Showing papers on "Receptive field published in 1985"

Selective attention gates visual processing in the extrastriate cortex.

Abstract: Single cells were recorded in the visual cortex of monkeys trained to attend to stimuli at one location in the visual field and ignore stimuli at another. When both locations were within the receptive field of a cell in prestriate area V4 or the inferior temporal cortex, the response to the unattended stimulus was dramatically reduced. Cells in the striate cortex were unaffected by attention. The filtering of irrelevant information from the receptive fields of extrastriate neurons may underlie the ability to identify and remember the properties of a particular object out of the many that may be represented on the retina.

Encoding of spatial location by posterior parietal neurons.

Abstract: The cortex of the inferior parietal lobule in primates is important for spatial perception and spatially oriented behavior. Recordings of single neurons in this area in behaving monkeys showed that the visual sensitivity of the retinotopic receptive fields changes systematically with the angle of gaze. The activity of many of the neurons can be largely described by the product of a gain factor that is a function of the eye position and the response profile of the visual receptive field. This operation produces an eye position-dependent tuning for locations in head-centered coordinate space.

Stimulus Specific Responses from Beyond the Classical Receptive Field: Neurophysiological Mechanisms for Local-Global Comparisons in Visual Neurons

Abstract: We perceive the visual world as a unitary whole, yet one of the guiding principles of nearly a half century of neurophysiological research since the early recordings by Hartline (1938) has been that the visual system consists of neurons that are driven by stimulation within small discrete portions of the total visual field. These classical receptive fields (CRFs) have been mapped with the excitatory responses evoked by a flashed or moving stimulus, usually a spot or bar of light. Most of the visual neurons, in turn, are organized in a series of maps of the visual field, at least 10 of which exist in the visual cortex in primates as well as additional topographic representations in the lateral geniculate body, pulvinar and optic tectum (Allman 1977, Newsome & Allman 1980, Allman & Kaas 1984). It has been widely assumed that perceptual functions that require the integration of inputs over large portions of the visual field must be either collective properties of arrays of neurons representing the visual field, or features of those neurons at the highest processing levels in the visual system, such as the cells in inferotemporal or posterior parietal cortex that typically possess very large receptive fields and do not appear to be organized in visuotopic maps. These assumptions have been based on the results of the many studies in which receptive fields were mapped with conventional stimuli, presented one at a time, against a featureless background. However, unlike the neurophysiologist's tangent screen, the natural visual scene is rich in features, and there is a growing body of evidence that in many visual neurons stimuli presented outside the CRF strongly and selectively influence neural responses to stimuli presented within the CRF. These results suggest obvious mechanisms for local-global comparisons within visuotopically organized structures. Such broad and specific surround mechanisms could participate in many functions that require the integration of inputs over wide regions of the visual space such as the perceptual constancies, the segregation of figure from ground, and depth perception through motion parallax. In the first section of this paper, we trace the historical development of the evidence of response selectivity for visual stimuli presented beyond the CRF; in the second, examine the anatomical pathways that sub serve these far-reaching surround mechanisms; and in the third, explore the possible relationships between these mechanisms and perception.

Direction- and velocity-specific responses from beyond the classical receptive field in the middle temporal visual area (MT).

Abstract: The true receptive field of more than 90% of neurons in the middle temporal visual area (MT) extends well beyond the classical receptive field (crf), as mapped with conventional bar or spot stimuli, and includes a surrounding region that is 50 to 100 times the area of the crf. These extensive surrounds are demonstrated by simultaneously stimulating the crf and the surround with moving stimuli. The surrounds commonly have directional and velocity-selective influences that are antagonistic to the response from the crf. The crfs of MT neurons are organized in a topographic representation of the visual field. Thus MT neurons are embedded in an orderly visuotopic array, but are capable of integrating local stimulus conditions within a global context. The extensive surrounds of MT neurons may be involved in figure–ground discrimination, preattentive vision, perceptual constancies, and depth perception through motion cues.

Contrast gain control in the cat's visual system

Abstract: We have examined the idea that the adaptation of cortical neurons to local contrast levels in a visual stimulus is functionally advantageous. Specifically, cortical cells may have large differential contrast sensitivity as a result of adjustments that center a limited response range around a mean level of contrast. To evaluate this notion, we measured contrast-response functions of cells in striate cortex while systematically adapting them to different contrast levels of stimulus gratings. For the majority of cortical neurons tested, the results of this basic experiment show that contrast-response functions shift laterally along a log-contrast axis so that response functions match mean contrast levels in the stimulus. This implies a contrast-dependent change in the gain of the cell's contrast-response relationship. We define this process as contrast gain control. The degree to which this contrast adjustment occurs varies considerably from cell to cell. There are no obvious differences regarding cell type (simple vs. complex) or laminar distribution. Contrast gain control is almost certainly a cortical function, since lateral geniculate cells and fibers exhibit only minimal effects. Tests presented in the accompanying paper (37) provide additional evidence on the cortical origin of the process. In another series of experiments, the effect of contrast adaptation on physiological estimates of contrast sensitivity was evaluated. Sustained adaptation to contrast levels as low as 3% was capable of nearly doubling the thresholds of most of the cells tested. Adaptation may therefore be an important factor in determinations of the contrast sensitivity of cortical neurons. We tested the spatial extent of the mechanisms responsible for these gain-control effects by attempting to adapt cells using both a large grating and a grating patch limited to that portion of a cell's receptive field from which excitatory discharges could be elicited directly (the central discharge region). Adaptation was found to be an exclusive property of the central region. This held even in the case of hypercomplex cells, which received strong influences from surrounding regions of the visual field. Finally, we measured the time course of contrast adaptation. We found the process to be rather slow, with a mean time constant of approximately 6 s. Once again, there was considerable variability in this value from cell to cell.

Segregation of efferent connections and receptive field properties in visual area V2 of the macaque

Abstract: V2 is a visual area of the macaque monkey which is at the second level in a recently proposed hierarchy of cortical visual areas1. Histochemical staining for cytochrome oxidase (CO) in V2 reveals a pattern of alternate thick and thin CO-rich stripes separated by CO-sparse interstripes2,3. These subregions receive distinct inputs from neurones in CO-rich and CO-sparse zones arrayed within the superficial layers of V1 (refs 4,5). Are output projections from V2 to higher visual areas also segregated? Using an anatomical double-label paradigm, we have now demonstrated that V2 cells projecting to two of its major target areas, MT and V4 (refs 6, 7), are arranged in stripe-like clusters which are largely segregated from one another and which are closely related to the pattern of CO stripes. Concomitant electrophysiological recordings from V2 indicate that groups of cells having similar receptive field properties are clustered within the subregions defined by these anatomical techniques.

Inferior olivary neurons in the awake cat: detection of contact and passive body displacement

Abstract: We have recorded from 306 neurons in the inferior olive of six alert cats. Most of the cats were trained to perform a simple task with the forelimb. We observed the neural responses to a wide variety of cutaneous and proprioceptive stimuli, as well as responses during spontaneous and learned active movements. Neurons responsive to somatosensory stimulation were found in all parts of the inferior olive, and they were roughly evenly divided between those responsive to cutaneous stimulation and those responsive to proprioceptive stimulation. In the dorsal accessory olive all neurons were responsive to somatosensory stimulation. In the medial accessory nucleus 88% and in the principal olive 74% of cells were responsive to somatosensory stimulation. Cells responsive to cutaneous stimulation usually had small receptive fields, commonly on the paw. These cells had low-threshold responses to one or more forms of cutaneous stimulation and typically fired one spike at the onset of the stimulus on 80% or more of stimulus applications. Cells responsive to proprioceptive stimulation most commonly responded to passive displacements of a limb. These cells were often very sensitive, responding to linear displacements of less than 1 cm in one specific direction. No cells in our sample responded reliably during active movement by the animal. Only 21% of cells responding to passive proprioceptive stimulation showed any modulation during active movement, and the modulation was weak. Likewise, cells responsive to cutaneous stimulation generally failed to respond when a similar stimulus was produced by an active movement by the animal. Exceptions to this were stimuli produced during exploratory movements or when the receptive field unexpectedly made contact with an object during active movement. Electrical stimulation applied in the inferior olive failed to evoke movements or to modify ongoing movement. Our results are consistent with the hypothesis that inferior olivary neurons function as somatic event detectors responding particularly reliably to unexpected stimuli.

Pulvinar nuclei of the behaving rhesus monkey: visual responses and their modulation

Abstract: We have examined the properties of neurons in three subdivisions of the pulvinar of alert, trained rhesus monkeys 1) an inferior, retinotopically mapped area (PI), 2) a lateral, retinotopically organized region (PL), and 3) a dorsomedial visual portion of the lateral pulvinar (Pdm), which has a crude retinotopic organization. We tested the neurons for visual responses to stationary and moving stimuli and for changes in these responses produced by behavioral manipulations. All areas contain cells sensitive to stimulus orientation as well as neurons selective for the direction of stimulus movement; however, the majority of cells in all three regions are either broadly tuned or nonselective for these attributes. Nearly all cells respond to stimulus onset, a significant number also give a response to stimulus termination, and rarely a cell gives only off responses. Nearly all cells increase their discharge rate to visual stimuli. Receptive fields in the two retinotopically mapped regions, PI and PL, have well-defined borders. The sizes of these receptive fields show a positive correlation with the eccentricity of the receptive fields. The receptive fields in the remaining region, Pdm, are frequently very large, but with these large fields excluded, show a similar correlation with eccentricity. All pulvinar cells tested (n = 20) were mapped in retinal coordinates; the receptive fields are positioned in relation to the retina. We found no cells with gaze-gated characteristics (2), nor cells mapped in a spatial coordinate system. The response latencies in PI and PL are shorter and less variable than the latencies in Pdm. Active use of a stimulus can produce an enhancement or attenuation of the visual response. Eye-movement modulation was found in all three subdivisions in about equal frequencies. Attentional modulation was common in Pdm and was rare in PI and PL. The modulation is spatially selective in Pdm and nonselective in PI for a small number of tested cells. These data demonstrate functional differences between Pdm and the other two areas and suggest that Pdm plays a role in selective visual attention, whereas PI and PL probably contribute to other aspects of visual perception.

Temporal and spatial integration in the rat SI vibrissa cortex

Abstract: Glass micropipettes were used to record the activity of 124 single units in the somatosensory vibrissa cortex (SI) of 16 rats in response to combined deflections of contralateral vibrissae. Compact multiangular electromechanical stimulators were used to stimulate individual vibrissal hairs alone or in combinations of two or three adjacent whiskers. Each whisker was stimulated independently to produce controlled temporal and spatial patterns of mechanical stimuli. Following displacement of a vibrissa, unit discharges to subsequent deflections of adjacent whiskers are reduced in a time-dependent fashion. Response suppression is strongest at short interdeflection intervals, i.e., 10-20 ms and decreases progressively during the 50-100 ms following the first deflection. In many cases this period also corresponds with a reduction in ongoing unit discharges. Response suppression was not observed for first-order neurons recorded in the trigeminal ganglion of barbiturate-anesthetized rats. In the cortex, the presence and/or degree of response suppression depends on a number of spatial factors. These include 1) the angular direction(s) in which the individual hairs are moved, 2) the sequence in which two whiskers are deflected, that is, which one is deflected first, 3) the particular combination of whiskers stimulated, and 4) the number (2 or 3) of vibrissae comprising the multiwhisker stimulus. Within a vertical electrode penetration, one particular whisker typically elicits the strongest excitatory and inhibitory effects; other, nearby vibrissae elicit variable (or no) excitation or inhibition. Excitatory and inhibitory subregions of a receptive field could thus be distributed asymmetrically around the maximally effective whisker. In these cases, the receptive fields displayed spatial orientations. Quantitative criteria were used to classify 30 cortical units on the basis of the distribution of inhibitory subregions on either side of the maximally effective whisker. Twenty-one of these cells had receptive fields (RFs) with symmetrical inhibitory side regions. Responses of the other nine units were strongly suppressed by a preceding deflection of a vibrissa on one side but relatively unaffected, or even slightly facilitated, by preceding deflection of the whisker on the other side.(ABSTRACT TRUNCATED AT 400 WORDS)

Intracortical facilitation among co-oriented, co-axially aligned simple cells in cat striate cortex.

Abstract: Most neurons in cat striate visual cortex show inhibitory effects when moving contours are presented beyond the limits of classic receptive field regions. Facilitatory effects are also present in about 40% of simple cells. Here, we report a highly specific form of this facilitation, mediated only by neurons possessing both an orientation tuning matched to the test unit, and a receptive field position aligned with its long axis. This finding illustrates one of the intracortical interconnection schemes hypothesized by Mitchison and Crick (1982). Periodic clustering in long, intrinsic axons may signify a neuron seeking specific functional interactions like these across columnar systems in both the spatial and orientation domains.

The morphology of physiologically identified GABAergic neurons in the somatic sensory part of the thalamic reticular nucleus in the cat

Abstract: Neurons with somatic sensory receptive fields were examined electrophysiologically in the thalamic reticular nucleus of the cat. All cells had receptive fields much larger than those of neurons in the ventral posterior nucleus and were driven by less readily defined somesthetic stimuli. Response latencies to peripheral or medial lemniscal stimulation were, on average, longer than in the ventral posterior nucleus and suggested activation of the reticular nucleus cells by collaterals of thalamocortical relay cell axons arising in the ventral posterior nucleus. When injected intracellularly with horseradish peroxidase, reticular nucleus cells displayed thin axons with intrareticular collaterals and diffuse branches through much of the ventral posterior and posterior thalamic nuclei. Dendrites ended in processes resembling synaptic terminals. Electron microscopic immunocytochemistry of the same part of the reticular nucleus revealed processes immunoreactive for glutamic acid decarboxylase and identifiable as both collateral axon terminals and presynaptic dendrites of GABAergic reticular nucleus cells. These synaptically linked reticular nucleus cells and, in addition, immunoreactive somata and presynaptic dendrites received synapses from at least three varieties of nonimmunoreactive profiles.

Synaptic connectivity of a local circuit neurone in lateral geniculate nucleus of the cat.

Abstract: Although receptive fields of relay cells in the lateral geniculate nucleus of the cat nearly match those of their retinal afferents1,2, only 10–20% of the synapses on these cells derive from the retina and are excitatory3,4. Many more (30–40%) are inhibitory and largely control the gating of retinogeniculate transmission3–7. These inhibitory synapses derive chiefly from two cell types: intrinsic local circuit neurones and cells in the adjacent perigeniculate nucleus5–7. It has been difficult to study the functional organization of these inhibitory pathways; most efforts have relied on indirect approaches6–12. Here we describe the use of direct techniques to study a local circuit neurone by iontophoresing horseradish peroxidase (HRP) into it, which completely labels the soma and processes of cells for subsequent light- and electron microscopic analysis. Although the response properties of the labelled cell are virtually indistinguishable from those of many relay cells6, its morphology is typical of ‘class 3’ neurones13 (see Fig. 1 legend), which are widely believed to be inter neu rones8–11 (but see ref. 12). Here, we refer to the cell as a ‘local circuit neurone’, which allows for the possibility of a projection axon, rather than as an ‘inter-neurone’, a term that commonly excludes a projection axon. We find that the labelled cell has a myelinated axon, but that the axon loses its myelin within 50 µm of the soma and has not yet been traced further. The dendrites of the labelled cell possess presynaptic terminals that act as intrinsic sources of inhibition on geniculate relay cells. We also characterize other morphological aspects of this inhibitory circuitry.

The post-natal development of cutaneous afferent fibre input and receptive field organization in the rat dorsal horn.

Abstract: The responses evoked in lumbar dorsal horn cells by both natural and electrical hind-limb skin stimulation were recorded in the spinal cord of rat pups aged 0-15 days under urethane anaesthesia. The input volley was recorded on the L4 dorsal root and consisted of two separate waves from birth. Latency and threshold measurements were consistent with these two waves being immature A (myelinated fibre) waves and C (non-myelinated fibre) waves. On the first 3 days of life background activity of cells in the dorsal horn was low and evoked discharges were sluggish. On electrical stimulation of the skin, neonatal dorsal horn cells frequently responded with only 1 or 2 impulses per input volley with long central delays of up to 20 ms. Synaptic linkage appeared weak and many cells failed to follow stimulation rates of 5 Hz. Natural skin stimulation showed that the majority of cells at days 0-3 responded to pinching the skin only. The development of responses evoked by C fibres in the dorsal horn was delayed compared to that of responses evoked by A fibres. Short and long latency responses corresponding to the early A and late C afferent input volleys could be recorded in the superficial laminae (I, II and III) of the dorsal horn from day 0, but in the deeper laminae only early short latency A responses were evoked until the age of day 7-8. After this time, a long latency C response also appeared and increased in strength with age. Convergence of low and high threshold inputs onto dorsal horn cells was rare at birth but increased gradually over the following two weeks. Receptive field areas, mapped by natural mechanical stimulation of skin, were large at birth and decreased in size with age. At birth the mean receptive field area was 14.2% of the total hind-limb area whereas at day 15 it was 3.6%. This fall in size was particularly marked in cells of the deep dorsal horn. Pinching or brushing the receptive field of many neonatal dorsal horn cells resulted in long-lasting after-discharges (30-90 s) which on days 0-3 could be more pronounced than the initial evoked response. The duration and amplitude of these responses decreased with age. Repetitive electrical skin stimulation of the receptive fields of these cells produced 'wind-up' and prolonged after-discharge. Ipsilateral, contralateral and distant inhibitory components to receptive fields were observed from day 0.(ABSTRACT TRUNCATED AT 400 WORDS)

Responsiveness and receptive field size of carp horizontal cells are reduced by prolonged darkness and dopamine.

Abstract: In the fish retina the interplexiform cells contain dopamine and provide a centrifugal pathway from the inner plexiform layer to horizontal cells of the outer plexiform layer. Dopamine application reduced the responsiveness and receptive field size of cone horizontal cells, as did a prolonged period of complete darkness. Other results suggest that the interplexiform cells may release dopamine after a prolonged period in the dark. The interplexiform-horizontal cell system may modify the strength of the antagonistic surrounds of retinal neurons as a function of time in the dark.

On the neuronal basis of figure-ground discrimination by relative motion in the visual system of the fly. 2: Figure-dectection cells, a new class of visual interneurones

Abstract: A new class of large-field tangential neurones (Figure Detection (FD-) cells) has been found and analysed in the lobula plate, the posterior part of the third visual ganglion, of the fly by combined extra-and intracellular recording as well as Lucifer Yellow injection. The FD-cells are likely to play a prominent role in figure-ground discrimination. Together with the Horizontal Cells, the output elements of the neuronal network underlying the optomotor course control reaction, they seem to be appropriate to account for the characteristic yaw torque response to relative motion. The FD-cells might thus compensate for the ldquodeficitsrdquo of the Horizontal Cells with respect to figureground discrimination (see Egelhaaf, 1985a). The FD-cells are directionally selective for either front-to-back (FD 1, FD 4) or back-to-front motion (FD 2, FD 3). Their excitatory receptive fields cover part of (FD 1, FD 2, FD 3) or the entire horizontal extent (FD 4) of the visual field of one eye. Their most important common property in the context of figureground discrimination is that they are more sensitive to relatively small objects than to spatially extended patterns. Their response to a small figure is much reduced by simultaneous large-field motion in front of the ipsi-as well as the contralateral eye. This large-field inhibition is either directionally selective or bidirectional, depending on the FD-cell under consideration. The main dendritic arborization of all FD-cells resides in the lobula plate. Their axonal projections lie in either the ipsi-or contralateral posterior optic foci and, thus, in the same area as the terminals of the Horizontal Cells. The FD-cells are, therefore, appropriate candidates for output elements of the optic lobes involved in figure-ground discrimination.

A complex-cell receptive-field model.

Abstract: The time course of the response of a single cortical neuron to counterphase-grating stimulation may vary as a function of stimulation parameters, as shown in the preceding paper (19). The poststimulus-time histograms of the response amplitudes against time are single or double peaked, and where double peaked, the two peaks are of equal or unequal amplitudes. Furthermore, the spatial-phase dependence of cortical complex-cell responses may be a function of spatial frequency, so that the receptive field appears to have linear spatial summation at some spatial frequencies and nonlinear spatial summation at others (19). In the first part of this paper, we analyze a model receptive field that displays this behavior, and in the second part experimental data are presented and analyzed with regard to the model. The model cortical receptive field in its simplest form contains (two rows) of geniculate X-cell-like, DOG (difference-of-Gaussians)-shaped, center-surround antagonistic, circular-input subunits. We propose nonlinear summation between these two subunits, by introducing a half-wave rectification stage before pooling. The model is tested for the responses it predicts for the application of counterphase-grating stimulation. This simple model predicts the appearance of three response forms as a function of counterphase-stimulation parameters. At periodic spatial frequencies the expected-response histogram has a single peak, whose amplitude has a sinusoidal dependence on spatial phase. At spatial frequencies halfway between these, the expected-response histogram has two equal peaks whose amplitudes have a full-wave rectified sinusoidal dependence on spatial phase. At all intermediate spatial frequencies the expected-response histogram has a "mixed" form; the histogram appears sometimes with one peak, sometimes with two equal peaks, and generally with two peaks of unequal amplitude, as a function of spatial phase. Null responses are expected to appear at specific spatial phases only for the periodic spatial frequencies that give "pure" response time courses as in paragraph 5 above, and not in the more common mixed response case of paragraph 6. The analysis procedure described in the preceding paper (19) is used, separating the odd and even Fourier components of the response histograms reflecting the receptive-field intrasubunit linear summation and intersubunit nonlinear summation, respectively. We propose that this model may be used as a working hypothesis for the analysis of these aspects of the various cortical receptive-field types. Experimental data are described and discussed in terms of the model.(ABSTRACT TRUNCATED AT 400 WORDS)

Induction of functional retinal projections to the somatosensory system.

Abstract: Optic axons can be induced to form permanent, retinotopic connections in the auditory (medial geniculate, MG) and somatosensory (ventrobasal, VB) nuclei of the Syrian hamster thalamus; this occurs when the principal targets of retinofugal axons are ablated in newborn hamsters and alternative terminal space is created by partial deafferentation of MG or VB1–3. The experimentally induced retinal projection to the somatosensory nucleus occurs by the stabilization of an early, normally transient projection4–5. The present study was undertaken to determine whether the anomalous, stabilized retino-VB projection is functional. Newborn hamsters were operated on to produce permanent retino-VB projections and when the animals were mature, neurophysiological recordings were made in the cortical targets of VB, the first and second somatosensory cortices (SI and SII, respectively). Visual stimulation within well-defined receptive fields reliably evoked multi-unit responses in SI and SII of operated, but not normal hamsters. The representations of the visual field in SI and SII showed a partially retinotopic organization. These results demonstrate that optic tract axons can form functional synapses in the thalamic somatosensory nucleus, and suggest that neural structures which normally process information specific to one sensory modality have the potential to mediate function for other modalities.

Termination patterns of individual X- and Y-cell axons in the visual cortex of the cat: projections to area 18, to the 17/18 border region, and to both areas 17 and 18.

Abstract: Horseradish peroxidase was injected intracellularly into single, physiologically identified X- and Y-cell geniculocortical axons that projected to area 18, to the 17/18 border region, or to both areas 17 and 18 via branching axons. The axon terminal fields in cortex were labeled anterogradely, and the cell bodies of the axons in the A-laminae, lamina C, and the medial interlaminar nucleus (MIN) of the dorsal lateral geniculate nucleus (LGN) were labeled retrogradely. The laminar projections in area 18 of eight Y-cells and one geniculate, non-Y-cell were analyzed. Most of the cells arborized densely within layer IVa and the lower 200 to 400 microns of layer III. Most provided little or no input to layer IVb or layer VI. Thus, the laminar projections of Y-cells to layer IV of area 18 were similar to those of their area 17 counterparts, although the input to layer III was greater and rose much higher in area 18 than in area 17. The terminal arbors in area 18 were two to three times larger in lateral extent than those in area 17. They spread over 2.0 to 2.8 mm2 of layer IV and occupied proportionately much greater regions of area 18 than the Y-cell arbors in area 17. This may partially account for the large receptive fields of cortical cells in area 18, and it indicates that a small region of area 18 may receive converging inputs from a relatively wide retinotopic region of the LGN. The terminal arbors were also highly asymmetric, generally being two to four times longer anteroposteriorly than mediolaterally. These asymmetric arbors may form the structural basis for the anisotropic organization of the retinotopic map in area 18. We recovered three cells (two Y, one X) whose axons arborized in the border zone between areas 17 and 18. One Y-cell axon had a receptive field located in the ipsilateral visual hemifield and it arborized in a small region restricted almost exclusively to the border zone. The other two cells had receptive fields on or adjacent to the vertical meridian, and they terminated on either side of the 17/18 border region as well as within it. Thus, geniculate afferents representing the ipsilateral hemifield or the vertical meridian appear to have different patterns of termination on and adjacent to the 17/18 border zone. Also, some X-cell input may invade area 18 in the region immediately adjacent to the border zone.(ABSTRACT TRUNCATED AT 400 WORDS)

Experience Alters the Spatial Tuning of Auditory Units in the Optic Tectum during A Sensitive Period in the Barn Owl

Abstract: The auditory spatial tuning of bimodal (auditory-visual) units in the optic tectum of the barn owl was altered by raising animals with one ear occluded. Changes in spatial tuning were assessed by comparing the location of a unit's auditory best area with that of its visual receptive field. As shown previously, auditory best areas are aligned with visual receptive fields in the tecta of normal birds (Knudsen, E. I. (1982) J. Neurosci. 2: 1177-1194). It was demonstrated in this study that, when birds were raised with one ear occluded, best areas and visual receptive fields were aligned only as long as the earplug was in place. When the earplug was removed, best areas and visual receptive fields became misaligned, indicating that a change in auditory spatial tuning had taken place during the period of occlusion. However, in a bird that received an earplug as an adult, no such alterations in auditory spatial tuning were observed; even after 1 year of monaural occlusion, auditory best areas and visual receptive fields were misaligned so long as the earplug was in place, and were aligned when the earplug was removed. These results suggest that exposure to abnormal localization cues modifies the auditory spatial tuning of tectal units only during a restricted, sensitive period early in development. After the earplug was removed from a juvenile bird that had been raised with an occluded ear, the initial misalignment between auditory best areas and visual receptive fields decreased gradually over a period of weeks. In contrast, when earplugs were removed from two adult birds that had been raised with monaural occlusions, auditory-visual misalignments persisted for as long as measurements were made, which was up to 1 year after earplug removal. These data indicate that auditory cues become permanently associated with locations in visual space during a critical period which draws to a close at about the age when the animal reaches adulthood. Horseradish peroxidase was injected into two optic tecta (in a single animal) that contained units with permanently altered auditory spatial tuning. The positions of retrogradely labeled cells in the external nucleus of the inferior colliculus (ICX) were the same as those observed in control birds (Knudsen, E. I., and P. F. Knudsen (1983) J. Comp. Neurol. 218: 187-196). Thus, the changes in spatial tuning were not due to a shift in the topographic projection from the ICX to the optic tectum.(ABSTRACT TRUNCATED AT 400 WORDS)

Capabilities of monkey cortical cells in spatial-resolution tasks

Abstract: The performance of individual neurons in monkey striate cortex has been examined in three spatial-resolution tasks by making microelectrode recordings from single cells in anaesthetized, paralyzed animals. The statistical reliability of responses from cells was used to estimate threshold levels of performance. For each task (resolution acuity for high-contrast gratings, discrimination of gratings varying in spatial frequency, and localization ability, i.e., discrimination of spatial phase), performance approaching psychophysical thresholds was obtained from single cortical cells. The receptive-field organization underlying localization performance was examined in detail by the use of a linear model that relates localization ability to the sensitivity of the receptive field to luminance contrast. Calculations from this model agree well with direct measurements of localization performance and are comparable with psychophysical measurements of hyperacuity. Though it has been suggested that cells with nonoriented receptive fields in cortical layer ivcβ may be responsible for recovering fine-grain spatial information, our calculations indicate that these cells are poorer at localization than many other cells in the cortex.

A comparison of visual responses of cat lateral geniculate nucleus neurones with those of ganglion cells afferent to them.

Abstract: We compared visual responses of cat lateral geniculate nucleus (l.g.n.) neurones with those of retinal ganglion cells providing their afferent inputs. Quantitative studies were made on twenty such pairs; eight X on-centre, seven Y on-centre, two X off-centre and three Y off-centre pairs. Receptive field centre locations of cell pairs with correlated activities were very closely superimposed, having a mean centre displacement of 1.6 minutes of arc for X cells and 11 minutes of arc for Y cells. With flashed spots and annuli, responses of l.g.n. cells were almost always smaller than those of their retinal afferents, with peaks and troughs in ganglion cell responses being faithfully followed in the geniculate neurones. This is consistent with almost all impulses from the l.g.n. cell being triggered by the afferent feeding its centre. With spots of different sizes and contrasts, modulation of responses by l.g.n. inhibition was obvious, but effects were complex. With moving bright-bar stimuli, although response histograms were clearly reshaped to some extent in the l.g.n., peak firing rates under different stimulus conditions were often merely attenuated by a constant factor for most l.g.n. cells in comparison with their retinal inputs. For velocity tuning curves, a few cell pairs showed selective attenuation at high speeds, while others showed it at low speeds. All the latter group appeared to have more than one major excitatory afferent. These changes in velocity tuning occurred across the X/Y classification, so that differences in velocity preference of the X and Y systems is more blurred in the l.g.n. than in the retina.

Direction-selective single units in the nucleus lentiformis mesencephali of the pigeon (Columba livia).

Abstract: The receptive field properties of single units within the nucleus lentiformis mesencephali (LM) of the pigeon were studied using electrophysiological methods. Previous studies have suggested that the avian LM may be homologous to the nucleus of the optic tract (NOT) in mammals. Single units in the pigeon LM are similar to mammalian NOT units in that they are direction-selective, mostly for horizontal directions, velocity-selective, have large visual receptive fields and respond preferentially to large stimuli with many visual contrasts. In contrast to most reports of NOT units of mammals, more than half of pigeon LM units prefer high velocities (>10°/s), a large proportion (0.37) prefer non-horizontal directions, and receptive fields that are retinotopically arranged within the LM. The response properties of pigeon LM units are compared to the response properties of units within the accessory optic nucleus (the nucleus of the basal optic root or nBOR). In the avian brain, nBOR neurons respond at low velocities (0.5−5°/s) and respond predominantly to vertical stimulus movement whereas LM units respond over a broader range of velocities (0.2−80°/s) and respond predominantly to horizontal movements. Thus, the LM and nBOR may play different roles in the control of compensatory eye movements.

Role of the cat substantia nigra pars reticulata in eye and head movements. I. Neural activity.

Abstract: 1. Single unit activity was recorded in the Substantia Nigra pars reticulata (SNpr) of cats trained to orient their gaze toward visual and/or auditory targets. 2. Cells in the SNpr have a steady high rate of spontaneous activity ranging from 35 to 120 spikes per second. The neurons respond to sensory stimuli or in relation to saccadic eye movements with a decrease or a cut-off of the spontaneous discharge. 3. Among 109 cells recorded in the SNPR 60 were responsive to visual stimuli (mean latency = 118 ms). Most of the receptive fields which were plotted were large encompassing part of the ipsilateral field. 4. Thirty nine (39) cells were responsive to auditory stimuli (mean latency = 81 ms). A majority of these cells showed a better response for stimuli located in the contralateral hemifield. 5. In a few cells, the sensory responses were modulated by the subsequent orienting behavior of the animals. 6. Thirty one (31) cells showed a response in relation to saccades. These units typically stopped discharging between 50 and 300 ms prior to the onset of the saccade. 39% of these units also responded in relation to spontaneous saccades in the dark. 61% of the saccadic cells also responded to sensory stimuli in the absence of saccades. Six (6) cells were found to respond to active head movements. 7. These results are discussed in the framework of the role that the basal ganglia might have in the selection of the sensory stimuli that trigger orienting behaviors.

The mapping of visual space onto foveal striate cortex in the macaque monkey

Abstract: Visuotopic maps of foveal striate cortex have been obtained by means of single cell recordings from four hemispheres in two awake, behaving macaque monkeys. The numbers of successful separate striate penetration sites in the four hemispheres were 42, 58, 81, and 61, for a total of 242. The resolution of the maps is 10 min of visual angle, nearly an order of magnitude finer than previous maps. No striate receptive field center was found more than 5 min into the ipsilateral visual field. The four maps were sufficiently compatible with one another that they could be combined into one. There are only minor magnification differences between the right and left hemispheres and between the upper and lower quadrants. There is a vertical/horizontal magnification anisotropy of about 1.5:1 in central foveal cortex (0 to 20 min). The composite map can be approximated by the complex logarithmic equation, w = 7.7 * ln (x + iy + 0.33), where w is expressed in millimeters and x and y are expressed in degrees.

Development of spatial receptive-field organization and orientation selectivity in kitten striate cortex

Abstract: The functional organization of the receptive field of neurons in striate cortex of kittens from 8 days to 3 mo of age was studied by extracellular recordings. A quantitative dual-stimulus technique was used, which allowed for analysis of both enhancement and suppression zones in the receptive field. Furthermore the development of orientation selectivity was studied quantitatively in the same cells. Already in the youngest kittens the receptive fields were spatially organized like adult fields, with a central zone and adjacent flanks that responded in opposite manner to the light stimulus. The relative suppression in the subzones was as strong as in adult cells. Both simple and complex cells were found from 8 days. The receptive fields were like magnified adult fields. The width of the dominant discharge-field zone and the distance between the positions giving maximum discharge and maximum suppression decreased with age in the same proportions. The decrease could be explained by a corresponding decrease of the receptive-field-center size of retinal ganglion cells. Forty percent of the cells were orientation selective before 2 wk, and the fraction increased to 94% at 4 wk. Cells whose responses could be attenuated to at least half of the maximal response by changes of slit orientation were termed orientation selective. The half-width of the orientation-tuning curves narrowed during the first 5 wk, and this change was most marked in simple cells. The ability of the cells to discriminate between orientations in statistical terms was weak in the youngest kittens due to a large response variability, and showed a more pronounced development than the half-width did. The orientation-tuning curves were fitted by an exponential function, which showed the shape to be adultlike in all age groups. Two kittens were dark reared until recording at 1 mo of age. The spatial receptive-field organization and the orientation selectivity in these kittens were similar to normal-reared kittens at 1 mo. The responsivity of the cells of the dark-reared kittens was lower, and the latency before firing was longer than in the normal-reared kittens of the same age, and these response properties were more similar to those in 1- to 2-wk-old normal kittens. Our results indicate that the spatial organization of the receptive field is innate in most cells and that visual experience is unnecessary for the organization to be maintained and for the receptive-field width to mature during the first month postnatally.(ABSTRACT TRUNCATED AT 400 WORDS)