Neuronal responses in ventroposterolateral nucleus of thalamus in monkeys (Macaca mulatta) during active touch of gratings.

Responses of single neurons were recorded from the ventroposterolateral nucleus (VPL) of the thalamus while a monkey stroked its fingertips over gratings. Monkeys were trained to stroke the gratings with consistent downward applied force and velocity of hand motion. Neurons were selected with receptive fields on the glabrous digits. Average firing rate was computed for a range of grating groove widths; groove width corresponded to roughness. Force and velocity were measured. VPL responses were compared to previously reported responses in primary somatosensory cortex (SI) under identical stimulus conditions, and to reports of peripheral afferent fiber responses to passively applied gratings. VPL responses more closely resembled those of peripheral afferent fibers than those of SI in important respects: lack of independent responses to roughness, force, and velocity; high temporal and force fidelity; and response patterns that closely followed the shape of elevated metal strips used to separate pairs of gratings. The presence in cortex of response patterns not seen in the thalamus, such as response independence and negative correlations to groove width, suggests that they stem from cortical processing.     

Neuronal Responses in Ventroposterolateral Nucleus of Thalamus in Monkeys (Macaca mulatta) during Active Touch of Gratings

Responses of single neurons were recorded from the ventroposterolateral nucleus (VPL) of the thalamus while a monkey stroked its fingertips over gratings. Monkeys were trained to stroke the gratings with consistent downward applied force and velocity of hand motion. Neurons were selected with receptive fields on the glabrous digits. Average firing rate was computed for a range of grating groove widths; groove width corresponded to roughness. Force and velocity were measured. VPL responses were compared to previously reported responses in primary somatosensory cortex (SI) under identical stimulus conditions, and to reports of peripheral afferent fiber responses to passively applied gratings. VPL responses more closely resembled those of peripheral afferent fibers than those of SI in important respects: lack of independent responses to roughness, force, and velocity; high temporal and force fidelity; and response patterns that closely followed the shape of elevated metal strips used to separate pairs of gratings. The presence in cortex of response patterns not seen in the thalamus, such as response independence and negative correlations to groove width, suggests that they stem from cortical processing.     

The Detection of Orientation Continuity and Discontinuity by Cat V1 Neurons

The orientation tuning properties of the non-classical receptive field (nCRF or “surround”) relative to that of the classical receptive field (CRF or “center”) were tested for 119 neurons in the cat primary visual cortex (V1). The stimuli were concentric sinusoidal gratings generated on a computer screen with the center grating presented at an optimal orientation to stimulate the CRF and the surround grating with variable orientations stimulating the nCRF. Based on the presence or absence of surround suppression, measured by the suppression index at the optimal orientation of the cells, we subdivided the neurons into two categories: surround-suppressive (SS) cells and surround-non-suppressive (SN) cells. When stimulated with an optimally oriented grating centered at CRF, the SS cells showed increasing surround suppression when the stimulus grating was expanded beyond the boundary of the CRF, whereas for the SN cells, expanding the stimulus grating beyond the CRF caused no suppression of the center response. For the SS cells, strength of surround suppression was dependent on the relative orientation between CRF and nCRF: an iso-orientation grating over center and surround at the optimal orientation evoked strongest suppression and a surround grating orthogonal to the optimal center grating evoked the weakest or no suppression. By contrast, the SN cells showed slightly increased responses to an iso-orientation stimulus and weak suppression to orthogonal surround gratings. This iso-/orthogonal orientation selectivity between center and surround was analyzed in 22 SN and 97 SS cells, and for the two types of cells, the different center-surround orientation selectivity was dependent on the suppressive strength of the cells. We conclude that SN cells are suitable to detect orientation continuity or similarity between CRF and nCRF, whereas the SS cells are adapted to the detection of discontinuity or differences in orientation between CRF and nCRF.     

视觉诱发电位(VEP)方位效应与刺激位置的关系

以人视觉诱发电位(VEP)反应为指标,在视野的不同位置测定了VEP对四个方位的闪烁方波光栅刺激(时间频率2.9Hz,空间频率1.4c/deg,对比度0.94)的反应幅度。在距中央凹20°视角同心圆的八个刺激位置上,VEP反应幅度对与向心线垂直方位的光栅刺激(同心圆的切线方向),有统计意义上的优势。这一规律在垂直、水平向心线上尤为明显。从总体上未发现VEP反应幅度与刺激光栅方位有着明显的关系。这说明在人视野周边区,VEP反应幅度与光栅方位和向心线的夹角(偏向角)相关,而与光栅的绝对方位无关。在相同的刺激条件下,中央区的VEP反应幅度与刺激光栅方位之间也未发现明显关系。     

Are receptive fields of the visual cortex detectors or spatial frequency filters?

Besides its principal maximum, the spatial frequency characteristic curve of the complex visual cortical receptive field of curarized cats also has additional maxima and also negative regions, as predicted by the theory of piecewise Fourier analysis. Comparison of responses of the complex receptive field to sinusoidal gratings completely and incompletely contained in the field and comparison of responses to sinusoidal and square-wave gratings indicate that the receptive field, as a spatial frequency filter, has linear properties. The response of the complex receptive field rises with an increase in the number of periods of the sinusoidal grating. Several periods of optimal frequency match the complex field. Receptive fields tuned to a broad band of spatial frequencies were found in neuron columns. The results confirm the view that complex receptive fields are spatial frequency filters and not detectors.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 11, No. 5, pp. 403–411, September–October, 1979.     

Effect of stimulus intensity on unit responses in the cat frontal cortex

Unit activity of the frontal cortex during changes in stimulus intensity in the near-threshold range (15–16 dB above the threshold for the combined evoked potential) was investigated by an extracellular recording method in acute experiments on cats anesthetized with chloralose (70 mg/kg). Comparative analysis of unit responses in specific (SI) and nonspecific projection areas revealed basically similar changes in pattern during an increase in stimulus intensity: A decrease in the latent period, an increase in the total frequency and the phasic character of the discharge, and an increase in the probability of response. However, a relatively stable latent period and probability of response were observed in specific projection neurons for a stimulus intensity of 3–5 threshold units, whereas for the nonspecific projection neurons it was observed for a stimulus intensity of 10–15 threshold units. All sensory projections in the frontal cortex are formed by two inputs: short-latency low-threshold and long-latency high-threshold. Analysis of modality-dependent differences in the threshold of sensitivity and the latent period of response of the polysensory neurons suggests that stimuli of different modalities converge directly on cortical neurons.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 8, No. 6, pp. 606–612, November–December, 1976.     

Changes in responsiveness of parietal association cortex neurons to stimulation of thalamic association nuclei during development of inhibition

Reactions of neurons of area 7 in the parietal association cortex to paired stimulation of the dorsolateral and posteriolateral thalamic nuclei were investigated using methods of extra- and intracellular recording. It was found that conditioning stimulation of association nuclei caused inhibition or facilitation of the test response; in some cases, the effect was indeterminant. Inhibition frequently led to other types of reactions. Inhibition of response to the test stimulus was pronounced during the initial period of IPSP, but facilitation of the response was possible during the repolarization phase. There was no inhibition of test responses when the chloride component of the IPSP was reversed. It is suggested that inhibition of responses to the test stimulus takes place in the presence of an inward chloride current and that facilitation of responses to such a stimulus may be due to increased effectiveness of late EPSPS.I. I. Mechnikov State University, Odessa. Translated from Neurofiziologiya, Vol. 24, No. 2, pp. 198–207, March–April, 1992.     

Selectivity of visual cortical neurons of rabbits to magnitude of moving stimuli

Unit responses of the rabbit visual cortex were investigated in relation to size of visual stimuli moving in their receptive field. With an increase in size of the stimulus in a direction perpendicular to the direction of movement ("width" of the stimulus) an initial increase in the intensity of the unit response through spatial summation of excitory effects is followed by a decrease through lateral inhibition. This inhibition is observed between zones of the receptive field which behave as activating when tested by a stimulus of small size. Each neuron has its own "preferred" size of stimuli evoking its maximal activation. No direct correlation is found between the "preferred" stimulus size and the size of the receptive field. With a change in stimulus size in the direction of movement ("length" of the stimulus) the responses to stimuli of optimal size may be potentiated through mutual facilitation of the effects evoked by the leading and trailing edges of the stimulus and weakened in response to stimuli of large size. The selective behavior of the neurons with respect to stimulus size is intensified in the case of coordinated changes in their length and width. It is postulated that the series of neurons responding to stimuli of different "preferred" dimensions may constitute a system classifying stimuli by their size.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 6, pp. 636–644, November–December, 1972.     

Movement sensitivities of cells in the fly's medulla

Summary Intracellular recordings were made in the medullae of intact, restrained females ofCalliphora vicina that faced a hemispherical, minimum-distortion surface upon which moving patterns and spots were projected from the rear (Fig. 2). In the distal medulla, noisy hyperpolarizations to light, most likely recorded in terminals of laminar (L) cells, had flicker-like oscillations to moving gratings of 15° spatial wavelength but not of 2.5° spatial wavelength (Fig. 3). Medullary (M) cells penetrated distally responded to grating movements with similar but depolarizing oscillations, in one cell 180° out of phase with a nearby laminar response (Figs. 4–6).A characteristic movement response recorded from most medullary cells consisted of abrupt, maintained nondirectional depolarizations in response to movements of gratings, often with directional ripple or spikes superimposed. When directions of movement reversed, there were brief repolarizations, but when movements stopped, depolarizations decayed away more slowly (Figs. 7 and 8). Magnitude of responses increased with increasing speeds of both 15° and 2.5° gratings (Figs. 9–11). In some cells, there were delayed decays of responses after stopping (Fig. 12). Still other cells seemed to receive inhibition from other, characteristically responding cells (Fig. 13).Receptive fields tested were simple and usually large, with only a suggestion of surround inhibition (Fig. 14). In general, intensity and position were interchangeable over a cell's receptive field (Figs. 15 and 16). Moving edges and dark spots elicited responses primarily within receptive field centers (Figs. 18–20).It is argued that waveforms of characteristic movement responses can be explained by multiplicative inputs from L- and M-cells to movement detectors (Figs. 21–26).Abbreviations L cells laminar (monopolar) cells - M cells medullary cells     

Peculiarities of summation processes in visual-sensitive neurons in the pretectal area of cat

Summation processes occurring in single neurons of the pretectal area in response to either moving or stationary light stimuli were studied in acute experiments on cats. In most neurons studied (85%), gradual increase of the angular size of stimulus resulted in clearly defined summation. In neurons lacking directional sensitivity (nondirectional neurons) the stimulus movement in two opposite directions caused, as a rule, similar and symmetrical changes in the number of spikes, whereas under the same conditions direction-sensitive neurons, in addition to symmetrical development of summation processes, could exhibit substantial differences in the summation curves. The responses to a preferred movement direction could be significantly inhibited or facilitated, while the responses to a non-preferred direction remained stable or changed reciprocally. Neuronal responses to different directions of the movement of stimulus might change independently of each other. This was also the case whenon andoff responses of theon—off neurons to stationary stimuli were compared. It is concluded that neurons of the pretectal area have a complex infrastructure of receptive fields that significantly influences the integration of incoming information.Neirofiziologiya/Neurophysiology, Vol. 25, No. 5, pp. 376–382, September–October, 1993.     

Neuronal response in the cat parietal association cortex to conditioned and non-conditioned acoustic stimuli

Neuronal response in the cat association cortex (area 5) to conditioned and non-conditioned acoustic stimulation was investigated. Numbers of neurons responding to a conditioned acoustic stimulus according to the traditional reflex pattern were twice as high. Numbers of inhibitory neuronal responses to the stimulus increased when instrumental reflex occurred. Neurons were found which only reacted to a conditioned acoustic stimulus in the absence of conditioned reflex movement occurring with instrumental food reflex. Although findings do not exclude the possibility of this cortical area contributing to the analysis of sensory signals and evaluation of their biological significance, it might be supposed that its main functional property lies in its involvement in the process of initiating behavioral response to a conditioned response.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 5, pp. 637–645, September–October, 1988.     

Patterns in the discharge of simple and complex visual cortical cells

The activity of visual cortical neurons (area 17) was recorded in anaesthetized cats in response to sinusoidal drifting gratings. The statistical structure of the discharge of simple and complex cells has been studied as a function of the various parameters of a drifting grating: spatial frequency, orientation, drifting velocity and contrast. For simple cells it has been found that the interspike interval distributions in response to drifting gratings of various spatial frequencies differ only by a time scale factor. They can be reduced to a unique distribution by a linear time transformation. Variations in the spatial frequency of the grating induce variations in the mean firing rate of the cell but leave unchanged the statistical structure of the discharge. On the contrary, the statistical structure of the simple cell activity changes when the contrast or the velocity of the stimulus is varied. For complex cells it has been found that the invariance property described above for simple cells is not valid. Complex cells present in their activity in response to visual stimuli two different firing patterns: spikes organized in clusters and spikes that do not show this organization ('isolated spikes'). The clustered component is the only component of the complex cell discharge that is tuned for spatial frequency and orientation, while the isolated spike component is correlated with the contrast of the stimulus.     

Unit responses of the secondary somatosensory cortex during defensive conditioning in cats

Unit responses in the secondary somatosensory cortex during the formation and extinction of a defensive conditioned reflex to acoustic stimulation were investigated in chronic experiments on cats. In 21 of 28 neurons tested during defensive conditioning the firing pattern changed in accordance with the character of responses to electric shock reinforcement. Two types of conditioned-reflex unit responses were distinguished: excitatory and inhibitory. Most neurons responding to the conditioned stimulus by activation did so during the first 50 msec, which was 80–100 msec before the conditioned motor response. Considerable variability of the unit responses was observed during conditioning. By the time of stabilization of the conditioned-reflex connections the unit response to the conditioned stimulus was stable in form. The pattern of extinction of the conditioned unit activity was expressed as a decrease in the discharge frequency in responses of excitatory type and disinhibition of activity in the case of inhibitory responses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev, Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 232–238, May–June, 1977.     

Dependence of unit responses of the catlateral geniculate body on photic stimulus contrast

Unit responses in the lateral geniculate body of cats to photic stimuli of different contrast were investigated. The number of spikes in the initial phase of the responses (the first 30–45 msec) was found not to change at first, but then to decrease with an increase in the intensity of background illumination. The background intensity starting from which the response diminishes was shown to increase with an increase in the intensity of the test stimulus. The unit response is a linear function of the logarithm of stimulus contrast if the contrast is changed through variation of the intensity of the test stimulus. If contrast increases on account of a decrease in the intensity of background illumination the responses first increase and then remain unchanged. The range of contrasts within which the response is a linear function is narrowed if the intensity of the test stimulus is reduced. Counting the number of spikes in different time intervals of the response (t) showed that the greater the value of t (within the first 70–90 msec of the response) the steeper the curve of the number of spikes as a function of contrast. The Weber-Fechner law applies in the receptive field of the lateral geniculate body. The results are compared with those of some psychophysiological experiments.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 9, No. 3, pp. 267–274, May–June, 1977.     

Spectral Characteristics of Phase Sensitivity and Discharge Rate of Neurons in the Ascending Tectofugal Visual System

Drifting gratings can modulate the activity of visual neurons at the temporal frequency of the stimulus. In order to characterize the temporal frequency modulation in the cat’s ascending tectofugal visual system, we recorded the activity of single neurons in the superior colliculus, the suprageniculate nucleus, and the anterior ectosylvian cortex during visual stimulation with drifting sine-wave gratings. In response to such stimuli, neurons in each structure showed an increase in firing rate and/or oscillatory modulated firing at the temporal frequency of the stimulus (phase sensitivity). To obtain a more complete characterization of the neural responses in spatiotemporal frequency domain, we analyzed the mean firing rate and the strength of the oscillatory modulations measured by the standardized Fourier component of the response at the temporal frequency of the stimulus. We show that the spatiotemporal stimulus parameters that elicit maximal oscillations often differ from those that elicit a maximal discharge rate. Furthermore, the temporal modulation and discharge-rate spectral receptive fields often do not overlap, suggesting that the detection range for visual stimuli provided jointly by modulated and unmodulated response components is larger than the range provided by a one response component.     

Orientation selectivity and spatial frequency characteristics of receptive fields of occipital cortical neurons in cats

Spatial frequency characteristics of receptive fields of occipital cortical neurons were investigated in cats during presentation of visual stimuli consisting of gratings in four or eight standard orientations. The maximal increase in discharge frequency of the neurons was observed when the grating was presented in one particular orientation, which was taken to be optimal for those particular neurons. Responses of some neurons to presentation of gratings in nonoptimal orientations were less than optimal; inhibition of activity below the spontaneous discharge level was observed in other cells in this case. Maximal inhibition was observed to the orientation perpendicular to optimal. Inhibition of unit activity evoked by presentation of gratings in the nonoptimal orientation was shown to be a function of spatial frequency.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 227–232, May–June, 1981.     

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.     

Stationary pattern adaptation and the early components in human visual evoked potentials

Pattern-onset visual evoked potentials were elicited from humans by sinusoidal gratings of 0.5, 1, 2 and 4 cpd (cycles/degree) following adaptation to a blank field or one of the gratings. The wave forms recorded after blank field adaptation showed an early positive component, P0, which decreased in amplitude with spatial frequency, whereas the immediately succeeding negative component, N1, increased in amplitude with spatial frequency. P0 and N1 components of comparable size were recorded at 1 cpd. Stationary pattern adaptation to a grating of the same spatial frequency as the test grating significantly reduced N1 amplitude at 4, 2 and 1 cpd. The N1 component elicited at 4 cpd was attenuated in log-linear fashion as the spatial frequency of the adaptation grating increased. P0, on the other hand, was unaffected by stationary pattern adaptation at all combinations of test and adapting spatial frequencies, although P0 amplitude is known to be attenuated by adaptation to a drifting grating. Since N1, but not P0, was significantly attenuated following adaptation and testing at 1 cpd, it was concluded that the neurons generating these components are functionally distinct. The use of a common adaptation grating discounted the possibility that N1, but not P0, was affected due to a difference in the rates of retinal image modulation caused by eye movements made while viewing adaptation gratings of different spatial frequencies. The neurons generating N1 were adapted at a lower rate of retinal image modulation than that apparently required for adaptation of the neurons generating P0, which suggests a difference between these neurons in the rate of stimulus modulation necessary for activation.     

Analysis of spike discharge of a neuron population in the cat motor cortex during a conditioned change of posture reflex

Spike responses of area 4 neurons in the projection area of the contralateral forelimb to acoustic stimulation (1 sec), which became the conditioned stimulus after training, and to dropping of the platform beneath the test limb, which served as reinforcing stimulus, were studied in trained and untrained cats. Responses only of those neurons which were activated during a passive movement caused by dropping of the platform were studied. In trained animals the number of these neurons which responded to the conditioned stimulus if a reflex occurred was 100%, and in the absence of conditioned-reflex movements to the conditioned stimulus it was 70%, much greater than the number of neurons responding to the same acoustic stimulus in untrained animals (45%). On peristimulus histograms of responses of the test neuron population in untrained and trained animals to acoustic stimulation (in the absence of movements) only the initial spike response with a latent period of under 50 msec and a duration of up to 100 msec could be clearly distinguished. In the presence of reflex movement multicomponent spike responses were observed: an initial spike response and early and late after-responses linked with performance of conditioned-reflex limb flexion. Early after-responses 100–200 msec in duration, appearing after a latent period of 100–150 msec, were linked to the time of application of the conditioned stimulus, whereas the appearance and duration of late after-responses were determined by the time of onset of conditioned-reflex movement. The magnitude of the neuronal response to reinforcement in trained animals does not depend on the appearance of the conditioned movement.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 93–102, January–February, 1985.     

Unit responses of the inferior colliculi to changes in the intensity of an amplitude-modulated signal

Unit responses of the inferior colliculi of anesthetized rats to amplitude-modulated sounds during a change in the carrier intensity were investigated. The following unit response characteristics were assessed: the number of spikes in the response, the range of reproduction of the modulation frequency, the response duration, and the pattern of the spike response relative to the envelope of the amplitude-modulated stimulus. The following parameters of the stimulus were varied: carrier intensity (usually of optimal frequency or noise), frequency of modulation (from 2 to 100 Hz), and carrier frequency. With a decrease in the intensity of the carrier in the case of monotonic neurons, and also with an increase or decrease in the intensity of the carrier relative to its optimal level in nonmonotonic neurons, the following changes in the discharge were regularly observed: the number of spikes in the response and its duration were reduced down to the appearance of only one initial response, the range of reproduction of the rhythm of modulation was narrowed, and the response pattern was sharply modified.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR. I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 4, pp. 355–366, July–August, 1973.