Drifting Sine Wave Gratings Research Articles

RPG: A low-cost, open-source, high-performance solution for displaying visual stimuli

BackgroundThe development of new high throughput approaches for neuroscience such as high-density silicon probes and 2-photon imaging have led to a renaissance in visual neuroscience. However, generating the stimuli needed to evoke activity in the visual system still represents a non-negligible difficulty for experimentalists. While several widely used software toolkits exist to deliver such stimuli, they all suffer from some shortcomings. Primarily, the hardware needed to effectively display such stimuli comes at a significant financial cost, and secondly, triggering and/or timing the stimuli such that it can be accurately synchronized with other devices requires the use of legacy hardware, further hardware, or bespoke solutions. ResultsHere we present RPG (Raspberry Pi Gratings), a Python package written for the Raspberry Pi, which overcomes these issues. Specifically, the Raspberry Pi is a low-cost, credit card sized computer with general purpose input/output pins, allowing RPG to be triggered to deliver stimuli and to provide real-time feedback on stimulus timing. RPG delivers stimuli at 60 frames per second and the feedback of frame timings is accurate to 10s of microseconds. Comparison with existing method(s)With respect to the accuracy of frame timings, the performance of RPG is at least as accurate as commonly used packages. However, the inbuilt ability to trigger stimuli and the real-time feedback of frame timings will be extremely useful for certain experiments. ConclusionsRPG provides a simple to use Python interface that is capable of generating drifting sine wave gratings, Gabor patches and displaying raw images/video.

Spatial clustering of orientation preference in primary visual cortex of the large rodent agouti.

SummaryAll rodents investigated so far possess orientation-selective neurons in the primary visual cortex (V1) but – in contrast to carnivores and primates – no evidence of periodic maps with pinwheel-like structures. Theoretical studies debating whether phylogeny or universal principles determine development of pinwheels point to V1 size as a critical constraint. Thus, we set out to study maps of agouti, a big diurnal rodent with a V1 size comparable to cats'. In electrophysiology, we detected interspersed orientation and direction-selective neurons with a bias for horizontal contours, corroborated by homogeneous activation in optical imaging. Compatible with spatial clustering at short distance, nearby neurons tended to exhibit similar orientation preference. Our results argue against V1 size as a key parameter in determining the presence of periodic orientation maps. They are consistent with a phylogenetic influence on the map layout and development, potentially reflecting distinct retinal traits or interspecies differences in cortical circuitry.

The speed of neuronal adaptation: A perspective through the visual cortex.

The speed of neuronal adaptation: A perspective through the visual cortex.

Spectral receptive field properties of visually active neurons in the caudate nucleus

Recent studies stress the importance of the caudate nucleus in visual information processing. Although the processing of moving visual signals depends upon the capability of a system to integrate spatial and temporal information, no study has investigated the spectral receptive field organization of the caudate nucleus neurons yet. Therefore, we tested caudate neurons of the feline brain by extracellular single-cell recording applying drifting sinewave gratings of various spatial and temporal frequencies, and reconstructed their spectral receptive fields by plotting their responsiveness as a function of different combinations of spatial and temporal frequencies. The majority of the caudate cells (74%) exhibited peak tuning, which means that their spatio-temporal frequency response profile had a characteristic region of increased activity with a single maximum in the spatio-temporal frequency domain. In one-quarter of the recorded caudate neurons ridge tuning was found, where the region of increased activity, forming an elongated ridge of maximal sensitivity parallel or angled to the spatial or the temporal frequency axis, indicating temporal (16%), spatial (5%) or speed (5%) tuning, respectively. The velocity preference of the ridge tuned caudate nucleus neurons is significantly lower than that of the peak tuned neurons. The peak tuned neuron could encode high velocities, while the ridge tuned neurons were responsible for the detection of moderate and lower velocities. Based upon our results, we suggest that the wide variety of spatio-temporal frequency response profiles might represent different functional neuronal groups within the caudate nucleus that subserve different behaviors to meet various environmental requirements.

Area Summation in Human Visual System: Psychophysics, fMRI, and Modeling

Contextual modulation is a fundamental feature of sensory processing, both on perceptual and on single-neuron level. When the diameter of a visual stimulus is increased, the firing rate of a cell typically first increases (summation field) and then decreases (surround field). Such an area summation function draws a comprehensive profile of the receptive field structure of a neuron, including areas outside the classical receptive field. We investigated area summation in human vision with psychophysics and functional magnetic resonance imaging (fMRI). The stimuli were drifting sine wave gratings similar to those used in previous macaque single-cell area summation studies [corrected]. A model was developed to facilitate comparison of area summation in fMRI to area summation in psychophysics and single cells. The model consisted of units with an antagonistic receptive field structure found in single cells in the primary visual cortex. The receptive field centers of the model neurons were distributed in the region of the visual field covered by a single voxel. The measured area summation functions were qualitatively similar to earlier single-cell data. The model with parameters derived from psychophysics captured the spatial structure of the summation field in the primary visual cortex as measured with fMRI. The model also generalized to a novel situation in which the neural population was displaced from the stimulus center. The current study shows that contextual modulation arises from similar spatially antagonistic and overlapping excitatory and inhibitory mechanisms, both in single cells and in human vision.

Cooling in cat visual cortex: stability of orientation selectivity despite changes in responsiveness and spike width

Cooling is one of several reversible methods used to inactivate local regions of the brain. Here the effect of cooling was studied in the primary visual cortex (area 17) of anaesthetized and paralyzed cats. When the cortical surface temperature was cooled to about 0 °C, the temperature 2 mm below the surface was 20 °C. The lateral spread of cold was uniform over a distance of at least ∼700 μm from the cooling loop. When the cortex was cooled the visually evoked responses to drifting sine wave gratings were strongly reduced in proportion to the cooling temperature, but the mean spontaneous activity of cells decreased only slightly. During cooling the strongest effect on the orientation tuning curve was on the peak response and the orientation bandwidth did not change, suggesting a divisive mechanism. Our results show that the cortical circuit is robust in the face of cooling and retains its essential functionality, albeit with reduced responsiveness. The width of the extracellular spike waveform measured at half height increased by 50% on average during cooling in almost all cases and recovered after re-warming. The increase in spike width was inversely correlated with the change in response amplitude to the optimal stimulus. The extracellular spike shape can thus be used as a reliable and fast method to assess whether changes in the responses of a neuron are due to direct cooling or distant effects on a source of its afferents.

Spatial and temporal visual properties of the neurons in the intermediate layers of the superior colliculus

Although the visual perception depends on the integration of spatial and temporal information, no knowledge is available concerning the responsiveness of neurons in the intermediate layers of the superior colliculus (SCi) to extended visual grating stimuli. Accordingly, we set out to investigate the responsiveness of these neurons in halothane-anesthetized cats to drifting sinewave gratings at various spatial and temporal frequencies. The SCi units responded optimally to gratings of low spatial frequencies (none of the analyzed SCi units exhibited maximal activity to spatial frequencies higher than 0.3 c/deg) and exhibited low spatial resolution and narrow spatial frequency tuning. On the other hand, the SCi neurons preferred high temporal frequencies and exhibited high temporal resolution. Thus, the SCi neurons seem to be good spatio–temporal filters of visual information in the low spatial and high temporal frequency domain. Based upon the above summarized results we suggest that the SCi units can detect large contours moving at high velocities well, but are unable to distinguish small details. This is in line with the generally held view that the SCi could possess visuomotor function, such as organizing the complex, sensory-guided oculomotor and skeletomotor responses during the self-motion of the animal.

The processing of feature discontinuities for different cue types in primary visual cortex

This study examines whether neurons in the primary visual cortex (V1) of the cat (also referred to as area 17) are sensitive to boundaries that are delineated by a difference in features other than luminance contrast. Most research on this issue has concentrated on the responses to texture borders (e.g. ‘illusory contours’) and has found neurons that are sensitive to such borders in V2 and to a lesser extent in V1. Here neurons in cat area 17 (V1) were exposed to borders that were oblique to the orientation preference of the neuron and that were created by differences in phase, orientation or direction of motion of two drifting sinewave gratings. Nearby phase borders evoked increased firing in 15 out of 98 neurons, orientation borders in 18 out of 98, and direction borders in 15 out of 70 neurons recorded in area 17 (V1) of anesthetized cats. The firing rates of these neurons were enhanced when a feature border was presented close to their receptive field, partly independent of the cue involved. Control experiments with a contrast border showed that the enhanced firing was due to a release of suppression rather than facilitation. A conceptual model is presented that can describe the data and uncovers a peculiarity of the phase domain compared to the orientation and direction domain. The model unifies the knowledge gained here about orientation-specific center–surround interactions, contextual effects, and end-stopping. The data and model suggest that these phenomena are part of a single mechanism that enables the brain to detect feature discontinuities across a range of features.

Complex motion selectivity in PMLS cortex following early lesions of primary visual cortex in the cat

In the cat, the analysis of visual motion cues has generally been attributed to the posteromedial lateral suprasylvian cortex (PMLS) (Toyama et al., 1985; Rauschecker et al., 1987; Rauschecker, 1988; Kim et al., 1997). The responses of neurons in this area are not critically dependent on inputs from the primary visual cortex (VC), as lesions of VC leave neuronal response properties in PMLS relatively unchanged (Spear & Baumann, 1979; Spear, 1988; Guido et al., 1990b). However, previous studies have used a limited range of visual stimuli. In this study, we assessed whether neurons in PMLS cortex remained direction-selective to complex motion stimuli following a lesion of VC, particularly to complex random dot kinematograms (RDKs). Unilateral aspiration of VC was performed on post-natal days 7-9. Single unit extracellular recordings were performed one year later in the ipsilateral PMLS cortex. As in previous studies, a reduction in the percentage of direction selective neurons was observed with drifting sinewave gratings. We report a previously unobserved phenomenon with sinewave gratings, in which there is a greater modulation of firing rate at the temporal frequency of the stimulus in animals with a lesion of VC, suggesting an increased segregation of ON and OFF sub-regions. A significant portion of neurons in PMLS cortex were direction selective to simple (16/18) and complex (11/16) RDKs. However, the strength of direction selectivity to both stimuli was reduced as compared to normals. The data suggest that complex motion processing is still present, albeit reduced, in PMLS cortex despite the removal of VC input. The complex RDK motion selectivity is consistent with both geniculo-cortical and extra-geniculate thalamo-cortical pathways in residual direction encoding.

Cortical locus of coherent motion deficits in deaf poor readers

Samar and Parasnis [Samar, V. J., & Parasnis, I. (2005). Dorsal stream deficits suggest hidden dyslexia among deaf poor readers: correlated evidence from reduced perceptual speed and elevated coherent motion detection thresholds. Brain and Cognition, 58, 300–311.] reported that correlated measures of coherent motion detection and perceptual speed predicted reading comprehension in deaf young adults. Because deficits in coherent motion detection have been associated with dyslexia in the hearing population, and because coherent motion detection is strongly dependent on extrastriate cortical area MT, these results are consistent with the claim that hidden dyslexia occurs within the deaf population and is associated with deficits in MT. However, coherent motion detection can also be influenced by subcortical deficits in both magnocellular and parvocellular pathways. To confirm the putative cortical locus of coherent motion perception deficits, we measured contrast thresholds for detecting the direction of movement of drifting sine wave gratings in the same participant group as [Samar, V. J., & Parasnis, I. (2005). Dorsal stream deficits suggest hidden dyslexia among deaf poor readers: correlated evidence from reduced perceptual speed and elevated coherent motion detection thresholds. Brain and Cognition, 58, 300–311.], under stimulus conditions that selectively biased for input from the subcortical magnocellular and parvocellular pathways, respectively. Contrast thresholds were not related to reading comprehension performance under either the magnocellular or parvocellular conditions. Furthermore, the previously reported correlations among reading comprehension, coherent motion thresholds, and perceptual speed remained significant even after contrast thresholds and non-verbal IQ were controlled in partial correlation analyses. In addition, coherent motion detection thresholds were found to correlate specifically with a reading-IQ discrepancy score, one commonly used indicator of dyslexia. These results provide direct psychophysical evidence that the previously reported deficit in coherent motion detection in deaf poor readers does not involve subcortical pathway deficits, but rather is associated with a cortical deficit likely involving area MT. They also strengthen the argument for the existence of hidden dyslexia in the deaf adult population.

Functional Response Properties of Neurons in the Dorsomedial Visual Area of New World Monkeys (Callithrix jacchus)

The dorsomedial visual area (DM), a subdivision of extrastriate cortex located near the dorsal midline, is characterized by heavy myelination and a relative emphasis on peripheral vision. To date, DM remains the least understood of the three primary targets of projections from the striate cortex (V1) in New World monkeys. Here, we characterize the responses of DM neurons in anaesthetized marmosets to drifting sine wave gratings. Most (82.4%) cells showed bidirectional sensitivity, with only 6.9% being strongly direction selective. The distribution of orientation sensitivity was bimodal, with a distinct population (corresponding to over half of the sample) formed by neurons with very narrow selectivity. When compared with a sample of V1 units representing a comparable range of eccentricities, DM cells revealed a preference for much lower spatial frequencies, and higher speeds. End inhibition was extremely rare, and the responses of many cells summated over distances as large as 30 degrees. Our results suggest clear differences between DM and the two other main targets of V1 projections, the second (V2) and middle temporal (MT) areas, with cells in DM emphasizing aspects of visual information that are likely to be relevant for motor control.

Visual Warning Signals Optimized for Human Perception: What the Eye Sees Fastest

This study aimed to answer the question of how to design a visual warning signal that is most easily seen and produces the quickest reaction time. This is a classic problem of bionic optimization—if one knows the properties of the receiver one can most easily find a suitable solution. Because the peak of the spatio-temporal contrast sensitivity function of the human visual system occurs at non-zero spatial and temporal frequencies, it is likely that movement enhances the detectability of threshold visual signals. Earlier studies employing extended drifting sinewave gratings bear out this prediction. We have studied the ability of human observers to detect threshold visual signals for both moving and stationary stimuli. We used discrete, localized signals such as might be employed in aerospace or automotive warning signal displays. Moving stimuli show a superior detectability to non-moving stimuli of the same integrated energy. Moving stimuli at threshold detectability are seen faster than non-moving threshold stimuli. Under some conditions the speed advantage is over 0.25 seconds. Similar advantages have also been shown to occur for suprathreshold signals.

Temporal frequency and velocity-like tuning in the pigeon accessory optic system.

Neurons in the accessory optic system (AOS) and pretectum are involved in the analysis of optic flow and the generation of the optokinetic response. Previous studies found that neurons in the pretectum and AOS exhibit direction selectivity in response to large-field motion and are tuned in the spatiotemporal domain. Furthermore, it has been emphasized that pretectal and AOS neurons are tuned to a particular temporal frequency, consistent with the "correlation" model of motion detection. We examined the responses of neurons in the nucleus of the basal optic root (nBOR) of the AOS in pigeons to large-field drifting sine wave gratings of varying spatial (SF) and temporal frequencies (TF). nBOR neurons clustered into two categories: "Fast" neurons preferred low SFs and high TFs, and "Slow" neurons preferred high SFs and low TFs. The fast neurons were tuned for TF, but the slow nBOR neurons had spatiotemporally oriented peaks that suggested velocity tuning (TF/SF). However, the peak response was not independent of SF; thus we refer to the tuning as "apparent velocity tuning" or "velocity-like tuning." Some neurons showed peaks in both the fast and slow regions. These neurons were TF-tuned at low SFs, and showed velocity-like tuning at high SFs. We used computer simulations of the response of an elaborated Reichardt detector to show that both the TF-tuning and velocity-like tuning shown by the fast and slow neurons, respectively, may be explained by modified versions of the correlation model of motion detection.

The Accessory Optic System Contributes to the Spatio-Temporal Tuning of Motion-Sensitive Pretectal Neurons

The nucleus of the basal optic root (nBOR) of the accessory optic system (AOS) and the pretectal nucleus lentiformis mesencephali (LM) are involved in the analysis of optic flow that results from self-motion and are important for oculomotor control. These neurons have large receptive fields and exhibit direction selectivity to large moving stimuli. In response to drifting sine wave gratings, LM and nBOR neurons are tuned to either low spatial/high temporal frequencies (SF, TF) or high SF/low TF stimuli. Given that velocity = TF/SF, these are referred to as "fast" and "slow" neurons, respectively. There is a heavy projection from the AOS to the pretectum, although its function is unknown. We recorded the directional and spatio-temporal tuning of LM units in pigeons before and after nBOR was inactivated by tetrodotoxin injection. After nBOR inactivation, changes in direction preference were observed for only one of 18 LM units. In contrast, the spatio-temporal tuning of LM units was dramatically altered by nBOR inactivation. Two major effects were observed. First, in response to motion in the preferred direction, most (82%) neurons showed a substantially reduced (mu = -67%) excitation to low SF/high TF gratings. Second, in response to motion in the anti-preferred direction, most (63%) neurons showed a dramatically reduced (mu = -78%) inhibition to high SF/low TF gratings. Thus the projection from the nBOR contributes to the spatio-temporal tuning rather than the directional tuning of LM neurons. We propose a descriptive model whereby LM receives inhibitory and excitatory input from "slow" and "fast" nBOR neurons, respectively.

Fast and slow neurons in the nucleus of the basal optic root in pigeons

The nucleus of the basal optic root (nBOR) is involved in the generation of the optokinetic response. Previous studies showed that most nBOR neurons exhibit direction-selectivity in response to largefield motion. We investigated the responses of pigeon nBOR neurons to drifting sine wave gratings of varying spatial and temporal frequency (SF,TF). Two groups of neurons were revealed. The first group preferred gratings of low SF (mean, 0.07 cycles per degree (cpd)) and high TF (mean, 0.76 Hz) (‘fast’ stimuli). The second group preferred gratings of high SF (mean, 0.56 cpd) and lower TF (mean, 0.33 Hz) (‘slow’ stimuli). Previous studies have demonstrated fast and slow neurons in pretectal nucleus lentiformis mesencephali, which is also involved in the generation of the optokinetic response.

Visual Warning Signals Optimized for Human Perception: What the Eye Sees Fastest

This study aimed to answer the question of how to design a visual warning signal that is most easily seen and produces the quickest reaction time. This is a classic problem of bionic optimization—if one knows the properties of the receiver one can most easily find a suitable solution. Because the peak of the spatio‐temporal contrast sensitivity function of the human visual system occurs at non‐zero spatial and temporal frequencies, it is likely that movement enhances the detectability of threshold visual signals. Earlier studies employing extended drifting sinewave gratings bear out this prediction. We have studied the ability of human observers to detect threshold visual signals for both moving and stationary stimuli. We used discrete, localized signals such as might be employed in aerospace or automotive warning signal displays. Moving stimuli show a superior detectability to non‐moving stimuli of the same integrated energy. Moving stimuli at threshold detectability are seen faster than non‐moving threshold stimuli. Under some conditions the speed advantage is over 0.25 seconds. Similar advantages have also been shown to occur for suprathreshold signals.

Residual binocular interactions in the striate cortex of monkeys reared with abnormal binocular vision.

We investigated the nature of residual binocular interactions in the striate cortex (V1) of monkey models for the two most common causes of visual dysfunction in young children, specifically anisometropia and strabismus. Infant rhesus monkeys were raised wearing either anisometropic spectacle lenses that optically defocused one eye or ophthalmic prisms that optically produced diplopia and binocular confusion. Earlier psychophysical investigations had demonstrated that all subjects exhibited permanent binocular vision deficits and, in some cases, amblyopia. When the monkeys were adults, the responses of individual V1 neurons were studied with the use of microelectrode recording techniques while the animals were anesthetized and paralyzed. The manner in which the signals from the two eyes were combined in individual cells was investigated by dichoptically stimulating both eyes simultaneously with drifting sine wave gratings. In both lens- and prism-reared monkeys, fewer neurons had balanced ocular dominances and greater numbers of neurons were excited by only one eye. However, many neurons that appeared to be monocular exhibited clear binocular interactions during dichoptic stimulation. For the surviving binocular neurons, the maximum binocular response amplitudes were lower than normal; fewer neurons, particularly complex cells, were sensitive to relative interocular spatial phase disparities; and the remaining disparity-sensitive neurons exhibited lower degrees of binocular interaction. In prism-reared monkeys, an unusually high proportion of complex cells exhibited binocular suppression during dichoptic stimulation. Binocular contrast summation experiments showed that for both cooperative and antagonistic binocular interactions, contrast signals from the two eyes were combined by individual neurons in a normal linear fashion in both lens- and prism-reared monkeys. The observed binocular deficits appear to reflect a reduction in functional inputs from one eye and/or spatial imprecision in the monocular receptive fields rather than an aberrant form of binocular interaction. In the prism-reared monkeys, the predominance of suppression suggests that inhibitory connections were, however, less susceptible to diplopia and confusion than excitatory connections. Overall, there were many parallels between V1 physiology in our monkey models and the residual vision of humans with anisometropia or strabismus.

Binocular spatial phase tuning characteristics of neurons in the macaque striate cortex.

We employed microelectrode recording techniques to study the sensitivity of individual neurons in the striate cortex of anesthetized and paralyzed monkeys to relative interocular image disparities and to determine the effects of basic stimulus parameters on these cortical binocular interactions. The visual stimuli were drifting sine wave gratings. After the optimal stimulus orientation, spatial frequency, and direction of stimulus movement were found, the cells' disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic grating pairs. No attempts were made to assess absolute position disparities or horizontal disparities relative to the horopter. The majority (approximately 70%) of simple cells were highly sensitive to interocular spatial phase disparities, particularly neurons with balanced ocular dominances. Simple cells typically demonstrated binocular facilitation at the optimal phase disparity and binocular suppression for disparities 180 degrees away. Fewer complex cells were phase selective (approximately 40%); however, the range of disparity selectivity in phase-sensitive complex cells was comparable with that for simple cells. Binocular interactions in non-phase-sensitive complex cells were evidenced by binocular response amplitudes that differed from responses to monocular stimulation. The degree of disparity tuning was independent of a cell's optimal orientation or the degree of direction tuning. However, disparity-sensitive cells tended to have narrow orientation tuning functions and the degree of disparity tuning was greatest for the optimal stimulus orientations. Rotating the stimulus for one eye 90 degrees from the optimal orientation usually eliminated binocular interactions. The effects of phase disparities on the binocular response amplitude were also greatest at the optimal spatial frequency. Thus a cell's sensitivity to absolute position disparities reflects its spatial tuning characteristics, with cells sensitive to high spatial frequencies being capable of signaling very small changes in image disparity. On the other hand, stimulus contrast had relatively little effect on a cell's disparity tuning, because response saturation occurred at the same contrast level for all relative interocular phase disparities. Thus, as with orientation tuning, a cell's optimal disparity and the degree of disparity selectivity were invariant with contrast. Overall, the results show that sensitivity to interocular spatial phase disparities is a common property of striate neurons. A cell's disparity tuning characteristics appear to largely reflect its monocular receptive field properties and the interocular balance between excitatory and inhibitory inputs. However, distinct functional classes of cortical neurons could not be discriminated on the basis of disparity sensitivity alone.

Stimulus-dependent modulation of spike burst length in cat striate cortical cells.

Burst activity, defined by groups of two or more spikes with intervals of < or = 8 ms, was analyzed in responses to drifting sinewave gratings elicited from striate cortical neurons in anesthetized cats. Bursting varied broadly across a population of 507 simple and complex cells. Half of this population had > or = 42% of their spikes contained in bursts. The fraction of spikes in bursts did not vary as a function of average firing rate and was stationary over time. Peaks in the interspike interval histograms were found at both 3-5 ms and 10-30 ms. In many cells the locations of these peaks were independent of firing rate, indicating a quantized control of firing behavior at two different time scales. The activity at the shorter time scale most likely results from intrinsic properties of the cell membrane, and that at the longer scale from recurrent network excitation. Burst frequency (bursts per s) and burst length (spikes per burst) both depended on firing rate. Burst frequency was essentially linear with firing rate, whereas burst length was a nonlinear function of firing rate and was also governed by stimulus orientation. At a given firing rate, burst length was greater for optimal orientations than for nonoptimal orientations. No organized orientation dependence was seen in bursts from lateral geniculate nucleus cells. Activation of cortical contrast gain control at low response amplitudes resulted in no burst length modulation, but burst shortening at optimal orientations was found in responses characterized by supersaturation. At a given firing rate, cortical burst length was shortened by microinjection of gamma-aminobutyric acid (GABA), and bursts became longer in the presence of N-methyl-bicuculline, a GABA(A) receptor blocker. These results are consistent with a model in which responses are reduced at nonoptimal orientations, at least in part, by burst shortening that is mediated by GABA. A similar mechanism contributes to response supersaturation at high contrasts via recruitment of inhibitory responses that are tuned to adjacent orientations. Burst length modulation can serve as a form of coding by supporting dynamic, stimulus-dependent reorganization of the effectiveness of individual network connections.

Directionally selective motion detection in the sustaining fibers of the crayfish optic nerve: linear and nonlinear mechanisms.

1. Directional selectivity of crayfish sustaining fibers was examined with drifting sine wave gratings and with intracellular and extracellular recordings. Directionality was measured for variations in stimulus contrast, spatial frequency, and temporal frequency. 2. Sustaining fibers exhibit directional selectivity in the magnitude of the compound postsynaptic potential (PSP), the impulse frequency modulation response, and the mean firing rate. The mean synaptic potential is insensitive to direction. The directionality of the mean impulse rate appears to arise by rectification in the voltage-to-impulse transduction. 3. The preferred directions of three identified sustaining fibers are similar to those of head-down optomotor neurons to which these sustaining fibers project. 4. The modulatory response, elicited by gratings drifting in the preferred direction, increased linearly with contrast until saturation (typically at a contrast of 0.5), where maximum directional selectivity obtains. 5. The magnitude of the directional response is a band-pass function of spatial and temporal frequency and exhibits reversal of directionality (i.e., aliasing) at high spatial and temporal frequencies. The results imply a spatial sampling interval of 4.5 degrees and a temperature-dependent inhibitory delay of 40-90 ms. The PSP modulation response shares several features with that of neighboring tangential (Tan1) neurons. 6. A qualitative model is proposed for the transformation of a phase-sensitive, linear directional response to a phase-insensitive and nonlinear time-averaged response, based on the functional connections from Tan1 neurons to sustaining fibers to optomotor neurons. The model includes a threshold rectification, a synaptic band-pass filter, and differences in temporal phase among converging modulatory signals.