the receptive-field size. The spatial structure of receptive fields in simple cells was studied using single narrow-bar stimuli flashed at different locations in the receptive field. The resulting line-weighting function exhibited alternating regions of ON and OFF responses having a characteristic spatial period or wavelength, lambda. Spatial subunit structure in complex cells was determined by flashing two bars simultaneously in the receptive field. The response as a function of bar separation was again a wavelike function having a spatial wavelength, lambda. Values of the optimal displacement for direction selectivity, Dopt, showed a clear relationship with the spatial wavelength, lambda, for a given unit. Dopt was also correlated to a somewhat lesser degree with receptive-field size. Generally, the ratio of Dopt to lambda was approximately 1/10 to 1/4, in agreement with theoretical predictions by Marr and Poggio. Taken together with the findings of Movshon et al., these results indicate a systematic relationship between Dopt and the spatial frequency of a sinusoidal grating, which is optimal for that cell. Such a relationship is consistent with the results of human psychophysical experiments on apparent motion.

Animals↗

A time-based stereoscopic depth mechanism in the visual cortex.

We propose a new depth mechanism, operating in the visual cortex, which uses temporal as well as spatial cues. By varying the timing of input to the two eyes, and the position of stimuli on the two receptive fields, we show that both temporal and spatial disparities influence the binocular responses of single cortical cells.

Animals↗

Sensitivity of cat primary auditory cortex (AI) neurons to the direction and rate of frequency modulation.

Responses of 65 single auditory cortex (AI) neurons to frequency-modulated (FM) sweeps with different rates and direction of frequency change were examined quantitatively. Most units responded differentially depending on the characteristics of the FM sweep stimulus. Sixty-five percent of the units encountered responded at least twice as well for one direction of the FM sweep as for the other direction. Of these direction selective neurons, 67% preferred downward-directed FM sweeps (i.e. changing from high to low frequencies) while only 33% preferred upward-directed FM sweeps. The preference for downward-directed FM sweeps was especially clear in EI cells. In addition, cortical neurons often displayed sensitivity to the rate of frequency modulation (speed sensitivity).

Acoustic Stimulation↗

Responses of single neurones in cat auditory cortex to time-varying stimuli: frequency-modulated tones of narrow excursion.

In the primary auditory cortex of cats anaesthetized with nitrous oxide, single neurones were examined with respect to their responses to tone bursts and linear modulations of the frequency of an on-going continuous tone. Using FM ramps of 2.0 kHz excursion and varying centre frequency, each of 39 neurones was examined for its preference for the direction of frequency change of a ramp whose centre frequency was varied in and around the neurone's response area. Direction preference was strictly associated with the slopes of the cell's spike count-versus-frequency function over the frequency range covered by the ramp. Preferences for upward- and downward-directed ramps were associated with the low- and high-frequency slopes of the spike count function, respectively. The strength of the cell's direction preference was associated with the relative steepness of the spike count function over the frequency range covered by the ramp. The timing of discharges elicited by the frequency modulations was found to be the sum of the cell's latent period for tone bursts plus the time after ramp onset that the stimulus frequency fell within the neurone's response area. The implications of these data for the processing of narrow and broad frequency-modulated ramps are discussed.

Animals↗

Some neural mechanisms in the cat's auditory cortex underlying sensitivity to combined tone and wide-spectrum noise stimuli.

In the auditory cortex of nitrous oxide-anesthetized, muscle-relaxed cats, single neurons were studied for their responsiveness to pure tones that were mixed acoustically with simultaneously gated wide-spectrum noise bursts presented using a calibrated sealed stimulating system. The intensities of both the tone and the noise were systematically varied, with a view to ascertaining the sensitivity of cortical cells to a characteristic frequency tone delivered in the presence of a noise mask. Neurons for which wide-spectrum noise provided a net excitatory influence typically displayed a 'strong-signal capture' effect; that is, the cell's responses were dominated by whichever of the two elements of the combined stimulus was the more effective when tested separately. These cells generally had monotonic tone rate intensity functions. Most of the cells that were suppressed by the noise displayed nonmonotonic pure tone rate intensity functions. When nonmonotonic cells were studied with the combined stimulus, the noise was found to produce an intensity-dependent suppression of their tone-evoked responses that could not be overcome by elevating the tone intensity. In contrast, for the minority of monotonic neurons whose tone-evoked responses were suppressed by noise, that suppression could be overcome by raising the tone intensity. None of the cells in the sample responded in a sustained fashion to continuous noise. In each of 11 cases examined, the effect of a continuous noise mask was to elevate tone thresholds and to prolong latent periods for tones; the magnitude of both of these effects depended on the intensity of the continuous noise mask.

Animals↗

Somatosensory cortical map changes following digit amputation in adult monkeys.

The cortical representations of the hand in area 3b in adult owl monkeys were defined with use of microelectrode mapping techniques 2-8 months after surgical amputation of digit 3, or of both digits 2 and 3. Digital nerves were tied to prevent their regeneration within the amputation stump. Successive maps were derived in several monkeys to determine the nature of changes in map organization in the same individuals over time. In all monkeys studied, the representations of adjacent digits and palmar surfaces expanded topographically to occupy most or all of the cortical territories formerly representing the amputated digit(s). With the expansion of the representations of these surrounding skin surfaces (1) there were severalfold increases in their magnification and (2) roughly corresponding decreases in receptive field areas. Thus, with increases in magnification, surrounding skin surfaces were represented in correspondingly finer grain, implying that the rule relating receptive field overlap to separation in distance across the cortex (see Sur et al., '80) was dynamically maintained as receptive fields progressively decreased in size. These studies also revealed that: the discontinuities between the representations of the digits underwent significant translocations (usually by hundreds of microns) after amputation, and sharp new discontinuous boundaries formed where usually separated, expanded digital representations (e.g., of digits 1 and 4) approached each other in the reorganizing map, implying that these map discontinuities are normally dynamically maintained. Changes in receptive field sizes with expansion of representations of surrounding skin surfaces into the deprived cortical zone had a spatial distribution and time course similar to changes in sensory acuity on the stumps of human amputees. This suggests that experience-dependent map changes result in changes in sensory capabilities. The major topographic changes were limited to a cortical zone 500-700 micron on either side of the initial boundaries of the representation of the amputated digits. More distant regions did not appear to reorganize (i.e., were not occupied by inputs from surrounding skin surfaces) even many months after amputation. The representations of some skin surfaces moved in entirety to locations within the former territories of representation of amputated digits in every monkey studied. In man, no mislocation errors or perceptual distortions result from stimulation of surfaces surrounding a digital amputation.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Alterations in response properties in the lateral and dorsal terminal nuclei of the cat accessory optic system following visual cortex lesions.

The response properties of cells in the lateral (LTN) and dorsal (DTN) terminal nuclei of the accessory optic system (AOS) were examined in 14 cats which underwent unilateral visual cortex ablation. Following decortication, single units in the LTN and DTN no longer showed the high degree of binocular convergence characteristic of the intact animal, but instead LTN and DTN units became almost completely dominated by the contralateral eye. In addition, responsivity of LTN and DTN cells to high stimulus velocities was abolished by removal of cortical input. This decrement in high velocity response was observed in both the excitatory and the inhibitory components of the velocity response profile. While the incidence of direction selective neurons in both the LTN or the DTN was not affected by decortication, the distribution of preferred and nonpreferred directions was dramatically altered in the LTN, and to a lesser extent in the DTN. In the LTN, there was a severe reduction in the number of cells which displayed maximal excitation for upward stimulus motion. Instead, most LTN units in the decorticate cat preferred downward directed stimulus motion. In the DTN, most units still preferred horizontal stimulus motion as in the intact animal, but the overall distribution of preferred directions displayed a clear downward vertical vector component. In other respects, such as receptive field size and position in visual space, on/off responses, and resting discharge rate, LTN and DTN units appeared unaffected by cortical lesions. These experiments demonstrate that the cortical input to the LTN and DTN plays a highly significant role in the formation of response properties of cells located in these nuclei. The results presented in this report indicate that the visual cortex is a major source of ipsilateral eye input, high velocity responses, and upward direction selectivity for the AOS units examined in these experiments.

Animals↗

Electrophysiology of lateral and dorsal terminal nuclei of the cat accessory optic system.

Visual responses were examined quantitatively in 96 units in the lateral (LTN) and dorsal (DTN) terminal nuclei of the cat accessory optic system (AOS). The receptive fields of LTN and DTN cells were quite large, with an average diameter of approximately 60 degrees. Individual cell receptive fields, which could be as small as 30 degrees vertically by 15 degrees horizontally or as large as 100 by 100 degrees, always included the area centralis. Large, moving textured stimuli provoked optimal modulation in these cells. In response to a 100 by 80 degrees random-dot pattern moving at a constant velocity, nearly all cells in both the LTN and DTN displayed a high degree of direction selectivity. Directional response profiles were subjected to a vector analysis that generated two quantities proportional to the direction and magnitude of the major excitatory (E vectors) and inhibitory (I vectors) responses of individual cells. Directional vectors of the LTN displayed a strikingly bimodal distribution: E vectors of individual LTN cells pointed either upward (25 of 49) or downward (23 of 49). I vectors also pointed either up or down in a direction opposite to that of the E vector for the same cell. E and I vectors in both LTN and DTN units were separated by approximately 180 degrees. With few exceptions, E vectors of DTN cells pointed in a horizontal-medial direction, while DTN I vectors pointed in a horizontal-lateral direction. A relatively broad range of stimulus velocities (0.8-102.4 degrees/s) evoked maximal excitation in individual LTN units. The majority of LTN cells, however, achieved maximal excitation at velocities between 0.8 and 12.8 degrees/s. The deepest inhibition was elicited over a range of velocities from 0.2 to 102.4 degrees/s, with two major peaks at 0.8 and 12.8 degrees/s. A similar range of velocity sensitivity was observed in DTN cells: maximal excitation was obtained for stimulus velocities from 1.6 to 102.4 degrees/s, with most DTN cells showing the greatest excitatory response between 6.4 and 12.8 degrees/s. A broad range of inhibitory velocity tuning was also observed in DTN units, with most cells exhibiting the deepest inhibitory modulation at 25.6 degrees/s. The majority of LTN and DTN units were driven most effectively through the eye contralateral to the recording site. Nonetheless, a large percentage of LTN (78%) and DTN (93%) cells could be driven to some extent through both eyes. Despite this conspicuous ipsilateral eye influence, no units were found in either the LTN or the DTN that were driven solely through the ipsilateral eye.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Basic functional organization of second auditory cortical field (AII) of the cat.

The spatial representations of response characteristics to acoustic stimuli within the ventral primary auditory cortex (AI) and the second auditory cortical field (AII) were mapped in the left hemispheres of 14 anesthetized adult cats using closely spaced (200-500 micron) microelectrode penetrations to record cortical responses from small cell clusters. Responses were activated by 200-ms tone bursts (1/s). The obtained response characteristics were l) frequency selectivity, i.e., characteristic frequency (CF) for narrowly tuned responses and center frequency (arithmetic mean of bandwidth 10-15 dB above threshold) for broadly tuned responses; 2) threshold at CF; 3) bandwidth 10 dB above threshold (Q10); 4) binaural interaction class. CF maps. In AI a strictly tonotopically organized CF map was observed with a frequency gradient from high to low along a rostral/caudal axis. At the dorsal end of AII a 0.5- to 1-mm-wide stripe with clear but not strict tonotopic organization was seen, paralleling the organization of AI. In the ventral part of AII a tonotopic organization concordant with AI was found to be still present. However, it was significantly blurred by the strong variability of the CFs, local low-frequency islands, and increasing bandwidth of receptive fields. Q10 maps. In general, narrowly tuned responses were obtained in AI with increasing bandwidth toward the AI/AII border. In AII, broadly tuned responses were usually obtained with a further decrease of the Q10 value along a dorsal/ventral axis. The Q10 at the transition from AI to AII can be described by Q10 = 1.6 X CF0.36 X Q10 in AI is usually higher and Q10 in AII is usually lower than this transition value. Threshold. The sensitivity of AI responses to tonal stimuli was slightly greater than in AII. Across the AI/AII border, a drop in the sensitivity of 10-15 dB was usually observed. The variability in the thresholds of adjacent locations was much higher in AII than in AI. Binaural interaction. Aggregates of neurons with similar binaural properties in ventral AI were "band" shaped and ran roughly orthogonal to isofrequency contours. The course of the binaural bands showed major variations with possible interruptions. The most ventral AI band was EI (contralateral excitation, ipsilateral inhibition) in about 70% of the investigated cortices. In AII binaural aggregates had polymorphic shapes. The sizes and principal orientations of these binaural patches varied widely in different cortices.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Competitive neuronal interactions underlying amblyopia.

Amblyopia, a Greek word meaning "blunt sight" refers to a common clinical condition characterised by lowered visual capacities in one or both eyes. Several different types of amblyopia have been defined and in the most common forms, strabismic and anisometropia amblyopia, one eye finds itself at a severe disadvantage relative to its counterpart. The evidence to be presented in this article will show that the reduction of visual capacities in one eye results from competition between binocular inputs rather than simply reflecting disuse of one eye. In addition, evidence will be represented to show that this binocular competition occurs in the visual cortex rather than in the periphery. In fact, there appear to be at least two separate levels of binocular competition within the visual cortex. Under most circumstances, it is possible to induce or to reverse amblyopia only during a well-defined sensitive period early in the life of the organism. With special efforts, however, prolongation of this sensitive period can be achieved and plasticity of cortical ocular dominance maintained into adulthood.

Amblyopia↗

A sustained input to the direction-selective mechanism in cat striate cortex neurons.

Direction-selective neurons in cat striate cortex were tested with bar-shaped stimuli, sequentially flashed at spatially displaced positions chosen to elicit maximal direction selectivity. Temporally overlapping flash exposures of prolonged duration (400-1000 ms) were employed at a series of onset asynchronies to explore the nature of temporal tuning of the direction-selective mechanism. In most neurons studied, direction selectivity was found to be supported by a surprisingly broad range of stimulus onset asynchronies, which was greater for longer exposure durations. These findings imply the existence of a sustained input to the direction-selective mechanism, in spite of the relatively transient nature of most cortical neurons' step responses. A model is described to illustrate how different front-end temporal filters can affect the dependence of two-flash direction selectivity on stimulus onset asynchrony. The versions of the model which successfully predict the form of the observed responses are those which combine inputs from sustained and transient filters.

Animals↗

Neuronal activity in primate visual cortex assessed by immunostaining for the transcription factor Zif268.

It is now well established that environmental signals mediated via neurotransmitters and hormones can induce responses in cells which involve a cascade of receptors, G proteins, and second messengers. These in turn can induce transcription factors which regulate long-term changes in gene expression. It has been proposed that the stimulus-transcription coupling properties of these DNA-binding proteins can be exploited to visualize activated neurons by way of immunostaining. We have used standard immunohistochemical techniques to detect the expression of one specific transcription factor, Zif268, in the visual cortex (area 17, V1) of vervet monkeys (Cercopithecus aethiops). Immunopositive neurons were present in large numbers throughout the visual cortex of the normal animal, being concentrated in layers 2/3 and 6 and at moderate levels in 4C beta and 5. To determine if Zif268 expression was affected by visual stimulation in the monkey, we restricted light input to one eye with the aim of revealing ocular-dominance columns in striate cortex. We found that short-term monocular deprivation induced either by enucleation, intravitreal TTX injection, or eyelid suturing resulted in dramatic changes in Zif268 levels, revealing vertically oriented columns of reduced Zif268 staining interdigitated with columns of normal expression. Furthermore, these columns were discernible after just 2 h of monocular blockade. A comparison of the ocular-dominance pattern obtained with Zif268 immunostaining and cytochrome oxidase histochemistry in long-term monocularly deprived animals showed a coincident reduction of both markers along columns that were precisely aligned in adjacent sections, indicating that Zif268 expression is restricted to cortical regions of high metabolic activity. Simultaneous immunostaining for Zif268 and the calcium-binding proteins calbindin and parvalbumin showed a negative correlation, suggesting that the Zif268 protein may be expressed selectively within excitatory neurons. A similar approach with immunostaining for neurofilament and microtubule-associated proteins (SMI-32 and MAP2) revealed pyramidal neurons which were regularly found to contain a Zif268-positive nucleus. Furthermore, confocal images of lucifer yellow filled neurons possessing Zif268-positive nuclei all showed pyramidal morphology. Taken together, these results point to activity-dependent expression of Zif268 within a subset of excitatory neurons.

Animals↗

Vernier acuity of neurones in cat visual cortex.

The ability of human observers to detect Vernier breaks of as little as 5 s arc has been termed hyperacuity as this distance is substantially less than the angular separation of the bars of the highest spatial frequency of grating (approximately 1 arc min) that can be detected. Although the visual cortex is a likely candidate for the location of detectors involved in this performance, it is not known whether there are cells sensitive enough to detect deviations from co-linearity that are small compared with their spatial resolution (defined in terms of the highest spatial frequency that the cell can detect). We report here the results of physiological experiments on single units in area 17 of the cat visual cortex in which we studied the effect of introducing a Vernier break into a bar stimulus moved across the receptive field of the cell at a constant velocity. Our results show that the responses of most simple and complex cells are significantly reduced by the introduction of a Vernier break that is substantially smaller than the spatial resolution of the cell. The most sensitive cells in our sample could discriminate Vernier offsets of 3-6 arc min with a reliability of approximately 70%. This was much smaller than their spatial resolution, which was in the range 25-30 arc min. We interpret these results in terms of mechanisms that could underly the orientation selectivity of cortical neurones and suggest how our results relate to human Vernier acuity.

Animals↗