Temporal Tuning Properties Research Articles

A chromatic feature detector in the retina signals visual context changes.

The retina transforms patterns of light into visual feature representations supporting behaviour. These representations are distributed across various types of retinal ganglion cells (RGCs), whose spatial and temporal tuning properties have been studied extensively in many model organisms, including the mouse. However, it has been difficult to link the potentially nonlinear retinal transformations of natural visual inputs to specific ethological purposes. Here, we discover a nonlinear selectivity to chromatic contrast in an RGC type that allows the detection of changes in visual context. We trained a convolutional neural network (CNN) model on large-scale functional recordings of RGC responses to natural mouse movies, and then used this model to search in silico for stimuli that maximally excite distinct types of RGCs. This procedure predicted centre colour opponency in transient suppressed-by-contrast (tSbC) RGCs, a cell type whose function is being debated. We confirmed experimentally that these cells indeed responded very selectively to Green-OFF, UV-ON contrasts. This type of chromatic contrast was characteristic of transitions from ground to sky in the visual scene, as might be elicited by head or eye movements across the horizon. Because tSbC cells performed best among all RGC types at reliably detecting these transitions, we suggest a role for this RGC type in providing contextual information (i.e. sky or ground) necessary for the selection of appropriate behavioural responses to other stimuli, such as looming objects. Our work showcases how a combination of experiments with natural stimuli and computational modelling allows discovering novel types of stimulus selectivity and identifying their potential ethological relevance.

Long-term recordings from area V4 neurons and an accurately-predicting deep convolutional energy model reveal spatial, chromatic and temporal tuning properties under naturalistic conditions

Long-term recordings from area V4 neurons and an accurately-predicting deep convolutional energy model reveal spatial, chromatic and temporal tuning properties under naturalistic conditions

An uncertainty principle for neural coding: Conjugate representations of position and velocity are mapped onto firing rates and co-firing rates of neural spike trains.

The hippocampal system contains neural populations that encode an animal's position and velocity as it navigates through space. Here, we show that such populations can embed two codes within their spike trains: a firing rate code () conveyed by within‐cell spike intervals, and a co‐firing rate code (R˙) conveyed by between‐cell spike intervals. These two codes behave as conjugates of one another, obeying an analog of the uncertainty principle from physics: information conveyed in comes at the expense of information in R˙, and vice versa. An exception to this trade‐off occurs when spike trains encode a pair of conjugate variables, such as position and velocity, which do not compete for capacity across and R˙. To illustrate this, we describe two biologically inspired methods for decoding and R˙, referred to as sigma and sigma‐chi decoding, respectively. Simulations of head direction and grid cells show that if firing rates are tuned for position (but not velocity), then position is recovered by sigma decoding, whereas velocity is recovered by sigma‐chi decoding. Conversely, simulations of oscillatory interference among theta‐modulated “speed cells” show that if co‐firing rates are tuned for position (but not velocity), then position is recovered by sigma‐chi decoding, whereas velocity is recovered by sigma decoding. Between these two extremes, information about both variables can be distributed across both channels, and partially recovered by both decoders. These results suggest that populations with different spatial and temporal tuning properties—such as speed versus grid cells—might not encode different information, but rather, distribute similar information about position and velocity in different ways across and R˙. Such conjugate coding of position and velocity may influence how hippocampal populations are interconnected to form functional circuits, and how biological neurons integrate their inputs to decode information from firing rates and spike correlations.

Sensory cortex is optimized for prediction of future input.

Neurons in sensory cortex are tuned to diverse features in natural scenes. But what determines which features neurons become selective to? Here we explore the idea that neuronal selectivity is optimized to represent features in the recent sensory past that best predict immediate future inputs. We tested this hypothesis using simple feedforward neural networks, which were trained to predict the next few moments of video or audio in clips of natural scenes. The networks developed receptive fields that closely matched those of real cortical neurons in different mammalian species, including the oriented spatial tuning of primary visual cortex, the frequency selectivity of primary auditory cortex and, most notably, their temporal tuning properties. Furthermore, the better a network predicted future inputs the more closely its receptive fields resembled those in the brain. This suggests that sensory processing is optimized to extract those features with the most capacity to predict future input.

Roles for Coincidence Detection in Coding Amplitude-Modulated Sounds.

Many sensory neurons encode temporal information by detecting coincident arrivals of synaptic inputs. In the mammalian auditory brainstem, binaural neurons of the medial superior olive (MSO) are known to act as coincidence detectors, whereas in the lateral superior olive (LSO) roles of coincidence detection have remained unclear. LSO neurons receive excitatory and inhibitory inputs driven by ipsilateral and contralateral acoustic stimuli, respectively, and vary their output spike rates according to interaural level differences. In addition, LSO neurons are also sensitive to binaural phase differences of low-frequency tones and envelopes of amplitude-modulated (AM) sounds. Previous physiological recordings in vivo found considerable variations in monaural AM-tuning across neurons. To investigate the underlying mechanisms of the observed temporal tuning properties of LSO and their sources of variability, we used a simple coincidence counting model and examined how specific parameters of coincidence detection affect monaural and binaural AM coding. Spike rates and phase-locking of evoked excitatory and spontaneous inhibitory inputs had only minor effects on LSO output to monaural AM inputs. In contrast, the coincidence threshold of the model neuron affected both the overall spike rates and the half-peak positions of the AM-tuning curve, whereas the width of the coincidence window merely influenced the output spike rates. The duration of the refractory period affected only the low-frequency portion of the monaural AM-tuning curve. Unlike monaural AM coding, temporal factors, such as the coincidence window and the effective duration of inhibition, played a major role in determining the trough positions of simulated binaural phase-response curves. In addition, empirically-observed level-dependence of binaural phase-coding was reproduced in the framework of our minimalistic coincidence counting model. These modeling results suggest that coincidence detection of excitatory and inhibitory synaptic inputs is essential for LSO neurons to encode both monaural and binaural AM sounds.

Spectrotemporal response properties of core auditory cortex neurons in awake monkey.

So far, most studies of core auditory cortex (AC) have characterized the spectral and temporal tuning properties of cells in non-awake, anesthetized preparations. As experiments in awake animals are scarce, we here used dynamic spectral-temporal broadband ripples to study the properties of the spectrotemporal receptive fields (STRFs) of AC cells in awake monkeys. We show that AC neurons were typically most sensitive to low ripple densities (spectral) and low velocities (temporal), and that most cells were not selective for a particular spectrotemporal sweep direction. A substantial proportion of neurons preferred amplitude-modulated sounds (at zero ripple density) to dynamic ripples (at non-zero densities). The vast majority (>93%) of modulation transfer functions were separable with respect to spectral and temporal modulations, indicating that time and spectrum are independently processed in AC neurons. We also analyzed the linear predictability of AC responses to natural vocalizations on the basis of the STRF. We discuss our findings in the light of results obtained from the monkey midbrain inferior colliculus by comparing the spectrotemporal tuning properties and linear predictability of these two important auditory stages.

Temporal Tuning Properties along the Human Ventral Visual Stream

Both our environment and our behavior contain many spatiotemporal regularities. Preferential and differential tuning of neural populations to these regularities can be demonstrated by assessing rate dependence of neural responses evoked during continuous periodic stimulation. Here, we used functional magnetic resonance imaging to measure regional variations of temporal sensitivity along the human ventral visual stream. By alternating one face and one house stimulus, we combined sufficient low-level signal modulation with changes in semantic meaning and could therefore drive all tiers of visual cortex strongly enough to assess rate dependence. We found several dissociations between early visual cortex and middle- and higher-tier regions. First, there was a progressive slowing down of stimulation rates yielding peak responses along the ventral visual stream. This finding shows the width of temporal integration windows to increase at higher hierarchical levels. Next, for fixed rates, early but not higher visual cortex responses additionally depended on the length of stimulus exposure, which may indicate increased persistence of responses to short stimuli at higher hierarchical levels. Finally, attention, which was recruited by an incidental task, interacted with stimulation rate and shifted tuning peaks toward lower frequencies. Together, these findings quantify neural response properties that are likely to be operational during natural vision and that provide putative neurofunctional substrates of mechanisms that are relevant in several psychophysical phenomena as masking and the attentional blink. Moreover, they illustrate temporal constraints for translating the deployment of attention into enhanced neural responses and thereby account for lower limits of attentional dwell time.

Information processing in the primate visual system

Information processing in the primate visual system

Temporal tuning characteristics of perceptual templates

External noise presented in temporal contiguity with a target impairs perceptual performance, reflecting the temporal tuning of the perceptual template. Deriving the temporal weights of the perceptual template, however, requires an observer model that segregates the impact of non-linearities and intrinsic inefficiencies of the observer in order to account for the impact of external noise in various temporal configurations. We showed that the perceptual template model successfully accounts for temporal masking functions measured with a wide range of temporal configurations of external noise, and estimates the temporal characteristics of the perceptual template. This was first demonstrated in estimating the temporal tuning characteristics of the perceptual template in a foveal Gabor orientation identification task. The same procedure was then used to compare the temporal tuning characteristics of the perceptual template in pre- and simultaneous cuing of spatial attention in a peripheral Gabor orientation identification task. In both experiments, four non-overlapping temporal regions of external noise, each occurring at different temporal intervals from the target display, were combined in 10 different temporal configurations. Psychometric functions were measured in the 10 external noise temporal configurations along with a zero noise condition. The PTM model provides a full account of all the psychometric functions. The estimated full width of the perceptual template at half-height is about 80 ms in experiment 1, 67 ms in the pre-cuing condition and about 90 ms in the simultaneous cuing condition in experiment 2. Manipulations of the temporal configurations of the external noise coupled with the PTM thus provide a method to characterize the temporal tuning properties of perceptual templates with an intrinsically coherent structure.

The predictive power of spectrotemporal receptive fields at the single spike train level

Event Abstract Back to Event The predictive power of spectrotemporal receptive fields at the single spike train level Auditory neurons encode the acoustic features of complex sounds according to tuning properties that make them particularly responsive to specific spectral and temporal cues. Reverse correlation models of tuning such as spectrotemporal receptive fields (STRFs) estimate the spectral and temporal tuning properties of neurons from responses to complex sounds. The accuracy of a STRF in capturing the response properties of a neuron is assessed by using the STRF to predict responses to novel sounds and comparing those predictions to peristimulus time histograms (PSTHs) of the actual responses. A PSTH averages the neural response over multiple trials and has lower temporal precision than individual spikes. Because an animal can identify a complex sound after hearing it only once, neural responses at the single spike train level, as opposed to averaged over many trials, may be highly relevant to perception. Therefore, accurate models of neural tuning should predict neural responses to a sound at the single spike train level. We asked how accurately STRFs predict individual responses to complex communication sounds. Songbirds reliably discriminate among the unique songs of individual birds. Songbird auditory midbrain neurons reliably produce spike trains that share precise temporal structure across trials. These neurons encode spectrotemporal features of songs, and their STRFs predict responses at the PSTH level well. We tested the accuracy of STRFs in predicting neural responses at the single spike train level. We recorded the responses of single auditory midbrain neurons in male zebra finches to song from 20 other males. We modeled single spike trains from those songs using a linear-nonlinear-Poisson cascade model (LNP). Using an inhomogeneous Poisson process, simulated spike trains were generated based on a STRF convolved with a song spectrogram. Simulated spike trains share temporal structure with actual spike trains, but are less reliable across trials and are therefore less temporally precise. We quantified the temporal precision of spike trains across the actual and simulated trials, and between actual and simulated spike trains from the same songs. The k-means neurometric applies the k-means clustering algorithm to Euclidean distances between spike trains as defined in the van Rossum spike timing metric. Spike trains were sorted into k groups, where k is the number of unique stimuli used to generate spike trains. Accuracy in discriminating among songs was calculated as the percentage of spike trains that cluster into the appropriate group. As a population, actual responses of MLd neurons to songs demonstrate highly accurate k-means discriminability. STRF-based LNP simulations typically fail to capture the stimulus discriminating capacity of actual neurons. This is largely due to the temporal imprecision generated by the STRF-based model. Simulated spike train clusters were more scattered than actual spike trains, and clusters were generally closer under the simulated condition. The deficiency of the STRF-based model may be due in part to an inability to resolve rapid temporal features of the stimulus. We propose methods for supplementing and improving the STRF-based LNP model, such as incorporating parameters that capture the neural response to amplitude modulations within a stimulus. Conference: Computational and systems neuroscience 2009, Salt Lake City, UT, United States, 26 Feb - 3 Mar, 2009. Presentation Type: Poster Presentation Topic: Poster Presentations Citation: (2009). The predictive power of spectrotemporal receptive fields at the single spike train level. Front. Syst. Neurosci. Conference Abstract: Computational and systems neuroscience 2009. doi: 10.3389/conf.neuro.06.2009.03.245 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 03 Feb 2009; Published Online: 03 Feb 2009. Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Google Google Scholar PubMed Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Stimulus-Dependent Auditory Tuning Results in Synchronous Population Coding of Vocalizations in the Songbird Midbrain

Physiological studies in vocal animals such as songbirds indicate that vocalizations drive auditory neurons particularly well. But the neural mechanisms whereby vocalizations are encoded differently from other sounds in the auditory system are unknown. We used spectrotemporal receptive fields (STRFs) to study the neural encoding of song versus the encoding of a generic sound, modulation-limited noise, by single neurons and the neuronal population in the zebra finch auditory midbrain. The noise was designed to match song in frequency, spectrotemporal modulation boundaries, and power. STRF calculations were balanced between the two stimulus types by forcing a common stimulus subspace. We found that 91% of midbrain neurons showed significant differences in spectral and temporal tuning properties when birds heard song and when birds heard modulation-limited noise. During the processing of noise, spectrotemporal tuning was highly variable across cells. During song processing, the tuning of individual cells became more similar; frequency tuning bandwidth increased, best temporal modulation frequency increased, and spike timing became more precise. The outcome was a population response to song that encoded rapidly changing sounds with power and precision, resulting in a faithful neural representation of the temporal pattern of a song. Modeling responses to song using the tuning to modulation-limited noise showed that the population response would not encode song as precisely or robustly. We conclude that stimulus-dependent changes in auditory tuning during song processing facilitate the high-fidelity encoding of the temporal pattern of a song.

Spatial and temporal visual properties of single neurons in the suprageniculate nucleus of the thalamus

The spatial and temporal visual sensitivity to drifting sinusoidal gratings was studied in 105 neurons of the suprageniculate nucleus of the feline thalamus. Extracellular single-unit recordings were performed in halothane-anesthetized, immobilized, artificially ventilated cats. Most suprageniculate nucleus cells were strongly sensitive to the direction of drifting gratings. The suprageniculate nucleus units had a clear preference for very low spatial frequencies with a mean of 0.05 cycle/deg. The spatial resolution was also very low with a mean of 0.16 cycle/deg. Most of the cells displayed low-pass spatial tuning characteristics, while the remainder of the units were band-pass tuned. The suprageniculate nucleus units were extremely narrowly tuned, to spatial frequencies with a mean spatial bandwidth of 1.07 octaves. A majority of the units responded optimally to high temporal frequencies, with a mean of 8.53Hz. The temporal frequency tuning functions predominantly revealed a band-pass character, with a mean temporal bandwidth of 1.66 octaves. These results demonstrate that the neurons in the suprageniculate nucleus display particular spatial and temporal characteristics. The spatial and temporal tuning properties of the suprageniculate nucleus neurons are very similar to those of the superior colliculus and the anterior ectosylvian cortex, structures that provide the main visual afferentation toward the suprageniculate nucleus. This suggests their common function in motion perception, and especially in the recording of movements of the visual environment relative to the body, and the related behavioral action.

Contrast and Temporal Frequency-Related Adaptation in the Pretectal Nucleus of the Optic Tract

In mammals, many cells in the retino-geniculate-cortical pathway adapt during stimulation with high contrast gratings. In the visual cortex, adaptation to high contrast images reduces sensitivity at low contrasts while only moderately affecting sensitivity at high contrasts, thus generating rightward shifts in the contrast response functions (contrast gain control). Similarly, motion adaptation at particular temporal frequencies (TFs) alters the temporal tuning properties of cortical cells. For the first time in any species, this paper investigates the influence of motion adaptation on both the contrast and TF responses of neurons in the retino-pretectal pathway by recording from direction-selective neurons in the nucleus of the optic tract (NOT) of the marsupial wallaby, Macropus eugenii. This species is of interest because its NOT receives almost all input directly from the retina, with virtually none from the visual cortex (unlike cats and primates). All NOT cells show changes in their contrast response functions after adaptation, many revealing contrast gain control. Contrast adaptation is direction-dependent, preferred directions producing the largest changes. The lack of cortical input suggests that contrast adaptation is generated independently from the cortex in the NOT or retina. Motion adaptation also produces direction-selective effects on the TF tuning of NOT neurons by shifting the location of the optimum TF. Cells that show strong adaptation to contrast also tend to show large changes in TF tuning, suggesting similar intracellular mechanisms. The data are discussed in terms of the generality of contrast adaptation across mammalian species and across unconnected brain regions within the same species.

Spatial and temporal visual properties of single neurons in the feline anterior ectosylvian visual area.

The spatial and temporal visual sensitivity to drifting sinusoidal gratings was studied in 75 neurons of the feline anterior ectosylvian visual area (AEV). Extracellular single-unit recordings were performed in halothane-anesthetized (0.6%), immobilized, artificially ventilated cats. Most cells were strongly sensitive to the direction of drifting gratings. The mean value of the direction tuning widths was approximately 90 deg. Most of the cells (69 of the 75 cases) displayed rather narrowly tuned band-pass characteristics in the low spatial frequency range, with a mean optimal spatial frequency of 0.2 cycles/degree (c/deg). The mean spatial bandwidth was 1.4 octaves. The remainder of the units was low-pass tuned. A majority of the units responded optimally to high temporal frequencies (mean 6.3 Hz), although some cells did exhibit preferences for every examined temporal frequency between 0.6 Hz and 10.8 Hz. The temporal frequency-tuning functions mostly revealed a band-pass character with a mean temporal bandwidth of 1.1 octaves. Our results demonstrate that the neurons along the anterior ectosylvian sulcus display particular spatial and temporal characteristics. The AEV neurons, with their preference for low spatial frequencies and with their fine spatial and temporal tuning properties, seem to be candidates for special tasks in motion perception.

Frequency and periodicity are represented in orthogonal maps in the human auditory cortex: evidence from magnetoencephalography

Timbre and pitch are two independent perceptual qualities of sounds closely related to the spectral envelope and to the fundamental frequency of periodic temporal envelope fluctuations, respectively. To a first approximation, the spectral and temporal tuning properties of neurons in the auditory midbrain of various animals are independent, with layouts of these tuning properties in approximately orthogonal tonotopic and periodotopic maps. For the first time we demonstrate by means of magnetoencephalography a periodotopic organization of the human auditory cortex and analyse its spatial relationship to the tonotopic organization by using a range of stimuli with different temporal envelope fluctuations and spectra and a magnetometer providing high spatial resolution. We demonstrate an orthogonal arrangement of tonotopic and periodotopic gradients. Our results are in line with the organization of such maps in animals and closely match the perceptual orthogonality of timbre and pitch in humans.

Dopamine inhibits mammalian photoreceptor Na+,K+-ATPase activity via a selective effect on the alpha3 isozyme.

The rat retina contains dopaminergic interplexiform cells that send processes to the outer plexiform layer where dopamine is released in a light-dependent manner. We report herein that physiologically relevant concentrations of dopamine inhibited ouabain-sensitive photoreceptor oxygen consumption in dark- and light-adapted rat retinas and inhibited Na+,K+-ATPase specific activity (EC 3.6.1.37) in a rat rod outer-inner segment preparation. Experiments with the selective D1 agonist fenoldopam or D2 agonist quinpirole and experiments with dopamine plus either the D1 antagonist SCH23390 or D2/D4 antagonist clozapine showed that the inhibition of oxygen consumption and enzyme activity were mediated by D2/D4-like receptors. The amphetamine-induced release of dopamine, monitored by the inhibition of oxygen consumption, was blocked by L-2-amino-4-phosphonobutyric acid and kynurenic acid. Pharmacological and biochemical experiments determined that the IC50 values of ouabain for the alpha1-low and alpha3-high ouabain affinity isozymes of photoreceptor Na+,K+-ATPase were approximately 10(-5) and approximately 10(-7) M, respectively, and that the D2/D4-like mediated inhibition of Na+,K+-ATPase was exclusively selective for the alpha3 isozyme. The dopamine-mediated inhibition of alpha3 first occurred at 5 nM, was maximal at 100 microM (-47%), had an IC50 value of 382 +/- 23 nM, and exhibited negative cooperativity (Hill coefficient, 0.27). Prior homogenization of the rod outer-inner segment completely prevented the long-lasting inhibition, suggesting that the effect was coupled to a second messenger. Although the physiological significance of our findings to photoreceptor function is unknown, we hypothesize that these results may have relevance for the temporal tuning properties of rods.

Brain-stem modulation of the response properties of cells in the cat's perigeniculate nucleus.

Transmission through the lateral geniculate nucleus is facilitated following activation of the cholinergic input from the brain stem, which is thought to reflect activity patterns seen during arousal. One of the underlying mechanisms is the suppression of inhibitory circuits local to the lateral geniculate nucleus. However, evidence exists that some visually driven inhibitory inputs to geniculate nucleus. preserved or even enhanced under conditions of arousal, and during electrical activation of the parabrachial region of the brain stem. We have therefore reexamined the effect of brain-stem activation on the visual responses of one group of local inhibitory inputs to geniculate relay cells, those emanating from the adjacent perigeniculate nucleus. We recorded single perigeniculate cells in anesthetized, paralyzed cats. Axons innervating the lateral geniculate and perigeniculate nuclei from the parabrachial region of the brain stem were electrically activated, and the effect of this activation was assessed on both spontaneous and visually evoked responses. Visual stimulation consisted of sinusoidally modulated sine-wave gratings of varying spatial and temporal frequency. For the great majority of perigeniculate cells (32 of 40), brain-stem activation inhibited spontaneous activity, while one cell was excited, three showed a mixed effect and four were unaffected. Nevertheless, the responses of most cells (30 of 40) were facilitated when brain-stem activation was paired with certain spatio-temporal patterns of visual stimulation. Spatial tuning curves were constructed for 17 cells and temporal tuning curves for 14, before and during parabrachial activation. The responses of any one cell could be facilitated, unchanged, or suppressed, depending on the visual stimulus used. In some cases, this substantially modified the cell's spatial and temporal tuning properties. We conclude that activation of the brain stem disinhibits geniculate relay cells in the absence of visual stimulation, but it has the potential to enhance either the magnitude or specificity of visually driven inhibition arising from the perigeniculate nucleus.

Effects of membrane voltage on receptive field properties of lateral geniculate neurons in the cat: contributions of the low-threshold Ca2+ conductance.

1. Thalamic relay cells, including those of the lateral geniculate nucleus, display a low-threshold spike (LT spike), which is a large depolarization due to an increased Ca2+ conductance. Typically riding the crest of each LT spike is a burst of from two to seven action potentials, which we refer to as the LT burst. The LT spike is voltage dependent, because if the cell's resting membrane potential is more depolarized than roughly -60 mV, the LT spike is inactivated, but if more hyperpolarized, the spike is deinactivated and can be activated by a depolarization, such as from an afferent excitatory postsynaptic potential (EPSP). Thalamic relay cells thus display two response modes: a relay or tonic mode, when the cell is depolarized and LT spikes are inactivated, leading to tonic firing of action potentials; and a burst mode, when the cell is hyperpolarized and tends to respond with LT spikes and their associated bursts of action potentials. 2. We were interested in the contribution of the LT spike on the transmission of visually evoked signals through geniculate relay cells to visual cortex. We recorded intracellularly from geniculate cells in an anesthetized, paralyzed, in vivo cat preparation to study the effects of membrane voltage, and thus the presence or absence of LT spikes, on responses to drifting sine-wave gratings. We monitored the visually evoked responses of 14 geniculate neurons (6 X, 7 Y, and 1 unclassified) at different membrane potentials at which LT spikes were inactivated or deinactivated. 3. Changing membrane voltage during visual stimulation switched the response mode of every cell between the relay and burst modes. In the burst mode, LT spikes occurred in phase with the visual stimulus and not at rhythmic intervals uncorrelated to visual stimuli. To any given stimulus cycle, the cell responded usually with an LT burst or a tonic response, and rarely was more than one LT burst evoked by a stimulus cycle. Occasionally a single cycle evoked both an LT burst and tonic response, but always the LT burst occurred first. 4. The spatial tuning characteristics of the cells did not differ dramatically as a function of membrane potential, because the tuning of the LT bursts was quite similar to that of the tonic response component. Although we did not obtain complete temporal tuning properties, we did note that hyperpolarized cells responded reliably with LT bursts at several temporal frequencies. 5. A consistent difference was seen between the LT burst and tonic response components in terms of response linearity.(ABSTRACT TRUNCATED AT 400 WORDS)

Spatial and temporal contrast sensitivity of neurones in areas 17 and 18 of the cat's visual cortex.

1. We have examined the spatial and temporal tuning properties of 238 cortical neurones, recorded using conventional techniques from acutely prepared anaesthetized cats. We determined spatial and temporal frequency tuning curves using sinusoidal grating stimuli presented to each neurone's receptive field by a digital computer on a cathode ray tube. 2. We measured tuning curves either by determining response amplitude as a function of spatial or temporal frequency, or by measuring contrast sensitivity (the inverse of the contrast of the grating that just elicited a detectable response). The two measures give very similar tuning curves in all cases. 3. We recorded from 184 neurones in area 17; of these 156 had receptive fields within 5 degrees of the area centralis. The range of preferred spatial frequency for these neurones was 0.3--3 c/deg, and their spatial frequency tuning band widths varied from 0.7 to 3.2 octaves at half-amplitude. The most common band width was roughly 1.3 octaves. Simple and complex cells in area 17 did not differ in their distributions of preferred spatial frequency, although complex cells were, on average, slightly less selective for spatial frequency than simple cells. 4. We recorded from fifty-four neurones from area 18, and performed several experiments in which we recorded from corresponding portions of both area 17 and area 18 in the same electrode penetration. Neurones in area 18 preferred spatial frequencies that were, on average, one third as high as those preferred by area 17 neurones at the same retinal eccentricity. Thus the range of preferred spatial frequency in area eighteen cells having receptive fields within 5 deg of the area centralis was between less than 0.1 and 0.5 c/deg. The distributions of optimum spatial frequency in the two areas were practically non-overlapping at eccentricities as high as 15 deg, the greatest eccentricity we examined. Neurones in area 18 were about as selective for spatial frequency as were neurones in area 17. 5. We determined temporal frequency tuning characteristics for some neurones from each area, using gratings that moved steadily across the screen. Neurones from area 17 all responded well to low temporal frequencies, and less well to higher frequencies (in excess of, usually, 2 or 4 Hz). In contrast, neurones recorded from area 18 sometimes had similar tuning properties, but more commonly showed a pronounced reduction in response as the temporal frequency was moved either above or below some optimum value (usually 2--8 Hz). 6. We conclude from these results that areas 17 and 18 act in parallel to process different aspects of the visual information relayed from the retina via the lateral geniculate complex. Some or all of the differences between the areas may be attributable to the predominance of Y cell input to area 18 and the predominance of X cell input to area 17...