Monocular focal retinal lesions induce short-term topographic plasticity in adult cat visual cortex

Electrophysiological recording in primary visual cortex (VI) was performed both prior to and in the hours immediately following the creation of a discrete retinal lesion in one eye with an argon laser. Lesion projection zones (LPZs; 21-64 mm2) were defined in the visual cortex by mapping the extent of the lesion onto the topographic representation in cortex. There was no effect on neuronal responses to the unlesioned eye or on its topographic representation. However, within hours of producing the retinal lesion, receptive fields obtained from stimulation of the lesioned eye were displaced onto areas surrounding the scotoma and were enlarged compared with the corresponding field obtained through the normal eye. The proportion of such responsive recording sites increased during the experiment such that 8-11 hours post-lesion, 56% of recording sites displayed neurons responsive to the lesioned eye. This is an equivalent proportion to that previously reported with long-term recovery (three weeks to three months). Responsive neurons were evident as far as 2.5 mm inside the border of the LPZ. The reorganization of the lesioned eye representation produced binocular disparities as great as 15 degrees, suggesting interactions between sites in VI up to 5.5 mm apart.     

Inhomogeneous retino-cortical mapping is supported and stabilized with correlation-learning during self-motion

In primates, the area of primary visual cortex representing a fixed area of visual space decreases with increasing eccentricity. We identify visual situations to which this inhomogeneous retino-cortical mapping is well adapted and study their relevance during natural vision and development. We assume that cortical activations caused by stationary objects during self-motion along the direction of gaze travel on average with constant speed across the cortical surface, independent of retinal eccentricity. This is the case if the distribution of objects corresponds to an ellipsoid with the observer in its center. We apply the resulting flow field to train a simple network of pulse coding neurons with Hebbian learning and demonstrate that the density of learned receptive field centers is in close agreement with primate retino-cortical magnification. In addition, the model reproduces the increase of receptive field size and the decrease of its peak sensitivity with increasing eccentricity. Our results suggest that self-motion may have played an important role in the evolution of the visual system and that cortical magnification can be refined and stabilized by Hebbian learning mechanisms in ontogenesis under natural viewing conditions.     

Object segmentation model: analytical results and biological implications

 A simple, biologically motivated neural network for segmentation of a moving object from a visual scene is presented. The model consists of two parts: an object selection model which employs a scaling approach for receptive field sizes, and a subsequent network implementing a spotlight by means of multiplicative synapses. The network selects one object out of several, segments the rough contour of the object, and encodes the winner object's position with high accuracy. Analytical equations for the performance level of the network, e.g., for the critical distance of two objects above which they are perceived as separate, are derived. The network preferentially chooses the object with the largest angular velocity and the largest angular width. An equation for the velocity and width preferences is presented. Additionally it is shown that for certain neurons of the model, flat receptive fields are more favourable than Gaussian ones. The network exhibits performances similar to those known from amphibians. Various electrophysiological and behavioral results – e.g., the distribution of the diameters of the receptive fields of tectal neurons, of the tongue-projecting salamander Hydromantes italicus and the range of optimal prey velocities for prey catching – can be understood on the basis of the model. Received: 7 December 2000 / Accepted: 13 February 2001     

Characteristics of dynamic reorganization of receptive fields of cortical and geniculate visual neurons of cat in response to changes in photic stimulation parameters

The effect of parameters of local photic stimulation of different points of the receptive field on the characteristics of dynamic reorganization of receptive fields of cortical and geniculate visual neurons within microintervals of time observed previously was studied in computer-controlled experiments on unanesthetized, curarized cats. Dependence on the degree of widening of the receptive field and the temporal characteristics of this process on the background illumination level, intensity, contrast, area, duration, energy, and orientation of a local rectangular or circular photic stimulus flashing in random order at 100 points of the tested part of the visual field was studied. It was concluded that the background illumination level and the intensity, size, duration, and orientation of the stimulus have a specific effect on dynamic reorganization of the receptive field. The effects of all the parameters studied on the dynamics of the receptive field were shown to be nonlinear functions with optimal values that differed for different cells. The possible functional role of this effect and also the probability that it may participate in information coding in the visual system are discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 15, No. 4, pp. 339–346, July–August, 1983.     

An architectural hypothesis for direction selectivity in the visual cortex: the role of spatially asymmetric intracortical inhibition

Within a linear field approach, an architectural model for simple cell direction selectivity in the visual cortex is proposed. The origin of direction selectivity is related to recurrent intracortical interactions with a spatially asymmetric character along the axis of stimulus motion. No explicit asymmetric temporal mechanisms are introduced or adopted. The analytical investigation of network behavior, carried out under the assumption of a linear superposition of geniculate and intracortical contributions, shows that motion sensitivity of the resulting receptive fields emerges as a dynamic property of the cortical network without any feed-forward direction selectivity bias. A detailed analysis of the effects of the architectural characteristics of the cortical network on directionality and velocity-response curves was conducted by systematically varying the model's parameters. Received: 8 May 1998 / Accepted in revised form: 10 November 1998     

Recursive features of circular receptive fields

The distribution of excitability in retinal receptive fields may be well approximated by functions with recursive features. Physiological data do not exclude an implementation of recursive structures in the visual system. It is the most remarkable advantage of a recursive visual system, that cortical receptive fields tuned to different spatial frequencies will have an identical neuronal circuitry. Structural consequences for retina, LGN and visual cortex are discussed.     

Synaptic integration fields and associative plasticity of visual cortical cells in vivo

Two major constraints in connectivity decide the spatial extent of visual cortical receptive fields, both during development and adult functioning: 1) feedforward input, extrinsic to visual cortex, is organized in an orderly projection to form a point-to-point mapping of the retina onto the cortical mantle and constitutes the core of the minimal discharge field (MDF) after amplification by local intracortical circuits; and 2) a second type of connectivity consists of long distance horizontal intracortical connections which are thought to favor the binding of local visual operations occurring simultaneously in different parts of the visual field. We review here possible factors, intrinsic to the considered cortical cell, that may control the effectiveness of post-synaptic integration. Their expression during sensory recognition could depend on the spatio-temporal distribution of active inputs onto the target cell dendrite and on their interplay with non-linearities of the membrane properties. On the basis of intracellular recordings in cat area 17, we have observed that peripheral responses (excitatory and inhibitory) could be boosted by coincident postsynaptic depolarization. This effect is lost if the response and the postsynaptic depolarization are mismatched by 1000 ms, suggesting that temporal correlation of central and peripheral responses could regulate in a non-linear manner the gain of center-surround interactions. This temporal selectivity is compatible with up-regulation of composite potentials by the transient voltage-dependent activation of slowly inactivating conductances in visual cortical neurons. A direct consequence of these different considerations is that the receptive field (RF) of neurons in visual pathways should not be considered as a static hardwired window probing the outer environment, but as an active filter which may continuously adapt and be updated as a function of global context and past experience.     

Receptive field dynamics of neurons in the cat visual cortex and lateral geniculate body

Spike responses of single neurons in the primary visual cortex and lateral geniculate body to random presentation of local photic stimuli in different parts of the receptive field of the cell were studied in acute experiments on curarized cats. Series of maps of receptive fields with time interval of 20 msec obtained by computer enabled the dynamics of the excitatory and inhibitory zones of the field to be assessed during development of on- and off-responses to flashes. Receptive fields of all cortical and lateral geniculate body neurons tested were found to undergo regular dynamic reorganization both after the beginning and after the end of action of the photic stimulus. During the latent period of the response no receptive field was found in the part of the visual field tested, but later a small zone of weak responses appeared only in the center of the field. Gradually (most commonly toward 60–100 msec after application of the stimulus) the zone of the responses widened to its limit, after which the recorded field began to shrink, ending with complete disappearance or disintegration into separate fragments. If two bursts of spikes were generated in response to stimulation, during the second burst the receptive field of the neuron changed in the same way. The effects described were clearly exhibited if the level of background illumination, the intensity of the test bars, their contrast with the background, duration, angles subtended, and orientation were varied, although the rate and degree of reorganization of the receptive field in this case changed significantly. The functional importance of the effect for coding of information about the features of a signal by visual cortical neurons is discussed.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 14, No. 6, pp. 622–630, November–December, 1982.     

Combined Hebbian development of geniculocortical and lateral connectivity in a model of primary visual cortex

We present a network model of visual map development in layer 4 of primary visual cortex. Our model comprises excitatory and inhibitory spiking neurons. The input to the network consists of correlated spike trains to mimick the activity of neurons in the lateral geniculate nucleus (LGN). An activity-driven Hebbian learning mechanism governs the development of both the network's lateral connectivity and feedforward projections from LGN to cortex. Plasticity of inhibitory synapses has been included into the model so as to control overall cortical activity. Even without feedforward input, Hebbian modification of the excitatory lateral connections can lead to the development of an intracortical orientation map. We have found that such an intracortical map can guide the development of feedforward connections from LGN to cortical simple cells so that the structure of the final feedforward orientation map is predetermined by the intracortical map. In a scenario in which left- and right-eye geniculocortical inputs develop sequentially one after the other, the resulting maps are therefore very similar, provided the intracortical connectivity remains unaltered. This may explain the outcome of so-called reverse lid-suture experiments, where animals are reared so that both eyes never receive input at the same time, but the orientation maps measured separately for the two eyes are nevertheless nearly identical. Received: 20 December 1999 / Accepted in revised form: 9 June 2000     

Wide-field, motion-sensitive neurons and matched filters for optic flow fields

 The receptive field organization of a class of visual interneurons in the fly brain (vertical system, or VS neurons) shows a striking similarity to certain self-motion-induced optic flow fields. The present study compares the measured motion sensitivities of the VS neurons (Krapp et al. 1998) to a matched filter model for optic flow fields generated by rotation or translation. The model minimizes the variance of the filter output caused by noise and distance variability between different scenes. To that end, prior knowledge about distance and self-motion statistics is incorporated in the form of a “world model”. We show that a special case of the matched filter model is able to predict the local motion sensitivities observed in some VS neurons. This suggests that their receptive field organization enables the VS neurons to maintain a consistent output when the same type of self-motion occurs in different situations. Received: 14 June 1999 / Accepted in revised form: 20 March 2000     

A computer simulation model of the afferent part of the visual foveation system

It is argued that the duplicity notion of visual perception refers to two separate neural systems, the foveation and identification system. A computer simulation model for the afferent part of the foveation system was developed. It consists of a receptive matrix and a retinal, geniculate, and cortical network. Within the networks neurons were substituted by threshold gates. The retinal information of the model can be controlled by the geniculate network and is analyzed with regard to stimulus orientation and movement by hypercolumns of cortical gates. The implemented columnar organization preserves retinocortical continuity. Retinal and cortical information together provide the necessary prerequisites for the efferent part of the foveation system.Supported by a grant from the Deutsche Forschungsgemeinschaft     

Selective visual attention in a neurocomputational model of phase oscillators

In order to understand the dynamic property of covert selective visual attention, which is different from the proposed mechanism of the spotlight metaphor, a two-layered network of phase oscillators was developed. The first layer is related to the hippocampus and controls attention focus formation. The second layer is related to the visual cortex, and each cortical oscillator in it simulates an assembly of cells coding for a particular stimulus in the sense of feature binding. Selective visual attention is interpreted as the result of the emergent synchronization of hippocampus oscillators and a part of cortical oscillators. Numerical experiments are presented to illustrate attention focus formation and attention shifting from one set of stimuli to another. From a neurocomputational point of view, our results demonstrate that attention is an emergent property of the dynamical cell assemblies responding to the whole visual field. Received: 2 January 1998 / Accepted in revised form: 10 November 1998     

An intracellular study of space and time representation in primary visual cortical receptive fields

In contrast with previous knowledge based on extracellular recordings, the recent development of intracellular techniques in vivo (sharp electrode or ‘blind patch’) ideally allows experimenters to analyze and dissect the contribution of feedforward and lateral connectivity in the functional expression of a synaptic ‘integration field’. We will present recent data which demonstrate that the visual receptive field of cortical neurons described at the level of subthreshold synaptic events extends over much larger regions of the visual field than previously thought, and that the capacity of cells to amplify subthreshold responses depends on the immediate past history of their membrane potential. Our data suggest that visual cortical receptive fields should not be considered as a fixed entity but more as a dynamic field of integration and association. Two types of dynamics can be argued for: 1) the spatial structure of the minimal discharge field (defined by suprathreshold activation of the cell) can be profoundly reorganized at least during development and most probably during selective phases of learning under the control of activity-dependent mechanisms. Adaptive changes in visual responses are thought to reflect long-lasting potentiation and/or depression of synaptic efficacies conveying ON- and OFF-center information; and 2) during sensory processing, reconfiguration of synaptic weights may be achieved on a much faster time-scale and linke to non-linear properties of the postsynaptic membrane as well as that of recruited networks. Association of information available in the central part of the receptive field (RF) and of input coming from the reputedly ‘unresponsive’ regions surrounding it, or arising simultaneously from different parts of the visual field, might be suppressive in certain cases and capable of boosting hidden responses in other cases, depending on the global stimulus configuration.     

Recognition of partly occluded patterns: a neural network model

 Human beings are often able to read a letter or word partly occluded by contaminating ink stains. However, if the stains are completely erased and the occluded areas of the letter are changed to white, we usually have difficulty in reading the letter. In this article I propose a hypothesis explaining why a pattern is easier to recognize when it is occluded by visible objects than by invisible opaque objects. A neural network model is constructed based on this hypothesis. The visual system extracts various visual features from the input pattern and then attempts to recognize it. If the occluding objects are not visible, the visual system will have difficulty in distinguishing which features are relevant to the original pattern and which are newly generated by the occlusion. If the occluding objects are visible, however, the visual system can easily discriminate between relevant and irrelevant features and recognize the occluded pattern correctly. The proposed model is an extended version of the neocognitron model. The activity of the feature-extracting cells whose receptive fields cover the occluding objects is suppressed in an early stage of the hierarchical network. Since the irrelevant features generated by the occlusion are thus eliminated, the model can recognize occluded patterns correctly, provided the occlusion is not so large as to prevent recognition even by human beings. Received: 21 February 2000 / Accepted in revised form: 11 September 2000     

Traveling Waves and the Processing of Weakly Tuned Inputs in a Cortical Network Module

Recent studies have shown that local cortical feedback can havean important effect on the response of neurons in primary visualcortex to the orientation of visual stimuli. In this work, westudy the role of the cortical feedback in shaping thespatiotemporal patterns of activity in cortex. Two questionsare addressed: one, what are the limitations on the ability ofcortical neurons to lock their activity to rotatingoriented stimuli within a single receptive field? Two, can thelocal architecture of visual cortex lead to the generation ofspontaneous traveling pulses of activity? We study theseissues analytically by a population-dynamic model of ahypercolumn in visual cortex. The order parameter thatdescribes the macroscopic behavior of the network is thetime-dependent population vector of the network. We firststudy the network dynamics under the influence of a weakly tunedinput that slowly rotates within the receptive field. We showthat if the cortical interactions have strong spatialmodulation, the network generates a sharply tuned activityprofile that propagates across the hypercolumn in a path thatis completely locked to the stimulus rotation. The resultantrotating population vector maintains a constant angular lagrelative to the stimulus, the magnitude of which grows with thestimulus rotation frequency. Beyond a critical frequency thepopulation vector does not lock to the stimulus but executes aquasi-periodic motion with an average frequency that is smallerthan that of the stimulus. In the second part we consider thestable intrinsic state of the cortex under the influence of isotropic stimulation. We show that if the local inhibitoryfeedback is sufficiently strong, the network does not settleinto a stationary state but develops spontaneous travelingpulses of activity. Unlike recent models of wave propagation incortical networks, the connectivity pattern in our model isspatially symmetric, hence the direction of propagation ofthese waves is arbitrary. The interaction of these waves withan external-oriented stimulus is studied. It is shown that thesystem can lock to a weakly tuned rotating stimulus if thestimulus frequency is close to the frequency of the intrinsic wave.     

Binding and segmentation of multiple objects through neural oscillators inhibited by contour information

 Temporal correlation of neuronal activity has been suggested as a criterion for multiple object recognition. In this work, a two-dimensional network of simplified Wilson-Cowan oscillators is used to manage the binding and segmentation problem of a visual scene according to the connectedness Gestalt criterion. Binding is achieved via original coupling terms that link excitatory units to both excitatory and inhibitory units of adjacent neurons. These local coupling terms are time independent, i.e., they do not require Hebbian learning during the simulations. Segmentation is realized by a two-layer processing of the visual image. The first layer extracts all object contours from the image by means of “retinal cells” with an “on-center” receptive field. Information on contour is used to selectively inhibit Wilson-Cowan oscillators in the second layer, thus realizing a strong separation among neurons in different objects. Accidental synchronism between oscillations in different objects is prevented with the use of a global inhibitor, i.e., a global neuron that computes the overall activity in the Wilson-Cowan network and sends back an inhibitory signal. Simulations performed in a 50×50 neural grid with 21 different visual scenes (containing up to eight objects + background) with random initial conditions demonstrate that the network can correctly segment objects in almost 100% of cases using a single set of parameters, i.e., without the need to adjust parameters from one visual scene to the next. The network is robust with reference to dynamical noise superimposed on oscillatory neurons. Moreover, the network can segment both black objects on white background and vice versa and is able to deal with the problem of “fragmentation.” The main limitation of the network is its sensitivity to static noise superimposed on the objects. Overcoming this problem requires implementation of more robust mechanisms for contour enhancement in the first layer in agreement with mechanisms actually realized in the visual cortex. Received: 25 October 2001 / Accepted: 26 February 2003 / Published online: 20 May 2003 Correspondence to: Mauro Ursino (e-mail: mursino@deis.unibo.it, Tel.: +39-051-2093008, Fax: +39-051-2093073)     

Rapid activity-dependent sprouting of optic fibers into a local area denervated by application of beta-bungarotoxin in goldfish tectum

The retinotectal projection is known to be capable of extensive long-term expansion of connections, but it is not known how fast such changes can occur or what triggers sprouting of terminals. We studied sprouting of optic fibers into an area denervated by local microinjection of beta-bungarotoxin (β-BTX), a specific presynaptic neurotoxin with phospholipase A₂ activity that destroys nerve terminals at the neuromuscular junction. After injection of 0.1 pmol of β-BTX, the optic terminals fired spontaneously with decreasing amplitude and became silent within 1 to 2 h. Outside the injection zone, the retinotectal map was normal, so the silent zone was associated with a scotoma in the visual field. Horseradish peroxidase (HRP) staining of the entire optic nerve showed a denervated region at the injection site with beaded, degenerating fibers at its edge. Between 3 and 9 days later, optic units were recorded within the injection zone whose receptive fields lay just outside the scotoma in the visual field, indicating that intact surrounding terminals had sprouted into the area. These sprouts made functional connections, as indicated by field potential recordings and current source-density analysis. At this time, HRP staining also demonstrated retinal innervation within the injection zone. By 12 days, normal maps with no scotoma were recorded and HRP staining was normal at the injection site, indicating that the β-BTX-damaged fibers had regenerated to reclaim their tectal sites. The results show that the retinotectal projection of goldfish is very dynamic, since intact optic fibers can sprout into adjacent vacant postsynaptic territory within 2 to 3 days, much faster than previously reported. In a final experiment, we showed that this sprouting is activity-dependent, since it could be prevented by blocking retinal activity with intraocular tetrodotoxin (TTX) during the first 2 days postinjection, even though TTX block of activity does not block regeneration in this system. One possible mechanism for this rapidly triggered sprouting is that arachidonic acid liberated by β-BTX acts as a sprouting factor to attract surrounding healthy fibers into the denervated region but requires activity at the terminals to be effective. © 1996 John Wiley & Sons, Inc.