Quantitative determination of orientational and directional components in the response of visual cortical cells to moving stimuli

The response characteristic of visual cortical cells to moving oriented stimuli consists mainly of directional (D) and orientational (O) components superimposed to a spontaneous activity (S). Commonly used polar plot diagrams reflect the maximal responses for different orientations and directions of stimulus movement with a periodicity of 360 degrees in the visual field. Fast Fourier analysis (FFT) is applied to polar plot data in order to determine the intermingled S, D, and O components. The zero order gain component of the spectrum corresponds to a (virtual) spontaneous activity. The first order component is interpreted as the strength of the direction selectivity and the second order component as the strength of the orientation specificity. The axes of the preferred direction and optimal orientation are represented by the respective phase values. Experimental data are well described with these parameters and relative changes of the shape of a polar plot can be detected with an accuracy better than 1%. The results are compatible with a model of converging excitatory and inhibitory inputs weighted according to the zero to second order components of the Fourier analysis. The easily performed quantitative determination of the S, D, and O components allows the study of pharmacologically induced changes in the dynamic response characteristics of single visual cortical cells.     

Responses of medullary neurons to moving visual stimuli in the common toad

Summary Intracellular recording and labeling of cells from the toad's (Bufo bufo spinosus) medulla oblongata in response to moving visual (and tactual) stimuli yield the following results. (i) Various response types characterized by extracellular recording in medullary neurons were also identified intracellularly and thus assigned to properties of medullary cell somata. (ii) Focussing on monocular small-field and cyclic bursting properties, somata of such neurons were recorded most frequently in the medial reticular formation and in the branchiomotor column but less often in the lateral reticular formation. (iii) Visual object disrimination established in pretectal/tectal networks is increased in its acuity in 4 types of medullary small-field neurons. The excitatory and inhibitory inputs to these neurons evoked by moving visual objects suggest special convergence likely to increase the filter properties. (iv) Releasing conditions, temporal pattern, and refractoriness of cyclic bursting neurons resemble membrane characteristics of vertebrate and invertebrate neurons known to play a role in premotor/motor activity. (v) Integrating functions of medullary cells have an anatomical correlate in the extensive arborizations of their dendritic trees; 5 morphological types of medullary neurons have been distinguished.Abbreviations A stripe moving in antiworm configuration - (W) moving in worm configuration - S square - BMC branchiomotor column - EPSP excitatory postsynaptic potential - IPSP inhibitory postsynaptic potential - RetF medullary reticular formation - RF receptive field - M neurons response properties of medullary neurons - T neurons classes of tectal neurons - TH neurons classes of thalamic/pretectal neurons - tr.tb.d. tractus tecto-bulbaris directus - tr.tbs.c. tractus tecto-bulbaris et spinalis cruciatus     

Responses of medullary neurons to moving visual stimuli in the common toad

The concept of coded 'command releasing systems' proposes that visually specialized descending tectal (and pretectal) neurons converge on motor pattern generating medullary circuits and release--in goal-specific combination--specific action patterns. Extracellular recordings from medullary neurons of the medial reticular formation of the awake immobilized toad in response to moving visual stimuli revealed the following main results. (i) Properties of medullary neurons were distinguished by location, shape, and size of visual receptive fields (ranging from relatively small to wide), by trigger features of various moving configural stimulus objects (including prey- and predator-selective properties), by tactile sensitivity, and by firing pattern characteristics (sluggish, tonic, warming-up, and cyclic). (ii) Visual receptive fields of medullary neurons and their responses to moving configural objects suggest converging inputs of tectal (and pretectal) descending neurons. (iii) In contrast to tectal monocular 'small-field' neurons, the excitatory visual receptive fields of comparable medullary neurons were larger, ellipsoidally shaped, mostly oriented horizontally, and not topographically mapped in an obvious fashion. Furthermore, configural feature discrimination was sharper. (iv) The observation of multiple properties in most medullary neurons (partly showing combined visual and cutaneous sensitivities) suggests integration of various inputs by these cells, and this is in principle consistent with the concept of command releasing systems. (v) There is evidence for reciprocal tectal/medullary excitatory pathways suitable for premotor warming-up. (vi) Cyclic bursting of many neurons, spontaneously or as a post-stimulus sustaining event, points to a medullary premotor/motor property.     

How to unconfound the directional and orientational information in visual neuron's response

When drifting bars or gratings are used as visual stimuli, information about orientation specificity (which has a period of 180°) and direction specificity (which has a period of 360°) is inherently confounded in the response of visual cortical neurons, which have long been known to be selective for both the orientation of the stimulus and the direction of its movement. It is essential to unconfound or separate these two components of the response as they may respectively contribute to form and motion perception, two of the main streams of information processing in the mammalian brain. Wörgötter and Eysel (1987) recently proposed the Fourier transform technique as a method of unconfounding the two components, but their analysis was incomplete. Here we formally develop the mathematical tools for this method to calculate the peak angles, bandwidths, and relative strengths, the three most important elements of a tuning curve, of both the orientational and the directional components, based on the experimentally-recorded neuron's response polar-plot. It will be shown that, in the 1-D Fourier decomposition of the polar-plot along its angular dimension, (1) the odd harmonics contain only the directional component, while the even harmonics are contributed to by both the orientational and the directional components; (2) the phases and the amplitudes of all the harmonics are related, respectively, to the peak angle and the bandwidth of the individual component. The basic assumption used here is that the two components are linearly additive; this in turn is immediately testable by the method itself.     

Representation of moving stimuli by somatosensory neurons.

The findings obtained in neurophysiological and psychophysical investigations using tactile stimuli that move at constant velocity across the skin are reviewed. For certain neurons in the postcentral gyrus of the cerebral cortex (S-I) of macaque monkeys, direction of stimulus motion is a "trigger feature" i.e., moving tactile stimuli evoke vigorous discharge activity in these neurons only if the stimuli are moved in a particular direction across the receptive field. This directional selectivity is maximal when stimulus velocity is between 5 and 50 cm/sec, and falls off rapidly at lower or higher velocities. The capacity for human subjects to correctly identify the direction of stimulus motion on the skin exhibits a similar dependence on stimulus velocity. The similar effects of velocity on neural and psychophysical measures of directional sensitivity support the idea that direction of stimulus motion on the skin can only be recognized if the moving stimulus optimally activates the group of S-I neurons for which that directions of simulus motion is the trigger feature.     

Intracellular activity of morphologically identified neurons of the grass frog's optic tectum in response to moving configurational visual stimuli

Summary In the grass frogRana temporaria, various classes of tectal neurons were identified by means of intracellular recording and iontophoretic staining using potassium-citrate/Co³⁺-lysine-filled micropipettes, which have been defined previously by extracellular recording methods. Class T5(1) neurons had receptive fields (RF) of 33°±5° diameter. In response to a moving 8°×8° square (S), a 2°×16° worm-like (W), or a 16°×2° antiworm-like (A) moving stripe, these cells showed excitatory postsynaptic potentials (EPSPs) and spikes which were interrupted occasionally by small inhibitory postsynaptic potentials (IPSPs). The excitatory responses (R) were strongest towards the square (R_S) and less to the worm (R_W). For the antiworm (R_A) the responses were smallest or equal to the worm stimulus yielding the relationship R_S>R_WR_A. Some of these cells were identified as pear-shaped or large ganglionic neurons, whose somata were located in the tectal cell layer 8. The somata of other large ganglionic neurons were found in layer 7 and the somata of other pear-shaped neurons at the top of layer 6, both displaying T5(1) properties. Class T5(2) neurons (RF=34°±3°) responded with large EPSPs and spikes, often interrupted by small IPSPs, when their RF was traversed by the square stimulus. The excitatory activity was somewhat less to the worm stimulus, whereas no activity at all, or only IPSPs, were recorded in response to the antiworm-stimulus; thus yielding the relationship for the excitatory activity R_S>R_W>R_A 0. Such a cell was identified as pyramidal neuron; the soma was located at the top of layer 6, with the long axon travelling into layer 7 to the medulla oblongata. Class T5(3) neurons (RF=29°±6°) showing EPSPs and spikes according to the relationship R_S>R_A>R_W have been identified as large ganglionic neurons. Their somata were located in layer 8. Class T5(4) neurons (RF=24±7°) responded only to the square stimulus with EPSPs and spikes, sometimes interrupted by IPSPs and yielding the relationship R_S>R_AR_W0. The somata of these large ganglionic or pear-shaped neurons were located in layer 8. Class T1(1) neurons (RF=30°–40°) were most responsive to stimuli moving at a relatively long distance in the binocular visual field, and have been identified as pear-shaped neurons. Their somata were located in layer 6.Further neurons are described and morphologically identified which have not yet been classified by extracellular recording methods. For example,IPSP neurons (RF=20°–30°) responded (R) with IPSPs only according to the relationship R_S>R_A R_W. The somata of these pear-shaped neurons were located in layer 6.The properties of tectal cells in response to electrical stimulation of the optic tract and to brisk changes of diffuse illumination suggest certain neuronal connectivity patterns. The results support the idea ofintegrative functional units (assemblies) of connected cells which are involved in various perceptual processes, such as configurational prey selection expressed by T5(2) prey-selective neurons.Abbreviations A antiworm-like 16°×2° stripe stimulus with long axis perpendicular to the direction of movement - W wormlike 2°×16° stripe stimulus with long axis oriented parallel to the direction of movement - S square 8°×8° moving stimulus - ERF excitatory receptive field - IRF inhibitory receptive field - RF receptive field - EPSP excitatory postsynaptic potential - IPSP inhibitory postsynaptic potential     

Varying response of caudate neurons to visual stimuli in awake cats

Activity in 62 caudate nucleus neurons produced during presentation of visual stimuli was recorded during experiments on awake cats. Response of a sensory pattern, associated with a photic stimulus falling within a certain section of the visual field was observed in 52% of the neurons tested as against only 11% manifesting motor response related to eye movement guided towards a target. About a quarter of the cells responded to biologically significant stimuli, producing a nonspecific response, i.e., not specifically related to the nature of the visual stimuli presented. Several different response patterns could be recorded from a single unit. A hypothesis that more than one parallel pathway for afferent visual inferences on the caudate nucleus may exist is presented on the basis of findings from this research.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 3, May–June, pp. 372–378, 1989.     

Effects of sombrevine on orientational tuning of visual cortex neurons in the cat

Changed orientational tuning (OT) in 58 visual cortex units was investigated during acute experiments on immobilized cats under light short-lasting sombrevine-induced anesthesia. A 47.6±5.6° alteration in the preferred orientation of 60% of cells occurred following sombrevine injection but no change occurred at any stage of anesthesia in the remainder. The latter group showed a preference for horizontal and vertical orientations, less pronounced in the former category. "Stable" neurons also displayed less acute tuning and more selective detection in comparison with "unstable" units. Breadth of orientational tuning consistently changed by an average of 65.2±6.7° in 55% of neurons, while tuning deteriorated in 31% and sharpened in 24% of cells. No regular change in tuning band occurred in the remainder. Background firing rate and evoked spike activity declined by 58% and 35%, respectively under anesthesia in 2/3 of the cells tested. Tuning bandwidth of unit firing rate had generally recovered within 20–40 min after administering the anesthetic (i.e., as the anesthesia wore off).Higher Nervous Activity and Neurophysiology Research Institute. Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 6, pp. 812–820, November–December, 1989.     

Tuning for cruciform stimuli in visual cortex neurons of cat

In the primary visual cortex of an immobilized awake cat, nearly one-third of the neurons studied (8 out of 22) were found to respond to flashing cruciform light stimuli 1.5–4 times better than to single stimulations with the strips of preferred orientation. It is suggested that such neurons can detect angles and line intersections.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 362–364, September–October, 1993.     

Dynamics of orientational tuning of feling visual cortex neurons under the effects of sombrevine

Dynamics of orientational tuning in 59 primary visual cortex neurons were investigated before and after sombrevine-induced anesthesia during acute experiments on immobilized cats using temporal slice techniques. A dynamic shift in preferred orientation of a flashing light strip, during which peak amplitude of spike discharges was noted (at an angle of between 22 and 157°) occurred as response developed in two-thirds of the cells. We had previously named this effect "scanning the orientational range" [9]. Scanning declined significantly in 45% of the sample, culminating in complete disappearance of this effect in some cells following sombrevine action. Scanning intensified in 30%, while dynamics of tuning remained unchanged in 25% of units. Sombrevine administration induced change in the preferred stimulus orientation of 60% of the neurons (referred to as "unstable" cells) and remained constant in "stable" cells (= 40%). Dynamic changes in preferred stimulus orientation were 2.5 times as high as those of stable cells in the waking state. The scanning effect declined significantly in 60% of "unstable" neurons under the action of anesthesia and remained unchanged in not more than 6%. At the same time, orientational tuning did not alter in the "stable" cell group in 46% of units, either declining (25%) or increasing (29%) in the remaining scanning ranges.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 22, No. 1, pp. 107–113, January–February, 1990.     

A model of the human visual system in its response to certain classes of moving stimuli

The purpose of this study is to construct a functional model of the human visual system in its response to certain classes of moving stimuli.Experimental data are presented describing the interdependence of the input variables, temporal frequency, spatial period, etc., for two constant response states, viz. threshold motion response and threshold flicker response. On the basis of these data, two basic units are isolated, a vertical (V) unit and a horizontal (H) unit. The H-unit is identified with the Reichardt multiplier (Reichardt and Varju, 1959), and the V-unit with the de Lange filter (de Lange, 1954).A definition of the general motion response of the H-units is obtained, and this is then reduced to an expression which may be applied directly to the observed motion response data. By this method, Thorson's simplification of the Reichardt scheme (Thorson, 1966) is adopted for the H-unit and total and relative (population) weighting factors, associated with the H-unit output, are defined.In order to reconcile the theoretical square-wave threshold motion response with the experimental data, Thorson's simplification is modified with the introduction of a low-pass filter on the output. The amended scheme is shown to predict a (temporal) frequency-dependent phase-sensitivity. This prediction is tested experimentally, and its validity indicated.     

Population coding of visual stimuli by cortical neurons tuned to more than one dimension

Neurons in the visual cortex are typically selective to a number of stimulus dimensions. Thus, there is a basic ambiguity in relating the response level of a single neuron to the stimulus values. It is shown that a multi-dimensional stimulus may be coded reliably by an ensemble of neurons, using a weighted average population coding model. Each neurons' contribution to the population signal for each dimension is the product of its response magnitude and its preferred value for that dimension. The sum of the products was normalized by the sum of the ensemble responses. Simulation results show that the representation accuracy increases as the square root of the number of units irrespective of the number of dimensions. Comparison of a specific 2D case of this population code for orientation and spatial frequency to behavioral discrimination levels yields that 10³–10⁴ neurons are needed to reach psychophysical performance. Introduction of each additional dimension requires about 1.7 times the number of neurons in the ensemble to reach the same level of accuracy. This result suggests that neurons may be selective for only 3 to 5 dimensions. It also provides another rationale for the existence of parallel processing streams in vision.     

Postsynaptic excitation and in hibition in neurons of the turtle general cortex in response to moving stimuli

Responses of the general cortex to moving stimuli were studied in turtles. The evoked potential, the synaptic nature of its individual components, and the mechanisms of their generation were analyzed. The evoked potential had a negative-positive sequence. The negative part consisted of a slow negative wave on which fast negative complexes were superposed. These components reflected EPSPs of afferent nature generated on dendrites of the principal neurons. The first fast negative complex was followed by a rhythmic discharge superposed on the slow negative and positve waves. The negative waves of the rhythmic discharge were shown to reflect EPSPs and the positive waves IPSPs, probably generated on dendrites of cortical neurons. The rhythmic EPSP — IPSPs are evidently generated by a feedback mechanism, whereas the positive wave reflects dendritic IPSPs of the principal neurons.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 249–256, May–June, 1977.     

Properties of neurons sensitive to moving black stimuli in lateral suprasylvian area of the cat cortex

Neurons sensitive to visual stimulation in the lateral suprasylvian area of the cortex were investigated in cats with pretrigeminal brain section. About 25% of the neuron population responding to visual stimulation were shown to be highly sensitive to moving black objects. These neurons were called black-sensitive. Neurons of this group had a low level of spontaneous activity and were mainly directionally sensitive. Some of them exhibited summation of responses during successive enlargement of the stimulus. An important distinguishing feature of these neurons was a change in the temporal structure of their response after contrast reversal of the stimulus.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 15, No. 1, pp. 16–21, January–February, 1983.     

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.     

Convergence of visual and auditory stimuli on the neurons of the visual cortex in the conscious rabbit

We investigated the visual-cortex neurons of the conscious rabbit during simultaneous stimulation with a clicking sound and a light flash (complex) and during separate application of these stimuli. We tested the development of the reflex with time and of the sound-light association during prolonged rhythmic application of the sound and light. Fifty visual-cortex neurons were studied; 20% of the cells responded with a specific phased reaction and 16% exhibited a specific response to the complex different from the responses to each of its components. Development of a sound-light association was observed in 18% of the cells and a temporal reflex was induced in 25%. In most cases, the conditioned reaction evoked was similar to some informational element in the neuronal response to the complex.M. V. Lomonosov Moscow State University. Institute of Cybernetics, Academy of Sciences of the GruzSSR. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 391–398, July–August, 1970.     

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.