Response histogram shapes and tuning curves: The predicted responses of several cortical cell types to drifting gratings stimuli

The responses to visual stimuli of simple cortical cells show linear spatial summation within and between their receptive field subunits. Complex cortical cells do not show this linearity. We analyzed the simulated responses to drifting sinusoidal grating stimuli of simple and of several types of complex cells. The complex cells, whose responses are seen to be half-wave rectified before pooling, have receptive fields consisting of two or more DOG (difference-of-Gaussians) shaped subunits. In both cases of stimulation by contrast-reversal gratings or drifting gratings, the cells' response as a function of spatial frequency is affected by the subunit distances 2 and the stimulation frequency . Furthermore, an increased number of subunits (a larger receptive field) yields a narrower peak tuning curve with decreased modulation depth for many of the spatial frequencies. The average and the peak response tuning curves are compared for the different receptive field types.     

Single-cell responses in the ectostriatum of the zebra finch

The responses of single cells to computer-generated spots, bars, gratings, and motion-in-depth stimuli were studied in the ectostriatum and the adjacent neostriatum of the zebra finch, Taeniopygia guttata. No differences in neuronal properties could be detected between ectostriatum and neostriatum. The receptive fields of ectostriatal neurons are large, often extending over the entire visual field of the contralateral eye, and have oddly defined borders. The centers of the receptive fields, located in the foveal region, generally yielded better responses than the periphery, and exhibited different subdivisions. Neurons responded selectively to moving bars, preferring those moving parallel to their longest axis. An SDO (sensitivity, direction, orientation) analysis of responses to sinusoidal gratings showed that all orientations were equally represented by ectostriatal neurons, while there was a slight preference for forward and upward movements. The neurons also showed preferences for gratings of a particular spatial frequency, and responded vigorously to stimuli moving towards the eye (looming). Our results indicate that the ectostriatum is involved in both detecting displacement of the surround and in stimulus identification. By comparison with results obtained in the extrastriate cortex of mammals, it is concluded that the homology of the ectostriatum with the extrastriate cortex of mammals, which was proposed on the basis of hodological findings, is supported by our study.Abbreviations Di index of directionality - HW HH half-width at half-height - PLLS posterolateral lateral suprasylvian cortex - PMLS posterior medial lateral suprasylvian area - PSTH poststimulus time histogram - SDO sensitivity, direction, orientation     

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

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

The responses to illusory contours of neurons in cortex areas 17 and 18 of the cats

Responses to illusory contours (ICs) were sampled from neurons in cortical areas 17 and 18 of the anesthetized cats. For ICs sensitive cells, the differences of receptive field properties were compared when ICs and real contour stimuli were applied. Two hundred orientation or direction selective cells were studied. We find that about 42 percent of these cells were the ICs sensitive cells. Although their orientation or direction tuning curves to ICs bar and real bars were similar, the response modes (especially latency and time course) were different. The cells’ responses to ICs were independent of the spatial phases of sinusoidal gratings, which composed the ICs. The cells’ optimal spatial frequency to composing gratings the ICs was much higher than the one to moving gratings. Therefore, these cells really responded to the ICs rather than the line ends of composing gratings. For some kinds of velocity-tuning cells, the optimal velocity to moving ICs bar was much lower than the optimal velocity to moving bars. The present results demonstrate that some cells in areas 17 and 18 of cats have the ability to respond to ICs and have different response properties of the receptive fields to ICs and luminance boundaries via different neural mechanisms.     

Binaural disparity cues available to the barn owl for sound localization

1. Bilateral recording of cochlear potentials was used to measure the variations in interaural time differences (ITDs) and interaural intensity differences (IIDs) as a free-field auditory stimulus was moved to different positions around a barn owl's head. 2. ITD varied smoothly with stimulus azimuth across a broad frequency range. 3. ITD varied minimally with stimulus elevation, except at extreme angles from the horizontal. 4. IID varied with both stimulus elevation and stimulus azimuth. Lower frequencies were more sensitive to variations in azimuth, whereas higher frequencies were more sensitive to variations in elevation. 5. The loci of spatial coordinates that form iso-IID contours and iso-ITD contours form a non-orthogonal grid that relates binaural disparity cues to sound location.     

Side peak suppression in responses of an across-frequency integration model to stimuli of varying bandwidth as demonstrated analytically and by implementation

Multiplication-like sound localization models are subjected to phase ambiguities for high-frequency tonal stimuli as multiplication creates several equivalent response peaks in tuning curves. By increasing the bandwidth of the stimulus, phase ambiguities can be reduced, which is often referred to as side peak suppression. In this study we present a Jeffress-based sound localization model, and determine side peak suppression analytically. The results were verified with an implementation of the same model, and compared to physiological data of barn owls. Three types of stimuli were analyzed: pure-tone stimuli, two-tone complexes with varying frequency distances, and noise signals with variable bandwidths. As an additional parameter we also determined the half-width of the main response peak to examine the scaling of tuning curves in azimuth. Results showed that side peak suppression did not only depend on bandwidth, but also on the center frequency and the distance of the side peak to the main response peak. In particular, the analytical model predicted that side peak suppression is a function of relative bandwidth, whereas half-width is inversely proportional to center frequency, with a proportionality factor depending on relative bandwidth. The analytical approach and the implementation yielded equivalent tuning curves (deviation < 1 %). Moreover, the electrophysiological data recorded in barn owls closely matched the predicted tuning curves.     

初级视皮层神经元响应反应与撤反应的朝向选择性

朝向选择性是初级视皮层(17区或V1)神经元的基本性质,在图形感知中起着关键作用.同时这些神经元对于持续时间大于100 ms的视觉刺激具有清晰的响应反应(Onset responses)和撤反应(Offset responses).以往的研究只关注响应反应的朝向选择性,而忽视了对撤反应的朝向选择性研究.我们比较了响应与撤反应的朝向调谐性质,大多数细胞的撤反应与响应反应基本上具有相似的最优朝向,而撤反应的朝向调谐宽度有窄于响应反应的趋势,撤反应的最优延迟普遍滞后于响应反应的最优延迟.撤反应的朝向选择性略强于响应反应和具有显著长的反应延迟提示,皮层内的反馈输入可能在形成撤反应的朝向选择性中起着作用.本研究揭示了撤反应的朝向选择性在刺激朝向的连续表征和主体在形状知觉的后期对朝向的精细区分中起着作用.     

Sensitivity to spectral interaural intensity difference cues in space-specific neurons of the barn owl

Barn owls use interaural intensity differences to localize sounds in the vertical plane. At a given elevation the magnitude of the interaural intensity difference cue varies with frequency, creating an interaural intensity difference spectrum of cues which is characteristic of that direction. To test whether space-specific cells are sensitive to spectral interaural intensity difference cues, pure-tone interaural intensity difference tuning curves were taken at multiple different frequencies for single neurons in the external nucleus of the inferior colliculus. For a given neuron, the interaural intensity differences eliciting the maximum response (the best interaural intensity differences) changed with the frequency of the stimulus by an average maximal difference of 9.4±6.2 dB. The resulting spectral patterns of these neurally preferred interaural intensity differences exhibited a high degree of similarity to the acoustic interaural intensity difference spectra characteristic of restricted regions in space. Compared to stimuli whose interaural intensity difference spectra matched the preferred spectra, stimuli with inverted spectra elicited a smaller response, showing that space-specific neurons are sensitive to the shape of the spectrum. The underlying mechanism is an inhibition for frequency-specific interaural intensity differences which differ from the preferred spectral pattern. Collectively, these data show that space-specific neurons are sensitive to spectral interaural intensity difference cues and support the idea that behaving barn owls use such cues to precisely localize sounds.Abbreviations ABI average binaural intensity - HRTF head-related transfer function - ICx external nucleus of the inferior colliculus - IID interaural intensity difference - ITD interaural time difference - OT optic tectum - RMS root mean square - VLVp nucleus ventralis lemnisci laterale, pars posterior     

Two spatio-temporal filters in human vision

1. We have studied visual detection of a circular target moving across a spatially and/or temporally modulated background. Illumination, I _t , for threshold detection of the target has been measured as a function of background modulation frequency and changes in I _t associated with background modulation provide a means of determining the frequency response characteristics of visual channels. 2. Temporal frequency responses obtained with temporally modulated, spatially uniform backgrounds have pass-band characteristics and the temporal frequency for peak response increases with increase in mean background illumination. These temporal frequency responses resemble those of the de Lange (1954) filter, but the latter incorporates the incremental thresholds for steady backgrounds. 3. The amplitude of this temporal response saturates at low (40%) background modulation, decreases to zero as the target velocity falls to zero, and is maximum for a circular target of diameter 2°. 4. The spatial characteristics of this temporal filter were measured with a background field consisting of alternate steady and flickering bars. The resulting spatial frequency curve peaks at 1 cycle deg^-1 for all background illuminations and is independent of the background grating orientation. This spatial response differs significantly from the IMG spatial functions observed with a background grating (Barbur and Ruddock, 1980). 5. The spatial and temporal responses reviewed above exhibit similar parametric variations and we therefore associate them with a single spatiotemporal filter, ST2. 6. A second temporal response, with low-pass frequency characteristics, was observed with a background field consisting of two matched gratings, presented in spatial and temporal antiphase. This response has parametric properties similar to those of the IMG spatial response described previously by Barbur and Ruddock (1980), thus we associated the two sets of data with a single spatio-temporal filter, ST1. 7. We show that the ST2 responses can be obtained by combining ST1 responses, and we present a network incorporating the two filters. 8. We review other psychophysical studies which imply the activity of two spatio-temporal filters with properties of the kind revealed in our studies. We argue that filter ST1 has properties equivalent to those of X-type and filter ST2 has properties equivalent to those of Y-type electrophysiological mechanisms.     

Mass-action view of single-cell responses to stimulation of the receptive field and/or beyond: exemplification with data from the rabbit primary visual cortex

Whereas single cells in the visual cortex prefer moving light bars, mass-action responses are evoked better by diffuse luminance changes. This discrepancy was investigated by quantitatively comparing the the response properties of individual cells with those a representative group of cells. The latter responses were derived from the single-cell responses, which were obtained from recording in the rabbit. These quantitative estimates of mean responses resolve the discrepancy between the single-cell domain and the mass-action domain: from the single-cell point of view, a properly oriented moving-bar stimulus is much more effective than a diffuse-light stimulus. The corresponding mass-action response to one common moving-bar stimulus, however, is as small as the mean response to a diffuse-light stimulus (which may even be presented at retinotopically non-corresponding sites). The peak intensities of these mass responses are even much stronger with the diffuse-light stimuli. The same conclusions are valid for the cat, as could be verified from published data. The restrictions of the local receptive field concept that may be implied by the mass-action view of cortical activity and the potential functional relevance of mass activities area discussed.     

On the variety of spatial frequency selectivities shown by neurons in area 17 of the cat

The amplitudes of the responses of over 300 neurons in area 17 of the cat were examined as a function of the spatial frequency of moving sinusoidal gratings. The optimal spatial frequency and the bandwidth of the tuning curves were determined. The bandwidth varied considerably from neuron to neuron. Neurons optimally responsive to high spatial frequencies tended to have narrower tuning curves than those responsive to lower frequencies. Neurons with narrow spatial frequency tuning curves also tended to have narrow orientation tuning curves. These observations suggest that linear spatial summation tends to occur over a relatively constant area of visual field despite marked differences in each neuron's optimal spatial frequency, a prediction of one model of visual analysis. There was little difference in either the optimal spatial frequencies or the bandwidths of tuning for different functional classes of neuron. Neurons with broad tuning curves tended to be restricted to lamina IV and its environs, being concentrated in the deep part of lamina II-III and the upper part of lamina IV ab. Neurons with very low optimal spatial frequencies were uncommon and tended to be found either at the border of laminae II-III and IV or in lamina V. These laminar distributions are discussed with respect to the laminar differences in the projection of l.g.m. X- and Y-cells to the visual cortex.     

Neural coding of 3D features of objects for hand action in the parietal cortex of the monkey.

In our previous studies of hand manipulation task-related neurons, we found many neurons of the parietal association cortex which responded to the sight of three-dimensional (3D) objects. Most of the task-related neurons in the AIP area (the lateral bank of the anterior intraparietal sulcus) were visually responsive and half of them responded to objects for manipulation. Most of these neurons were selective for the 3D features of the objects. More recently, we have found binocular visual neurons in the lateral bank of the caudal intraparietal sulcus (c-IPS area) that preferentially respond to a luminous bar or place at a particular orientation in space. We studied the responses of axis-orientation selective (AOS) neurons and surface-orientation selective (SOS) neurons in this area with stimuli presented on a 3D computer graphics display. The AOS neurons showed a stronger response to elongated stimuli and showed tuning to the orientation of the longitudinal axis. Many of them preferred a tilted stimulus in depth and appeared to be sensitive to orientation disparity and/or width disparity. The SOS neurons showed a stronger response to a flat than to an elongated stimulus and showed tuning to the 3D orientation of the surface. Their responses increased with the width or length of the stimulus. A considerable number of SOS neurons responded to a square in a random dot stereogram and were tuned to orientation in depth, suggesting their sensitivity to the gradient of disparity. We also found several SOS neurons that responded to a square with tilted or slanted contours, suggesting their sensitivity to orientation disparity and/or width disparity. Area c-IPS is likely to send visual signals of the 3D features of an object to area AIP for the visual guidance of hand actions.     

Response properties of visual interneurons to motion stimuli in the praying mantis,Tenodera aridifolia

Intracellular responses of motion-sensitive visual interneurons were recorded from the lobula complex of the mantis, Tenodera aridifolia. The interneurons were divided into four classes according to the response polarity, spatial tuning, and directional selectivity. Neurons of the first class had small, medium, or large receptive fields and showed a strong excitation in response to a small-field motion such as a small square moving in any direction (SF neurons). The second class neurons showed non-directionally selective responses: an excitation to a large-field motion of gratings in any direction (ND neurons). Most ND neurons had small or medium-size receptive fields. Neurons of the third class had large receptive fields and exhibited directionally selective responses: an excitation to a large-field motion of gratings in preferred direction and an inhibition to a motion in opposite, null direction (DS neurons). The last class neurons had small receptive fields and showed inhibitory responses to a moving square and gratings (I neurons). The functional roles of these neurons in prey recognition and optomotor response were discussed.     

An automated paradigm for Drosophila visual psychophysics

Background

Mutations that cause learning and memory defects in Drosophila melanogaster have been found to also compromise visual responsiveness and attention. A better understanding of attention-like defects in such Drosophila mutants therefore requires a more detailed characterization of visual responsiveness across a range of visual parameters.

Methodology/Principal Findings

We designed an automated behavioral paradigm for efficiently dissecting visual responsiveness in Drosophila. Populations of flies walk through multiplexed serial choice mazes while being exposed to moving visuals displayed on computer monitors, and infra-red fly counters at the end of each maze automatically score the responsiveness of a strain. To test our new design, we performed a detailed comparison between wild-type flies and a learning and memory mutant, dunce ¹. We first confirmed that the learning mutant dunce ¹ displays increased responsiveness to a black/green moving grating compared to wild type in this new design. We then extended this result to explore responses to a wide range of psychophysical parameters for moving gratings (e.g., luminosity, contrast, spatial frequency, velocity) as well as to a different stimulus, moving dots. Finally, we combined these visuals (gratings versus dots) in competition to investigate how dunce ¹ and wild-type flies respond to more complex and conflicting motion effects.

Conclusions/Significance

We found that dunce ¹ responds more strongly than wild type to high contrast and highly structured motion. This effect was found for simple gratings, dots, and combinations of both stimuli presented in competition.     

Response characteristics of wide-field non-habituating non-directional motion detecting neurons in the optic lobe of the locust,Locusta migratoria

1. Extracellular recordings from wide-field nonhabituating non-directional (ND) motion detecting neurons in the second optic chiasma of the locust Locusta migratoria are presented. The responses to various types of stepwise moving spot and bar stimuli were monitored (Fig. 1)
2. Stepwise motion in all directions elicited bursts of spikes. The response is inhibited at stimulus velocities above 5°/s. At velocities above 10°/s the ND neurons are slightly more sensitive to motion in the horizontal direction than to motion in the vertical direction (Fig. 2). The ND cells have a preference for small moving stimuli (Fig. 3).
3. The motion response has two peaks. The latency of the second peak depends on stimulus size and stimulus velocity. Increasing the height from 0.1 to 23.5° of a 5°/s moving bar results in a lowering of this latency time from 176 to 130 ms (Fig. 4). When the velocity from a single 0.1° spot is increased from 1 to 16°/s, the latency decreases from 282 to 180 ms (Figs. 5–6).
4. A change-of-direction sensitivity is displayed. Stepwise motion in one particular direction produces a continuous burst of spike discharges. Reversal or change in direction leads to an inhibition of the response (Fig. 7).
5. It shows that non-directional motion perception of the wide-field ND cells can simply be explained by combining self-and lateral inhibition.

     

Coding of sound location and frequency in the auditory midbrain of diurnal birds of prey,families accipitridae and falconidae

Summary The coding of sound frequency and location in the avian auditory midbrain nucleus (nMLD) was examined in three diurnal raptors: the brown falcon (Falco berigora), the swamp harrier (Circus aeruginosus) and the brown goshawk (Accipiter fasciatus). Previously this nucleus has been studied with free field stimuli in only one other species, the barn owl (Tyto alba).We found some parallels between the organisation of nMLD in the diurnal raptors and that reported in the barn owl in that the central region of nMLD was tonotopically organised and contained cells that did not encode location, and the lateral region (nMLDl) contained cells which were sensitive to stimulus position. However, unlike the barn owl, which has units with circumscribed receptive fields, cells sensitive to stimulus location had large receptive fields which were restricted in azimuth but not in elevation (hemifield units). Such cells could not provide an acoustic space map in which both azimuthal and elevational dimensions were represented, but there was a tendency for units with contralateral borders to be found superficially, and those with ipsilateral borders to be found deep, in nMLDl. Hemifield units displayed receptive field properties consistent with the directional properties of the tympana in the presence of sound transmission through the interaural canal, if there is a central mechanism which is sensitive to interaural intensity differences.Abbreviations nMLD nucleus mesencephalicus lateralis pars dorsalis - SPL sound pressure level re 20 Pa - nMLDl lateral region of nMLD - ICC central nucleus of the inferior colliculus - ICX external nucleus of the inferior colliculus - IID interaural intensity difference - EI excitatory inhibitory     

Perceptual “Read-Out” of Conjoined Direction and Disparity Maps in Extrastriate Area MT

Cortical neurons are frequently tuned to several stimulus dimensions, and many cortical areas contain intercalated maps of multiple variables. Relatively little is known about how information is “read out” of these multidimensional maps. For example, how does an organism extract information relevant to the task at hand from neurons that are also tuned to other, irrelevant stimulus dimensions? We addressed this question by employing microstimulation techniques to examine the contribution of disparity-tuned neurons in the middle temporal (MT) visual area to performance on a direction discrimination task. Most MT neurons are tuned to both binocular disparity and the direction of stimulus motion, and MT contains topographic maps of both parameters. We assessed the effect of microstimulation on direction judgments after first characterizing the disparity tuning of each stimulation site. Although the disparity of the stimulus was irrelevant to the required task, we found that microstimulation effects were strongly modulated by the disparity tuning of the stimulated neurons. For two of three monkeys, microstimulation of nondisparity-selective sites produced large biases in direction judgments, whereas stimulation of disparity-selective sites had little or no effect. The binocular disparity was optimized for each stimulation site, and our result could not be explained by variations in direction tuning, response strength, or any other tuning property that we examined. When microstimulation of a disparity-tuned site did affect direction judgments, the effects tended to be stronger at the preferred disparity of a stimulation site than at the nonpreferred disparity, indicating that monkeys can selectively monitor direction columns that are best tuned to an appropriate conjunction of parameters. We conclude that the contribution of neurons to behavior can depend strongly upon tuning to stimulus dimensions that appear to be irrelevant to the current task, and we suggest that these findings are best explained in terms of the strategy used by animals to perform the task.     

The responses to illusory contours of neurons in cortex areas 17 and 18 of the cats

Responses to illusory contours (ICs) were sampled from neurons in cortical areas 17 and 18 of the anesthetized cats. For ICs sensitive cells, the differences of receptive field properties were compared when ICs and real contour stimuli were applied. Two hundred orientation or direction selective cells were studied. We find that about 42 percent of these cells were the ICs sensitive cells. Although their orientation or direction tuning curves to ICs bar and real bars were similar, the response modes (especially latency and time course) were different. The cells' responses to ICs were independent of the spatial phases of sinusoidal gratings, which composed the ICs. The cells' optimal spatial frequency to composing gratings the ICs was much higher than the one to moving gratings. Therefore, these cells really responded to the ICs rather than the line ends of composing gratings. For some kinds of velocity-tuning cells, the optimal velocity to moving ICs bar was much lower than the optimal velocity to moving     

Perceptual “Read-Out” of Conjoined Direction and Disparity Maps in Extrastriate Area MT

Cortical neurons are frequently tuned to several stimulus dimensions, and many cortical areas contain intercalated maps of multiple variables. Relatively little is known about how information is “read out” of these multidimensional maps. For example, how does an organism extract information relevant to the task at hand from neurons that are also tuned to other, irrelevant stimulus dimensions? We addressed this question by employing microstimulation techniques to examine the contribution of disparity-tuned neurons in the middle temporal (MT) visual area to performance on a direction discrimination task. Most MT neurons are tuned to both binocular disparity and the direction of stimulus motion, and MT contains topographic maps of both parameters. We assessed the effect of microstimulation on direction judgments after first characterizing the disparity tuning of each stimulation site. Although the disparity of the stimulus was irrelevant to the required task, we found that microstimulation effects were strongly modulated by the disparity tuning of the stimulated neurons. For two of three monkeys, microstimulation of nondisparity-selective sites produced large biases in direction judgments, whereas stimulation of disparity-selective sites had little or no effect. The binocular disparity was optimized for each stimulation site, and our result could not be explained by variations in direction tuning, response strength, or any other tuning property that we examined. When microstimulation of a disparity-tuned site did affect direction judgments, the effects tended to be stronger at the preferred disparity of a stimulation site than at the nonpreferred disparity, indicating that monkeys can selectively monitor direction columns that are best tuned to an appropriate conjunction of parameters. We conclude that the contribution of neurons to behavior can depend strongly upon tuning to stimulus dimensions that appear to be irrelevant to the current task, and we suggest that these findings are best explained in terms of the strategy used by animals to perform the task.