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T J Zumbroich

Publications and source records attributed to T J Zumbroich.

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Postnatal development of corticocortical efferents from area 17 in the cat's visual cortex.

We are interested in the postnatal development of corticocortical connections in the cat's visual cortex. In this study, we injected the anterograde tracer 3H-proline into visual cortical area 17 of kittens, aged 4-70 d, and adult cats to visualize the distribution of terminals of the association projections to areas 18, 19, 21a, and the lateral suprasylvian visual cortex. The density of anterograde label was quantified using computerized image analysis. There was dense labeling at topographically appropriate locations in area 18 in animals of all ages. In 4- and 8-d-old kittens, other extrastriate areas (19, 21a and the lateral suprasylvian cortex) contained only sparse label, localized in a few solitary axons; these areas were densely labeled in animals aged 12 d or more. In kittens aged 4-20 d there was considerable, widespread label within fibers located in the white matter, and many of these axons lay underneath regions of extrastriate, and also striate, cortex that were almost certainly not destined to be persistently innervated by cells at the injection site. This pattern of extensive white matter label was not seen in animals older than 20 d. In each extrastriate region, from the earliest age at which we identified dense cortical innervation from area 17, the terminals were distributed in clusters. At first these patches were mainly in infragranular layers, but later, during the second and third postnatal weeks, they began to appear in more superficial laminae. By 70 d, an adult-like distribution of terminals was found in each extrastriate area: most fibers appeared to end in layers II and III in areas 18, 19, and 21a and centered on layer IV in the medial bank of the middle suprasylvian sulcus in adult cats. We suggest that the development of ipsilateral association projections from area 17 to extrastriate cortex is a 2-stage process. First, cells at a particular point in area 17 send immature fibers in a nonspecific fashion through white matter towards a very wide area of extrastriate cortex. Second, corticocortical axons penetrate extrastriate cortex mainly in patches at topographically appropriate regions and grow to their targets in a specific fashion.

Aging↗

Development of stimulus selectivity and functional organization in the suprasylvian visual cortex of the cat.

We have recorded from single neurons in the medial bank of the middle suprasylvian sulcus (PMLS) of anaesthetized and paralysed cats aged between nine days and eight weeks. Visual responses were assessed qualitatively, by using conventional projected stimuli, and quantitatively for drifting, high-contrast gratings of optimum spatial and temporal frequencies, but varying in orientation and direction of drift. At 9 days of age, some cells in the PMLS were spontaneously active but in three long penetrations only one visually responsive neuron was isolated. Between 9 and 15 days there was a rapid increase in the proportion of responsive units, which first appeared in small clusters in the lower layers (IV, V, VI). During the second and third postnatal weeks, spontaneous activity and the strength of visual responses increased to adult levels, and the proportion of cells showing rapid habituation to visual stimulation decreased. Even before two weeks of age, at least 85% of responsive cells in the PMLS were selective, by quantitative criteria, for image motion along one particular axis, and a majority of these were clearly direction-selective (responding to movement in one direction significantly more strongly than to that in the opposite). By the end of the third postnatal week the proportion of units with strong direction preference reached adult levels. The selective cells were initially more broadly 'tuned', on average, for the direction of motion of a grating (mean half-width in animals of 10-12 days was 32.6 degrees), but the sharpness of tuning improved to reach the adult level (ca. 23 degrees) during the third postnatal week. In animals younger than three weeks a slightly smaller proportion of cells than in adults (but always more than one third of all visually responsive cells) responded to stationary, contrast-modulated gratings. The majority of these cells showed clear selectivity for the orientation of a flashed grating. A few 'non-selective' cells were found in the youngest animals but by the end of the third postnatal week virtually all cells responsive to stationary gratings displayed orientation selectivity. There was always good agreement between the preferred orientations for stationary and drifting gratings. Even before two weeks of age, when responsive cells occurred only in small clusters, there was a clear tendency for neighbouring neurons to have similar or opposite preferred directions, just as in adult cats. By 2-3 weeks of age there were clear progressive shifts in stimulus preference along oblique or tangential tracks.(ABSTRACT TRUNCATED AT 400 WORDS)

Age Factors↗

Visual receptive field properties in the posterior suprasylvian cortex of the cat: a comparison between the areas PMLS and PLLS.

Receptive field (RF) properties of single units were examined in two visual areas in the cat's postero-medial and postero-lateral suprasylvian cortex (PMLS/PLLS). Electrophysiological recordings were made in the paralyzed and anesthetized preparation in corresponding regions of the medial and lateral banks of the lateral suprasylvian sulcus (LS). In both areas, cell samples were obtained from within the same range of A/P co-ordinates. In both samples, cells responded best to moving stimuli, had large RF's, and did not differ with respect to the distributions of their ocular dominance. Binocularity was equally high, but units in PLLS showed significantly less binocular summation. Cells in both areas preferred high velocities, and a high percentage was direction selective, with directions up and away from the vertical meridian being most common. While in PMLS the preferred direction of a unit usually could be classified into a "radial" vs a "circular" category when RF position was taken into account, this was not possible in PLLS. Units in PLLS also had significantly higher spontaneous activity, higher optimal velocity, and larger RF sizes. In PLLS the investigation of background-foreground interactions revealed a large variety of phase-dependent responses. This is in contrast to the clear preponderance of in-phase inhibition and antiphase facilitation effects in PMLS. The results indicate important differences of RF properties for the two areas, but do not yet suggest a clear functional differentiation. These differences reflect in some respects the RF properties of cells in the structures that provide the main afferents to the two areas.

Action Potentials↗

Stimulus selectivity and functional organization in the lateral suprasylvian visual cortex of the cat.

1. We have examined the responses of cells in the postero-medial and postero-lateral lateral suprasylvian areas (p.m.l.s./p.l.l.s.) in the medial and lateral banks of the middle suprasylvian sulcus of the anaesthetized, paralysed cat. 2. Visual responses were assessed qualitatively (for projected spot and bar stimuli) and quantitatively (for drifting, high-contrast gratings of optimum spatial and temporal frequencies, but varying in orientation and direction of drift). There was excellent agreement between qualitative and quantitative estimates of preferred direction of motion. 3. Comparison of responses to motion in the preferred direction and the opposite direction confirmed that the percentage of units with strong directional preference is higher in this region of cortex than in other cortical and subcortical visual structures so far investigated. 4. Cells were comparatively well 'tuned' for the direction of motion of a grating: on average the half-width at half-amplitude for the variation in response around the principal preferred direction was 23.2 deg for p.m.l.s. and 25.3 deg for p.l.l.s., thus falling within the range found for complex cells in area 17. In this and other aspects of their direction selectivity, neurones in p.m.l.s. and p.l.l.s. were very similar to each other. 5. Quantitative analysis of the direction-response functions revealed them frequently to be more complex than previously described. For many cells there were subsidiary response peaks in addition to the main peak at the principal preferred direction. Often there was an accessory peak 180 deg from the optimum direction (i.e. directional preference was incomplete), but there was also a clear tendency for responses to be specifically elevated for directions orthogonal to the principal direction. 6. Tests with stationary, contrast-modulated gratings of the optimum spatial frequency, but differing in orientation, revealed that neurones responsive to such stimuli (though limited to about half the population) were orientation selective, their preferred orientations for stationary and drifting gratings being very similar. 7. The functional architecture of the lateral suprasylvian cortex was studied by means of single and multiple penetrations at different angles to the cortical surface: in most penetrations in p.m.l.s. the preferred directions of cells tended to shift progressively in small steps across the cortex, at a maximum rate of about 360 deg mm-1. However, there were occasional 180 deg differences between neighbouring recording sites or between cells recorded at different depths within a single radial column.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Spatial and temporal selectivity in the suprasylvian visual cortex of the cat.

We recorded from single units in the medial and lateral banks of the posterolateral suprasylvian visual cortex (PMLS/PLLS) of the cat. The responses to drifting high-contrast gratings of optimum orientation and direction of motion, but varying in spatial and temporal frequency, were examined quantitatively for a sample of cells, whose receptive fields covered a wide range of eccentricities. The optimum spatial frequencies (average about 0.2 cycles/deg) were low compared to the values reported for striate cortex but similar to those for area 18. The mean spatial bandwidth (about 2 octaves) was slightly broader than that of cells in other cortical visual areas. The cut-off spatial frequencies ("acuities") covered a wide range, from 0.05 to 2.1 cycles/deg, similar to those of cells in area 18. Responses to drifting sinusoidal gratings were usually dominated by an unmodulated elevation of discharge, although some modulation occurred at the temporal frequency of drift, especially at low spatial frequencies. Modulated responses were relatively stronger in PMLS than in PLLS. For those cells that responded to flashed stimuli, stationary, contrast-modulated gratings presented at different spatial positions typically evoked small responses at the fundamental frequency (dependent on spatial phase) and a larger component at the second harmonic of temporal frequency, with no overall "null-position." The optimum spatial frequency was usually higher than would be predicted by simple summation within the dimensions of the receptive field. Thus, neurons in PMLS and PLLS, like complex cells in areas 17 and 18, behave nonlinearly and their spatial selectivity is determined by "subunits" smaller than their receptive fields. The range of preferred temporal frequencies ranged from less than 2.5 Hz to more than 10 Hz. In their temporal selectivity neurons in PMLS resembled cells in area 17, with little attenuation at low temporal frequencies, whereas there was a tendency for cells in PLLS to prefer higher temporal frequencies, as is common in area 18.

Animals↗

Differences of visual field representation in the medial and lateral banks of the suprasylvian cortex (PMLS/PLLS) of the cat.

We have studied the orderliness of representation of visual space in the medial and lateral banks of the middle suprasylvian sulcus. Penetrations were made either parallel to the sulcus, in one bank or the other, or vertical, thus crossing the sulcus between the postero-medial (PMLS) and posterolateral (PLLS) divisions of this area. In some cases we found clear evidence for topographical order in the representation of the visual field with a tendency (greater in PMLS than in PLLS) for the receptive fields of cells recorded deeper in the walls of the sulcus to lie closer to the area centralis, but along many penetrations the receptive fields were so large and so scattered that no retinotopic arrangement could be discerned. In PMLS the receptive fields of the majority of units we studied were centered below and close to the horizontal meridian, whereas in PLLS they were distributed over both the upper and lower visual fields with an over-representation of the upper field. Receptive fields were significantly larger in PLLS (mean field area = 442.2 deg2) than in PMLS (mean area = 154.4 deg2); there was also less clear correlation between receptive field size and eccentricity in PLLS (correlation coefficient = +0.25) than in PMLS (corr. coeff. = +0.72). Analysis of the distance between the receptive field centres of consecutively recorded units demonstrated that the mean scatter in both PMLS and PLLS amounts to about half the average receptive field diameter. In summary the topographical representation of visual space is less orderly in PLLS, and may involve a wider area of the visual field. These findings may relate to the segregated visual cortical and extrageniculate thalamic connections that the medial and lateral banks of the LS receive.

Animals↗