Responses Of Complex Cells Research Articles

Insensitivity of V1 complex cell responses to small shifts in the retinal image of complex patterns.

An important role for neurons in the early visual system is to convey information about the structure of visual stimuli. However, neuronal responses show substantial variation across presentations of the same stimulus. In awake monkeys, it has been assumed that a great deal of this variation is related to the scatter in eye position (inducing scatter in the retinal position of the stimulus). Here we investigate the implied consequence of this assumption, i.e., that the scatter variation in eye position degrades the decodability of the neural response. We recorded from 50 complex cells in primary visual cortex of fixating monkeys while different complex stimuli were presented. Three types of retinal shifts were considered: natural scatter in the fixation, systematic fixation point shift, and systematic stimulus position shift. The stimulus pattern accounts for >50% of the response variance, always six times that accounted for by the scatter in eye position during fixation. The retinal location of a stimulus had to be shifted by 10-12 min of arc, an amount almost two times larger than the smallest picture element, before the responses changed systematically. Nonetheless, changes of the stimulus at the single pixel level often gave rise to discriminable responses. Thus complex cells convey information about the spatial structure of a stimulus, independent of rigid stimulus displacements on the order of the receptive field size or smaller.

Encoding of binocular disparity by complex cells in the cat's visual cortex.

To examine the roles that complex cells play in stereopsis, we have recorded extracellularly from isolated single neurons in the striate cortex of anesthetized paralyzed cats. We measured binocular responses of complex cells using a comprehensive stimulus set that encompasses all possible combinations of positions over the receptive fields for the two eyes. For a given position combination, stimulus contrast could be the same for the two eyes (2 bright or 2 dark bars) or opposite (1 bright and 1 dark). These measurements provide a binocular receptive field (RF) profile that completely characterizes complex cell responses in a joint domain of left and right stimulus positions. Complex cells typically exhibit a strong selectivity for binocular disparity, but are only broadly selective for stimulus position. For most cells, selectivity for disparity is more than twice as narrow as that for position. These characteristics are highly desirable if we assume that a disparity sensor should exhibit position invariance while encoding small changes in stimulus depth. Complex cells have nearly identical binocular RFs for bright and dark stimuli as long as the sign of stimulus contrast is the same for the two eyes. When stimulus contrast is opposite, the binocular RF also is inverted such that excitatory subregions become suppressive. We have developed a disparity energy model that accounts for the behavior of disparity-sensitive complex cells. This is a hierarchical model that incorporates specific constraints on the selection of simple cells from which a complex cell receives input. Experimental data are used to examine quantitatively predictions of the model. Responses of complex cells generally agree well with predictions of the disparity energy model. However, various types of deviations from the predictions also are found, including a highly elongated excitatory region beyond that supported by a single energy mechanism. Complex cells in the visual cortex appear to provide a next level of abstraction in encoding information for stereopsis based on the activity of a group of simple-type subunits. In addition to exhibiting narrow disparity tuning and position invariance, these cells seem to provide a partial solution to the stereo correspondence problem that arises in complex natural scenes. Based on their binocular response properties, these cells provide a substantial reduction in the complexity of the correspondence problem.

Biology and adoptive cell therapy of Epstein-Barr virus-associated lymphoproliferative disorders in recipients of marrow allografts.

Epstein-Barr virus (EBV) is an ubiquitous herpesvirus which is carried as a latent infection of B lymphocytes and salivary gland epithelial cells in over 90% of normal adults. Latently infected EBV-transformed B cells circulate at low frequency in the blood for the life of the host. These transformed B cells stimulate a heterogeneous and complex host cell response, ultimately leading to the development and maintenance of high frequencies of HLA-restricted T cells specific for the EBV-encoded nuclear antigens EBNA2-EBNA6 and the latency membrane proteins LMP-1 and LMP-2. Responses to latent EBV-encoded proteins are hierarchical with responses to certain epitopes predominating, dependent upon the HLA genotype of the host. Profound suppression of T-cell immunity may permit the emergence of polyclonal, oligoclonal or monoclonal EBV antigen-expressing lymphoproliferative disorders or malignant B-cell lymphomas expressing these latent EBV antigens. Adoptive transfer of small numbers of peripheral blood mononuclear cells or HLA-partially matched T cells from in vitro expanded EBV-specific T-cell lines derived from a seropositive marrow donor has induced durable regressions of bulky, widely metastatic monoclonal EBV lymphomas in a high proportion of cases. This review describes the current state of knowledge and hypothesis regarding the biology and immunology of EBV infection in the normal host, the features of donor, host and virus which contribute to the development of EBV-associated lymphoproliferative diseases and the mechanisms whereby they are controlled by adoptive transfer of immune T cells.

Responses of Complex Cells in Cat Area 17 to Apparent Motion of Random Pixel Arrays

The characteristics of directionally selective cells in area 17 of the cat are studied using moving random pixel arrays (RPAs) with 50% white and 50% black pixels. The apparent motion stimulus is similar to that used in human psychophysics [Fredericksen et al. (1993). Vision Research, 33, pp. 1193–1205]. We compare motion sensitivity measured with single-step pixel lifetimes and unlimited pixel lifetimes. A motion stimulus with a single-step pixel lifetime contains directional motion energy primarily at one combination of spatial displacement and temporal delay. We recorded the responses of complex cells to different combinations of displacement and delay to describe their spatio-temporal correlation characteristics. The response to motion of RPAs with unlimited lifetime is strongest along the preferred speed line in a delay vs displacement size diagram. When using an RPA with a single-step pixel lifetime, the cells are responsive to a much smaller range of spatial displacements and temporal delays of the stimulus. The maximum displacement that still gives a directionally selective response is larger when the preferred speed of the cell is higher. It is on average about three times smaller than the receptive field size. © 1997 Elsevier Science Ltd. All rights reserved.

Computational models of visual neurons specialised in the detection of periodic and aperiodic oriented visual stimuli: bar and grating cells.

Computational models of periodic- and aperiodic-pattern selective cells, also called grating and bar cells, respectively, are proposed. Grating cells are found in areas V1 and V2 of the visual cortex of monkeys and respond strongly to bar gratings of a given orientation and periodicity but very weakly or not at all to single bars. This non-linear behaviour, which is quite different from the spatial frequency filtering behaviour exhibited by the other types of orientation-selective neurons such as the simple cells, is incorporated in the proposed computational model by using an AND-type non-linearity to combine the responses of simple cells with symmetric receptive field profiles and opposite polarities. The functional behaviour of bar cells, which are found in the same areas of the visual cortex as grating cells, is less well explored and documented in the literature. In general, these cells respond to single bars and their responses decrease when further bars are added to form a periodic pattern. These properties of bar cells are implemented in a computational model in which the responses of bar cells are computed as thresholded differences of the responses of corresponding complex (or simple) cells and grating cells. Bar and grating cells seem to play complementary roles in resolving the ambiguity with which the responses of simple and complex cells represent oriented visual stimuli, in that bar cells are selective only for form information as present in contours and grating cells only respond to oriented texture information. The proposed model is capable of explaining the results of neurophysiological experiments as well as the psychophysical observation that the perception of texture and the perception of form are complementary processes.

Responses of Complex Cells in Area 17 of the Cat to Bi-vectorial Transparent Motion

We examined the responses to transparent motion of complex cells in cat area 17 which show directional selectivity to moving random pixel arrays (RPAs). The response to an RPA moving in the cell's preferred direction is inhibited when a second RPA is transparently moving in another direction. The inhibition by the second pattern is quantified as a function of its direction. The response to a pattern moving in the preferred direction is never completely suppressed, not even when a second pattern is moving transparently in the opposite direction.To the extent that supra-spontaneous firing rates signal the presence of the optimal velocity vector, these cells therefore still signal the presence of this line-label stimulus despite additional opposing, or otherwise directed, motion components. The results confirm previous suggestions that, for the computation of motion energy in cat area 17 complex cells, a full opponent stage is not plausible. Furthermore, we show that the response to a combination of two motion vectors can be predicted by the average of the responses to the individual components. Copyright © 1996 Elsevier Science Ltd.

Studies on the response of carrot cells to a Sclerotinia sclerotiorum elicitor: induction of the expression of an extracellular glycoprotein mRNA.

A heat-released elicitor (HRE) obtained from Sclerotinia sclerotiorum mycelium induced a transient increase in the activity of phenylalanine-ammonia lyase (PAL) in a carrot cell culture. Differential display reactions carried out on total RNA extracted from cells exposed for 7 h to the HRE revealed a complex cell response involving the induction and repression of several mRNAs. One of these elicitor-induced mRNAs encodes a 42.6-kDa protein. Northern (RNA) blot analysis and reverse-transcription polymerase chain reaction studies showed that noninduced cells have a basal expression of the 42.6-kDa protein mRNA that was stimulated five- to 10-fold by treatment with the elicitor.

Function of GABAA inhibition in specifying spatial frequency and orientation selectivities in cat striate cortex.

Responses of simple and complex cells in cat striate cortex were studied with moving sine-wave gratings before and during application of the GABAA receptor antagonist bicuculline methiodide. Both simple and complex cells exhibited a broadening of their spatial frequency tuning functions under bicuculline. This was especially evident at spatial frequencies lower than the ones the cell was responding to before the drug administration. The effects cannot be explained by response saturation and could be reversed by cessation of the iontophoresis. The results indicate that the band-pass response characteristics of the spatial frequency response functions of striate cells derive largely from intracortical inhibition. The findings have implications also for the orientation selectivity of cortical cells. Since many geniculate cells are tuned for stimulus orientation at higher spatial frequencies, suppression of the low-spatial-frequency component would remove some of the orientation non-specific response in striate cortical cells and contribute to their orientation selectivity.

Position-specific adaptation in complex cell receptive fields of the cat striate cortex

1. Responses of complex cells in cat striate cortex were studied with flashed light slit stimuli. The responses to slits flashed in different positions in the receptive field were assessed quantitatively before and after periods of prolonged stimulation of one small region of the receptive field. This type of prolonged stimulation resulted in reduced responsivity over a limited zone within the complex cell receptive field. 2. The adaptation-induced responsivity decrement was generally observed in both the ON and OFF response profiles but could also be restricted to one or the other. In general, the magnitude of the response decrements was greatest in the ON response profiles. The adaptation-induced response decrement did not necessarily spread throughout the receptive field but was restricted to a small region surrounding the adapted receptive field position (RFP). Adaptation spread equally widely across the ON and OFF response profiles despite the smaller adaptation effects in the OFF profile. 3. The adaptation effects from repeated stimulation at a single RFP did not spread symmetrically across the receptive field, and a given cell's preferred direction of motion indicated the direction of the asymmetric spread of the adaptation. RFPs that would be stimulated by a light slit originating at the point of adaptation and moving in the preferred direction (preferred side) showed greater adaptation-induced response decrements than did RFPs that would be stimulated by a light slit moving in the opposite direction from the point of adaptation (nonpreferred side). There was significant enhancement of responses at some RFPs on the non-preferred side of the point of adaptation. This asymmetric spread of adaptation could be caused by adaptation of inhibitory connections that contribute to complex cell direction selectivity. 4. The asymmetry of adaptation was significantly different for the ON and OFF response profiles. The asymmetric spread of adaptation for the ON response profile was similar to that observed previously in simple cells with greater decrements in the preferred direction side of the point of adaptation. However, the OFF response profiles showed less directional asymmetry in the spread of adaptation and showed greater decrements at RFPs in the nonpreferred direction side of the point of adaptation. 5. The similarity between the spread of adaptation in simple and complex cells suggests that the adaptation in these cells is occurring through a common mechanism. The directional asymmetry of the spread of adaptation is likely due to a local postsynaptic mechanism of adaptation rather than presynaptic transmitter depletion.

A computational model of the analysis of some first-order and second-order motion patterns by simple and complex cells.

Although spatio-temporal gradient schemes are widely used in the computation of image motion, algorithms are ill conditioned for particular classes of input. This paper addresses this problem. Motion is computed as the space-time direction in which the difference in image illuminance from the local mean is conserved. This method can reliably detect motion in first-order and some second-order motion stimuli. Components of the model can be identified with directionally asymmetric and directionally selective simple cells. A stage in which we compute spatial and temporal derivatives of the difference between image illuminance and the local mean illuminance using a truncated Taylor series gives rise to a phase-invariant output reminiscent of the response of complex cells.

Predicting responses of nonlinear neurons in monkey striate cortex to complex patterns

The overwhelming majority of neurons in primate visual cortex are nonlinear. For those cells, the techniques of linear system analysis, used with some success to model retinal ganglion cells and striate simple cells, are of limited applicability. As a start toward understanding the properties of nonlinear visual neurons, we have recorded responses of striate complex cells to hundreds of images, including both simple stimuli (bars and sinusoids) as well as complex stimuli (random textures and 3-D shaded surfaces). The latter set tended to give the strongest response. We created a neural network model for each neuron using an iterative optimization algorithm. The recorded responses to some stimulus patterns (the training set) were used to create the model, while responses to other patterns were reserved for testing the networks. The networks predicted recorded responses to training set patterns with a median correlation of 0.95. They were able to predict responses to test stimuli not in the training set with a correlation of 0.78 overall, and a correlation of 0.65 for complex stimuli considered alone. Thus, they were able to capture much of the input/output transfer function of the neurons, even for complex patterns. Examining connection strengths within each network, different parts of the network appeared to handle information at different spatial scales. To gain further insights, the network models were inverted to construct "optimal" stimuli for each cell, and their receptive fields were mapped with high-resolution spots. The receptive field properties of complex cells could not be reduced to any simpler mathematical formulation than the network models themselves.

Interactive effects among several stimulus parameters on the responses of striate cortical complex cells

1. Although neurons within the visual system are often described in terms of their responses to particular patterns such as bars and edges, they are actually sensitive to many different stimulus features, such as the luminances making up the patterns and the duration of presentation. Many different combinations of stimulus parameters can result in the same neuronal response, raising the problem of how the nervous system can extract information about visual stimuli from such inherently ambiguous responses. It has been shown that complex cells transmit significant amounts of information in the temporal modulation of their responses, raising the possibility that different stimulus parameters are encoded in different aspects of the response. To find out how much information is actually available about individual stimulus parameters, we examined the interactions among three stimulus parameters in the temporally modulated responses of striate cortical complex cells. 2. Sixteen black and white patterns were presented to two awake monkeys at each of four luminance-combinations and five durations, giving a total of 320 unique stimuli. Complex cells were recorded in layers 2 and 3 of striate cortex, with the stimuli centered on the receptive fields as determined by mapping with black and white bars. 3. An analysis of variance (ANOVA) was applied to these data with the three stimulus parameters of pattern, the luminance-combinations, and duration as the independent variables. The ANOVA was repeated with the magnitude and three different aspects of the temporal modulation of the response as the dependent variables. For the 19 neurons studied, many of the interactions between the different stimulus parameters were statistically significant. For some response measures the interactions accounted for more than one-half of the total response variance. 4. We also analyzed the stimulus-response relationships with the use of information theoretical techniques. We defined input codes on the basis of each stimulus parameter alone, as well as their combinations, and output codes on the basis of response strength, and on three measures of temporal modulation, also taken individually and together. Transmitted information was greatest when the response of a neuron was interpreted as a temporally modulated message about combinations of all three stimulus parameters. The interaction terms of the ANOVA suggest that the response of a complex cell can only be interpreted as a message about combinations of all three stimulus parameters.(ABSTRACT TRUNCATED AT 400 WORDS)

Acquired thermotolerance following heat shock protein synthesis prevents impairment of mitochondrial ATPase activity at elevated temperatures in Saccharomyces cerevisiae

The complex molecular response of cells to sudden temperature changes is a well-characterized phenomenon. Although it is clear that the induction of heat shock proteins provides protection from heat in all of the organisms so far tested, very little is known about the role that this set of proteins plays in cellular homeostasis. Recently, putative roles for hsp60 and hsp70-like proteins have been proposed in Saccharomyces cerevisiae. hsp70-like proteins have been shown to be necessary for translocation of precursor polypeptides into mitochondria and endoplasmic reticulum, while hsp60 is required for the assembly of precursor polypeptides into oligomeric complexes following incorporation into the mitochondrial matrix. In this paper, we report that a brief temperature shock (44 °C) impairs coupling of oxidative phosphorylation in S. cerevisiae as measured indirectly by the CI-CCP/oligomycin assay. Furthermore, at high temperature oligomycin stimulates rather than inhibits oxygen uptake under nonthermotolerant conditions. Pretreatment of cells for a short period of time at 37 °C, prior to exposure to higher temperatures rescues the capacity to maintain coupling between oxidative phosphorylation and electron transport. Inhibition of cytoplasmic RNA or protein synthesis during heat shock prevents the protection of this mitochondrial activity. We propose that one of the roles of the induction of heat shock proteins (or related activities) is to protect mitochondrial ATPase activity under conditions of further increase in temperature.

Temporal encoding of two-dimensional patterns by single units in primate primary visual cortex. I. Stimulus-response relations

1. Previously we developed a new approach for investigating visual system neuronal activity in which single neurons are considered to be communication channels transmitting stimulus-dependent codes in their responses. Application of this approach to the stimulus-response relations of inferior temporal (IT) neurons showed that these carry stimulus-dependent information in the temporal modulation as well as in the strength of their responses. IT cortex is a late station in the visual processing stream. Presumably the neuronal properties arise from the properties of the inputs. However, the discovery that IT neuronal spike trains transmit information in stimulus-dependent temporally modulated codes could not be assumed to be true for those earlier stations, so the techniques used in the earlier study were applied to single-striate cortical neurons in the studies reported here. 2. Single-striate cortical neurons were recorded from three awake, fixating rhesus monkeys. The neurons were stimulated by two sets of patterns. The first set was made up of 128 black-and-white patterns based on a complete, orthogonal set of two-dimensional Walsh-Hadamard functions. These stimuli appear as combinations of black-and-white rectangles and squares, and they fully span the range of all possible black-and-white pictures that can be constructed in an 8 x 8 grid. Except for the stimulus that appeared as an all-white or all-black square, each stimulus had equal areas of white and black. The second stimulus set was made up of single bars constructed in the same 8 x 8 grid as the Walsh stimuli. These were presented both as black against a gray background and white against a gray background. The stimuli were centered on the receptive field, and each member of the stimulus set was presented once before any stimulus appeared again. 3. The responses of 21 striate cortical neurons were recorded and analyzed. Two were identified as simple cells and the other 19 as complex cells according to the criteria originally used by Hubel and Wiesel. The stimulus set elicited a wide variety of response strengths and patterns from each neuron. The responses from both the bars and the Walsh set could be used to differentiate and classify simple and complex cells. 4. The responses of both simple and complex cells showed striking stimulus-related strength and temporal modulation. For all of the complex cells there were instances where the responses to a stimulus and its contrast-reversed mate were substantially different in response strength or pattern, or both.(ABSTRACT TRUNCATED AT 400 WORDS)

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 responses 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.

Responses of simple and complex cells to compound sine-wave gratings

Responses of simple and complex cells to compound sine-wave gratings

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.

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.

Computer Modeling of Batteries from Nonlinear Circuit Elements

Circuit analogs for a single battery cell have previously been composed of resistors, capacitors, and inductors. This work introduces a nonlinear circuit model for cell behavior. The circuit is configured around the PIN junction diode, whose charge‐storage behavior has features similar to those of electrochemical cells. A user‐friendly integrated circuit simulation computer program has reproduced a variety of complex cell responses including electrical isolation effects causing capacity loss, as well as potentiodynamic peaks and discharge phenomena hitherto thought to be thermodynamic in origin. However, in this work, they are shown to be simply due to spatial distribution of stored charge within a practical electrode.

A complex-cell receptive-field model.

The time course of the response of a single cortical neuron to counterphase-grating stimulation may vary as a function of stimulation parameters, as shown in the preceding paper (19). The poststimulus-time histograms of the response amplitudes against time are single or double peaked, and where double peaked, the two peaks are of equal or unequal amplitudes. Furthermore, the spatial-phase dependence of cortical complex-cell responses may be a function of spatial frequency, so that the receptive field appears to have linear spatial summation at some spatial frequencies and nonlinear spatial summation at others (19). In the first part of this paper, we analyze a model receptive field that displays this behavior, and in the second part experimental data are presented and analyzed with regard to the model. The model cortical receptive field in its simplest form contains (two rows) of geniculate X-cell-like, DOG (difference-of-Gaussians)-shaped, center-surround antagonistic, circular-input subunits. We propose nonlinear summation between these two subunits, by introducing a half-wave rectification stage before pooling. The model is tested for the responses it predicts for the application of counterphase-grating stimulation. This simple model predicts the appearance of three response forms as a function of counterphase-stimulation parameters. At periodic spatial frequencies the expected-response histogram has a single peak, whose amplitude has a sinusoidal dependence on spatial phase. At spatial frequencies halfway between these, the expected-response histogram has two equal peaks whose amplitudes have a full-wave rectified sinusoidal dependence on spatial phase. At all intermediate spatial frequencies the expected-response histogram has a "mixed" form; the histogram appears sometimes with one peak, sometimes with two equal peaks, and generally with two peaks of unequal amplitude, as a function of spatial phase. Null responses are expected to appear at specific spatial phases only for the periodic spatial frequencies that give "pure" response time courses as in paragraph 5 above, and not in the more common mixed response case of paragraph 6. The analysis procedure described in the preceding paper (19) is used, separating the odd and even Fourier components of the response histograms reflecting the receptive-field intrasubunit linear summation and intersubunit nonlinear summation, respectively. We propose that this model may be used as a working hypothesis for the analysis of these aspects of the various cortical receptive-field types. Experimental data are described and discussed in terms of the model.(ABSTRACT TRUNCATED AT 400 WORDS)