Cortical areas in visual awareness.
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Frontal Lobe↗
Biomedical subjects
D A Pollen
Publications and source records attributed to D A Pollen.
Altered expression of transforming growth factor-beta in Alzheimer's disease.
We compared immunohistochemical expression of the transforming growth factor-betas (TGF-beta 1, TGF-beta 2, and TGF-beta 3) using brain tissue from patients with nondominantly inherited Alzheimer's disease (NDAD) (n = 9), autosomal dominantly inherited Alzheimer's disease with linkage to 14q24.3 (FAD-14) (n = 4), and cognitively normal controls (n = 10) to determine whether their pathologic changes are associated with an altered distribution of the TGF-betas. We found increased expression of TGF-beta 2 in large, tangle-bearing neurons with widespread staining of glia in NDAD and FAD-14 patients compared with control cases. This result was confirmed with sandwich ELISA assays of brain tissue, which showed TGF-beta 2 levels in AD and NDAD to average 3.2 times the average level of control cases. Despite proximity of TGF-beta 1 and TGF-beta 3 to the sites of susceptibility loci on chromosomes 19 and 14, we did not find that TGF-beta 1 and TGF-beta 3 were selectively altered in any AD subtypes. However, selective induction of TGF-beta 2 may occur in NDAD and FAD-14.
Aged↗
Interneuronal interaction between members of quadrature phase and anti-phase pairs in the cat's visual cortex.
Interactions between adjacent simple cells recorded simultaneously from the same microelectrode placement were studied by correlational analysis. The receptive fields of pairs of such cells exhibit either 90 degrees (quadrature phase) or 180 degrees (anti-phase) phase relationships. We now show that the majority of quadrature phase pair members do not receive common input from the immediately precedent stage along the visual pathway, nor do these cells interact with each other. The anti-phase pairs show relatively strong mutual inhibition. These results suggest that each of the physically adjacent phase-related simple cells receives excitatory input from a distinct group of pre-cortical cells, and that mutual inhibitions between members of anti-phase pairs are used to construct the inhibitory subzones of these cells. We propose a model which incorporates these new results and provides a parsimonious explanation for the construction of both quadrature phase and anti-phase pairs.
Animals↗
Spatial and temporal frequency selectivity of neurons in visual cortical area V3A of the macaque monkey.
Response properties of neurons in V3A were studied at a retinal eccentricity of 2-4 deg. The distributions of spatial frequency bandwidths and orientation bandwidths were similar to those of neurons in V1. Peaks of spatial frequency tuning curves ranged from 0.35 to 8.0 c/deg with a mean of 1.75 c/deg. Most V3A cells showed lowpass or, less often, broad bandpass temporal frequency selectivity. The mean direction selectivity index was 0.41. The response properties of cells in V3A differed most from those in V1 with respect to the larger receptive field widths in V3A averaging about 4 deg, the consequent larger number of cycles of the preferred grating that fall within the receptive field, and the previously reported profound response suppression incurred when patches of the preferred grating are extended both within and beyond the classical receptive field. The response properties of cells in V3A differed most from those in V3 in that V3A neurons are much less selective to the speed and direction of stimulus motion than are neurons in V3. The overall response properties of cells in V3A are consistent with anatomical evidence that places this cortical area in the visual pathway from V3A to V4 and then to IT.
Animals↗
Responses of simple and complex cells to compound sine-wave gratings.
We have studied the responses of simple and complex cells in the primary visual cortex of the cat to rigidly drifting compound sine-wave gratings as a function of the phase offset between fundamental and harmonic frequencies that both fell within the passband of the cell. Simple cells show phase-dependent increases and decreases in peak and mean response which are predictable on the basis of a cell's line weighting function. However, the amplitudes and phases of the base and harmonic frequencies in the response are, in general, not well predicted by the relationships of these same components in the compound grating stimuli. These distortions are shown to be largely a consequence of the rectification that follows linear summation at the simple cell stage. Such distortions are, in principle, correctable when the responses of a second simple cell, as part of a 180 deg phase pair, are taken into account. Complex cells typically showed a strong nonlinear response component at the difference frequency of drifting compound gratings. This was sometimes accompanied by a linear response component at one, or both, of the separate stimulus frequencies. Information about the absolute phases of the frequency components of a compound grating is not preserved in the nonlinear response of complex cells; however, information about the local phase difference between the gratings is preserved. In effect, the nonlinear component of the complex cell response is proportional to the time-varying signal envelope that results from the mutual interference of stimulus frequencies that fall in the cell's spatial receptive field and frequency passband.
Animals↗
Diversity of complex cell responses to even- and odd-symmetric luminance profiles in the visual cortex of the cat.
We have tested the hypothesis that complex cell receptive fields are made up of subfields which, for a given cell, have either exclusively even or exclusively odd symmetry. To do this we have measured the response of complex cells in the visual cortex of the cat to members of pairs of spatially limited even-symmetric stimuli (single light and dark bars) and pairs of odd-symmetric stimuli ("light-dark" and "dark-light" double bars) successively drifting across their receptive fields. The strength of a cell's response was estimated by measuring the sum of all spikes produced by a stimulus. Some complex cells respond about equally to single light and dark bars; others respond appreciably more to either the light or dark bar. The central tendency of average response histograms was estimated by measuring the mean with respect to position across the width of the receptive field. Many complex cells show distinct spatial offsets between the mean for narrow single light and that for dark bars as well as between means to double bars of opposite phase. Combined offset plots were constructed with the spatial offsets between means for single light bars and single dark bars along the x axis and the offsets between means to double bars of opposite phase along the y axis. There is significant scatter in the combined offset points; some falling at the origin, some at significant distances from the origin along the axes, and others well within each of the four quadrants. These diverse localizations in the offset plots rule out the simple models of complex cell spatial substructure described above and, therefore, imply considerable heterogeneity within the population of complex cells.
Animals↗
Response suppression by extending sine-wave gratings within the receptive fields of neurons in visual cortical area V3A of the macaque monkey.
Even though there are many more cycles of the "optimal" grating extending across the receptive fields of cells in V3A than of cells in V1 and V2, the spatial frequency bandwidths in V3A are no narrower than in V1 or V2. Thus, the inputs to V3A cells are not combined in a phase coherent manner across the entire receptive field. Moreover, the defined receptive fields of cells in V3A are generally surrounded by suppressive regions which are, on average, much stronger than those found for neurons in V1 and V2. Even within the classical receptive field, most neurons in V3A respond far more vigorously to a limited patch of a few cycles of a grating at the preferred spatial frequency than to wider grating stimuli. This intra-receptive field suppression demonstrates a new level of response complexity, and suggests that V3A cells may antagonistically combine nonlinear mechanisms that themselves encode stimulus energy over a restricted region of space and spatial-frequency.
Animals↗
Spatial and temporal frequency selectivity of neurones in visual cortical areas V1 and V2 of the macaque monkey.
The spatial and temporal frequency selectivity of 148 neurones in the striate cortex, V1, and of 122 neurones in the second visual cortical area, V2, of the macaque monkey were studied using sine-wave gratings of suprathreshold contrast drifting over the receptive field at the preferred orientation and direction. Neurones in V1 and V2 were selective for different but partially overlapping ranges of the spatial frequency spectrum. At retinal eccentricities of 2-5 deg from the fovea, the spatial frequency preferences for neurones ranged from 0.5 to 8.0 cycles/deg in V1 and from 0.2 to 2.1 cycles/deg in V2 and were on average almost 2 octaves lower in V2 than in V1. Spatial frequency full band widths in the two cortical areas were in the range 0.8-3.0 octaves, with a mean value of 1.8 octaves, in the parafoveal representation of both V1 and V2, and 1.4 and 1.6 octaves respectively in the foveal representation of V1 and V2. Most neurones in V1 and some in V2 responded well at temporal frequencies up to 5.6-8.0 Hz before their responses dropped off at still higher frequencies. In V1, 68% of the neurones exhibited low-pass temporal tuning characteristics and 32% were very broadly tuned, with a mean temporal frequency full band width of 2.9 octaves. However, in V2 only 30% of the neurones showed low-pass temporal selectivity and 70% of the cells had bandpass temporal characteristics, with a mean full band width of 2.1 octaves. In V2 the minimal overlap of bandpass tuning curves across the temporal frequency spectrum suggests that there are at least two distinct bandpass temporal frequency mechanisms as well as neurones with low-pass temporal frequency tuning at each spatial frequency. A matrix of spatial and temporal frequency combinations was employed as stimuli for neurones with bandpass temporal frequency selectivity in both V1 and V2. The resultant spatio-temporal surfaces provided evidence that a neurone's preference for spatial frequency is essentially independent of the test temporal frequency; however, in V2 there was some tendency for temporal frequency peaks to shift slightly towards lower frequencies when non-optimum values of spatial frequency either above or below the preferred value were tested. Neurones with pronounced directional selectivity were encountered over a wide range of spatial frequencies, although in both cortical areas there was a tendency for an increased incidence of directional selectivity among neurones which were selective for lower spatial frequencies and higher temporal frequencies.
Animals↗
Use of Gabor elementary functions to probe receptive field substructure of posterior inferotemporal neurons in the owl monkey.
The large receptive fields of inferotemporal neurons in the owl monkey were studied with visual stimuli whose luminance profiles were one-dimensional Gabor functions, i.e. sinusoidal gratings within Gaussian envelopes. The members of one set of such patterns all had a full bandwidth at half-amplitude of 0.8 octaves, but different center frequencies and spatial extents. These spatially restricted stimuli were ideal for determining whether one or more than one spatial frequency band projected onto discrete subsections of the neuron's receptive field. The other set of Gabor stimuli comprised sine waves within Gaussian envelopes of constant size, but with different center frequencies and hence different bandwidths. These stimuli allowed assessment of the neuron's spatial frequency selectivity across the full breadth of its receptive field. Results suggest that only one orientation band and one spatial frequency band provide an input onto each inferotemporal neuron under our experimental conditions. The preferred spatial frequencies found (0.2-0.6 c/deg) were all in the very low spatial frequency range for this animal. Calculations show that about 3.5-7.0 full cycles of the optimal grating usually cover the full width of the receptive field, but the observed spatial frequency tuning is not nearly as sharp as that which would be predicted according to phase coherent linear summation. Moreover, at the preferred spatial frequency, the peak response to gratings in the constant aperture series was generally less than the response to the same preferred spatial frequency in the constant relative bandwidth series. These results suggest either incomplete phase coherent summation from contributing subgroups, non-linear processing, or both.
Animals↗
Spatial computation performed by simple and complex cells in the visual cortex of the cat.
Simple and complex cells have been tested with drifting sine-wave and square-wave gratings. Despite the known differences in the response pattern of each cell type to drifting sine-wave gratings, the tuning curves for square-wave gratings for both cell types show a similar secondary response band peaking at one-third the preferred spatial frequency as determined from sine-wave studies. These results establish that both cell types respond predominantly to the third harmonic of square-wave gratings in this frequency range. At the simple cell stage, all the information required to specify the amplitude and phase for channel at a given orientation, direction and spatial frequency can be conveyed by four cells for a given subsection of visual space. At the complex cell stage, the cell's mean firing rate appears to represent the amplitude of a local Fourier coefficient, but phase information is not conveyed in the action potential code.
Animals↗
Phase relationships between adjacent simple cells in the visual cortex.
Adjacent simple cells recorded and "isolated" simultaneously from the same microelectrode placement were usually tuned to the same orientation and spatial frequency. The responses of the members of these "spatial frequency pairs" to drifting sine-wave gratings were cross-correlates. Within the middle range of the spatial frequency selectivity curves, the responses of the paired cells differed in phase by approximately 90 percent. This phase relationship suggests that adjacent simple cells tuned to the same spatial frequency and orientation represent paired sine and cosine filters in terms of their processing of afferent spatial inputs and truncated sine and cosine filters in terms of the output of simple cells.
Animals↗
Intracortical microstimulation of neurons in the visual cortex of the cat.
The response of visual cortex neurons to local intracortical microstimulation was measured in the anesthetized cat. When the recording microelectrode was very close (about 20 micrometers) to the tip of the stimulating electrode, threshold currents as low as 10 micro A were capable of firing neurons. Over a 20-fold range in distance from the site of stimulation, an 80-fold increase in threshold current was observed. The mean latency of activation for 30 neurons tested with intracortical stimulation was 2.88 +/- 0.45 msec. The majority of these cells were probably synaptically activated. The mean threshold current for these neurons was 0.55 +/- 0.12 mA (N = 30). These values were significantly smaller than the thresholds found previously when stimulating electrodes were located on the pia-arachnoid surface of the visual cortex.
Animals↗
Relationship between spatial frequency selectivity and receptive field profile of simple cells.
1. Receptive fields of simple cells in area 17 of the cat were mapped with stationary stimuli. Spatial frequency selectivities of the same cells were measured with drifting sinusoidal gratings. 2. The reconstructed field profile (inverse Fourier transform of selectivity curve) shows qualitative agreement with the mapped profile, and suggests the existence of additional side-lobes in the field. The side-lobes may correspond to the 'unresponsive regions' investigated by Maffei & Fiorentini (1976). 3. Our data suggest that the simple cell may perform approximately linear spatial summation of inputs to the visual system. However, the output of the simple cell is generally non-linear as reflected in its truncated responses to gratings.
Animals↗
Spatial periodicities of periodic complex cells in the visual cortex cluster at one-half octave intervals.
Within individual penetrations in the visual cortex, spatial periodicities of periodic complex cells differ by either one-half or one octave. When data are pooled from neurons subserving the central visual area in many cats, the results indicate that spatial periodicities cluster at one-half octave intervals over a 2 1/2-octave range (0.95 to 5.4 cyc/deg). Thus a relatively small number of such channels spaced at regular intervals along a logarithmic scale within each orientation column may suffice for this stage of spatial processing.
Animals↗
Spatial frequency selectivity of periodic complex cells in the visual cortex of the cat.
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Action Potentials↗
Periodic complex cells in cortical area 19 of the cat.
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Action Potentials↗
Responses of single neurons to electrical stimulation of the surface of the visual cortex.
We have recorded from single neurons in the visual cortex of the cat while stimulating the cortical surface with the same type of electrodes and the same parameters of stimulation which have produced phosphenes in conscious man. Parameters of stimulation which will permit excitation of single cortical neurons with little risk of producing afterdischarges are described. The patterns of excitation of single neurons during surface stimulation and the effects of stimulation on neuroglial cells have also been studied. Surface stimulation also produced marked alterations in cortical excitability as tested with visually presented stimuli. Mechanisms by which single neurons are excited by surface stimulation are also considered.
Action Potentials↗
Responses of complex cells in the visual cortex of the cat as a function of the length of moving slits.
(1) As a step towards specifying the spatial selectivity characteristics of complex cells with spatially periodic substructures, we have studied single cell responses to narrow slits of variable length moved across the receptive field in the preferred direction. In general, the length-response curves were linear over a considerable and sometimes full range until an optimal slit length was reached. (2) In those cells in which the rate of rise of the slit length-response functions decreased before the optimal length was reached, at least 3 factors contribute to the shape of the curve. First, the receptive field shapes of some complex cells are more ovoid or rounded than rectangular, and the summation of responses from excitatory zones of varying optimal lengths itself results in a nonlinear slit length-response function at long slit lengths. Second, central regions may contribute more to cell response than do more lateral regions along the length dimension. Third, a nonlinearity in the slit length-response curve may occur in the upper range of slit lengths as a saturation effect because discharge rates may reach 600/sec, which appears to be close to a limiting firing rate. (3) Some cells believed to be complex during preliminary receptive field testing showed weak inhibitory regions beyond the region of the optimal slit length. Many of these cells also displayed periodic average response histograms to moving slits. The extent and magnitude of the inhibition were variable from cell to cell. In terms of receptive field properties, these cells and 'regular' complex cells seem part of a continuum.
Animals↗