High Spatial Frequency Gratings Research Articles

Retinal inputs that drive optomotor responses of mice under mesopic conditions

Optomotor responses are a popular way to assess sub-cortical visual responses in mice. We studied photoreceptor inputs into optomotor circuits using genetically-modified mice lacking the exocytotic calcium sensors synaptotagmin 1 (Syt1) and 7 (Syt7) in rods or cones. We also tested mice that in which cone transducin, GNAT2, had been eliminated. We studied spatial frequency sensitivity under mesopic conditions by varying the spatial frequency of a grating rotating at 12 deg/s and contrast sensitivity by varying luminance contrast of 0.2c/deg gratings. We found that eliminating Syt1 from rods reduced responses to a low spatial frequency grating (0.05c/deg) consistent with low resolution in this pathway. Conversely, eliminating the ability of cones to respond to light (by eliminating GNAT2) or transmit light responses (by selectively eliminating Syt1) showed weaker responses to a high spatial frequency grating (3c/deg). Eliminating Syt7 from the entire optomotor pathway in a global knockout had no significant effect on optomotor responses. We isolated the secondary rod pathway involving transmission of rod responses to cones via gap junctions by simultaneously eliminating Syt1 from rods and GNAT2 from cones. We found that the secondary rod pathway is sufficient to drive robust optomotor responses under mesopic conditions. Finally, eliminating Syt1 from both rods and cones almost completely abolished optomotor responses, but we detected weak responses to large, bright rotating gratings that are likely driven by input from intrinsically photosensitive retinal ganglion cells.

Association between retinal and cortical visual electrophysiological impairments in schizophrenia.

Electrophysiological impairments in the magnocellular visual system have been reported among patients with schizophrenia, but previous theories proposed that these deficits may begin in the retina. We therefore sought to evaluate the potential contribution of the retina by comparing retinal and cortical visual electrophysiological impairments between patients with schizophrenia and healthy controls. We recruited patients with schizophrenia and age- and sex-matched healthy controls. We recorded the P100 amplitude and latency using electroencephalography (EEG) while projecting low (0.5 cycles/degree) or high (15 cycles/degree) spatial frequency gratings at a temporal frequency of 0 Hz or 8 Hz. We compared the P100 results with previous results for retinal ganglion cell activity (N95) in these participants. We analyzed data using repeated-measures analysis of variance and correlation analyses. We recruited 21 patients with schizophrenia and 29 age- and sex-matched healthy controls. Results showed decreased P100 amplitude and increased P100 latency among patients with schizophrenia compared with healthy controls (p < 0.05). Analyses reported the main effects of spatial and temporal frequency but no interaction effects of spatial or temporal frequency by group. Moreover, correlation analysis indicated a positive association between P100 latency and previous retinal results for N95 latency in the schizophrenia group (p < 0.05). Alterations in the P100 wave among patients with schizophrenia are consistent with the deficits in early visual cortical processing shown in the literature. These deficits do not seem to correspond to an isolated magnocellular deficit but appear to be associated with previous retinal measurements. Such an association emphasizes the role of the retina in the occurrence of visual cortical abnormalities in schizophrenia. Studies with coupled electroretinography-EEG measurements are now required to further explore these findings. https://clinicaltrials.gov/ct2/show/NCT02864680.

Impaired P100 among regular cannabis users in response to magnocellular biased visual stimuli

Regular cannabis using causes vision impairment by affecting human retinal neurotransmission. However, studies less considered its impact on the subsequent visual cortical processing, key feature for the integration of the visual signal in brain. We aimed at investigating this purpose in regular cannabis users using spatial frequencies and temporal frequencies filtered visual stimuli.We recruited 45 regular cannabis users and 25 age-matched controls. We recorded visual evoked potentials during the projection of low spatial frequency (0.5 cycles/degree) or high spatial frequency gratings (15 cycles/degree), which were presented statically (0 Hz) or dynamically (8 Hz). We analyzed the amplitude, latency, and area under the curve of both P100 and N170, best EEG markers for early visual processing. Data were compared between groups by repeated measures ANCOVA.Results showed a significant decrease in P100 amplitude among regular cannabis users in low spatial frequency (F(1,67) = 4.43; p = 0.04) and in dynamic condition (F(1,67) = 4.35; p = 0.04). Analysis also reported a decrease in P100 area under the curve in regular cannabis users to low spatial frequency (F(1,67) = 4.31; p = 0.04) and in dynamic condition (F(1,67) = 7.65; p < 0.01). No effect was found on P100 latency, N170 amplitude, latency, or area under the curve.We found alteration of P100 responses to low spatial frequency and dynamic stimuli in regular cannabis users. This result could be interpreted as a preferential magnocellular impairment where such deficit could be linked to glutamatergic dysfunction. As mentioned in the literature, visual and electrophysiological anomalies in schizophrenia are related to a magnocellular dysfunction. Further studies are needed to clarify electrophysiological deficits in both populations. Clinical trials registrationElectrophysiological Study of the Functioning of Magnocellular Visual Pathway in Regular Cannabis Users (CAUSA MAP). [NCT 02864680; ID 2013-A00097–38]. https://clinicaltrials.gov/ct2/show/NCT02864680?cond=Cannabis&cntry=FR&draw=2&rank=1

Hemispheric Asymmetry in Visual Processing: An ERP Study on Spatial Frequency Gratings

A hemispheric asymmetry for the processing of global versus local visual information is known. In this study, we investigated the existence of a hemispheric asymmetry for the visual processing of low versus high spatial frequency gratings. The event-related potentials were recorded in a group of healthy right-handed volunteers from 30 scalp sites. Six types of stimuli (1.5, 3 and 6 c/deg gratings) were randomly flashed 180 times in the left and right upper hemifields. The stimulus duration was 80 ms, and the interstimulus interval (ISI) ranged between 850 and 1000 ms. Participants paid attention and responded to targets based on their spatial frequency and location. The C1 and P1 visual responses, as well as a later selection negativity and a P300 component of event-related potentials (ERPs), were quantified and subjected to repeated-measure analyses of variance (ANOVAs). Overall, the performance was faster for the right visual field (RVF), thus suggesting a left hemispheric advantage for the attentional selection of local elements. Similarly, the analysis of the mean area amplitude of the C1 (60–110 ms) sensory response showed a stronger attentional effect (F+L+ vs. F−L+) at the left occipital areas, thus suggesting the sensory nature of this hemispheric asymmetry.

Unconscious processing of coarse visual information during anticipatory threat.

Rapid detection of threats has been proposed to rely on automatic processing of their coarse visual features. However, it remains unclear whether such a mechanism is restricted to detection of threat cues, or whether it reflects a broader sensitivity to even neutral coarse visual information features during states of threat. We used a backward masking task in which participants discriminated the orientation of subliminally presented low (3 cpd) and high (6 cpd) spatial frequency gratings, under threat (of shock) and safe conditions. Visual awareness of the gratings was assessed objectively using an additional localization task. When participants were unaware of the gratings, above chance and improved discrimination of low-spatial frequency gratings was observed under threat compared to safe trials. These findings demonstrate unconscious processing of neutral coarse visual information during threat state, supporting the view that automatic threat detection may rely on a general facilitation of coarse features irrespective of threat content.

Visuocortical changes during a freezing-like state in humans

An adaptive response to threat requires optimized detection of critical sensory cues. This optimization is thought to be aided by freezing - an evolutionarily preserved defensive state of immobility characterized by parasympathetically mediated fear bradycardia and regulated by the amygdala-periaqueductal grey (PAG) circuit. Behavioral observations in humans and animals have suggested that freezing is also a state of enhanced visual sensitivity, particularly for coarse visual information, but the underlying neural mechanisms remain unclear. We induced a freezing-like state in healthy volunteers using threat of electrical shock and measured threat-related changes in both stimulus-independent (baseline) and stimulus-evoked visuocortical activity to low-vs. high-spatial frequency gratings, using functional MRI. As measuring immobility is not feasible in MRI environments, we used fear bradycardia and amygdala-PAG coupling in inferring a freezing-like state. An independent functional localizer and retinotopic mapping were used to assess the retinotopic specificity of visuocortical modulations. We found a threat-induced increase in baseline (stimulus-independent) visuocortical activity that was retinotopically nonspecific, which was accompanied by increased connectivity with the amygdala. A positive correlation between visuocortical activity and fear bradycardia (while controlling for sympathetic activation), and a concomitant increase in amygdala-PAG connectivity, confirmed the specificity of these findings for the parasympathetically dominated freezing-like state. Visuocortical responses to gratings were retinotopically specific, but did not differ between threat and safe conditions across participants. However, individuals who exhibited better discrimination of low-spatial frequency stimuli showed reduced stimulus-evoked V1 responses under threat. Our findings suggest that a defensive state of freezing involves an integration of preparatory defensive and perceptual changes which may be regulated by a common mechanism involving the amygdala.

Color weight photometry

We study the “color weight” for a number of rather different paradigms. In well researched heterochromatic photometry methods we find that the “weights” determined by settings of naive observers are closely determined by the CIE luminance functional. This is very different for tasks that involve mid- and high-level aspects of perception. In several cases we find equipollence for the display red, green and blue channels. Moreover, in such cases the very nonlinear maximum-rule fits the data rather better than a linear functional. These findings are of interest when photometry needs to be applied for stimuli that are different from the high temporal and low spatial frequency gratings typical for flicker photometry. These results are relevant for science, ergonomics and art.

Neuronal basis of perceptual learning in striate cortex.

It is well known that, in humans, contrast sensitivity training at high spatial frequency (SF) not only leads to contrast sensitivity improvement, but also results in an improvement in visual acuity as assessed with gratings (direct effect) or letters (transfer effect). However, the underlying neural mechanisms of this high spatial frequency training improvement remain to be elucidated. In the present study, we examined four properties of neurons in primary visual cortex (area 17) of adult cats that exhibited significantly improved acuity after contrast sensitivity training with a high spatial frequency grating and those of untrained control cats. We found no difference in neuronal contrast sensitivity or tuning width (Width) between the trained and untrained cats. However, the trained cats showed a displacement of the cells’ optimal spatial frequency (OSF) to higher spatial frequencies as well as a larger neuronal signal-to-noise ratio (SNR). Furthermore, both the neuronal differences in OSF and SNR were significantly correlated with the improvement of acuity measured behaviorally. These results suggest that striate neurons might mediate the perceptual learning-induced improvement for high spatial frequency stimuli by an alteration in their spatial frequency representation and by an increased SNR.

Greater magnocellular saccadic suppression in high versus low autistic tendency suggests a causal path to local perceptual style.

Saccadic suppression—the reduction of visual sensitivity during rapid eye movements—has previously been proposed to reflect a specific suppression of the magnocellular visual system, with the initial neural site of that suppression at or prior to afferent visual information reaching striate cortex. Dysfunction in the magnocellular visual pathway has also been associated with perceptual and physiological anomalies in individuals with autism spectrum disorder or high autistic tendency, leading us to question whether saccadic suppression is altered in the broader autism phenotype. Here we show that individuals with high autistic tendency show greater saccadic suppression of low versus high spatial frequency gratings while those with low autistic tendency do not. In addition, those with high but not low autism spectrum quotient (AQ) demonstrated pre-cortical (35–45 ms) evoked potential differences (saccade versus fixation) to a large, low contrast, pseudo-randomly flashing bar. Both AQ groups showed similar differential visual evoked potential effects in later epochs (80–160 ms) at high contrast. Thus, the magnocellular theory of saccadic suppression appears untenable as a general description for the typically developing population. Our results also suggest that the bias towards local perceptual style reported in autism may be due to selective suppression of low spatial frequency information accompanying every saccadic eye movement.

Neural oscillatory deficits in schizophrenia predict behavioral and neurocognitive impairments.

Paying attention to visual stimuli is typically accompanied by event-related desynchronizations (ERD) of ongoing alpha (7–14 Hz) activity in visual cortex. The present study used time-frequency based analyses to investigate the role of impaired alpha ERD in visual processing deficits in schizophrenia (Sz). Subjects viewed sinusoidal gratings of high (HSF) and low (LSF) spatial frequency (SF) designed to test functioning of the parvo- vs. magnocellular pathways, respectively. Patients with Sz and healthy controls paid attention selectively to either the LSF or HSF gratings which were presented in random order. Event-related brain potentials (ERPs) were recorded to all stimuli. As in our previous study, it was found that Sz patients were selectively impaired at detecting LSF target stimuli and that ERP amplitudes to LSF stimuli were diminished, both for the early sensory-evoked components and for the attend minus unattend difference component (the Selection Negativity), which is generally regarded as a specific index of feature-selective attention. In the time-frequency domain, the differential ERP deficits to LSF stimuli were echoed in a virtually absent theta-band phase locked response to both unattended and attended LSF stimuli (along with relatively intact theta-band activity for HSF stimuli). In contrast to the theta-band evoked responses which were tightly stimulus locked, stimulus-induced desynchronizations of ongoing alpha activity were not tightly stimulus locked and were apparent only in induced power analyses. Sz patients were significantly impaired in the attention-related modulation of ongoing alpha activity for both HSF and LSF stimuli. These deficits correlated with patients’ behavioral deficits in visual information processing as well as with visually based neurocognitive deficits. These findings suggest an additional, pathway-independent, mechanism by which deficits in early visual processing contribute to overall cognitive impairment in Sz.

An early origin for detailed perception in Autism Spectrum Disorder: biased sensitivity for high-spatial frequency information.

Autistics demonstrate superior performances on several visuo-spatial tasks where local or detailed information processing is advantageous. Altered spatial filtering properties at an early level of visuo-spatial analysis may be a plausible perceptual origin for such detailed perception in Autism Spectrum Disorder. In this study, contrast sensitivity for both luminance and texture-defined vertically-oriented sine-wave gratings were measured across a range of spatial frequencies (0.5, 1, 2, 4 & 8 cpd) for autistics and non-autistic participants. Contrast sensitivity functions and peak frequency ratios were plotted and compared across groups. Results demonstrated that autistic participants were more sensitivity to luminance-defined, high spatial frequency gratings (8 cpd). A group difference in peak distribution was also observed as 35% of autistic participants manifested peak sensitivity for luminance-defined gratings of 4 cpd, compared to only 7% for the comparison group. These findings support that locally-biased perception in Autism Spectrum Disorder originates, at least in part, from differences in response properties of early spatial mechanisms favouring detailed spatial information processing.

Attending to global versus local stimulus features modulates neural processing of low versus high spatial frequencies: an analysis with event-related brain potentials.

Spatial frequency (SF) selection has long been recognized to play a role in global and local processing, though the nature of the relationship between SF processing and global/local perception is debated. Previous studies have shown that attention to relatively lower SFs facilitates global perception, and that attention to relatively higher SFs facilitates local perception. Here we recorded event-related brain potentials (ERPs) to investigate whether processing of low versus high SFs is modulated automatically during global and local perception, and to examine the time course of any such effects. Participants compared bilaterally presented hierarchical letter stimuli and attended to either the global or local levels. Irrelevant SF grating probes flashed at the center of the display 200 ms after the onset of the hierarchical letter stimuli could either be low or high in SF. It was found that ERPs elicited by the SF grating probes differed as a function of attended level (global versus local). ERPs elicited by low SF grating probes were more positive in the interval 196–236 ms during global than local attention, and this difference was greater over the right occipital scalp. In contrast, ERPs elicited by the high SF gratings were more positive in the interval 250–290 ms during local than global attention, and this difference was bilaterally distributed over the occipital scalp. These results indicate that directing attention to global versus local levels of a hierarchical display facilitates automatic perceptual processing of low versus high SFs, respectively, and this facilitation is not limited to the locations occupied by the hierarchical display. The relatively long latency of these attention-related ERP modulations suggests that initial (early) SF processing is not affected by attention to hierarchical level, lending support to theories positing a higher level mechanism to underlie the relationship between SF processing and global versus local perception.

Influence of spherical aberration, stimulus spatial frequency, and pupil apodisation on subjective refractions.

To test competing hypotheses (Stiles Crawford pupil apodising or superior imaging of high spatial frequencies by the central pupil) for the pupil size independence of subjective refractions in the presence of primary spherical aberration. Subjective refractions were obtained with a variety of test stimuli (high contrast letters, urban cityscape, high and low spatial frequency gratings) while modulating pupil diameter, levels of primary spherical aberration and pupil apodisation. Subjective refractions were also obtained with low-pass and high-pass stimuli and using 'darker' and 'sharper' subjective criteria. Subjective refractions for stimuli containing high spatial frequencies focus a near paraxial region of the pupil and are affected only slightly by level of Seidel spherical aberration, degree of pupil apodisation and pupil diameter, and generally focused a radius of about 1-1.5mm from the pupil centre. Low spatial frequency refractions focus a marginal region of the pupil, and are significantly affected by level of spherical aberration, amount of pupil apodisation, and pupil size. Clinical refractions that employ the 'darker' or 'sharper' subjective criteria bias the patient to use lower or higher spatial frequencies, respectively. In the presence of significant levels of spherical aberration, the pupil size independence of subjective refractions occurs with or without Stiles Crawford apodisation for refractions that optimise high spatial frequency content in the image. If low spatial frequencies are optimised by a subjective refraction, spherical refractive error varies with spherical aberration, pupil size, and level of apodisation. As light levels drop from photopic to scotopic, therefore, we expect a shift from pupil size independent to pupil size dependent subjective refractions. Emphasising a 'sharper' criterion during subjective refractions will improve image quality for high spatial frequencies and generate pupil size independent refractions.

Effects of stimulus spatial frequency, size, and luminance contrast on orientation tuning of neurons in the dorsal lateral geniculate nucleus of cat

It is generally thought that orientation selectivity first appears in the primary visual cortex (V1), whereas neurons in the lateral geniculate nucleus (LGN), an input source for V1, are thought to be insensitive to stimulus orientation. Here we show that increasing both the spatial frequency and size of the grating stimuli beyond their respective optimal values strongly enhance the orientation tuning of LGN neurons. The resulting orientation tuning was clearly contrast-invariant. Furthermore, blocking intrathalamic inhibition by iontophoretically administering γ-aminobutyric acid (GABA)A receptor antagonists, such as bicuculline and GABAzine, slightly but significantly weakened the contrast invariance. Our results suggest that orientation tuning in the LGN is caused by an elliptical classical receptive field and orientation-tuned surround suppression, and that its contrast invariance is ensured by local GABAA inhibition. This contrast-invariant orientation tuning in LGN neurons may contribute to the contrast-invariant orientation tuning seen in V1 neurons.

Objective and Subjective Refractive Error Measurements in Monkeys

To better understand the functional significance of refractive-error measures obtained using common objective methods in laboratory animals, we compared objective and subjective measures of refractive error in adolescent rhesus monkeys. The subjects were 20 adolescent monkeys. Spherical-equivalent spectacle-plane refractive corrections were measured by retinoscopy and autorefraction while the animals were cyclopleged and anesthetized. The eye's axial dimensions were measured by A-Scan ultrasonography. Subjective measures of the eye's refractive state, with and without cycloplegia, were obtained using psychophysical methods. Specifically, we measured spatial contrast sensitivity as a function of spectacle lens power for relatively high spatial frequency gratings. The lens power that produced the highest contrast sensitivity was taken as the subjective refraction. Retinoscopy and autorefraction consistently yielded higher amounts of hyperopia relative to subjective measurements obtained with or without cycloplegia. The subjective refractions were not affected by cycloplegia and on average were 1.42 ± 0.61 D and 1.24 ± 0.62 D less hyperopic than the retinoscopy and autorefraction measurements, respectively. Repeating the retinoscopy and subjective measurements through 3 mm artificial pupils produced similar differences. The results show that commonly used objective methods for assessing refractive errors in monkeys significantly overestimate the degree of hyperopia. It is likely that multiple factors contributed to the hyperopic bias associated with these objective measurements. However, the magnitude of the hyperopic bias was in general agreement with the "small-eye artifact" of retinoscopy.

Consequences of Magnocellular Dysfunction on Processing Attended Information in Schizophrenia

Schizophrenia is associated with perceptual and cognitive dysfunction including impairments in visual attention. These impairments may be related to deficits in early stages of sensory/perceptual processing, particularly within the magnocellular/dorsal visual pathway. In the present study, subjects viewed high and low spatial frequency (SF) gratings designed to test functioning of the parvocellular/magnocellular pathways, respectively. Schizophrenia patients and healthy controls attended to either the low SF (magnocellularly biased) or high SF (parvocellularly biased) gratings. Functional magnetic resonance imaging (fMRI) and recordings of event-related potentials (ERPs) were carried out during task performance. Patients were impaired at detecting low-frequency targets. ERP amplitudes to low-frequency gratings were diminished, both for the early sensory-evoked components and for the attend minus unattend difference component (the selection negativity), which is regarded as a neural index of feature-selective attention. Similarly, fMRI revealed that activity in extrastriate visual cortex was reduced in patients during attention to low, but not high, SF. In contrast, activity in frontal and parietal areas, previously implicated in the control of attention, did not differ between patients and controls. These findings suggest that impaired sensory processing of magnocellularly biased stimuli lead to impairments in the effective processing of attended stimuli, even when the attention control systems themselves are intact.

When and where is binocular rivalry resolved in the visual cortex?

Anatomically "early" visual areas including striate cortex (V1) are known to play a role in the resolution of binocular rivalry. However, the time course of such activity and its relationship with subjective perception are unclear. The present study used an intermittent stimulation design to measure event-related potentials (ERPs) associated with perception during binocular rivalry and during physical alternations of the same stimuli. When gratings of high or low spatial frequencies were presented in physical alternation, the amplitude of the earliest cortical ERP component (the C1, at 60-100 ms) was larger for high spatial frequencies. When the same two stimuli competed during rivalry, however, C1 amplitudes were equivalent for perceptions of high versus low spatial frequency. These findings suggest that rivalry is not resolved before or during the initial cortical response at 60-100 ms. At longer latencies (approximately 130-160 ms), however, occipital ERPs with similar topographies and estimated sources as the C1 differed according to perception in both the rivalry and physical alternation conditions. These results suggest that during rivalry, neural activity coupled with the dominant percept first emerges in anatomically early visual cortex, but at delayed latencies.

VEP contrast sensitivity responses reveal reduced functional segregation of mid and high filters of visual channels in Autism

Despite the vast amount of behavioral data showing a pronounced tendency in individuals with autism spectrum disorder (ASD) to process fine visual details, much less is known about the neurophysiological characteristics of spatial vision in ASD. Here, we address this issue by assessing the contrast sensitivity response properties of the early visual-evoked potentials (VEPs) to sine-wave gratings of low, medium and high spatial frequencies in adults with ASD and in an age- and IQ-matched control group. Our results show that while VEP contrast responses to low and high spatial frequency gratings did not differ between ASD and controls, early VEPs to mid spatial frequency gratings exhibited similar response characteristics as those to high spatial frequency gratings in ASD. Our findings show evidence for an altered functional segregation of early visual channels, especially those responsible for processing mid- and high-frequency spatial scales.

Differences in early sensory-perceptual processing in synesthesia: A visual evoked potential study

Synesthesia is a condition where stimulation of a single sensory modality or processing stream elicits an idiosyncratic, yet reliable perception in one or more other modalities or streams. Various models have been proposed to explain synesthesia, which have in common aberrant cross-activation of one cortical area by another. This has been observed directly in cases of linguistic-color synesthesia as cross-activation of the ‘color area’, V4, by stimulation of the grapheme area. The underlying neural substrates that mediate cross-activations in synesthesia are not well understood, however. In addition, the overall integrity of the visual system has never been assessed and it is not known whether wider differences in sensory-perceptual processing are associated with the condition. To assess whether fundamental differences in perceptual processing exist in synesthesia, we utilised high-density 128-channel electroencephalography (EEG) to measure sensory-perceptual processing using stimuli that differentially bias activation of the magnocellular and parvocellular pathways of the visual system. High and low spatial frequency gratings and luminance-contrast squares were presented to 15 synesthetes and 15 controls. We report, for the first time, early sensory-perceptual differences in synesthetes relative to non-synesthete controls in response to simple stimuli that do not elicit synesthetic color experiences. The differences are manifested in the early sensory components of the visual evoked potential (VEP) to stimuli that bias both magnocellular and parvocellular responses, but are opposite in direction, suggesting a differential effect on these two pathways. We discuss our results with reference to widespread connectivity differences as a broader phenotype of synesthesia.

Diffusion study in tailored gratings recorded in photopolymer glass with high refractive index species

We report results on the temporal evolution of the diffraction efficiency of volume holographic gratings recorded in a photopolymer glass incorporating Zr-based high refractive index species (HRIS) at molecular level. We record high spatial frequency gratings in this material with diffraction efficiencies near 100%. A two-component diffusion model is introduced for the evolution of refractive index modulation in darkness. Diffusion parameters for the Zr-based HRIS and monomer components have been determined. Codirectional diffusion of both components is demonstrated. The results show the feasibility for tailoring in this photomaterial holographic gratings with high diffraction efficiency over a wide range of spatial frequencies.