A ring model for spatiotemporal properties of simple cells in the visual cortex

 A neural model is proposed for the spatiotemporal properties of simple cells in the visual cortex. In the model, several cortical cells are arranged on a ring, with mutual excitatory or inhibitory connections. The cells also receive excitatory inputs either from lagged and nonlagged cells of the lateral geniculate nucleus in one setting or from nonlagged cells in the other. Computer simulation shows that the cortical cells have spatiotemporally inseparable receptive fields in the former setting and separable fields in the latter; spatial profiles at a given time in the spatiotemporal fields are described with a Gabor function whose phase parameter varies regularly from 0 to 2π with rotation along the ring; the inseparable cells have directional selectivity as observed physiologically. Received: 13 November 1995 / Accepted in revised form: 1 July 1997     

Orientation-selective neurons in the squirrel visual cortex

The organization of receptive fields of neurons sensitive to orientation of visual stimuli was investigated in the squirrel visual cortex. Neurons with mutually inhibitory on- and off-areas of the receptive field, with partially and completely overlapping excitatory and inhibitory mechanisms, were distinguished. Neurons of the second group are most typical. They exhibit orientation selectivity within the excitatory area of the receptive field because, if the stimulus widens in the zero direction, perpendicular to the preferred direction, lateral inhibition is much stronger than if it widens in the preferred direction. Additional inhibitory areas (outside the excitatory area) potentiate this inhibition and increase selectivity. It is suggested that there is no strict separation of simple (with separate excitatory and inhibitory mechanisms in the receptive field) and complex (with overlapping of these mechanisms) neurons in the squirrel visual cortex.A. N. Severtsov Institute of Evolutionary Morphology and Ecology of Animals, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 11, No. 6, pp. 540–549, November–December, 1979.     

Transformation of peripheral inputs by the first-order lateral line brainstem nucleus

Extracellular recording techniques were used to record the responses of medial nucleus cells and posterior lateral line nerve fibers in mottled sculpin, Cottus bairdi, and goldfish, Carassius auratus, to a 50-Hz dipole source (vibrating sphere). Responses were characterized in terms of (1) receptive fields that relate responsiveness (spike rate and phase-locking) to the location of the source along the length of the fish, (2) input-output functions that relate responsiveness to vibration amplitude for a fixed source location, and (3) peri-stimulus time histograms that relate responsiveness to time during a sustained period of vibration. Relative to posterior lateral line nerve fibers, medial nucleus cells in both species were similar in showing (1) lower spontaneous and evoked rates of spike activity, (2) greater degrees of adaptation, (3) greater heterogeneity in all response characteristics, and (4) evidence for inhibitory/excitatory interactions. Whereas receptive fields of nerve fibers in both species faithfully reflect both pressure gradient amplitudes (with rate changes) and directions (with phase-angle changes) in the stimulus field, receptive fields of medial nucleus were more difficult to relate to the stimulus field. Some, but not all, receptive fields could be modeled with excitatory center/inhibitory surround and inhibitory center/excitatory surround organizations. Accepted: 26 November 1997     

Characteristics of dorsal horn neurons expressing subliminal responses to sural nerve stimulation

The present study was designed (1) to characterize the subliminal responses of dorsal horn neurons to stimulation of the sural nerve, and (2) to correlate the type of response to this stimulus with the responses to natural mechanical stimulation of the skin. To accomplish this, intracellular and extracellular recordings were carried out in L6 and L7 dorsal horn neurons in the cat. The excitatory responses of each cell to electrical stimulation of the sural nerve and to mechanical stimulation of the skin were noted. Of 35 dorsal horn cells recorded intracellularly, 11 responded with impulses to sural nerve stimulation, 9 responded with excitatory postsynaptic potentials (EPSPs) but not impulses, and 15 had no excitatory responses to this stimulus. The type of response to sural nerve stimulation was strongly correlated with receptive field modality. Most cells receiving an input from high-threshold cutaneous mechanoreceptors responded with impulses or gave no excitatory response to sural nerve stimulation, whereas most cells that had only low-threshold mechanoreceptor input responded with EPSPs only or gave no response. In cells with only low-threshold (LT) mechanoreceptive input, response to sural nerve stimulation was highly correlated with receptive field locus. Those LT cells with no excitatory responses to sural nerve stimulation had receptive fields confined to the foot and/or toes, whereas those that gave EPSPs had more proximal receptive fields. The possible significance of these data with reference to changes observed after lesions, such as increased response to sural nerve stimulation, increased receptive field size, and somatotopic reorganization, is discussed.     

15th Annual spring brain conference,march 10–13, 2004,summary report

The present study was designed (1) to characterize the subliminal responses of dorsal horn neurons to stimulation of the sural nerve, and (2) to correlate the type of response to this stimulus with the responses to natural mechanical stimulation of the skin. To accomplish this, intracellular and extracellular recordings were carried out in L⁶ and L⁷ dorsal horn neurons in the cat. The excitatory responses of each cell to electrical stimulation of the sural nerve and to mechanical stimulation of the skin were noted.

Of 35 dorsal horn cells recorded intracellularly, 11 responded with impulses to sural nerve stimulation, 9 responded with excitatory postsynaptic potentials (EPSPs) but not impulses, and 15 had no excitatory responses to this stimulus. The type of response to sural nerve stimulation was strongly correlated with receptive field modality. Most cells receiving an input from high-threshold cutaneous mechanoreceptors responded with impulses or gave no excitatory response to sural nerve stimulation, whereas most cells that had only low-threshold mechanoreceptor input responded with EPSPs only or gave no response. In cells with only low-threshold (LT) mechanoreceptive input, response to sural nerve stimulation was highly correlated with receptive field locus. Those LT cells with no excitatory responses to sural nerve stimulation had receptive fields confined to the foot and/or toes, whereas those that gave EPSPs had more proximal receptive fields. The possible significance of these data with reference to changes observed after lesions, such as increased response to sural nerve stimulation, increased receptive field size, and somatotopic reorganization, is discussed.     

Inhibitory zones of receptive fields of the lateral geniculate body and visual cortex of the cat

Unit responses to moving strips were investigated. The organization of the inhibitory zones in the receptive fields of the lateral geniculate body and visual cortex of the cat was compared. The response in the receptive field of the lateral geniculate body was inhibited only during simultaneous stimulation of the excitatory and inhibitory zones of the field. Stimulation of the inhibitory zone in the receptive field of the visual cortex was effective for a long time (several hundreds of milliseconds) after stimulation of the excitatory zone. The inhibitory zones of the simple and complex receptive fields of the visual cortex differed significantly. An increase in the width of the strip above the optimal size reduced the inhibitory effect in the complex fields. This was not observed in the simple receptive fields. The functional and structural models of the receptive field of the visual cortex are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 5, No. 2, pp. 201–209, March–April, 1973.     

Prediction of orientation selectivity from receptive field architecture in simple cells of cat visual cortex

From the intracellularly recorded responses to small, rapidly flashed spots, we have quantitatively mapped the receptive fields of simple cells in the cat visual cortex. We then applied these maps to a feedforward model of orientation selectivity. Both the preferred orientation and the width of orientation tuning of the responses to oriented stimuli were well predicted by the model. Where tested, the tuning curve was well predicted at different spatial frequencies. The model was also successful in predicting certain features of the spatial frequency selectivity of the cells. It did not successfully predict the amplitude of the responses to drifting gratings. Our results show that the spatial organization of the receptive field can account for a large fraction of the orientation selectivity of simple cells.     

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.     

Brainstem lateral line responses to sinusoidal wave stimuli in the goldfish, Carassius auratus

Extracellular recordings were made from single lateral line units in the medial octavolateralis nucleus in the brainstem of goldfish, Carassius auratus. Units were defined as receiving lateral line input if they responded to the water motions generated by a stationary, sinusoidally oscillating sphere and/or a moving sphere but not to airborne sound and vibrations. Units which responded to airborne sound or vibrations were assumed to receive input from the inner ear and were not further investigated. Responses of lateral line units were quantified in terms of the number of evoked spikes and the degree of phase-locking to a 50 Hz vibrating sphere presented at various stationary locations along the side of the fish. Receptive fields were characterized based on spike rate, degree of phase-locking and average phase angle as a function of sphere location. Four groups of units were distinguished: 1, units with receptive fields comparable to those of primary afferents; 2, units with receptive fields which consisted of one excitatory and one inhibitory area; 3, units with receptive fields which consisted of more than two excitatory and/or inhibitory areas; 4, units with receptive fields which consisted of a single excitatory or a single inhibitory area. The receptive fields of most units were characterized by adjacent excitatory and inhibitory areas. This organization is reminiscent of excitatory-inhibitory receptive field organizations in other vertebrate sensory systems.     

The spatial substructure of visual receptive fields in the cat's superior colliculus

Although the direction selective properties of the superficial layer cells of the cat's superior colliculus have been extensively studied, the mechanisms underlying this property remain controversial. With the aim to understand the mechanism(s) underlying directional selectivity of collicular neurons we examined the substructure of their visual receptive fields. 1. The strength of cell responses and the direction selectivity indices varied in relation to the location of the tested region within the receptive field and the amplitude of stimulus movement. 2. Decrease of the amplitude of motion resulted in a decrease of direction selectivity index both in the group of direction-selective cells and in the group of cells classified as direction nonselective but with a directional bias. 3. The decrease of direction selectivity for small amplitude movement resulted mainly from increase in the magnitude of response in the nonpreferred direction of movement. 4. These results suggest that the receptive fields of most collicular cells are composed of subregions with different response profiles and indicate that inhibitory mechanisms dictate direction selectivity of collicular cells.     

Somatosensory cortical unit responses to stimulation of the vibrissae in cats

Characteristics of extra- and intracellular responses of 57 neurons in the vibrissal projection zone of the first somatosensory area of the cat cortex were investigated. The intensity of both excitatory and inhibitory unit responses was found to diminish during successive stimulation of different parts of the receptive fields in the direction from center toward periphery. Usually, when central parts of receptive fields were stimulated, inhibition in the unit responses was postexcitatory, whereas when peripheral parts were stimulated inhibition could precede excitation. The possibility of an increase in the role of interaction between excitatory and inhibitory processes arising in neurons in response to vibrissal stimulation with an increase in the distance from center to periphery of receptive fields of single cortical cells is discussed. Neurons found during one insertion of the microelectrode were seen to have common center for their receptive fields, but the diameters of the receptive fields of individual neurons could differ significantly. Moreover, during such vertical insertions responses of neurons with primary inhibition to the stimuli presented were recorded.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 2, pp. 124–130, March–April, 1980.     

Stereoscopic mechanisms: binocular responses of the striate cells of cats to moving light and dark bars

New knowledge concerning the internal structure and response properties of the receptive fields of striate cells calls for a fresh appraisal of their binocular interactions in the interest of a better understanding of the neural mechanisms underlying binocular depth discrimination. Binocular position-disparity response profiles were recorded from 71 simple and B-cells in response to moving light and dark bars. Predominantly excitatory (PE) cells (N = 48) had disparity response profiles that were spatially closely similar to their respective monocular responses. In addition, the centrally located excitatory subregions were flanked on one or both sides by non-specific inhibitory regions. PE cells with a preferred stimulus orientation within 30 degrees of the vertical (N = 17) showed binocular facilitations with maximal values that were always more than twice (mean 3.3) the sum of the two monocular responses to the same stimuli and generally greater than the facilitations shown by cells with orientations more than 30 degrees from the vertical (N = 29; mean 2.2 times the sum of the respective monocular responses). The strength of the binocular facilitation depended on the stimulus contrast, the facilitation decreasing with increasing contrast. The receptive-field disparity distribution of the 31 PE cells capable of making significant horizontal disparity discriminations has standard deviations of 0.37 degrees and 0.40 degrees, respectively. Predominantly inhibitory cells (PI) (N = 23) showed two basic types of disparity response profile: symmetric (N = 17) and asymmetric (N = 6). Uncertainty regarding the precise location of the binocular fixation point in the anaesthetized and paralysed preparation made it difficult to categorize PI cells adequately.     

Dopaminergic modulation of visual responses in toads II. Influences of apomorphine on retinal ganglion cells and tectal cells

The behavioral studies of Part I have shown in common toads that after systemic administration of the dopamine agonist apomorphine the prey-directed orienting turning movements are suppressed while prey snapping is facilitated. Part II focusses on retinal and tectal single cell responses to moving objects. (1) After systemic administration of apomorphine, the discharge rates of retinal class R2 and R3 ganglion cell fibres – recorded from the retino-tectal projection – speeded up in response to visual objects traversing their excitatory receptive fields. This enhancing effect was independent of the recording site in the retino-tectal map. (2) The diameters of the excitatory receptive fields of R2 and R3 neurons doubled their sizes. Probably, apomorphine enhances the center-dominated excitatory responses at the expense of the strength of the inhibitory surround. (3) The apomorphine-induced effects were fully developed 20–35 min after drug administration. (4) At the same time the discharge rates of T5.1 and T5.2 tectal neurons were reduced under apomorphine. The effect was independent of the recording site in the retino-tectal map. The diameters of the excitatory receptive fields of these tectal neurons were not influenced. (5) To changes in configurational stimulus features, the basic pattern of discrimination was maintained. (6) It is suggested that tectal output to the turn-generating motor network – mediated by T5.1 and T5.2 neurons – is modulated by a pretecto-tectal pathway which involves dopaminergic pretectal cells. (7) The enhanced snapping can be interpreted in terms of a modulation of reticular/hypoglossal structures by dopaminergic preoptic/hypothalamic/solitary systems.     

Excitatory and inhibitory receptive fields of tectal cells are differentially modified by magnocellular and parvocellular divisions of the pigeon nucleus isthmi

It has been known that magnocellular and parvocellular divisions of the pigeon nucleus isthmi exert excitatory and inhibitory actions on tectal cells, respectively. The present study shows that injection of N-methyl-D-aspartate into the parvocellular division results in an increase in responsive strength and extent of the inhibitory receptive fields, which expand into the excitatory receptive fields of tectal cells. This injection concurrently leads to a decrease in responsiveness and extent of the excitatory fields. On the other hand, injection of acetylcholine into the magnocellular division enhances visual responsiveness, although the excitatory field is not obviously changed in extent. Meanwhile, strength and extent of the inhibitory fields are decreased by acetylcholine. The excitatory and inhibitory fields are reduced in both strength and extent by magnocellular and parvocellular injection of lidocaine, respectively. It suggests that isthmic inputs from both parvocellular and magnocellular divisions converge onto the same tectal cells, and the magnocellular and parvocellular subnuclei can modulate excitatory and inhibitory receptive fields of tectal cells, respectively, with some interactions between both fields. Accepted: 1 March 2000     

Stimulus size selectivity and receptive field organization of ectostriatal neurons in the pigeon

The avian ectostriatum is the telencephalic recipient zone of the tectofugal pathway, which may be homologous to the colliculo-pulvinar-cortical pathway in mammals. The present paper studies the visual response properties and receptive field organization of ectostriatal cells in pigeons with extracellular recording and computer mapping techniques. The results indicate that 90% of ectostriatal cells prefer forward, downward and upward motion of visual stimuli at velocities in the range of 16-128 degrees s(-1). They respond optimally to small stimuli (1-4 degrees visual angle), mostly preferring a target of 2 degrees. Most cells (78.8%) have one excitatory receptive field that usually possesses one or two hotspots, some cells (13.5%) have two excitatory receptive fields each having one or two hotspots, and a few cells (7.7%) have one excitatory receptive field with an unresponsive region in the center. An inhibitory receptive field is not found surrounding the excitatory receptive field in the ectostriatal cells examined. These response properties and receptive field organization may reflect the possible roles of the ectostriatum in stimulus discrimination and visual cognition.     

Characteristics of inhibition in receptive fields of the cat visual cortex

Unit responses of neurons of zone 17 in the cat striate cortex to stripes of different widths were studied. Changes in the number of spikes during different time intervals (cuts) from the beginning of the response were analyzed in relation to stimulus area. Comparison of the results with those obtained by the study of receptive fields of the lateral geniculate body showed a significant difference in the dynamics of inhibition between cortical and geniculate receptive fields. Similar results were obtained when cortical unit responses to simultaneous and consecutive appearance of two stripes in the receptive field, one in the excitatory zone and the other at the inhibitory periphery, were studied. Evidence of the longer duration of cortical inhibition also was obtained by the same technique. When both stripes were placed in the excitatory center of the field another feature of cortical inhibition was revealed: its dependence on the order of stimulus application. If the order of stimulus application coincided with the optimal direction of movement of the stripe for the given field, the unit response to the next stimulus was strongly facilitated by the action of the stimulus applied previously. Application of stimuli in the opposite order invoked inhibition. The sensitivity of inhibition to the order of stimulus application was observed in the center of the field; it diminished toward the periphery, where application of the stimuli in any order evokes inhibition of the response.Medical Academy, Sofia, Bulgaria, I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 9, No. 4, pp. 339–346, July–August, 1977.     

Spatial and Feature-Based Attention in a Layered Cortical Microcircuit Model

Directing attention to the spatial location or the distinguishing feature of a visual object modulates neuronal responses in the visual cortex and the stimulus discriminability of subjects. However, the spatial and feature-based modes of attention differently influence visual processing by changing the tuning properties of neurons. Intriguingly, neurons'' tuning curves are modulated similarly across different visual areas under both these modes of attention. Here, we explored the mechanism underlying the effects of these two modes of visual attention on the orientation selectivity of visual cortical neurons. To do this, we developed a layered microcircuit model. This model describes multiple orientation-specific microcircuits sharing their receptive fields and consisting of layers 2/3, 4, 5, and 6. These microcircuits represent a functional grouping of cortical neurons and mutually interact via lateral inhibition and excitatory connections between groups with similar selectivity. The individual microcircuits receive bottom-up visual stimuli and top-down attention in different layers. A crucial assumption of the model is that feature-based attention activates orientation-specific microcircuits for the relevant feature selectively, whereas spatial attention activates all microcircuits homogeneously, irrespective of their orientation selectivity. Consequently, our model simultaneously accounts for the multiplicative scaling of neuronal responses in spatial attention and the additive modulations of orientation tuning curves in feature-based attention, which have been observed widely in various visual cortical areas. Simulations of the model predict contrasting differences between excitatory and inhibitory neurons in the two modes of attentional modulations. Furthermore, the model replicates the modulation of the psychophysical discriminability of visual stimuli in the presence of external noise. Our layered model with a biologically suggested laminar structure describes the basic circuit mechanism underlying the attention-mode specific modulations of neuronal responses and visual perception.     

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.