Sensitivity of striate neurons to Y-like figures: experiment and simulation

Under stimulation of the receptive fields (RF) of neurons in the cat area 17 by flashing Y-like figures of different shape and orientation, the sensitivity to these figures was revealed in 72% of the studied cells, while 62% of units were sensitive to cross-like figures as well. Tuning to Y-like figures was typically selective to their shape and orientation, but in some cases it was invariant to these features. Response magnitudes to single bar, Y-like figure and cross were positively correlated. Simulation showed that the disinhibition might be a sufficient mechanism for effective detection of Y-like figures in a classical receptive field.     

Simulation of the role of intracortical inhibition as a source of sensitivity to cross-like figures

Simulation of receptive fields of striate neurons sensitive to cross-like figures under the blockade of intracortical inhibition was performed. It was shown that without the inhibition, a neuron with convergence of signals from two orientation detectors widens its tuning to a cross in such a way that the tuning becomes invariant to the shape and orientation of the cross. Detector of a cross whose orientation is based on the disinhibition mechanisms becomes a bar orientation detector under conditions of inhibition blockade. Another scheme of receptive field is advanced, in which the inhibitory zones mask tuning to cross-like figures, but the blockade of inhibition unveils such sensitivity. We specified the features of the receptive fields (shape, localization, and weight of their zones), which, being applied in simulation, imitate properties of the real striate neurons sensitive to shape and orientation of a cross-like figures.     

Simulation of mechanisms of tuning to incomplete cross-like figures in neurons of visual cortex

We performed a digital simulation of the receptive fields (RF) of cat cortical neurons in the area 17 that are able to detect cross-like figures with partly masked central or peripheral fragments. It was shown that the reciprocal interaction between the RF center and periphery may produce sharp, selective, and pronounced tuning to a cross shape and orientation due to blocking the end-stopping inhibition in the RF by its side-disinhibitory zone. Under conditions of cooperative interaction between the RF center and periphery the sensitivity index (cross/bar response ratio) was typically lower than in the reciprocal model. Features of the model that are critical for sensitivity to cross in cases of small or incomplete figures (the shape, localization, and weight of the RF zones) are specified.     

Selective sensitivity of the cat striate neurons to cruciform and angular figures of various orientations

Selectivity and invariability of tuning were studied in 51 neurons of the primary visual cortex (area 17); cruciform and angular figures (CF and AF, respectively) of different configurations and orientations were presented in their receptive fields. Twenty-three neurons, or 45% of the studied cells, demonstrated selective sensitivity to these figures. Their responses considerably (2.38±0.36 times, on average) increased, as compared with those evoked by presentation of a single bar of preferred orientation. In the examined group, 2 cells demonstrated sensitivity both to the CF and AF. A wide range of detector properties related to the CF and AF analysis was found in the analyzed neuronal population. Detectors of configuration of these figures are described. Selective sensitivity to the angle between branches of these figures was observed in 17 neurons, and responses of 2 neurons among them showed invariability to orientation of these figures. Four cells were selective for orientation and were insensitive to configuration, and 4 other cells showed no specific sensitivity to either of these properties, but were sensitive to the appearance of a CF itself in their receptive field (these cells were regarded as invariant detectors of crossing nodes). Data inconsistent with the hierarchic principle of detection of the above properties are presented. Possible mechanisms and functional significance of selective sensitivity of striate neurons to the CF and AF are discussed.Neirofiziologiya/Neurophysiology, Vol. 27, No. 5/6, pp. 403–412, September–December, 1995.     

Neural network model of visual cortex for determining surface curvature from images of shaded surfaces

The visual system can extract information about shape from the pattern of light and dark surface shading on an object. Very little is known about how this is accomplished. We have used a learning algorithm to construct a neural network model that computes the principal curvatures and orientation of elliptic paraboloids independently of the illumination direction. Our chief finding is that receptive fields developed by units of such model network are surprisingly similar to some found in the visual cortex. It appears that neurons that can make use of the continuous gradations of shading have receptive fields similar to those previously interpreted as dealing with contours (i.e. 'bar' detectors or 'edge' detectors). This study illustrates the difficulty of deducing neuronal function within a network solely from receptive fields. It is also important to consider the pattern of connections a neuron makes with subsequent stages, which we call the 'projective field'.     

Structure of receptive fields of hyppocampal visually-sensitive neurons in the cat

We studied the structure of receptive fields (RF) in the visually sensitive CA1 and CA3 fields of the dorsal hippocampus in alert cats subjected to pretrigeminal section of the brain stem. Scanning with moving stimuli and mapping the whole RF area with point by point application of a stationary stimulus, we studied 76 RF neurons. We found no significant differences in the response characteristics of neurons of fields CA1 and CA3. Our data are a basis for classifying the neurons in the following groups: homogeneous RF structure of the RF (54%) and nonhomogeneous structure (28%) and weak reaction to the standard stimulus, but a strong reaction to movement (18%). The results showed that responses of some of the hippocampal neurons exhibit high specificity. Thus, 9% of the neurons with nonhomogeneous RF structure disclosed with static stimuli reacted variously to a change in contrast and contours of moving stimuli. The data presented indicate that visually sensitive neurons of the hippocamus have a relatively well-developed mechanism for processing visual sensory information and this obviously ensures participation of the limbic system in visually controlled behavior of the animal.Institute of Experimental Biology, Academy of Sciences of the Armenian SSR, Yerevan. Translated from Neirofiziologiya, Vol. 23, No. 2, pp. 160–167, March–April, 1991.     

Response of Cat Ventrolateral Spinal Axons to an Itch-Producing Stimulus (Cowhage)

A comparison was made between different categories of mechanically sensitive, ventrolateral spinal axons to assess their sensitivity to the itch-producing substance cowhage. Of 52 wide-dynamic-range (WDR) units, 17 had contralateral, 22 had ipsilateral, and 13 had bilateral receptive fields. Of the 5 low-threshold units, 1 had an ipsilateral receptive field and the remainder were bilateral. Among the high-threshold units, 10 were contralateral, 6 ipsilateral, and 5 bilateral. Although there was no evidence of cowhage sensitivity in either low- or high-threshold spinal axons, neurons with WDR properties were reactive to cowhage. WDR neurons were subclassified on the basis of their resting discharge pattern as having intermittent, continuous, or no resting discharge. WDR units with an intermittent pattern of resting discharge demonstrated a significant sensitivity to active cowhage and hence might be regarded as pruritogen-responsive spinal axons. Inactive cowhage was used as a control stimulus. In some WDR units with large receptive fields, there were observations suggesting convergence of chemoreceptive and mechanoreceptive inputs, which produced inhibitory as well as excitatory effects.     

Response of cat ventrolateral spinal axons to an itch-producing stimulus (cowhage).

A comparison was made between different categories of mechanically sensitive, ventrolateral spinal axons to assess their sensitivity to the itch-producing substance cowhage. Of 52 wide-dynamic-range (WDR) units, 17 had contralateral, 22 had ipsilateral, and 13 had bilateral receptive fields. Of the 5 low-threshold units, 1 had an ipsilateral receptive field and the remainder were bilateral. Among the high-threshold units, 10 were contralateral, 6 ipsilateral, and 5 bilateral. Although there was no evidence of cowhage sensitivity in either low- or high-threshold spinal axons, neurons with WDR properties were reactive to cowhage. WDR neurons were subclassified on the basis of their resting discharge pattern as having intermittent, continuous, or no resting discharge. WDR units with an intermittent pattern of resting discharge demonstrated a significant sensitivity to active cowhage and hence might be regarded as pruritogen-responsive spinal axons. Inactive cowhage was used as a control stimulus. In some WDR units with large receptive fields, there were observations suggesting convergence of chemoreceptive and mechanoreceptive inputs, which produced inhibitory as well as excitatory effects.     

Structural organization of receptive fields of lateral suprasylvian cortical neurons in cats

The substructural organization of receptive fields of lateral suprasylvian cortical neurons, sensitive to movement of visual stimuli, was investigated in cats. The experimental results showed that receptive fields of neurons in this cortical area, judging by responses to movement, consist mainly of cells with qualitatively different characteristics. With the unmasked method of presentation of a moving stimulus, a reduction in the amplitude of movement as a rule evoked a directional response of the cell, whereas with the masked method, and with the same amplitudes of movement, a nondirectional response appeared. The receptive fields of some neurons were particularly sensitive to movement of borders but did not respond to the body of the stimulus like receptive fields of neurons described in other visual structures. Heterogeneity of the substructural organization of receptive fields of lateral suprasylvian cortical neurons can be explained by convergence of inputs on the neuron and it is regarded as the basis of integrative mechanisms in this structure.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 17, No. 3, pp. 293–300, May–June, 1985.     

Border ownership from intracortical interactions in visual area v2

A border between two image regions normally belongs to only one of the regions; determining which one it belongs to is essential for surface perception and figure-ground segmentation. Border ownership is signaled by a class of V2 neurons, even though its value depends on information coming from well outside their classical receptive fields. I use a model of V2 to show that this visual area is able to generate the ownership signal by itself, without requiring any top-down mechanism or external explicit labels for figures, T junctions, or corners. In the model, neurons have spatially local classical receptive fields, are tuned to orientation, and receive information (from V1) about the location and orientation of borders. Border ownership signals that model physiological observations arise through finite range, intraareal interactions. Additional effects from surface features and attention are discussed. The model licenses testable predictions.     

Retinotopic organization of small-field-target-detecting neurons in the insect visual system

BACKGROUND: Despite having tiny brains and relatively low-resolution compound eyes, many fly species frequently engage in precisely controlled aerobatic pursuits of conspecifics. Recent investigations into high-order processing in the fly visual system have revealed a class of neurons, coined small-target-motion detectors (STMDs), capable of responding robustly to target motion against the motion of background clutter. Despite limited spatial acuity in the insect eye, these neurons display exquisite sensitivity to small targets. RESULTS: We recorded intracellularly from morphologically identified columnar neurons in the lobula complex of the hoverfly Eristalis tenax. We show that these columnar neurons with exquisitely small receptive fields, like their large-field counterparts recently described from both male and female flies, have an extreme selectivity for the motion of small targets. In doing so, we provide the first physiological characterization of small-field neurons in female flies. These retinotopically organized columnar neurons include both direction-selective and nondirection-selective classes covering a large area of visual space. CONCLUSIONS: The retinotopic arrangement of lobula columnar neurons sensitive to the motion of small targets makes a strong case for these neurons as important precursors in the local processing of target motion. Furthermore, the continued response of STMDs with such small receptive fields to the motion of small targets in the presence of moving background clutter places further constraints on the potential mechanisms underlying their small-target tuning.     

Neural action fields for optic flow based navigation: a simulation study of the fly lobula plate network

Optic flow based navigation is a fundamental way of visual course control described in many different species including man. In the fly, an essential part of optic flow analysis is performed in the lobula plate, a retinotopic map of motion in the environment. There, the so-called lobula plate tangential cells possess large receptive fields with different preferred directions in different parts of the visual field. Previous studies demonstrated an extensive connectivity between different tangential cells, providing, in principle, the structural basis for their large and complex receptive fields. We present a network simulation of the tangential cells, comprising most of the neurons studied so far (22 on each hemisphere) with all the known connectivity between them. On their dendrite, model neurons receive input from a retinotopic array of Reichardt-type motion detectors. Model neurons exhibit receptive fields much like their natural counterparts, demonstrating that the connectivity between the lobula plate tangential cells indeed can account for their complex receptive field structure. We describe the tuning of a model neuron to particular types of ego-motion (rotation as well as translation around/along a given body axis) by its 'action field'. As we show for model neurons of the vertical system (VS-cells), each of them displays a different type of action field, i.e., responds maximally when the fly is rotating around a particular body axis. However, the tuning width of the rotational action fields is relatively broad, comparable to the one with dendritic input only. The additional intra-lobula-plate connectivity mainly reduces their translational action field amplitude, i.e., their sensitivity to translational movements along any body axis of the fly.     

Spatial summation processes in the receptive fields of visually driven neurons of the cat's cortical area 21a

The spatial summation in receptive fields (RF) of single neurons in cat's extrastriate area 21a was investigated as a basic neurophysiological substrate for central integration processing of visual information. The results showed that the majority of investigated neurons changed their response patterns with gradual increase of applied stimulus size. In approximately 82% of cases the suppression of neuron discharges was observed when the length of the moving strip exceeded that of the RF. In some neurons the increased size of the moving stimulus leads to the changes in the RF substructure. Receptive fields of neurons recorded at the same microelectrode penetration depth showed a great variety of RF superpositions distributed in a spatially asymmetric manner. As a result, every single RF consists of multiple sub-regions within the RF, differing from each other by the number of superimposed RF-s (density factor). We suggest that such complex spatial organization of the RF provides the neurophysiological basis for central integration processing of the visual information.     

Organization of receptive fields of motion-senstitive neurons in the cat pulvinar

The distribution of 70 visually sensitive neurons in the cat pulvinar sensitive to motion in the receptive fields was studied. The experimental results showed that components with directional characteristics are present in the structure of these fields of both direction-selective and unselective neurons. In the receptive fields of direction-selective neurons the directional elements of the substructure have identical preferred directions, which coincide with the preferred directions of response to stimulus movement over the entire receptive field. The organization of receptive fields of direction-selective neurons of the visual association structure thus does not differ significantly from that of analogous fields of visual projection neurons. Directional elements of the receptive fields of direction-unselective neurons were found to have different preferred directions, thereby providing a basis for organization of the nondirectional response of the neuron to a stimulus moving across the entire receptive field.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 339–346, July–August, 1982.     

Responses of nervous elements of the visual cortex of the chipmunk Eutamias sibiricus to moving and stationary stimuli]

The receptive field organization of cortical units has been studied in experiments with testing by moving and stationary light spots. The size of the receptive fields varied from 3 degrees to 10 degrees. Receptive fields which were tested by a stationary light spot exhibited various types of organization. Some of the neurons produced extensive excitatory on- and off-responses to stimulation by a light spot. Neuronal excitation evoked by light decreased if the stimulus was near the field boundary. Some of the neurons produced either on- or off-responses in any point of the receptive field. A small part of neurons had receptive fields with on- and off-reactions in the center, and either on- or off-responses at the peripheral zones. Most of the neurons exhibited specialization with respect to high-speed motion.     

“Regular” visual receptive fields of neurons of the cat lateral geniculate body

The spatial organization of receptive fields (RF) of neurons was studied in the lateral geniculate body (LGB) of cats with pretrigeminal transection of the brainstem (without general anesthesia). Using systematic point testing of the entire RF area and adjacent regions, the RF configuration and distribution of the response types for a stable flickering stimulus throughout the RF area were determined. Only 40% (64 units of 160 studied) LGB neurons had simple RF configuration. Such RF of ellipsoid or round shape were called regular receptive fields, RRF. Most RRF (51, or about 80%) demonstrated spatially homogeneous organization with similar-type (on, off, oron-off) responses to stimulation of the entire RF area. The RRF of 13 neurons, i.e., about 20%, included subfields with qualitatively different responses to application of a stable flickering light spot. The position of subfields was asymmetrical in 8 neurons (13%), while a nearly concentric RF arrangement, with the center surrounded by an antagonistic area, was found only in 5 units (7%) with RRF. Nearly all neurons with heterogeneous RRF demonstrated directional selectivity to moving stimuli.Neirofiziologiya/Neurophysiology, Vol. 27, No. 5/6, pp. 413–424, September–December, 1995.     

A system of insect neurons sensitive to horizontal and vertical image motion connects the medulla and midbrain

Summary The response properties and gross morphologies of neurons that connect the medulla and midbrain in the butterfly Papilio aegeus are described. The neurons presented give direction-selective responses, i.e. they are excited by motion in the preferred direction and the background activity of the cells is inhibited by motion in the opposite, null, direction. The neurons are either maximally sensitive to horizontal motion or to slightly off-axis vertical upward or vertical downward motion, when tested in the frontal visual field. The responses of the cells are dependent on the contrast frequency of the stimulus with peak values at 5–10 Hz. The receptive fields of the medulla neurons are large and are most sensitive in the frontal visual field. Examination of the local and global properties of the receptive fields of the medulla neurons indicates that (1) they are fed by local elementary motion-detectors consistent with the correlation model and (2) there is a non-linear spatial integration mechanism in operation.     

Pattern-dependent response modulations in motion-sensitive visual interneurons--a model study

Even if a stimulus pattern moves at a constant velocity across the receptive field of motion-sensitive neurons, such as lobula plate tangential cells (LPTCs) of flies, the response amplitude modulates over time. The amplitude of these response modulations is related to local pattern properties of the moving retinal image. On the one hand, pattern-dependent response modulations have previously been interpreted as 'pattern-noise', because they deteriorate the neuron's ability to provide unambiguous velocity information. On the other hand, these modulations might also provide the system with valuable information about the textural properties of the environment. We analyzed the influence of the size and shape of receptive fields by simulations of four versions of LPTC models consisting of arrays of elementary motion detectors of the correlation type (EMDs). These models have previously been suggested to account for many aspects of LPTC response properties. Pattern-dependent response modulations decrease with an increasing number of EMDs included in the receptive field of the LPTC models, since spatial changes within the visual field are smoothed out by the summation of spatially displaced EMD responses. This effect depends on the shape of the receptive field, being the more pronounced--for a given total size--the more elongated the receptive field is along the direction of motion. Large elongated receptive fields improve the quality of velocity signals. However, if motion signals need to be localized the velocity coding is only poor but the signal provides--potentially useful--local pattern information. These modelling results suggest that motion vision by correlation type movement detectors is subject to uncertainty: you cannot obtain both an unambiguous and a localized velocity signal from the output of a single cell. Hence, the size and shape of receptive fields of motion sensitive neurons should be matched to their potential computational task.     

A model for the neuronal implementation of selective visual attention based on temporal correlation among neurons

We propose a model for the neuronal implementation of selective visual attention based on temporal correlation among groups of neurons. Neurons in primary visual cortex respond to visual stimuli with a Poisson distributed spike train with an appropriate, stimulus-dependent mean firing rate. The spike trains of neurons whose receptive fields donot overlap with the focus of attention are distributed according to homogeneous (time-independent) Poisson process with no correlation between action potentials of different neurons. In contrast, spike trains of neurons with receptive fields within the focus of attention are distributed according to non-homogeneous (time-dependent) Poisson processes. Since the short-term average spike rates of all neurons with receptive fields in the focus of attention covary, correlations between these spike trains are introduced which are detected by inhibitory interneurons in V4. These cells, modeled as modified integrate-and-fire neurons, function as coincidence detectors and suppress the response of V4 cells associated with non-attended visual stimuli. The model reproduces quantitatively experimental data obtained in cortical area V4 of monkey by Moran and Desimone (1985).     

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.