Responses of neurons of the extrastriate cortex of the cat brain to moving stimuli of different forms

We examined responses of neurons of the field 21b of the cat brain cortex to presentation of moving visual stimuli of different forms. Characteristics of the responses of about 54% of the studied neurons showed that in these cases configurations of the contours of moving stimuli were to a certain extent discriminated. Most neurons selectively reacting to changes in the form of the stimulus were dark-sensitive units (they generated optimum responses to presentation of dark visual stimuli on the light background). Detailed examination of the spatial infrastructure of receptive fields (RFs) of the neurons and comparison of this structure with the selectivity of neuronal responses showed that there is no significant correlation between static organization of the RF and responses of the neuron to the movements of stimuli of different forms. We hypothesize that the dynamic infrastructure of the RF and the combined activity of functional groups of neurons, whose RFs spatially overlap the RF of the neuron under study, play a definite role in the mechanisms responsible for neuronal discrimination of the form of the visual stimulus. Neirofiziologiya/Neurophysiology, Vol. 38, No. 1, pp. 61–71, January–February, 2006.     

Predictive Feedback Can Account for Biphasic Responses in the Lateral Geniculate Nucleus

Biphasic neural response properties, where the optimal stimulus for driving a neural response changes from one stimulus pattern to the opposite stimulus pattern over short periods of time, have been described in several visual areas, including lateral geniculate nucleus (LGN), primary visual cortex (V1), and middle temporal area (MT). We describe a hierarchical model of predictive coding and simulations that capture these temporal variations in neuronal response properties. We focus on the LGN-V1 circuit and find that after training on natural images the model exhibits the brain's LGN-V1 connectivity structure, in which the structure of V1 receptive fields is linked to the spatial alignment and properties of center-surround cells in the LGN. In addition, the spatio-temporal response profile of LGN model neurons is biphasic in structure, resembling the biphasic response structure of neurons in cat LGN. Moreover, the model displays a specific pattern of influence of feedback, where LGN receptive fields that are aligned over a simple cell receptive field zone of the same polarity decrease their responses while neurons of opposite polarity increase their responses with feedback. This phase-reversed pattern of influence was recently observed in neurophysiology. These results corroborate the idea that predictive feedback is a general coding strategy in the brain.     

Spatio-temporal organization of receptive fields of the cat striate cortex

The responses of cortical cells to gratings and bars were compared. The excitatory and inhibitory on-and off-zones of a simple cell are composed of on- and off-subfields of CGL. Any zone is formed by an opponent pair of subfields one of which gives an excitatory effect, the other — inhibitory. Such organization assumes the linear properties of a simple field. The deviations from linearity are due to spatial dis-placements of the subfields, heterogeneity of subfields, or the absence of one subfield in the opponent pair. Subfields may be both phasic and tonic, even in the same RF. Analysis of the most common type of a complex cell with modulated responses against unmodulated background shows that a mask eliminating stimulation of any half of the RF causes (according to the theory of filtres) increasing the bandwidth due to the increase or the appearance of responses to side low and high frequencies. The modulated components of the responses from both halves of the RF are out of phase. Analysis of this fact and the responses to thin bars suggests that a complex field is formed by linear and nonlinear subsystems converging onto output neuron. Other types of complex fields are organized by different combinations of subsystems. Limited in area by masking the RF responds to much higher spatial frequencies than the whole RF. The optimal frequency in two-dimensional spatial frequency characteristics of the RF does not change with orientation. Simple RFs and a part of complex RF calculate the amplitude and the phase of the stimulus, the other part of complex RFs (with unmodulated response) calculate only amplitude. Given all this, the RFs are grating filters of spatial frequency.     

Adaptability of the receptive fields of neurons of the posterior temporal cortex and their sensitivity to parameters of photic stimulation in the cat

Study of receptive fields (RFs) of neurones in the postero-temporal cortex (field 21) of alert cat at three levels of visual adaptation: light photopic, light mesopic and practically dark or extremely low scotopic adaptations--revealed invariance of the most part of the studied RFs to the level of visual adaptation. Reorganization of RFs, connected with change of background luminosity were observed only in 12% of visually activated neurones. Significant reduction of responses to optic stimulation is shown at increase of the level of luminosity in 75% of neurones, revealing adaptive reorganizations. It is suggested that these reorganizations may take place in analogy with neurones of the field 17 on account of different involvement of intracortical inhibitory mechanisms (and, probably, not only in the postero-temporal cortex, but also in structures which precede it in visual hierarchy). Study of neurones sensitivity in the field 21 to parameters of optic stimulation revealed their considerable invariance to the length and orientation of the optic stimulus moving through the RF (60% of cases). Testing of RF by a rhombic optic stimulus did not change neuronal reactions, the form and dimensions of RF did not significantly change.     

研究: 外膝状体神经元的亮度反应与感受野ON-OFF反应的关系

外膝体是视觉信息进入新皮层的主要通路,其编码亮度信息的神经机制还不清楚.我们采用随机呈现的连续快速变化(50 Hz)的均匀亮度刺激,显著地提高了猫外膝体神经元对均匀亮度的反应强度,通过反相关算法抽提出神经元的亮度反应函数.约81%的神经元的亮度反应函数为单调性上升或下降,有19%的神经元亮度反应函数为V型.通过分析这些神经元对亮度上升和下降的反应强度与感受野ON和OFF反应强度的关系,表明83%的神经元对亮度的反应模式是由其感受野ON-OFF反应的相对强度决定的,其余17%则与其感受野ON-OFF区的兴奋和抑制的变化相关.这些结果揭示了外膝体神经元编码亮度变化的机制.     

Types of receptive fields in the lateral geniculate body and their functional model

In the Type I receptive fields (RFs) changes of the luminance leads to a shift of the curve relating the response and the stimulus area along the abscissa, in the Type II RFs the maximum of a response does not shift with changes of the luminance (Types I and II on classification by Glezer et al., 1971, 1972). The transient responses were observed in the Type I RFs and sustained responses in the Type II RFs. In the Type I RFs variation of the stimulus area and intensity brings about the change in the temporal and spatial frequency characteristics. This is produced by functional reorganization of the RF. In the Type II RFs there is no functional reorganization. The data obtained indicate that the Type I RFs are non-linear. By contrast, the Type II RFs are linear systems. The analysis of the model has shown that the distinctions in the dynamic characteristics of the responses of RFs belonging to different types is mainly due to different time constants for excitation and inhibition as well as inhibition coefficients. Distinctions in the mode of dependence of the RF response on stimulus parameters have been found to result from different relationship between delay time and stimulus parameters as well as different forms of the spatial weighting functions. It is shown that the Type I RFs transmit higher frequency components of the image spectrum, i.e. they emphasise the temporal and spatial contrasts. The Type II RFs transmit low frequency components of the spectrum including information about the intensity of an input stimulus.     

Reorganization of Receptive Fields of Cortical Field 21a Neurons and Formation of Responses to Moving Visual Stimuli

Using extracellular recording of spike activity from single neurons of field 21a of the cat neocortex, we examined in detail the spatial organization of receptive fields (RFs) of such cells after conditions of presentation of an immobile blinking light spot (a static RF) and moving visual stimuli (dynamic RFs). As was shown, the excitability of different RF subfields of a group of neurons possessing homogeneous on–off organization of the static RF changes significantly depended on the contrast, shape, dimension, orientation, and direction of movement of the applied mobile visual stimulus. This is manifested in changes in the number of discharge centers and shifts of their spatial localization. A hypothesis on the possible role of synchronous activation of the neurons neighboring the cell under study in the formation of an additional neuronal mechanism providing specialization of neuronal responses is proposed.     

Receptive Fields of Visually Sensitive Neurons of the Extrastriate Associative Area 21b of the Cat Cerebral Cortex

Organization of the receptive fields (RFs) of neurons of the extrastriate associative region 21b of the cerebral cortex was studied in cats. Most neurons under study (63%) were “monocular,” while 37% of the cells were “binocular” units. Among 178 neurons examined in detail, heterogeneous RF functional organization was typical of about 76% of the units; point-to-point testing of the entire RF area by stationary stimuli resulted in the generation of various types of responses (on, off, or on-off). The rest of the neurons (24%) generated homogeneous responses. The dimension, form, and functional organization of RFs of the neurons under study depended to a certain extent on the parameters of visual stimuli used for the measurements. Examination of the influence of the visual space, which surrounded the RF, on responses of the neurons evoked by stimulation of the RF per se showed that darkening of the visual space adjacent to the RF inhibited neuronal responses to moving stimuli; in some cases the responses were totally suppressed. Analysis of spatial overlapping of the RF sequentially recorded in the course of each insertion of the electrode showed that the density of distribution of the overlapping RF areas of neighboring neurons with the RF of the examined neuron is irregular, and that the RF is of a mosaic nature. We hypothesize that the visual space surrounding the RF plays a significant role in the formation of responses of visually sensitive neurons to presentation of moving stimuli. Neirofiziologiya/Neurophysiology, Vol. 37, No. 3, pp. 223–234, May–June, 2005.     

Spatial structure of complex cell receptive fields measured with natural images

Neuronal receptive fields (RFs) play crucial roles in visual processing. While the linear RFs of early neurons have been well studied, RFs of cortical complex cells are nonlinear and therefore difficult to characterize, especially in the context of natural stimuli. In this study, we used a nonlinear technique to compute the RFs of complex cells from their responses to natural images. We found that each RF is well described by a small number of subunits, which are oriented, localized, and bandpass. These subunits contribute to neuronal responses in a contrast-dependent, polarity-invariant manner, and they can largely predict the orientation and spatial frequency tuning of the cell. Although the RF structures measured with natural images were similar to those measured with random stimuli, natural images were more effective for driving complex cells, thus facilitating rapid identification of the subunits. The subunit RF model provides a useful basis for understanding cortical processing of natural stimuli.     

Development of spatial coarse-to-fine processing in the visual pathway

The sequential analysis of information in a coarse-to-fine manner is a fundamental mode of processing in the visual pathway. Spatial frequency (SF) tuning, arguably the most fundamental feature of spatial vision, provides particular intuition within the coarse-to-fine framework: low spatial frequencies convey global information about an image (e.g., general orientation), while high spatial frequencies carry more detailed information (e.g., edges). In this paper, we study the development of cortical spatial frequency tuning. As feedforward input from the lateral geniculate nucleus (LGN) has been shown to have significant influence on cortical coarse-to-fine processing, we present a firing-rate based thalamocortical model which includes both feedforward and feedback components. We analyze the relationship between various model parameters (including cortical feedback strength) and responses. We confirm the importance of the antagonistic relationship between the center and surround responses in thalamic relay cell receptive fields (RFs), and further characterize how specific structural LGN RF parameters affect cortical coarse-to-fine processing. Our results also indicate that the effect of cortical feedback on spatial frequency tuning is age-dependent: in particular, cortical feedback more strongly affects coarse-to-fine processing in kittens than in adults. We use our results to propose an experimentally testable hypothesis for the function of the extensive feedback in the corticothalamic circuit.     

Changes in visual receptive fields with microstimulation of frontal cortex

The influence of attention on visual cortical neurons has been described in terms of its effect on the structure of receptive fields (RFs), where multiple stimuli compete to drive neural responses and ultimately behavior. We stimulated the frontal eye field (FEF) of passively fixating monkeys and produced changes in V4 responses similar to known effects of voluntary attention. Subthreshold FEF stimulation enhanced visual responses at particular locations within the RF and altered the interaction between pairs of RF stimuli to favor those aligned with the activated FEF site. Thus, we could influence which stimulus drove the responses of individual V4 neurons. These results suggest that spatial signals involved in saccade preparation are used to covertly select among multiple stimuli appearing within the RFs of visual cortical neurons.     

Formation and functional significance of the dark receptive fields of the cat visual cortex

Receptive fields (RFs) of single units in the 17th field of the visual cortex of immobilized cat were investigated under dark adaptation. The mean RF size was equal to 67 degrees and varied from 3 degrees up to 120 degrees. The RFs with centres located near gaze were from 3 degrees up to 120 degrees in dia, but with growth of excentricity the number of small RFs decreased, and in the region of 70 to 100 degrees from gaze only RFs with diameters equal to 100 degrees were found. The shape of "dark" RFs was either ellipsoidal (in most cases) or round. Detector properties (orientational, directional, size and velocity selectivity) of the "dark" RFs were significantly less manifest or absent. Under photopic light adaptation the same units reorganized their RFs to well known sizes and configuration. The hypothesis is discussed of the formation of local detector RF in the visual cortex in light adaptation by selective cortical inhibition which is activated in darkness only slightly. This view is an alternative to the commonly-accepted scheme of local cortical RF formation by the hierarchical and selective excitatory convergence.     

Binocular responses of neurons in the barn owl's visual Wulst

Binocular responses have been recorded extra-cellularly at 58 sites in the barn owl's (Tyto alba) visual Wulst. Neurons showed disparity tuning to stimulation with moving bars, moving sinewave gratings and a moving visual-noise stimulus. Responses to sinewave gratings as a function of disparity were cyclic, with the period of a cycle of the response being correlated to one cycle of the stimulus. Cyclic responses were also found when bars or noise were used as a stimulus, but, especially in response to visual noise, one response peak, the main peak, was different from the other peaks, the sidepeaks: usually, the main peak was either higher or narrower or both higher and narrower than the sidepeaks. When the responses to different spatial frequencies were compared, response maxima coincided at the main peak, but not at the other peaks. In analogy to auditory physiology the disparity at which the frequency-independent peak occurs is termed characteristic disparity. Spatial-frequency tuning revealed broad tuning, ranging from 1 to more than 3 octaves at 50% of the maximal response. Disparity tuning was broad at the onset of the response and sharpened later. The data are discussed within the framework of a model for the neural representation of visual disparity that was derived from a model proposed earlier for the representation of interaural time difference, the main cue for encoding sound-source azimuths in the barn owl.Abbreviations ITD interaural time difference - CD characteristic delay - RF receptive field     

Flexibility of spatial averaging in visual perception

The classical receptive field (RF) concept-the idea that a visual neuron responds to fixed parts and properties of a stimulus-has been challenged by a series of recent physiological results. Here, we extend these findings to human vision, demonstrating that the extent of spatial averaging in contrast perception is also flexible, depending strongly on stimulus contrast and uniformity. At low contrast, spatial averaging is greatest (about 11 min of arc) within uniform regions such as edges, as expected if the relevant neurons have orientation-selective RFs. At high contrast, spatial averaging is minimal. These results can be understood if the visual system is balancing a trade-off between noise reduction, which favours large areas of averaging, and detail preservation, which favours minimal averaging. Two distinct populations of neurons with hard-wired RFs could account for our results, as could the more intriguing possibility of dynamic, contrast-dependent RFs.     

Dynamic Spatial Organization of Receptive Fields of Neurons in the 21a Cortical Area

We studied changes in the spatial parameters of receptive fields (RFs) of visually sensitive neurons in the associative area 21a of the cat cortex under conditions of presentation of moving visual stimuli. The results of experiments demonstrated that these parameters are dynamic and depend, from many aspects, on the pattern of the stimulus used for their estimation. Angular lengths of the horizontal and vertical axes of the RFs measured in the case of movement of the visual stimuli exceeded many times those determined by presentation of stationary blinking stimuli. As is supposed, a visual stimulus, when moving along the field of vision, activates a certain number of the neurons synaptically connected with the examined cell and possessing RFs localized along the movement trajectory. As a result, such integrated activity of the neuronal group can change the excitation threshold and discharge frequency of the studied neuron. It seems probable that correlated directed activation of the neuronal groups represents a significant neurophysiological mechanism providing dynamic modifications of the RF parameters of visually sensitive neurons in the course of processes of visual perception and identification of moving objects within the field of vision.     

Effects of local adaptation of receptive field on sensitivity of the cat visual cortex neurons to cruciform images

The responses to flashing single light bars of different orientation and to cruciform images (CI) were compared in 9 neurons of the cat striate cortex possessing high specific sensitivity to CI, during local adaptation of various receptive field (RF) zones. In most neurons, a two- to threefold reduction in the response to CI with a constantly present bar of optimum or orthogonal orientation, if compared with a response to the figure consisting of two flashing bars, was found. Responses to the CI including an adaptation bar were often increased, if compared with those observed at usual orientation tuning. The role of a cross-orientation inhibition in the formation of a selective sensitivity to CI in the neurons of the visual cortex is discussed.Neirofiziologiya/Neurophysiology, Vol. 27, No. 2, pp. 134–139, March–April, 1995.     

A new visual area on the inferior wall of the cat cruciate sulcus

Properties of 187 neurons in the inferior wall of the cruciate sulcus, in an area where electrical stimulation evoked unidirectional saccadic eye movements, were investigated in waking cats. Of the total number 172 responded to visual stimulation. Neurons in the surface layers of the cortex responded to simple visual stimuli: light or dark spots or bars, both stationary and moving at speeds of around 30 deg/sec. These neurons showed no selectivity as regards stimulus orientation but sometimes behaved selectively toward the direction of their movements. In the intermediate layers the maximal neuronal response was obtained to a model of a bird flaping its wings. Neuronal responses in the depth of the cortex were characterized by selectivity to movement of stimuli toward or away from the animal in a certain part of the visual field, irrespective of whether a light stimulus was presented against a dark background or a dark stimulus against the light background. Responses to visual stimulation were exhibited only if the animal was in a state of activation, when the EEG showed desynchronization, and they were absent in a state of quite wakefulness. No responses were obtained to auditory or somatic stimulation. Responses to visual stimulation were not found in neurons of the medial wall of the brain beneath the cruciate sulcus, but responses were recorded to eye movements of definite size or orientation. It is postulated that at least two contiguous retinotopically organized zones exist in this part of the brain. Activity of one of them is connected with visual function, that of the other with eye movements.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 16, No. 6, pp. 766–773, November–December, 1984.     

Mechanism of directional sensitivity of receptive fields in the cat visual cortex

Responses of directional-sensitive neurons in area 17 of the cat's cortex were studied to presentation of two flashing bars of light, one located in the center of the field, the other in the inhibitory zone relative to the center. The order of activation of the stimuli and the time interval between them could be varied; presentation of two bars was thus the analog of a moving stimulus. The inhibitory off-zone located at the entrance to the field from the side of the optimal direction of movement was found to have an initial inhibitory phase, followed by a phase of disinhibition, and again by a second inhibitory phase. Presentation of the bars with different time intervals in cases when stimulation of the center of the field coincided in time with one of the inhibitory phases, led to inhibition of the response, but if stimulation coincided with the phase of disinhibition, it led to facilitation. Phases of disinhibition were not found in the inhibitory zone located at the entrance to the field along the course of a nonoptimal direction of movement. The importance of the temporal characteristics of inhibitory zones for the appearance of directional sensitivity of visual courtical neurons is discussed.