Dopamine transmission in the tail striatum: Regional variation and contribution of dopamine clearance mechanisms.

The striatum can be divided into four anatomically and functionally distinct domains: the dorsolateral, dorsomedial, ventral and the more recently identified caudolateral (tail) striatum. Dopamine transmission in these striatal domains underlies many important behaviours, yet little is known about this phenomenon in the tail striatum. Furthermore, the tail is divided anatomically into four divisions (dorsal, medial, intermediate and lateral) based on the profile of D1 and D2 dopamine receptor-expressing medium spiny neurons, something that is not seen elsewhere in the striatum. Considering this organisation, how dopamine transmission occurs in the tail striatum is of great interest. We recorded evoked dopamine release in the four tail divisions, with comparison to the dorsolateral striatum, using fast-scan cyclic voltammetry in rat brain slices. Contributions of clearance mechanisms were investigated using dopamine transporter knockout (DAT-KO) rats, pharmacological transporter inhibitors and dextran. Evoked dopamine release in all tail divisions was smaller in amplitude than in the dorsolateral striatum and, importantly, regional variation was observed: dorsolateral ≈ lateral > medial > dorsal ≈ intermediate. Release amplitudes in the lateral division were 300% of that in the intermediate division, which also exhibited uniquely slow peak dopamine clearance velocity. Dopamine clearance in the intermediate division was most dependent on DAT, and no alternative dopamine transporters investigated (organic cation transporter-3, norepinephrine transporter and serotonin transporter) contributed significantly to dopamine clearance in any tail division. Our findings confirm that the tail striatum is not only a distinct dopamine domain but also that each tail division has unique dopamine transmission characteristics. This supports that the divisions are not only anatomically but also functionally distinct. How this segregation relates to the overall function of the tail striatum, particularly the processing of multisensory information, is yet to be determined.

A filter-rectify-filter model of the tactile perception of 3D-printed, texture-defined form.

We show that a two-stage filter-rectify-filter (FRF) model, previously used to explain the visual perception of texture-defined form, can also account for the tactile perception of texture-defined form. This result is interesting because, first, relatively little is known about the neural mechanisms of tactile form perception, and second, the generalization of the model may reflect a canonical computation at work in both visual and somatosensory cortex. We 3D-printed test objects comprising a regular, rectangular array of raised, oriented bars measuring 0.75 × 0.75 × 3 mm (width × height × length) that were centre-to-centre spaced by 4 mm. Bars on the left-hand-side of a test object were horizontal, and those on the right were vertical, thus defining a texture boundary. We independently jittered the orientations of bars by drawing random numbers from a uniform distribution; across trials, we systematically increased jitter from 0° (i.e., no jitter) to ±90° (i.e., no boundary). Blindfolded participants (n = 25) used the preferred index finger pad to actively scan objects for 10 seconds before reporting the texture boundary's orientation (vertical or horizontal; randomised across trials). Results showed a threshold jitter of ±52.7° (i.e., the jitter at which the boundary orientation was only just discriminable). Computational modelling indicated that the first stage of the FRF model is a Gabor function tuned to spatial frequency = 0.23 cycles per mm with extent = 2.53 mm (full-width at half-maximum). We discuss this result with regard to neuronal receptive field structure in non-human primate somatosensory cortex.

Support-vector classification of low-dose nitrous oxide administration with multi-channel EEG power spectra.

Support-vector machines (SVMs) can potentially improve patient monitoring during nitrous oxide anaesthesia. By elucidating the effects of low-dose nitrous oxide on the power spectra of multi-channel EEG recordings, we quantified the degree to which these effects generalise across participants. In this single-blind, cross-over study, 32-channel EEG was recorded from 12 healthy participants exposed to 0, 20, 30 and 40% end-tidal nitrous oxide. Features of the delta-, theta-, alpha- and beta-band power were used within a 12-fold, participant-wise cross-validation framework to train and test two SVMs: (1) binary SVM classifying EEG during 0 or 40% exposure (chance = 50%); (2) multi-class SVM classifying EEG during 0, 20, 30 or 40% exposure (chance = 25%). Both the binary (accuracy 92%) and the multi-class (accuracy 52%) SVMs classified EEG recordings at rates significantly better than chance (p < 0.001 and p = 0.01, respectively). To determine the relative importance of frequency band features for classification accuracy, we systematically removed features before re-training and re-testing the SVMs. This showed the relative importance of decreased delta power and the frontal region. SVM classification identified that the most important effects of nitrous oxide were found in the delta band in the frontal electrodes that was consistent between participants. Furthermore, support-vector classification of nitrous oxide dosage is a promising method that might be used to improve patient monitoring during nitrous oxide anaesthesia.

Detection of sleep apnea from single-channel electroencephalogram (EEG) using an explainable convolutional neural network (CNN).

Sleep apnea (SA) is a common disorder involving the cessation of breathing during sleep. It can cause daytime hypersomnia, accidents, and, if allowed to progress, serious, chronic conditions. Continuous positive airway pressure is an effective SA treatment. However, long waitlists impede timely diagnosis; overnight sleep studies involve trained technicians scoring a polysomnograph, which comprises multiple physiological signals including multi-channel electroencephalography (EEG). Therefore, it is important to develop simplified and automated approaches to detect SA. In the present study, we have developed an explainable convolutional neural network (CNN) to detect SA events from single-channel EEG recordings which generalizes across subjects. The network architecture consisted of three convolutional layers. We tuned hyperparameters using the Hyperband algorithm, optimized parameters using Adam, and quantified network performance with subjectwise 10-fold cross-validation. Our CNN performed with an accuracy of 69.9%, and a Matthews correlation coefficient (MCC) of 0.38. To explain the mechanisms of our trained network, we used critical-band masking (CBM): after training, we added bandlimited noise to test recordings; we parametrically varied the noise band center frequency and noise intensity, quantifying the deleterious effect on performance. We reconciled the effects of CBM with lesioning, wherein we zeroed the trained network's 1st-layer filter kernels in turn, quantifying the deleterious effect on performance. These analyses indicated that the network learned frequency-band information consistent with known SA biomarkers, specifically, delta and beta band activity. Our results indicate single-channel EEG may have clinical potential for SA diagnosis.

Balanced, Orientation-Dependent Dichoptic Masking in Cortex of Visually Normal Humans Measured Using Electroencephalography (EEG).

In the human visual system, cerebral cortex combines left- and right-eye retinal inputs, enabling single, comfortable binocular vision. In visual cortex, the signals from each eye inhibit one another (interocular suppression). While this mechanism may be disrupted by e.g. traumatic brain injury, clinical assessments of interocular suppression are subjective, qualitative, and lack reliability. EEG is a potentially useful clinical tool for objective, quantitative assessment of binocular vision. In a cohort of normal participants, we measured occipital, visual evoked potentials (VEPs) in response to dichoptically-presented vertical and/or horizontal sine-wave gratings. Response amplitudes to orthogonal gratings were greater than that of parallel gratings, which were in turn greater than that of monocular gratings. Our results indicate that interocular suppression is (normally) balanced, orientation-tuned, and that suppression per se is reduced for orthogonal gratings. This objective measure of suppression may have application in clinical settings.

Towards the Classification of Error-Related Potentials using Riemannian Geometry.

The error-related potential (ErrP) is an event-related potential (ERP) evoked by an experimental participant's recognition of an error during task performance. ErrPs, originally described by cognitive psychologists, have been adopted for use in brain-computer interfaces (BCIs) for the detection and correction of errors, and the online refinement of decoding algorithms. Riemannian geometry-based feature extraction and classification is a new approach to BCI which shows good performance in a range of experimental paradigms, but has yet to be applied to the classification of ErrPs. Here, we describe an experiment that elicited ErrPs in seven normal participants performing a visual discrimination task. Audio feedback was provided on each trial. We used multi-channel electroencephalogram (EEG) recordings to classify ErrPs (success/failure), comparing a Riemannian geometry-based method to a traditional approach that computes time-point features. Overall, the Riemannian approach outperformed the traditional approach (78.2% versus 75.9% accuracy, p <0.05); this difference was statistically significant (p <0.05) in three of seven participants. These results indicate that the Riemannian approach better captured the features from feedback-elicited ErrPs, and may have application in BCI for error detection and correction.

Retinal Implantation of Electronic Vision Prostheses to Treat Retinitis Pigmentosa: A Systematic Review.

Purpose: Retinitis pigmentosa (RP) is a hereditary disease causing photoreceptor degeneration and permanent vision loss. Retinal implantation of a stimulating electrode array is a new treatment for RP, but quantification of its efficacy is the subject of ongoing work. This review evaluates vision-related outcomes resulting from retinal implantation in participants with RP. Methods: We searched MEDLINE and Embase for journal articles published since January 1, 2015. We selected articles describing studies of implanted participants that reported the postimplantation measurement of vision. We extracted study information including design, participants' residual vision, comparators, and assessed outcomes. To assess the risk of bias, we used signaling questions and a target trial. Results: Our search returned 425 abstracts. We reviewed the full text of 34 articles. We judged all studies to be at high risk of bias owing to the study design or experimental conduct. Regarding design, studies lacked the measures that typical clinical trials take to protect against bias (e.g., control groups and masking). Regarding experimental conduct, outcome measures were rarely comparable before and after implantation, and psychophysical methods were prone to bias (subjective, not forced choice, methods). The most common comparison found was between postimplantation visual function with the device powered off versus on. This comparison is at high risk of bias. Conclusions: There is a need for high-quality evidence of efficacy of retinal implantation to treat RP. Translational Relevance: For patients and clinicians to make informed choices about RP treatment, visual function restored by retinal implantation must be properly quantified and reported.

Liquid-Crystal Display (LCD) of achromatic, mean-modulated flicker in clinical assessment and experimental studies of visual systems.

Achromatic, mean-modulated flicker-wherein luminance increments and decrements of equal magnitude are applied, over time, to a test field-is commonly used in both clinical assessment of vision and experimental studies of visual systems. However, presenting flicker on computer-controlled displays is problematic; displays typically introduce luminance artifacts at high flicker frequency or contrast, potentially interfering with the validity of findings. Here, we present a battery of tests used to weigh the relative merits of two displays for presenting achromatic, mean-modulated flicker. These tests revealed marked differences between a new high-performance liquid-crystal display (LCD; EIZO ColorEdge CG247X) and a new consumer-grade LCD (Dell U2415b), despite displays' vendor-supplied specifications being almost identical. We measured displayed luminance using a spot meter and a linearized photodiode. We derived several measures, including spatial uniformity, the effect of viewing angle, response times, Fourier amplitude spectra, and cycle-averaged luminance. We presented paired luminance pulses to quantify the displays' nonlinear dynamics. The CG247X showed relatively good spatial uniformity (e.g., at moderate luminance, standard deviation 2.8% versus U2415b's 5.3%). Fourier transformation of nominally static test patches revealed spectra free of artifacts, with the exception of a frame response. The CG247X's rise and fall times depended on both the luminance from which, and to which, it responded, as is to be generally expected from LCDs. Despite this nonlinear behaviour, we were able to define a contrast and frequency range wherein the CG247X appeared largely artifact-free; the relationship between nominal luminance and displayed luminance was accurately modelled using a causal, linear time-invariant system. This range included contrasts up to 80%, and flicker frequencies up to 30 Hz. This battery of tests should prove useful to others conducting clinical assessment of vision and experimental studies of visual systems.

Low-cost, microcontroller-based, two-channel piezoelectric bender device for somatosensory experiments.

Understanding the joint encoding of multiple tactile stimulus features (e.g., spatial position, amplitude, and frequency of vibration) is a major goal of somatosensory neuroscience, and the development of experimental set-ups to probe joint encoding is important. We describe in detail a microcontroller-based, piezoelectric bender device for tactile experiments. The device comprises an Arduino Due microcontroller board with a 32-bit ARM Cortex-M3 RISC processor, and two 12-bit digital-to-analog converters, enabling precise, independent stimulation of adjacent epithelial points. Using laser doppler vibrometry, we developed a model of the benders' structural mechanics, which we implemented on the device. We used the device to delivered precise, reliable somatosensory stimulation in an experimental setting, recording electrophysiological responses in the peripheral nervous system of the Gisborne cockroach (Drymaplaneta semivitta) to sinusoidal vibration of tibial spines. We plotted tuning curves and derived bandwidths of multi-unit populations. We also stimulated rat facial vibrissae ex vivo. This microcontroller-based, low-cost, open-source system leverages a large developer community associated with Arduino, and may help speed advances in systems neuroscience.

Laminar Differences in Responses to Naturalistic Texture in Macaque V1 and V2.

Most single units recorded from macaque secondary visual cortex (V2) respond with higher firing rates to synthetic texture images containing "naturalistic" higher-order statistics than to spectrally matched "noise" images lacking these statistics. In contrast, few single units in V1 show this property. We explored how the strength and dynamics of response vary across the different layers of visual cortex by recording multiunit (defined as high-frequency power in the local field potential) and gamma-band activity evoked by brief presentations of naturalistic and noise images in V1 and V2 of anesthetized macaque monkeys of both sexes. As previously reported, recordings in V2 showed consistently stronger responses to naturalistic texture than to spectrally matched noise. In contrast to single-unit recordings, V1 multiunit activity showed a preference for images with naturalistic statistics, and in gamma-band activity this preference was comparable across V1 and V2. Sensitivity to naturalistic image structure was strongest in the supragranular and infragranular layers of V1, but weak in granular layers, suggesting that it might reflect feedback from V2. Response timing was consistent with this idea. Visual responses appeared first in V1, followed by V2. Sensitivity to naturalistic texture emerged first in V2, followed by the supragranular and infragranular layers of V1, and finally in the granular layers of V1. Our results demonstrate laminar differences in the encoding of higher-order statistics of natural texture, and suggest that this sensitivity first arises in V2 and is fed back to modulate activity in V1.SIGNIFICANCE STATEMENT The circuit mechanisms responsible for visual representations of intermediate complexity are largely unknown. We used a well validated set of synthetic texture stimuli to probe the temporal and laminar profile of sensitivity to the higher-order statistical structure of natural images. We found that this sensitivity emerges first and most strongly in V2 but soon after in V1. However, sensitivity in V1 is higher in the laminae (extragranular) and recording modalities (local field potential) most likely affected by V2 connections, suggesting a feedback origin. Our results show how sensitivity to naturalistic image structure emerges across time and circuitry in the early visual cortex.

Asymmetric Dichoptic Masking in Visual Cortex of Amblyopic Macaque Monkeys.

In amblyopia, abnormal visual experience leads to an extreme form of eye dominance, in which vision through the nondominant eye is degraded. A key aspect of this disorder is perceptual suppression: the image seen by the stronger eye often dominates during binocular viewing, blocking the image of the weaker eye from reaching awareness. Interocular suppression is the focus of ongoing work aimed at understanding and treating amblyopia, yet its physiological basis remains unknown. We measured binocular interactions in visual cortex of anesthetized amblyopic monkeys (female Macaca nemestrina), using 96-channel "Utah" arrays to record from populations of neurons in V1 and V2. In an experiment reported recently (Hallum et al., 2017), we found that reduced excitatory input from the amblyopic eye (AE) revealed a form of balanced binocular suppression that is unaltered in amblyopia. Here, we report on the modulation of the gain of excitatory signals from the AE by signals from its dominant fellow eye (FE). Using a dichoptic masking technique, we found that AE responses to grating stimuli were attenuated by the presentation of a noise mask to the FE, as in a normal control animal. Responses to FE stimuli, by contrast, could not be masked from the AE. We conclude that a weakened ability of the amblyopic eye to modulate cortical response gain creates an imbalance of suppression that favors the dominant eye.SIGNIFICANCE STATEMENT In amblyopia, vision in one eye is impaired as a result of abnormal early visual experience. Behavioral observations in humans with amblyopia suggest that much of their visual loss is due to active suppression of their amblyopic eye. Here we describe experiments in which we studied binocular interactions in macaques with experimentally induced amblyopia. In normal monkeys, the gain of neuronal response to stimulation of one eye is modulated by contrast in the other eye, but in monkeys with amblyopia the balance of gain modulation is altered so that the weaker, amblyopic eye has little effect while the stronger fellow eye has a strong effect. This asymmetric suppression may be a key component of the perceptual losses in amblyopia.

Altered Balance of Receptive Field Excitation and Suppression in Visual Cortex of Amblyopic Macaque Monkeys.

In amblyopia, a visual disorder caused by abnormal visual experience during development, the amblyopic eye (AE) loses visual sensitivity whereas the fellow eye (FE) is largely unaffected. Binocular vision in amblyopes is often disrupted by interocular suppression. We used 96-electrode arrays to record neurons and neuronal groups in areas V1 and V2 of six female macaque monkeys (Macaca nemestrina) made amblyopic by artificial strabismus or anisometropia in early life, as well as two visually normal female controls. To measure suppressive binocular interactions directly, we recorded neuronal responses to dichoptic stimulation. We stimulated both eyes simultaneously with large sinusoidal gratings, controlling their contrast independently with raised-cosine modulators of different orientations and spatial frequencies. We modeled each eye's receptive field at each cortical site using a difference of Gaussian envelopes and derived estimates of the strength of central excitation and surround suppression. We used these estimates to calculate ocular dominance separately for excitation and suppression. Excitatory drive from the FE dominated amblyopic visual cortex, especially in more severe amblyopes, but suppression from both the FE and AEs was prevalent in all animals. This imbalance created strong interocular suppression in deep amblyopes: increasing contrast in the AE decreased responses at binocular cortical sites. These response patterns reveal mechanisms that likely contribute to the interocular suppression that disrupts vision in amblyopes.SIGNIFICANCE STATEMENT Amblyopia is a developmental visual disorder that alters both monocular vision and binocular interaction. Using microelectrode arrays, we examined binocular interaction in primary visual cortex and V2 of six amblyopic macaque monkeys (Macaca nemestrina) and two visually normal controls. By stimulating the eyes dichoptically, we showed that, in amblyopic cortex, the binocular combination of signals is altered. The excitatory influence of the two eyes is imbalanced to a degree that can be predicted from the severity of amblyopia, whereas suppression from both eyes is prevalent in all animals. This altered balance of excitation and suppression reflects mechanisms that may contribute to the interocular perceptual suppression that disrupts vision in amblyopes.

Population representation of visual information in areas V1 and V2 of amblyopic macaques.

Amblyopia is a developmental disorder resulting in poor vision in one eye. The mechanism by which input to the affected eye is prevented from reaching the level of awareness remains poorly understood. We recorded simultaneously from large populations of neurons in the supragranular layers of areas V1 and V2 in 6 macaques that were made amblyopic by rearing with artificial strabismus or anisometropia, and 1 normally reared control. In agreement with previous reports, we found that cortical neuronal signals driven through the amblyopic eyes were reduced, and that cortical neurons were on average more strongly driven by the non-amblyopic than by the amblyopic eyes. We analyzed multiunit recordings using standard population decoding methods, and found that visual signals from the amblyopic eye, while weakened, were not degraded enough to explain the behavioral deficits. Thus additional losses must arise in downstream processing. We tested the idea that under monocular viewing conditions, only signals from neurons dominated by - rather than driven by - the open eye might be used. This reduces the proportion of neuronal signals available from the amblyopic eye, and amplifies the interocular difference observed at the level of single neurons. We conclude that amblyopia might arise in part from degradation in the neuronal signals from the amblyopic eye, and in part from a reduction in the number of signals processed by downstream areas.

Surround suppression supports second-order feature encoding by macaque V1 and V2 neurons.

Single neurons in areas V1 and V2 of macaque visual cortex respond selectively to luminance-modulated stimuli. These responses are often influenced by context, for example when stimuli extend outside the classical receptive field (CRF). These contextual phenomena, observed in many sensory areas, reflect a fundamental cortical computation and may inform perception by signaling second-order visual features which are defined by spatial relationships of contrast, orientation and spatial frequency. In the anesthetized, paralyzed macaque, we measured single-unit responses to a drifting preferred sinusoidal grating; low spatial frequency sinusoidal contrast modulations were applied to the grating, creating contrast-modulated, second-order forms. Most neurons responded selectively to the orientation of the contrast modulation of the preferred grating and were therefore second-order orientation-selective. Second-order selectivity was created by the asymmetric spatial organization of the excitatory CRF and suppressive extraclassical surround. We modeled these receptive field subregions using spatial Gaussians, sensitive to the modulation of contrast (not luminance) of the preferred carrier grating, that summed linearly and were capable of recovering asymmetrical receptive field organizations. Our modeling suggests that second-order selectivity arises both from elongated excitatory CRFs, asymmetrically organized extraclassical surround suppression, or both. We validated the model by successfully testing its predictions against conventional surround suppression measurements and spike-triggered analysis of second-order form responses. Psychophysical adaptation measurements on human observers revealed a pattern of second-order form selectivity consistent with neural response patterns. We therefore propose that cortical cells in primates do double duty, providing signals about both first- and second-order forms.

Human primary visual cortex (V1) is selective for second-order spatial frequency.

A variety of cues can differentiate objects from their surrounds. These include "first-order" cues such as luminance modulations and "second-order" cues involving modulations of orientation and contrast. Human sensitivity to first-order modulations is well described by a computational model involving spatially localized filters that are selective for orientation and spatial frequency (SF). It is widely held that first-order modulations are represented by the firing rates of simple and complex cells ("first-order" neurons) in primary visual cortex (V1) that, likewise, have spatially localized receptive fields that are selective for orientation- and SF. Human sensitivity to second-order modulations is well described by a filter-rectify-filter (FRF) model, with first- and second-order filters selective for orientation and SF. However, little is known about how neuronal activity in visual cortex represents second-order modulations. We tested the FRF model by using an functional (f)MRI-adaptation protocol to characterize the selectivity of activity in visual cortex to second-order, orientation-defined gratings of two different SFs. fMRI responses throughout early visual cortex exhibited selective adaptation to these stimuli. The low-SF grating was a more effective adapter than the high-SF grating, incompatible with the FRF model. To explain the results, we extended the FRF model by incorporating normalization, yielding a filter-rectify-normalize-filter model, in which normalization enhances selectivity for second-order SF but only for low spatial frequencies. We conclude that neurons in human visual cortex are selective for second-order SF, that normalization (surround suppression) contributes to this selectivity, and that the selectivity in higher visual areas is simply fed forward from V1.

Optical imaging of electrically evoked visual signals in cats: I. Responses to corneal and intravitreal electrical stimulation.

Microelectronic retinal prostheses have been shown to restore the perception of light to the blind through electrical stimulation. Conventional recording techniques such as recording electrode arrays on the visual cortex can give a basic understanding of the events that occur during such stimulation events, but their finite size and number limits the spatial resolution achievable with them. Optical imaging of intrinsic signals (OIS imaging) allows for greater resolution (approximately 50 microm) of the activity in the cortex. This can be used to facilitate a greater understanding of the complex neurophysiological events that allow prosthetic vision. This paper shows responses to visual and electrical stimulation of the retina, and demonstrates that OIS imaging may be an effective technique in further refining stimulation techniques and implant designs for retinal prostheses.

Optical imaging of electrically evoked visual signals in cats: II. ICA "harmonic filtering" noise reduction.

Optical imaging of intrinsic signals (OIS) is a tool for visualizing differential areas in the primary visual cortex devoted to visual functions such as ocular dominance and spatial orientation preferences. The OIS methodology was employed to verify visual cortical response to a retinal vision prosthesis whereby electrical stimulation is applied to the neural retina in order to elicit visual percepts. However, OIS recording is quite susceptible to cardiac and respiratory artifact, and inherent noise related to the measurement process. This complicates the identification of evoked signals using standard ensemble averaging based image processing. We therefore developed an independent component analysis (ICA) "harmonic filtering" technique to improve the signal-to-noise ratio. This technique is capable of reducing noise components, highlighting response signals to visual and electrical stimuli. Particularly, we demonstrated extraction of an ocular dominance map due to corneal stimulation and localized cortical activation due to intravitreal stimulation.

Retinal neurostimulator for a multifocal vision prosthesis.

A neurostimulator application-specific integrated circuit (ASIC) with scalable circuitry that can stimulate 14 channels, has been developed for an epi-retinal vision prosthesis. This ASIC was designed to allow seven identical units to be connected to control up to 98 channels, with the ability to stimulate 14 electrodes simultaneously. The neurostimulator forms part of a vision prosthesis, designed to restore vision to patients who have lost their sight due to retinal diseases such as retinitis pigmentosa and macular degeneration. For charge balance, the neurostimulator was designed to stimulate with current sources and sinks operating together, and with the ability to drive a hexagonal mosaic of electrodes to reduce the electrical crosstalk that occurs when multiple bipolar stimulation sites are active simultaneously. A hexagonal mosaic of electrodes surrounds each stimulation site and has been shown to effectively isolate each site, increasing the ability to inject localized independent charge into multiple regions simultaneously.

Psychophysics of prosthetic vision: III. stochastic rendering, the phosphene image, and perception.

For pt.I see ibid., p.Z004024-7. For pt.II see ibid., p.Z004558-61. This paper examines the rendering of luminous spots ("phosphenes") in the visual field, and their stochastic positioning as a means of anti-aliasing the resulting spotty image ("phosphene image"). We derive an equation concerning the correlations of pairs of phosphenes comprising the phosphene image, and show the relationship to the statistics governing the stochastic positioning. We present some examples where stochastic rendering assists the veridical perception of textures, and argue for its superiority as cf. ordered rendering. Our preliminary results suggest that it may be perceptually effective to manufacture disordered arrays of stimulating electrodes for intraocular implantation.

Psychophysics of prosthetic vision: II. stochastic sampling, the phosphene image, and noise.

Stimulation of the diseased retina via an intraocular electrode array is a proposed means of restoring some vision to the profoundly blind. A prosthetic device to this end would involve post-implantation calibration (analogous to cochlear implant fitting), wherein the subject indicates those discrete positions in the visual field where luminous percepts are elicited. This procedure would be a source of noise, because the indicated positions would only approximate the actual positions in the visual field. Put differently, the procedure introduces sampling jitter, and would therefore affect clinical outcomes such as mobility and reading speeds. The nature of this noise is the concern of the present paper; we derive an expression for the noise power spectrum as it relates to the statistical nature of the sampling jitter. We show that, generally, jitter has greater effect on higher spatial-frequencies, that is, those areas of the implantee's visual perception that represent fine detail are more prone to noise. More specifically, the noise spectrum depends on the characteristic function of the random variable describing the sampling jitter. Our results signal the need for experimental work that characterizes sampling jitter in implantees, plus the need for simulations that allow a better understanding of perception and the noisy phosphene image.