My research program centers around spatial vision and involves the use of behavioral, oculomotor, electrophysiological and functional MRI techniques in humans. My research has focused for many years on normal visual development as well as abnormal visual development in patients with strabismus, autism and cortical visual impairment. To inform this work, I also work with normal adults and animal models. I have published over 30 papers on the normal developmental process, the first in 1977. In addition, I have published over 30 papers on abnormal visual development over the last 20 years and am an expert in the diagnostic classification and pathophysiology of disorders of binocular vision and developmental disorders. I am thus in a position to translate basic science results into clinical applications. My laboratory has established numerous paradigms in which Visual Evoked Potentials have been used as objective measures of sensory and cognitive function. My group has published many papers in which we have related perceptual judgments to human brain activity. We have developed multiple generations of sophisticated and flexible instrumentation that we have shared with many other laboratories. Central to our research program is our unique integration of functional MRI and high-density EEG source imaging that allows us to relate brain electrical activity to identified visual areas with 1-2 cm accuracy in individual participants. The development of this technique involved the creation and management of a diverse team of experts in functional MRI, physics, biomedical engineering and visual physiology. I have completed training of 18 post-doctoral fellows, 14 of whom have gone on to faculty positions or full-time research positions. I am currently training three post-docs and three graduate students. I have served on 12 Ph.D. thesis committees at Stanford. As a Research Professor, I am able to devote most of my time to direct mentoring of my trainees.

Administrative Appointments

  • Committee on Research, Stanford University (2012 - 2015)

Honors & Awards

  • William A. Kettlewell Chair of Research in Visual Science, Smith-Kettlewell Eye Research Institute (1992)
  • Catherine Doyle Kettlewell Chair of Research in Visual Science, Smith-Kettlewell Eye Research Institute (2000)
  • Edridge Green Lecturer, Royal College of Ophthalmology, UK (2008)
  • Walt and Lilly Disney Award for Amblyopia, Research to Prevent Blindness (2008)
  • Trimble Medalist, British Isles Paediatric Ophthalmology Association (2016)
  • Inaugural Fellow, Association for Research in Vision and Ophthalmology (2009)

Boards, Advisory Committees, Professional Organizations

  • Director, Vision Sciences Society (2012 - 2016)
  • President, Vision Sciences Society (2016 - 2016)
  • Director, Vision4Children, UK (2010 - Present)

Professional Education

  • Ph.D, Stanford University, Physiological Psychology (1981)

Current Research and Scholarly Interests

Vision, development, functional imaging, systems analysis

2023-24 Courses

Stanford Advisees

Graduate and Fellowship Programs

All Publications

  • Poster Session I: Noninvasive neuromodulation of subcortical visual pathways with transcranial focused ultrasound. Journal of vision Ash, R., Mohammadjavadi, M., Butts Pauly, K., Norcia, A. 2023; 23 (15): 23


    Transcranial ultrasound stimulation (TUS) is an emerging tool to noninvasively modulate neural activity in deep brain areas. In preparation for our first in-human TUS studies, we targeted TUS to the lateral geniculate nucleus (visual thalamus) in a large mammal (sheep). Full-field light flash stimuli were presented with or without concomitant TUS in randomly interleaved trials. Similar to what has previously observed by Fry et al (Nature 1959) in cats, EEG visual-evoked potentials (VEPs) were reversibly suppressed by TUS to the LGN. No changes in VEPs were observed in sheep who received sham-TUS to a control site in the basal ganglia, ruling out potential transducer auditory-somatosensory confounds. Magnetic resonance acoustic radiation force imaging (MR-ARFI), a technique to measure the ultrasound focus in situ, showed a focal volume of microscopic displacement at the expected target. Excitingly, MR-ARFI predicted the suppressive effect on VEPs in individual subjects, suggesting that MR-ARFI can be used to confirm TUS targeting and estimate neurophysiological impact. We are now translating this paradigm into human, targeting TUS to the LGN while participants perform a contrast detection task with EEG recording of steady-state VEPs. MR-ARFI will be measured to evaluate targeting and estimate TUS dosage in each participant. This work provides the foundation for a dissection of the roles of different subcortical nuclei in different aspects of human vision.

    View details for DOI 10.1167/jov.23.15.23

    View details for PubMedID 38109625

  • Mouse visual cortex as a limited resource system that self-learns an ecologically-general representation. PLoS computational biology Nayebi, A., Kong, N. C., Zhuang, C., Gardner, J. L., Norcia, A. M., Yamins, D. L. 2023; 19 (10): e1011506


    Studies of the mouse visual system have revealed a variety of visual brain areas that are thought to support a multitude of behavioral capacities, ranging from stimulus-reward associations, to goal-directed navigation, and object-centric discriminations. However, an overall understanding of the mouse's visual cortex, and how it supports a range of behaviors, remains unknown. Here, we take a computational approach to help address these questions, providing a high-fidelity quantitative model of mouse visual cortex and identifying key structural and functional principles underlying that model's success. Structurally, we find that a comparatively shallow network structure with a low-resolution input is optimal for modeling mouse visual cortex. Our main finding is functional-that models trained with task-agnostic, self-supervised objective functions based on the concept of contrastive embeddings are much better matches to mouse cortex, than models trained on supervised objectives or alternative self-supervised methods. This result is very much unlike in primates where prior work showed that the two were roughly equivalent, naturally leading us to ask the question of why these self-supervised objectives are better matches than supervised ones in mouse. To this end, we show that the self-supervised, contrastive objective builds a general-purpose visual representation that enables the system to achieve better transfer on out-of-distribution visual scene understanding and reward-based navigation tasks. Our results suggest that mouse visual cortex is a low-resolution, shallow network that makes best use of the mouse's limited resources to create a light-weight, general-purpose visual system-in contrast to the deep, high-resolution, and more categorization-dominated visual system of primates.

    View details for DOI 10.1371/journal.pcbi.1011506

    View details for PubMedID 37782673

  • Stability of steady-state visual evoked potential contrast response functions. Psychophysiology Ash, R. T., Nix, K. C., Norcia, A. M. 2023: e14412


    Repetitive sensory stimulation has been shown to induce neuroplasticity in sensory cortical circuits, at least under certain conditions. We measured the plasticity-inducing effect of repetitive contrast-reversal-sweep steady-state visual-evoked potential (ssVEP) stimuli, hoping to employ the ssVEP's high signal-to-noise electrophysiological readout in the study of human visual cortical neuroplasticity. Steady-state VEP contrast-sweep responses were measured daily for 4 days (four 20-trial blocks per day, 20 participants). No significant neuroplastic changes in response amplitude were observed either across blocks or across days. Furthermore, response amplitudes were stable within-participant, with measured across-block and across-day coefficients of variation (CV = SD/mean) of 15-20 ± 2% and 22-25 ± 2%, respectively. Steady-state VEP response phase was also highly stable, suggesting that temporal processing delays in the visual system vary by at most 2-3 ms across blocks and days. While we fail to replicate visual stimulation-dependent cortical plasticity, we show that contrast-sweep steady-state VEPs provide a stable human neurophysiological measure well suited for repeated-measures longitudinal studies.

    View details for DOI 10.1111/psyp.14412

    View details for PubMedID 37614220

  • Progress in elementary school reading linked to growth of cortical responses to familiar letter combinations within visual word forms. Developmental science Wang, F., Kaneshiro, B., Toomarian, E. Y., Gosavi, R. S., Hasak, L. R., Moron, S., Nguyen, Q. T., Norcia, A. M., McCandliss, B. D. 2023: e13435


    Learning to read depends on the ability to extract precise details of letter combinations, which convey critical information that distinguishes tens of thousands of visual word forms. To support fluent reading skill, one crucial neural developmental process is one's brain sensitivity to statistical constraints inherent in combining letters into visual word forms. To test this idea in early readers, we tracked the impact of two years of schooling on within-subject longitudinal changes in cortical responses to three different properties of words: coarse tuning for print, and fine tuning to either familiar letter combinations within visual word forms or whole word representations. We then examined how each related to growth in reading skill. Three stimulus contrasts-words versus pseudofonts, words versus pseudowords, pseudowords versus nonwords-were presented while high-density EEG Steady-State Visual Evoked Potentials (SSVEPs, n = 31) were recorded. Internalization of abstract visual word form structures over two years of reading experience resulted in a near doubling of SSVEP amplitude, with increasing left lateralization. Longitudinal changes (decreases) in brain responses to such word form structural information were linked to the growth in reading skills, especially in rapid automatic naming of letters. No such changes were observed for whole word representation processing and coarse tuning for print. Collectively, these findings indicate that sensitivity to visual word form structure develops rapidly through exposure to print and is linked to growth in reading skill. RESEARCH HIGHLIGHTS: Longitudinal changes in cognitive responses to coarse print tuning, visual word from structure, and whole word representation were examined in early readers. Visual word form structure processing demonstrated striking patterns of growth with nearly doubled in EEG amplitude and increased left lateralization. Longitudinal changes (decreases) in brain responses to visual word form structural information were linked to the growth in rapid automatic naming for letters. No longitudinal changes were observed for whole word representation processing and coarse tuning for print.

    View details for DOI 10.1111/desc.13435

    View details for PubMedID 37465984

  • Different contrast response functions for sawtooth stimulation favoring ON vs OFF visual pathways in the upper and lower visual fields Scott, M., Yakovleva, A., Norcia, A. ASSOC RESEARCH VISION OPHTHALMOLOGY INC. 2023
  • Lexical and sublexical cortical tuning for print revealed by Steady-State Visual Evoked Potentials (SSVEPs) in early readers. Developmental science Wang, F., Nguyen, Q. T., Kaneshiro, B., Hasak, L., Wang, A. M., Toomarian, E. Y., Norcia, A. M., McCandliss, B. D. 2022: e13352


    There are multiple levels of processing relevant to reading that vary in their visual, sublexical and lexical orthographic processing demands. Segregating distinct cortical sources for each of these levels has been challenging in EEG studies of early readers. To address this challenge, we applied recent advances in analyzing high-density EEG using Steady-State Visual Evoked Potentials (SSVEPs) via data-driven Reliable Components Analysis (RCA) in a group of early readers spanning from kindergarten to second grade. Three controlled stimulus contrasts-familiar words versus unfamiliar pseudofonts, familiar words versus pseudowords, and pseudowords versus nonwords-were used to isolate coarse print tuning, lexical processing, and sublexical orthography-related processing, respectively. First, three overlapping yet distinct neural sources-left vOT, dorsal parietal, and primary visual cortex were revealed underlying coarse print tuning. Second, we segregated distinct cortical sources for the other two levels of processing: lexical fine tuning over occipito-tempopral/parietal regions; sublexical orthographic fine tuning over left occipital regions. Finally, exploratory group analyses based on children's reading fluency suggested that coarse print tuning emerges early even in children with limited reading knowledge, while sublexical and higher-level lexical processing emerge only in children with sufficient reading knowledge. Cognitive processes underlying coarse print tuning, sublexical, and lexical fine tuning were examined in beginning readers. Three overlapping yet distinct neural sources-left ventral occipito-temporal (vOT), left temporo-parietal, and primary visual cortex-were revealed underlying coarse print tuning. Responses to sublexical orthographic fine tuning were found over left occipital regions, while responses to higher-level linguistic fine tuning were found over occipito- temporal/parietal regions. Exploratory group analyses suggested that coarse print tuning emerges in children with limited reading knowledge, while sublexical and higher-level linguistic fine tuning effects emerge in children with sufficient reading knowledge. This article is protected by copyright. All rights reserved.

    View details for DOI 10.1111/desc.13352

    View details for PubMedID 36413170

  • Cognitive penetrability of scene representations based on horizontal image disparities. Scientific reports Chen, Y. D., Kaestner, M., Norcia, A. M. 2022; 12 (1): 17902


    The structure of natural scenes is signaled by many visual cues. Principal amongst them are the binocular disparities created by the laterally separated viewpoints of the two eyes. Disparity cues are believed to be processed hierarchically, first in terms of local measurements of absolute disparity and second in terms of more global measurements of relative disparity that allow extraction of the depth structure of a scene. Psychophysical and oculomotor studies have suggested that relative disparities are particularly relevant to perception, whilst absolute disparities are not. Here, we compare neural responses to stimuli that isolate the absolute disparity cue with stimuli that contain additional relative disparity cues, using the high temporal resolution of EEG to determine the temporal order of absolute and relative disparity processing. By varying the observers' task, we assess the extent to which each cue is cognitively penetrable. We find that absolute disparity is extracted before relative disparity, and that task effects arise only at or after the extraction of relative disparity. Our results indicate a hierarchy of disparity processing stages leading to the formation of a proto-object representation upon which higher cognitive processes can act.

    View details for DOI 10.1038/s41598-022-22670-7

    View details for PubMedID 36284130

  • Preferential Loss of Contrast Decrement Responses in Human Glaucoma. Investigative ophthalmology & visual science Norcia, A. M., Yakovleva, A., Jehangir, N., Goldberg, J. L. 2022; 63 (11): 16


    Purpose: The purpose of this study was to determine whether glaucoma in human patients produces preferential damage to OFF visual pathways, as suggested by animal experimental models, patient electroretinogram (ERG), and retinal imaging data.Methods: Steady-state visual evoked potentials (SSVEPs) were recorded monocularly from 50 patients with glaucoma and 28 age-similar controls in response to equal Weber contrast increments and decrements presented using 2.73 hertz (Hz) sawtooth temporal waveforms.Results: The eyes of patients with glaucoma were separated into mild (better than -6 decibel [dB] mean deviation; n = 28) and moderate to severe (worse than -6 dB mean deviation, n = 22) groups based on their Humphrey 24-2 visual field measurements. Response amplitudes and phases from the two glaucoma-severity groups were compared to controls at the group level. SSVEP amplitudes were depressed in both glaucoma groups, more so in the moderate to severe glaucoma group. The differences between controls and the moderate-severe glaucoma groups were more statistically reliable for decrements than for increments. Mean responses to decremental sawtooth stimuli were larger than those to increments in controls and in the mild glaucoma but not in the moderate to severe glaucoma group at the first harmonic. OFF/decrement responses at the first harmonic were faster in controls, but not in patients.Conclusions: The observed pattern of preferential loss of decremental responses in human glaucoma is consistent with prior reports of selective damage to OFF retinal ganglion cells in murine models and in data from human ERG and retinal imaging. These data motivate pursuit of SSVEP as a biomarker for glaucoma progression.

    View details for DOI 10.1167/iovs.63.11.16

    View details for PubMedID 36264656

  • PseudoSloan: A perimetric-complexity and area-controlled font for vision and reading research. Journal of vision Vildavski, V. Y., Lo Verde, L., Blumberg, G., Parsey, J., Norcia, A. M. 2022; 22 (10): 7


    Artificial orthographies have long been used in studies of verbal learning and reading. These orthographies, also known as pseudo or false fonts, are designed to match the letters of an existing alphabet on a range of visual features, isolating effects of orthography from those owing to lexical processing. In a parallel line of research, there has been much interest in the design of optotypes for measuring visual acuity that have good properties in terms of character complexity and graceful degradation under blur. Here we merge these two traditions by designing a fully scalable pseudofont, "PseudoSloan," that is based on the design rubric of the widely used Sloan optotypes. The font includes 26 Latin letters as well as two sets of letter-like symbols matching the Latin alphabet on a letter-by-letter basis. Quantitative matching of the pairs of Sloan and PseudoSloan glyphs is done on the basis of ink area and perimetric complexity. We provide the installable PseudoSloan font in TrueType and OpenType formats, plus a large number of PseudoSloan glyphs in .svg format that vary over wide ranges in their perimetric complexity and ink area (

    View details for DOI 10.1167/jov.22.10.7

    View details for PubMedID 36074477

  • Automatic retinal layer segmentation of visible-light optical coherence tomography images using deep learning Gopal, B., Zhang, T., Norcia, A., Goldberg, J. L., Dubra, A., Zhang, H., Soetikno, B. ASSOC RESEARCH VISION OPHTHALMOLOGY INC. 2022
  • Bilirubin-induced neurotoxicity and visuocortical dysfunction. Journal of perinatology : official journal of the California Perinatal Association Good, W. V., Wong, R. J., Norcia, A. M., Stevenson, D. K., Slagel, T., Hou, C., Bhutani, V. K. 2022

    View details for DOI 10.1038/s41372-022-01417-2

    View details for PubMedID 35618749

  • Behavioural and neural indices of perceptual decision-making in autistic children during visual motion tasks. Scientific reports Manning, C., Hassall, C. D., Hunt, L. T., Norcia, A. M., Wagenmakers, E., Evans, N. J., Scerif, G. 2022; 12 (1): 6072


    Many studies report atypical responses to sensory information in autistic individuals, yet it is not clear which stages of processing are affected, with little consideration given to decision-making processes. We combined diffusion modelling with high-density EEG to identify which processing stages differ between 50 autistic and 50 typically developing children aged 6-14years during two visual motion tasks. Our pre-registered hypotheses were that autistic children would show task-dependent differences in sensory evidence accumulation, alongside a more cautious decision-making style and longer non-decision time across tasks. We tested these hypotheses using hierarchical Bayesian diffusion models with a rigorous blind modelling approach, finding no conclusive evidence for our hypotheses. Using a data-driven method, we identified a response-locked centro-parietal component previously linked to the decision-making process. The build-up in this component did not consistently relate to evidence accumulation in autistic children. This suggests that the relationship between the EEG measure and diffusion-modelling is not straightforward in autistic children. Compared to a related study of children with dyslexia, motion processing differences appear less pronounced in autistic children. Exploratory analyses also suggest weak evidence that ADHD symptoms moderate perceptual decision-making in autistic children.

    View details for DOI 10.1038/s41598-022-09885-4

    View details for PubMedID 35414064

  • Dynamics of absolute and relative disparity processing in human visual cortex. NeuroImage Kaestner, M., Evans, M. L., Chen, Y. D., Norcia, A. M. 2022: 119186


    Cortical processing of binocular disparity is believed to begin in V1 where cells are sensitive to absolute disparity, followed by the extraction of relative disparity in higher visual areas. While much is known about the cortical distribution and spatial tuning of disparity-selective neurons, the relationship between their spatial and temporal properties is less well understood. Here, we use steady-state Visual Evoked Potentials and dynamic random dot stereograms to characterize the temporal dynamics of spatial mechanisms in human visual cortex that are primarily sensitive to either absolute or relative disparity. Stereograms alternated between disparate and non-disparate states at 2 Hz. By varying the disparity-defined spatial frequency content of the stereograms from a planar surface to corrugated ones, we biased responses towards absolute vs. relative disparities. Reliable Components Analysis was used to derive two dominant sources from the 128 channel EEG records. The first component (RC1) was maximal over the occipital pole. In RC1, first harmonic responses were sustained, tuned for corrugation frequency, and sensitive to the presence of disparity references, consistent with prior psychophysical sensitivity measurements. By contrast, the second harmonic, associated with transient processing, was not spatially tuned and was indifferent to references, consistent with it being generated by an absolute disparity mechanism. Thus, our results reveal a duplex coding strategy in the disparity domain, where relative disparities are computed via sustained mechanisms and absolute disparities are computed via transient mechanisms.

    View details for DOI 10.1016/j.neuroimage.2022.119186

    View details for PubMedID 35398280

  • Increasing neural network robustness improves match to macaque V1 eigenspectrum, spatial frequency preference and predictivity. PLoS computational biology Kong, N. C., Margalit, E., Gardner, J. L., Norcia, A. M. 2022; 18 (1): e1009739


    Task-optimized convolutional neural networks (CNNs) show striking similarities to the ventral visual stream. However, human-imperceptible image perturbations can cause a CNN to make incorrect predictions. Here we provide insight into this brittleness by investigating the representations of models that are either robust or not robust to image perturbations. Theory suggests that the robustness of a system to these perturbations could be related to the power law exponent of the eigenspectrum of its set of neural responses, where power law exponents closer to and larger than one would indicate a system that is less susceptible to input perturbations. We show that neural responses in mouse and macaque primary visual cortex (V1) obey the predictions of this theory, where their eigenspectra have power law exponents of at least one. We also find that the eigenspectra of model representations decay slowly relative to those observed in neurophysiology and that robust models have eigenspectra that decay slightly faster and have higher power law exponents than those of non-robust models. The slow decay of the eigenspectra suggests that substantial variance in the model responses is related to the encoding of fine stimulus features. We therefore investigated the spatial frequency tuning of artificial neurons and found that a large proportion of them preferred high spatial frequencies and that robust models had preferred spatial frequency distributions more aligned with the measured spatial frequency distribution of macaque V1 cells. Furthermore, robust models were quantitatively better models of V1 than non-robust models. Our results are consistent with other findings that there is a misalignment between human and machine perception. They also suggest that it may be useful to penalize slow-decaying eigenspectra or to bias models to extract features of lower spatial frequencies during task-optimization in order to improve robustness and V1 neural response predictivity.

    View details for DOI 10.1371/journal.pcbi.1009739

    View details for PubMedID 34995280

  • Transcranial ultrasound neuromodulation of the thalamic visual pathway in a large animal model and the dose‐response relationship with MR‐ARFI Scientific Reports Mohammadjavadi, M., Ash, R. T., Li, N., Gaur, P., Kubanek, J., Saenz, Y., Glover, G. H., Popelka, G. R., Norcia, A. M., Butts Pauly, K. 2022; 12: 19588
  • Stability and Plasticity of Steady-State Visual-Evoked Potential Contrast-Response Functions Ash, R., Nix, K., Lee, M., Pauly, K., Williams, N., Norcia, A. SPRINGERNATURE. 2021: 507
  • Symmetry Processing in the Macaque Visual Cortex. Cerebral cortex (New York, N.Y. : 1991) Audurier, P., Hejja-Brichard, Y., De Castro, V., Kohler, P. J., Norcia, A. M., Durand, J., Cottereau, B. R. 2021


    Symmetry is a highly salient feature of the natural world that is perceived by many species. In humans, the cerebral areas processing symmetry are now well identified from neuroimaging measurements. Macaque could constitute a good animal model to explore the underlying neural mechanisms, but a previous comparative study concluded that functional magnetic resonance imaging responses to mirror symmetry in this species were weaker than those observed in humans. Here, we re-examined symmetry processing in macaques from a broader perspective, using both rotation and reflection symmetry embedded in regular textures. Highly consistent responses to symmetry were found in a large network of areas (notably in areas V3 and V4), in line with what was reported in humans under identical experimental conditions. Our results suggest that the cortical networks that process symmetry in humans and macaques are potentially more similar than previously reported and point toward macaque as a relevant model for understanding symmetry processing.

    View details for DOI 10.1093/cercor/bhab358

    View details for PubMedID 34617100

  • Distinct neural sources underlying visual word form processing as revealed by steady state visual evoked potentials (SSVEP). Scientific reports Wang, F., Kaneshiro, B., Strauber, C. B., Hasak, L., Nguyen, Q. T., Yakovleva, A., Vildavski, V. Y., Norcia, A. M., McCandliss, B. D. 2021; 11 (1): 18229


    EEG has been central to investigations of the time course of various neural functions underpinning visual word recognition. Recently the steady-state visual evoked potential (SSVEP) paradigm has been increasingly adopted for word recognition studies due to its high signal-to-noise ratio. Such studies, however, have been typically framed around a single source in the left ventral occipitotemporal cortex (vOT). Here, we combine SSVEP recorded from 16 adult native English speakers with a data-driven spatial filtering approach-Reliable Components Analysis (RCA)-to elucidate distinct functional sources with overlapping yet separable time courses and topographies that emerge when contrasting words with pseudofont visual controls. The first component topography was maximal over left vOT regions with a shorter latency (approximately 180 ms). A second component was maximal over more dorsal parietal regions with a longer latency (approximately 260 ms). Both components consistently emerged across a range of parameter manipulations including changes in the spatial overlap between successive stimuli, and changes in both base and deviation frequency. We then contrasted word-in-nonword and word-in-pseudoword to test the hierarchical processing mechanisms underlying visual word recognition. Results suggest that these hierarchical contrasts fail to evoke a unitary component that might be reasonably associated with lexical access.

    View details for DOI 10.1038/s41598-021-95627-x

    View details for PubMedID 34521874

  • Global motion evoked potentials in autistic and dyslexic children: a cross-syndrome approach Toffoli, L., Scerif, G., Snowling, M. J., Norcia, A. M., Manning, C. SAGE PUBLICATIONS LTD. 2021: 578-579
  • A case-control study of visual, auditory and audio-visual sensory interactions in children with autism spectrum disorder. Journal of vision Norcia, A. M., Lee, A., Meredith, W. J., Kohler, P. J., Pei, F., Ghassan, S. A., Libove, R. A., Phillips, J. M., Hardan, A. Y. 2021; 21 (4): 5


    To assess the relative integrity of early visual and auditory processes in autism spectrum disorder (ASD), we used frequency-tagged visual and auditory stimulation and high-density electroencephalogram recordings of unimodal and dual-modality responses in a case-control design. To test for the specificity of effects on ASD, we recorded from a smaller group of children with attention-deficit hyperactivity disorder (ADHD). Horizontal 3 cycle per degree (cpd) gratings were presented at 5 Hz, and a random stream of /ba/, /da/, /ga/ syllables was presented at 6 Hz. Grating contrast response functions were measured unimodally and in the presence of a 64-dB auditory input. Auditory response functions were measured unimodally and in the presence of a 40% contrast grating. Children with ASD (n = 34) and ADHD (n = 13) showed a common lack of audio-visual interaction compared to typically developing children (n = 40) when measured at the first harmonic of the visual stimulus frequency. Both patient groups also showed depressed first harmonic responses at low contrast, but the ADHD group had consistently higher first-harmonic responses at high contrast. Children with ASD had a preferential loss of second-harmonic (transient) responses. The alteredtransient responses in ASD are likely to arise very early in the visual pathway and could thus have downstream consequences for many other visual mechanisms and processes. The alteration in audio-visual interaction could be a signature of a comorbid phenotype shared by ASD and ADHD, possibly due to alterations in attentional selection systems.

    View details for DOI 10.1167/jov.21.4.5

    View details for PubMedID 33830169

  • Visual motion and decision-making in dyslexia: Reduced accumulation of sensory evidence and related neural dynamics. The Journal of neuroscience : the official journal of the Society for Neuroscience Manning, C., Hassall, C. D., Hunt, L. T., Norcia, A. M., Wagenmakers, E. J., Snowling, M. J., Scerif, G., Evans, N. J. 2021


    Children with and without dyslexia differ in their behavioural responses to visual information, particularly when required to pool dynamic signals over space and time. Importantly, multiple processes contribute to behavioural responses. Here we investigated which processing stages are affected in children with dyslexia when performing visual motion processing tasks, by combining two methods that are sensitive to the dynamic processes leading to responses. We used a diffusion model which decomposes response time and accuracy into distinct cognitive constructs, and high-density EEG. 50 children with dyslexia (24 male) and 50 typically developing children (28 male) aged 6 to 14 years judged the direction of motion as quickly and accurately as possible in two global motion tasks (motion coherence and direction integration), which varied in their requirements for noise exclusion. Following our pre-registered analyses, we fitted hierarchical Bayesian diffusion models to the data, blinded to group membership. Unblinding revealed reduced evidence accumulation in children with dyslexia compared to typical children for both tasks. Additionally, we identified a response-locked EEG component which was maximal over centro-parietal electrodes which indicated a neural correlate of reduced drift-rate in dyslexia in the motion coherence task, thereby linking brain and behaviour. We suggest that children with dyslexia tend to be slower to extract sensory evidence from global motion displays, regardless of whether noise exclusion is required, thus furthering our understanding of atypical perceptual decision-making processes in dyslexia.SIGNIFICANCE STATEMENTReduced sensitivity to visual information has been reported in dyslexia, with a lively debate about whether these differences causally contribute to reading difficulties. In this large pre-registered study with a blind modelling approach, we combine state-of-the art methods in both computational modelling and EEG analysis to pinpoint the stages of processing that are atypical in children with dyslexia in two visual motion tasks that vary in their requirement for noise exclusion. We find reduced evidence accumulation in children with dyslexia across both tasks, and identify a neural marker, allowing us to link brain and behaviour. We show that children with dyslexia exhibit general difficulties with extracting sensory evidence from global motion displays, not just in tasks that require noise exclusion.

    View details for DOI 10.1523/JNEUROSCI.1232-21.2021

    View details for PubMedID 34782439

  • Motion Processing Deficits in Children With Cerebral Visual Impairment and Good Visual Acuity. Investigative ophthalmology & visual science Chandna, A., Nichiporuk, N., Nicholas, S., Kumar, R., Norcia, A. M. 2021; 62 (14): 12


    We sought to characterize neural motion processing deficits in children with cerebral visual impairment (CVI) who have good visual acuity using an objective, quantifiable method (steady-state visual evoked potentials [SSVEPs]).We recorded SSVEPs in response to three types of visual motion - absolute motion and more complex relative and rotary motion, comparing them to form-related vernier and contour responses. We studied a group of 31 children with CVI diagnosed via detailed clinical examinations and 28 age-matched healthy controls.Using measurements made at the appropriate response harmonics of the stimulation frequency, we found significant deficits in cerebral processing of relative and rotary motion but not of absolute motion in children with CVI compared with healthy controls. Vernier acuity, in keeping with good recognition acuity in both groups, was not different, nor were contour-related form responses.Deficits for complex motion but relative sparing of elementary motion and form-related signals suggests preferential damage to extra-striate visual motion areas in children with CVI. The fact that these preferential losses occur in the absence of significant acuity loss indicates that they are not secondary to reduced visual acuity, but rather are an independent vulnerability in CVI. These results corroborate parental and caregivers' reports of difficulties with tasks that involve motion perception in children with CVI.

    View details for DOI 10.1167/iovs.62.14.12

    View details for PubMedID 34779820

  • Global motion evoked potentials in autistic and dyslexic children: A cross-syndrome approach. Cortex; a journal devoted to the study of the nervous system and behavior Toffoli, L., Scerif, G., Snowling, M. J., Norcia, A. M., Manning, C. 2021; 143: 109-126


    Atypicalities in psychophysical thresholds for global motion processing have been reported in many neurodevelopmental conditions, including autism and dyslexia. Cross-syndrome comparisons of neural dynamics may help determine whether altered motion processing is a general marker of atypical development or condition-specific. Here, we assessed group differences in N2 peak amplitude (previously proposed as a marker of motion-specific processing) in typically developing (n = 57), autistic (n = 29) and dyslexic children (n = 44) aged 6-14 years, in two global motion tasks. High-density EEG data were collected while children judged the direction of global motion stimuli as quickly and accurately as possible, following a period of random motion. Using a data-driven component decomposition technique, we identified a reliable component that was maximal over occipital electrodes and had an N2-like peak at ~160 msec. We found no group differences in N2 peak amplitude, in either task. However, for both autistic and dyslexic children, there was evidence of atypicalities in later stages of processing that require follow up in future research. Our results suggest that early sensory encoding of motion information is unimpaired in dyslexic and autistic children. Group differences in later processing stages could reflect sustained global motion responses, decision-making, metacognitive processes and/or response generation, which may also distinguish between autistic and dyslexic individuals.

    View details for DOI 10.1016/j.cortex.2021.06.018

    View details for PubMedID 34399308

  • Disparity in Context: Understanding how monocular image content interacts with disparity processing in human visual cortex. NeuroImage Duan, Y. n., Thatte, J. n., Yaklovleva, A. n., Norcia, A. M. 2021: 118139


    Horizontal disparities between the two eyes' retinal images are the primary cue for depth. Commonly used Random dot stereograms (RDS) intentionally camouflage the disparity cue, breaking the correlations between monocular image structure and the depth map that are present in natural images. Because of the nonlinear nature of visual processing, it is unlikely that simple computational rules derived from RDS will be sufficient to explain binocular vision in natural environments. In order to understand the interplay between natural scene structure and disparity encoding, we used a depth-image-based-rendering technique and a library of natural 3D stereo pairs to synthesize two novel stereogram types in which monocular scene content was manipulated independent of scene depth information. The half-images of the novel stereograms comprised either random-dots or scrambled natural scenes each with the same depth maps as the corresponding natural scene stereograms. Using these stereograms in a simultaneous Event-Related Potential and behavioral discrimination task, we identified multiple disparity-contingent encoding stages between 100 ∼ 500 msec. The first disparity sensitive evoked potential was observed at ∼100 msec after an earlier evoked potential (between ∼50-100 msec) that was sensitive to the structure of the monocular half-images but blind to disparity. Starting at ∼150 msec, disparity responses were stereogram-specific and predictive of perceptual depth. Complex features associated with natural scene content are thus at least partially coded prior to disparity information, but these features and possibly others associated with natural scene content interact with disparity information only after an intermediate, 2D scene-independent disparity processing stage.

    View details for DOI 10.1016/j.neuroimage.2021.118139

    View details for PubMedID 33964460

  • Perceptual Decision-Making in Children: Age-Related Differences and EEG Correlates. Computational brain & behavior Manning, C. n., Wagenmakers, E. J., Norcia, A. M., Scerif, G. n., Boehm, U. n. 2021; 4 (1): 53–69


    Children make faster and more accurate decisions about perceptual information as they get older, but it is unclear how different aspects of the decision-making process change with age. Here, we used hierarchical Bayesian diffusion models to decompose performance in a perceptual task into separate processing components, testing age-related differences in model parameters and links to neural data. We collected behavioural and EEG data from 96 6- to 12-year-old children and 20 adults completing a motion discrimination task. We used a component decomposition technique to identify two response-locked EEG components with ramping activity preceding the response in children and adults: one with activity that was maximal over centro-parietal electrodes and one that was maximal over occipital electrodes. Younger children had lower drift rates (reduced sensitivity), wider boundary separation (increased response caution) and longer non-decision times than older children and adults. Yet, model comparisons suggested that the best model of children's data included age effects only on drift rate and boundary separation (not non-decision time). Next, we extracted the slope of ramping activity in our EEG components and covaried these with drift rate. The slopes of both EEG components related positively to drift rate, but the best model with EEG covariates included only the centro-parietal component. By decomposing performance into distinct components and relating them to neural markers, diffusion models have the potential to identify the reasons why children with developmental conditions perform differently to typically developing children and to uncover processing differences inapparent in the response time and accuracy data alone.

    View details for DOI 10.1007/s42113-020-00087-7

    View details for PubMedID 33604512

    View details for PubMedCentralID PMC7870772

  • Memory failure predicted by attention lapsing and media multitasking. Nature Madore, K. P., Khazenzon, A. M., Backes, C. W., Jiang, J., Uncapher, M. R., Norcia, A. M., Wagner, A. D. 2020


    With the explosion of digital media and technologies, scholars, educators and the public have become increasingly vocal about the role that an 'attention economy' has in our lives1. The rise of the current digital culture coincides with longstanding scientific questions about why humans sometimes remember and sometimes forget, and why some individuals remember better than others2-6. Here we examine whether spontaneous attention lapses-in the moment7-12, across individuals13-15 and as a function of everyday media multitasking16-19-negatively correlate with remembering. Electroencephalography and pupillometry measures of attention20,21 were recorded as eighty young adults (mean age, 21.7years) performed a goal-directed episodic encoding and retrieval task22. Trait-level sustained attention was further quantified using task-based23 and questionnaire measures24,25. Using trial-to-trial retrieval data, we show that tonic lapses in attention in the moment before remembering, assayed by posterior alpha power and pupil diameter, were correlated with reductions in neural signals of goal coding and memory, along with behavioural forgetting. Independent measures of trait-level attention lapsing mediated the relationship between neural assays of lapsing and memory performance, and between media multitasking and memory. Attention lapses partially account for why we remember or forget in the moment, and why some individuals remember better than others. Heavier media multitasking is associated with a propensity to have attention lapses and forget.

    View details for DOI 10.1038/s41586-020-2870-z

    View details for PubMedID 33116309

  • Neural sources of letter and Vernier acuity. Scientific reports Barzegaran, E., Norcia, A. M. 2020; 10 (1): 15449


    Visual acuity can be measured in many different ways, including with letters and Vernier offsets. Prior psychophysical work has suggested that the two acuities are strongly linked given that they both depend strongly on retinal eccentricity and both are similarly affected in amblyopia. Here we used high-density EEG recordings to ask whether the underlying neural sources are common as suggested by the psychophysics or distinct. To measure visual acuity for letters, we recorded evoked potentials to 3Hz alternations between intact and scrambled text comprised of letters of varying size. To measure visual acuity for Vernier offsets, we recorded evoked potentials to 3Hz alternations between bar gratings with and without a set of Vernier offsets. Both alternation types elicited robust activity at the 3Hz stimulus frequency that scaled in amplitude with both letter and offset size, starting near threshold. Letter and Vernier offset responses differed in both their scalp topography and temporal dynamics. The earliest evoked responses to letters occurred on lateral occipital visual areas, predominantly over the left hemisphere. Later responses were measured at electrodes over early visual cortex, suggesting that letter structure is first extracted in second-tier extra-striate areas and that responses over early visual areas are due to feedback. Responses to Vernier offsets, by contrast, occurred first at medial occipital electrodes, with responses at later time-points being more broadly distributed-consistent with feedforward pathway mediation. The previously observed commonalities between letter and Vernier acuity may be due to common bottlenecks in early visual cortex but not because the two tasks are subserved by a common network of visual areas.

    View details for DOI 10.1038/s41598-020-72370-3

    View details for PubMedID 32963270

  • Dynamics of Contrast Decrement and Increment Responses in Human Visual Cortex. Translational vision science & technology Norcia, A. M., Yakovleva, A., Hung, B., Goldberg, J. L. 2020; 9 (10): 6


    Purpose: The goal of the present experiments was to determine whether electrophysiologic response properties of the ON and OFF visual pathways observed in animal experimental models can be observed in humans.Methods: Steady-state visual evoked potentials (SSVEPs) were recorded in response to equivalent magnitude contrast increments and decrements presented within a probe-on-pedestal Westheimer sensitization paradigm. The probes were modulated with sawtooth temporal waveforms at a temporal frequency of 3 or 2.73 Hz. SSVEP response waveforms and response spectra for incremental and decremental stimuli were analyzed as a function of stimulus size and visual field location in 67 healthy adult participants.Results: SSVEPs recorded at the scalp differ between contrast decrements and increments of equal Weber contrast: SSVEP responses were larger in amplitude and shorter in latency for contrast decrements than for contrast increments. Both increment and decrement responses were larger for displays that were scaled for cortical magnification.Conclusions: In a fashion that parallels results from the early visual system of cats and monkeys, two key properties of ON versus OFF pathways found in single-unit recordings are recapitulated at the population level of activity that can be observed with scalp electrodes, allowing differential assessment of ON and OFF pathway activity in human.Translational Relevance: As data from preclinical models of visual pathway dysfunction point to differential damage to subtypes of retinal ganglion cells, this approach may be useful in future work on disease detection and treatment monitoring.

    View details for DOI 10.1167/tvst.9.10.6

    View details for PubMedID 32953246

  • Discovery and clinical translation of novel glaucoma biomarkers. Progress in retinal and eye research Beykin, G., Norcia, A. M., Srinivasan, V. J., Dubra, A., Goldberg, J. L. 2020: 100875


    Glaucoma and other optic neuropathies are characterized by progressive dysfunction and loss of retinal ganglion cells and their axons. Given the high prevalence of glaucoma-related blindness and the availability of treatment options, improving the diagnosis and precise monitoring of progression in these conditions is paramount. Here we review recent progress in the development of novel biomarkers for glaucoma in the context of disease pathophysiology and we propose future steps for the field, including integration of exploratory biomarker outcomes into prospective therapeutic trials. We anticipate that, when validated, some of the novel glaucoma biomarkers discussed here will prove useful for clinical diagnosis and prediction of progression, as well as monitoring of clinical responses to standard and investigational therapies.

    View details for DOI 10.1016/j.preteyeres.2020.100875

    View details for PubMedID 32659431

  • Time-resolved correspondences between deep neural network layers and EEG measurements in object processing. Vision research Kong, N. C., Kaneshiro, B., Yamins, D. L., Norcia, A. M. 2020; 172: 27–45


    The ventral visual stream is known to be organized hierarchically, where early visual areas processing simplistic features feed into higher visual areas processing more complex features. Hierarchical convolutional neural networks (CNNs) were largely inspired by this type of brain organization and have been successfully used to model neural responses in different areas of the visual system. In this work, we aim to understand how an instance of these models corresponds to temporal dynamics of human object processing. Using representational similarity analysis (RSA) and various similarity metrics, we compare the model representations with two electroencephalography (EEG) data sets containing responses to a shared set of 72 images. We find that there is a hierarchical relationship between the depth of a layer and the time at which peak correlation with the brain response occurs for certain similarity metrics in both data sets. However, when comparing across layers in the neural network, the correlation onset time did not appear in a strictly hierarchical fashion. We present two additional methods that improve upon the achieved correlations by optimally weighting features from the CNN and show that depending on the similarity metric, deeper layers of the CNN provide a better correspondence than shallow layers to later time points in the EEG responses. However, we do not find that shallow layers provide better correspondences than those of deeper layers to early time points, an observation that violates the hierarchy and is in agreement with the finding from the onset-time analysis. This work makes a first comparison of various response features-including multiple similarity metrics and data sets-with respect to a neural network.

    View details for DOI 10.1016/j.visres.2020.04.005

    View details for PubMedID 32388211

  • Natural music evokes correlated EEG responses reflecting temporal structure and beat. NeuroImage Kaneshiro, B. n., Nguyen, D. T., Norcia, A. M., Dmochowski, J. P., Berger, J. n. 2020: 116559


    The brain activity of multiple subjects has been shown to synchronize during salient moments of natural stimuli, suggesting that correlation of neural responses indexes a brain state operationally termed 'engagement'. While past electroencephalography (EEG) studies have considered both auditory and visual stimuli, the extent to which these results generalize to music-a temporally structured stimulus for which the brain has evolved specialized circuitry-is less understood. Here we investigated neural correlation during natural music listening by recording EEG responses from N=48 adult listeners as they heard real-world musical works, some of which were temporally disrupted through shuffling of short-term segments (measures), reversal, or randomization of phase spectra. We measured correlation between multiple neural responses (inter-subject correlation) and between neural responses and stimulus envelope fluctuations (stimulus-response correlation) in the time and frequency domains. Stimuli retaining basic musical features, such as rhythm and melody, elicited significantly higher behavioral ratings and neural correlation than did phase-scrambled controls. However, while unedited songs were self-reported as most pleasant, time-domain correlations were highest during measure-shuffled versions. Frequency-domain measures of correlation (coherence) peaked at frequencies related to the musical beat, although the magnitudes of these spectral peaks did not explain the observed temporal correlations. Our findings show that natural music evokes significant inter-subject and stimulus-response correlations, and suggest that the neural correlates of musical 'engagement' may be distinct from those of enjoyment.

    View details for DOI 10.1016/j.neuroimage.2020.116559

    View details for PubMedID 31978543

  • Corrigendum to "Neural dynamics underlying coherent motion perception in children and adults" [Dev. Cogn. Neurosci. 38 (August) (2019) 100670]. Developmental cognitive neuroscience Manning, C. n., Kaneshiro, B. n., Kohler, P. J., Duta, M. n., Scerif, G. n., Norcia, A. M. 2020; 41: 100748

    View details for DOI 10.1016/j.dcn.2019.100748

    View details for PubMedID 31999566

  • Relative contributions to vergence eye movements of two binocular cues for motion-in-depth. Scientific reports Giesel, M., Yakovleva, A., Bloj, M., Wade, A. R., Norcia, A. M., Harris, J. M. 2019; 9 (1): 17412


    When we track an object moving in depth, our eyes rotate in opposite directions. This type of "disjunctive" eye movement is called horizontal vergence. The sensory control signals for vergence arise from multiple visual cues, two of which, changing binocular disparity (CD) and inter-ocular velocity differences (IOVD), are specifically binocular. While it is well known that the CD cue triggers horizontal vergence eye movements, the role of the IOVD cue has only recently been explored. To better understand the relative contribution of CD and IOVD cues in driving horizontal vergence, we recorded vergence eye movements from ten observers in response to four types of stimuli that isolated or combined the two cues to motion-in-depth, using stimulus conditions and CD/IOVD stimuli typical of behavioural motion-in-depth experiments. An analysis of the slopes of the vergence traces and the consistency of the directions of vergence and stimulus movements showed that under our conditions IOVD cues provided very little input to vergence mechanisms. The eye movements that did occur coinciding with the presentation of IOVD stimuli were likely not a response to stimulus motion, but a phoria initiated by the absence of a disparity signal.

    View details for DOI 10.1038/s41598-019-53902-y

    View details for PubMedID 31758028

  • Differential Experience-Dependent Plasticity of Form and Motion Mechanisms in Anisometropic Amblyopia. Investigative ophthalmology & visual science Chen, S. I., Chandna, A., Nicholas, S., Norcia, A. M. 2019; 60 (13): 4109–19


    Purpose: We measure neural responses associated with form and motion processing in children with anisometropia before and after treatment with spectacles and occlusion.Methods: In this prospective, case-control treatment study, 10 children with anisometropia and amblyopia and 16 age-matched visually normal children participated. Steady-state visual evoked potentials (VEP) were recorded from electrodes over the occipital cortex. The visual stimulus comprised a horizontal bar grating into which Vernier offsets were introduced and withdrawn periodically at 3.75 Hz. The VEP amplitude at 3.75 Hz (first harmonic [1F]) and 7.5 Hz (second harmonic [2F]) were recorded to index the sensitivity of form/position-sensitive versus motion/transient-sensitive neural populations, respectively. Response amplitude at 1F and 2F were recorded over a series of 10 logarithmically spaced offset sizes before and after treatment. Main outcome measures are VEP amplitude versus displacement functions, interocular response amplitude differences.Results: After relaxing into spectacles (minimally-treated state), form/position-sensitive responses in the dominant/less ametropic eye of the children with anisometropia were larger and responses in the more ametropic eye were smaller than those of controls. Motion-transient responses were equal to those of controls in the less ametropic eye, but were smaller than controls in the more ametropic eye. After treatment, responses did not differ from those of controls.Conclusions: Form and motion responses are differentially susceptible to neural deprivation via optical blur. Form responses are more plastic than motion responses in minimally-treated children with anisometropic amblyopia. Most treatment effects occurred above threshold range, suggesting some treatment effects are not detected clinically.

    View details for DOI 10.1167/iovs.19-27005

    View details for PubMedID 31574139

  • Humans perceive binocular rivalry and fusion in a tristable dynamic state. The Journal of neuroscience : the official journal of the Society for Neuroscience Riesen, G., Norcia, A. M., Gardner, J. L. 2019


    Human vision combines inputs from the two eyes into one percept. Small differences 'fuse' together, while larger differences are seen 'rivalrously' from one eye at a time. These outcomes are typically treated as mutually exclusive processes, with paradigms targeting one or the other and fusion being unreported in most rivalry studies. Is fusion truly a default, stable state that only breaks into rivalry for non-fusible stimuli? Or are monocular and fused percepts three sub-states of one dynamical system? To determine whether fusion and rivalry are separate processes, we measured human perception of Gabor patches with a range of inter-ocular orientation disparities. Observers (10 female, 5 male) reported rivalrous, fused and uncertain percepts over time. We found a dynamic "tristable" zone spanning from 25-35 degrees of orientation disparity where fused, left- or right-eye dominant percepts could all occur. The temporal characteristics of fusion and non-fusion periods during tristability matched other bistable processes. We tested statistical models with fusion as a higher-level bistable process alternating with rivalry against our findings. None of these fit our data, but a simple bistable model extended to have three states reproduced many of our observations. We conclude that rivalry and fusion are multistable sub-states capable of direct competition, rather than separate bistable processes.SIGNIFICANCE STATEMENTWhen inputs to the two eyes differ, they can either fuse together or engage in binocular rivalry, where each eye's view is seen exclusively in turn. Visual stimuli have often been tailored to produce either fusion or rivalry, implicitly treating them as separate mutually-exclusive perceptual processes. We have found that some similar-but-different stimuli can result in both outcomes over time. Comparing various simple models with our results suggests that rivalry and fusion are not independent processes, but compete within a single multistable system. This conceptual shift is a step toward unifying fusion and rivalry, and understanding how they both contribute to the visual system's production of a unified interpretation of the conflicting images cast on the retina by real-world scenes.

    View details for DOI 10.1523/JNEUROSCI.0713-19.2019

    View details for PubMedID 31519817

  • EEGSourceSim: A framework for realistic simulation of EEG scalp data using MRI-based forward models and biologically plausible signals and noise. Journal of neuroscience methods Barzegaran, E., Bosse, S., Norcia, A. M. 2019: 108377


    BACKGROUND: Electroencephalography (EEG) is widely used to investigate human brain function. Simulation studies are essential for assessing the validity of EEG analysis methods and the interpretability of results.NEW METHOD: Here, we present a simulation environment for generating EEG data by embedding biologically plausible signal and noise into MRI-based forward models that incorporate individual-subject variability in structure and function.RESULTS: The package includes pipelines for the evaluation and validation of EEG analysis tools for source estimation, functional connectivity, and spatial filtering. EEG dynamics can be simulated using realistic noise and signal models with user specifiable signal-to-noise ratio (SNR). We also provide a set of quantitative metrics tailored to source estimation, connectivity and spatial filtering applications.COMPARISON WITH EXISTING METHOD(S): We provide a larger set of forward solutions for individual MRI-based head models than has been available previously. These head models are surface-based and include two sets of regions-of-interest (ROIs) that have been brought into registration with the brain of each individual using surface-based alignment - one from a whole brain and the other from a visual cortex atlas. We derive a realistic model of noise by fitting different model components to measured resting state EEG. We also provide a set of quantitative metrics for evaluating source-localization, functional connectivity and spatial filtering methods.CONCLUSIONS: The inclusion of a larger number of individual head-models, combined with surface-atlas based labeling of ROIs and plausible models of signal and noise, allows for simulation of EEG data with greater realism than previous packages.

    View details for DOI 10.1016/j.jneumeth.2019.108377

    View details for PubMedID 31381946

  • Contrast decrement vs increment responsiveness in glaucoma Norcia, A., Yakovleva, A., Goldberg, J. L. ASSOC RESEARCH VISION OPHTHALMOLOGY INC. 2019
  • Image Segmentation Based on Relative Motion and Relative Disparity Cues in Topographically Organized Areas of Human Visual Cortex. Scientific reports Kohler, P. J., Cottereau, B. R., Norcia, A. M. 2019; 9 (1): 9308


    The borders between objects and their backgrounds create discontinuities in image feature maps that can be used to recover object shape. Here we used functional magnetic resonance imaging to identify cortical areas that encode two of the most important image segmentation cues: relative motion and relative disparity. Relative motion and disparity cues were isolated by defining a central 2-degree disk using random-dot kinematograms and stereograms, respectively. For motion, the disk elicited retinotopically organized activations starting in V1 and extending through V2 and V3. In the surrounding region, we observed phase-inverted activations indicative of suppression, extending out to at least 6 degrees of retinal eccentricity. For disparity, disk activations were only found in V3, while suppression was observed in all early visual areas. Outside of early visual cortex, several areas were sensitive to both types of cues, most notably LO1, LO2 and V3B, making them additional candidate areas for motion- and disparity-cue combination. Adding an orthogonal task at fixation did not diminish these effects, and in fact led to small but measurable disk activations in V1 and V2 for disparity. The overall pattern of extra-striate activations is consistent with recent three-stream models of cortical organization.

    View details for DOI 10.1038/s41598-019-45036-y

    View details for PubMedID 31243297

  • Neural dynamics underlying coherent motion perception in children and adults. Developmental cognitive neuroscience Manning, C., Kaneshiro, B., Kohler, P. J., Duta, M., Scerif, G., Norcia, A. M. 2019; 38: 100670


    Motion sensitivity increases during childhood, but little is known about the neural correlates. Most studies investigating children's evoked responses have not dissociated direction-specific and non-direction-specific responses. To isolate direction-specific responses, we presented coherently moving dot stimuli preceded by incoherent motion, to 6- to 7-year-olds (n = 34), 8- to 10-year-olds (n = 34), 10- to 12-year-olds (n = 34) and adults (n = 20). Participants reported the coherent motion direction while high-density EEG was recorded. Using a data-driven approach, we identified two stimulus-locked EEG components with distinct topographies: an early component with an occipital topography likely reflecting sensory encoding and a later, sustained positive component over centro-parietal electrodes that we attribute to decision-related processes. The component waveforms showed clear age-related differences. In the early, occipital component, all groups showed a negativity peaking at ~300 ms, like the previously reported coherent-motion N2. However, the children, unlike adults, showed an additional positive peak at ~200 ms, suggesting differential stimulus encoding. The later positivity in the centro-parietal component rose more steeply for adults than for the youngest children, likely reflecting age-related speeding of decision-making. We conclude that children's protracted development of coherent motion sensitivity is associated with maturation of both early sensory and later decision-related processes.

    View details for DOI 10.1016/j.dcn.2019.100670

    View details for PubMedID 31228678

  • Autism sensory dysfunction in an evolutionarily conserved system. Proceedings. Biological sciences Vilidaite, G., Norcia, A. M., West, R. J., Elliott, C. J., Pei, F., Wade, A. R., Baker, D. H. 2018; 285 (1893): 20182255


    There is increasing evidence for a strong genetic basis for autism, with many genetic models being developed in an attempt to replicate autistic symptoms in animals. However, current animal behaviour paradigms rarely match the social and cognitive behaviours exhibited by autistic individuals. Here, we instead assay another functional domain-sensory processing-known to be affected in autism to test a novel genetic autism model in Drosophila melanogaster. We show similar visual response alterations and a similar development trajectory in Nhe3 mutant flies (total n = 72) and in autistic human participants (total n = 154). We report a dissociation between first- and second-order electrophysiological visual responses to steady-state stimulation in adult mutant fruit flies that is strikingly similar to the response pattern in human adults with ASD as well as that of a large sample of neurotypical individuals with high numbers of autistic traits. We explain this as a genetically driven, selective signalling alteration in transient visual dynamics. In contrast to adults, autistic children show a decrease in the first-order response that is matched by the fruit fly model, suggesting that a compensatory change in processing occurs during development. Our results provide the first animal model of autism comprising a differential developmental phenotype in visual processing.

    View details for DOI 10.1098/rspb.2018.2255

    View details for PubMedID 30963913

    View details for PubMedCentralID PMC6304042

  • Autism sensory dysfunction in an evolutionarily conserved system PROCEEDINGS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES Vilidaite, G., Norcia, A. M., West, R. H., Elliott, C. H., Pei, F., Wade, A. R., Baker, D. H. 2018; 285 (1893)
  • Tractography optimization using quantitative T1 mapping in the human optic radiation NEUROIMAGE Schurr, R., Duan, Y., Norcia, A. M., Ogawa, S., Yeatman, J. D., Mezer, A. A. 2018; 181: 645–58


    Diffusion MRI tractography is essential for reconstructing white-matter projections in the living human brain. Yet tractography results miss some projections and falsely identify others. A challenging example is the optic radiation (OR) that connects the thalamus and the primary visual cortex. Here, we tested whether OR tractography can be optimized using quantitative T1 mapping. Based on histology, we proposed that myelin-sensitive T1 values along the OR should remain consistently low compared with adjacent white matter. We found that complementary information from the T1 map allows for increasing the specificity of the reconstructed OR tract by eliminating falsely identified projections. This T1-filtering outperforms other, diffusion-based tractography filters. These results provide evidence that the smooth microstructural signature along the tract can be used as constructive input for tractography. Finally, we demonstrate that this approach can be applied in a case of multiple sclerosis, and generalized to the HCP-available MRI measurements. We conclude that multimodal MRI microstructural information can be used to eliminate spurious tractography results in the case of the OR.

    View details for PubMedID 29936310

  • Revisiting the functional significance of binocular cues for perceiving motion-in-depth. Nature communications Kohler, P. J., Meredith, W. J., Norcia, A. M. 2018; 9 (1): 3511


    Binocular differencing of spatial cues required for perceiving depth relationships is associated with decreased sensitivity to the corresponding retinal image displacements. However, binocular summation of contrast signals increases sensitivity. Here, we investigated this divergence in sensitivity by making direct neural measurements of responses to suprathreshold motion in human adults and 5-month-old infants using steady-state visually evoked potentials. Interocular differences in retinal image motion generated suppressed response functions and correspondingly elevated perceptual thresholds compared to motion matched between the two eyes. This suppression was of equal strength for horizontal and vertical motion and therefore not specific to the perception of motion-in-depth. Suppression is strongly dependent on the presence of spatial references in the image and highly immature in infants. Suppression appears to be the manifestation of a succession of spatial and interocular opponency operations that occur at an intermediate processing stage either before or in parallel with the extraction of motion-in-depth.

    View details for PubMedID 30158523

  • The neural basis of visual symmetry and its role in mid- and high-level visual processing ANNALS OF THE NEW YORK ACADEMY OF SCIENCES Bertamini, M., Silvanto, J., Norcia, A. M., Makin, A. J., Wagemans, J. 2018; 1426 (1): 111–26


    Symmetry is an important and prominent feature of the visual world. It has been studied as a basis for image segmentation and perceptual organization, but it also plays a role in higher level processes, such as face and object perception. Over the past decade, there has been progress in the study of the neural mechanisms of symmetry perception in humans and other animals. There is extended activity in the ventral stream, including the lateral occipital complex (LOC) and VO1; this activity starts in V3 and it occurs independently of the task (automatic response). Additionally, when the task requires processing of symmetry, the activation may emerge for objects that are symmetrical, even though they do not project a symmetrical image. There is also some evidence of hemispheric lateralization, especially for the LOC. We review the studies on the cortical basis of visual symmetry processing and its links to encoding of other aspects of the visual world, such as faces and objects.

    View details for PubMedID 29604083

  • Dynamics of human ON and OFF Visual Pathways Goldberg, J. L., Yakovleva, A., Hung, B., Norcia, A. ASSOC RESEARCH VISION OPHTHALMOLOGY INC. 2018
  • Measuring Integration Processes in Visual Symmetry with Frequency-Tagged EEG SCIENTIFIC REPORTS Alp, N., Kohler, P., Kogo, N., Wagemans, J., Norcia, A. 2018; 8: 6969


    Symmetry is a highly salient feature of the natural world which requires integration of visual features over space. The aim of the current work is to isolate dynamic neural correlates of symmetry-specific integration processes. We measured steady-state visual evoked potentials (SSVEP) as participants viewed symmetric patterns comprised of distinct spatial regions presented at two different frequencies (f1 and f2). We measured intermodulation components, shown to reflect non-linear processing at the neural level, indicating integration of spatially separated parts of the pattern. We generated a wallpaper pattern containing two reflection symmetry axes by tiling the plane with a two-fold reflection symmetric unit-pattern and split each unit-pattern diagonally into separate parts which could be presented at different frequencies. We compared SSVEPs measured for wallpapers and control patterns for which both images were equal in terms of translation and rotation symmetry but reflection symmetry could only emerge for the wallpaper pattern through integration of the image-pairs. We found that low-frequency intermodulation components differed between the wallpaper and control stimuli, indicating the presence of integration mechanisms specific to reflection symmetry. These results showed that spatial integration specific to symmetry perception can be isolated through a combination of stimulus design and the frequency tagging approach.

    View details for PubMedID 29725022

  • Electrophysiological Correlates of Motion Processing and Decision-Making in Children and Adults Manning, C., Scerif, G., Norcia, A. SAGE PUBLICATIONS LTD. 2018: 556–57
  • Towards a Machine-learning Approach for Sickness Prediction in 360 degrees Stereoscopic Videos Padmanaban, N., Ruban, T., Sitzmann, V., Norcia, A. M., Wetzstein, G. IEEE COMPUTER SOC. 2018: 1594–1603


    Virtual reality systems are widely believed to be the next major computing platform. There are, however, some barriers to adoption that must be addressed, such as that of motion sickness - which can lead to undesirable symptoms including postural instability, headaches, and nausea. Motion sickness in virtual reality occurs as a result of moving visual stimuli that cause users to perceive self-motion while they remain stationary in the real world. There are several contributing factors to both this perception of motion and the subsequent onset of sickness, including field of view, motion velocity, and stimulus depth. We verify first that differences in vection due to relative stimulus depth remain correlated with sickness. Then, we build a dataset of stereoscopic 3D videos and their corresponding sickness ratings in order to quantify their nauseogenicity, which we make available for future use. Using this dataset, we train a machine learning algorithm on hand-crafted features (quantifying speed, direction, and depth as functions of time) from each video, learning the contributions of these various features to the sickness ratings. Our predictor generally outperforms a naïve estimate, but is ultimately limited by the size of the dataset. However, our result is promising and opens the door to future work with more extensive datasets. This and further advances in this space have the potential to alleviate developer and end user concerns about motion sickness in the increasingly commonplace virtual world.

    View details for DOI 10.1109/TVCG.2018.2793560

    View details for Web of Science ID 000427682500022

    View details for PubMedID 29553929

  • Detection of Amblyopia Using Sweep VEP Vernier and Grating Acuity INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Hou, C., Good, W. V., Norcia, A. M. 2018; 59 (3): 1435–42


    Vernier and grating acuity can be measured with swept-parameter visual evoked potentials (sVEP). However, whether sVEP Vernier and grating acuities are comparable in predicting letter acuity has not been systematically evaluated. This study evaluated the validity and reliability of sVEP Vernier and grating acuity for the detection of amblyopia in adults.Three types of acuity were measured in 36 adults with amblyopia and 36 age-matched normal-vision controls. Letter acuity was measured with a logMAR chart. Both Vernier and grating acuity were estimated by sVEP and psychophysics for the same stimuli. Regression analyses were performed between the perceptual and electrophysiologic acuity measurements.SVEP Vernier and grating acuities were significantly correlated with their corresponding psychophysical acuities (P < 0.001). Both the sVEP Vernier (P < 0.0001) and grating (P < 0.01) acuities were also significantly correlated with letter acuity. However, Vernier acuity more precisely reflected the magnitude of the letter acuity loss than did grating acuity for both sVEP and psychophysical measures. Repeating sVEP grating acuity tests with different temporal frequencies and modulation types indicated good reliability of sVEP acuity measures.SVEP Vernier acuity has a 1:1 relationship with letter acuity, but sVEP grating acuity does not. SVEP Vernier acuity thus provides a better characterization of the magnitude of the amblyopic acuity loss than does sVEP grating acuity. Nonetheless, each of the sVEP measurements can be used to predict letter acuity and because they can be made without a behavioral response, they may be useful measures of visual function in pre- and nonverbal patients.

    View details for PubMedID 29625467

    View details for PubMedCentralID PMC5858252

  • Determinants of neural responses to disparity in natural scenes JOURNAL OF VISION Duan, Y., Yakovleva, A., Norcia, A. M. 2018; 18 (3): 21


    We studied disparity-evoked responses in natural scenes using high-density electroencephalography (EEG) in an event-related design. Thirty natural scenes that mainly included outdoor settings with trees and buildings were used. Twenty-four subjects viewed a series of trials composed of sequential two-alternative temporal forced-choice presentation of two different versions (two-dimensional [2D] vs. three-dimensional [3D]) of the same scene interleaved by a scrambled image with the same power spectrum. Scenes were viewed orthostereoscopically at 3 m through a pair of shutter glasses. After each trial, participants indicated with a key press which version of the scene was 3D. Performance on the discrimination was >90%. Participants who were more accurate also tended to respond faster; scenes that were reported more accurately as 3D also led to faster reaction times. We compared visual evoked potentials elicited by scrambled, 2D, and 3D scenes using reliable component analysis to reduce dimensionality. The disparity-evoked response to natural scene stimuli, measured from the difference potential between 2D and 3D scenes, comprised a sustained relative negativity in the dominant response component. The magnitude of the disparity-specific response was correlated with the observer's stereoacuity. Scenes with more homogeneous depth maps also tended to elicit large disparity-specific responses. Finally, the magnitude of the disparity-specific response was correlated with the magnitude of the differential response between scrambled and 2D scenes, suggesting that monocular higher-order scene statistics modulate disparity-specific responses.

    View details for PubMedID 29677337

  • Dynamics of perceptual decisions about symmetry in visual cortex NEUROIMAGE Kohler, P. J., Cottereau, B. R., Norcia, A. M. 2018; 167: 316–30


    Neuroimaging studies have identified multiple extra-striate visual areas that are sensitive to symmetry in planar images (Kohler et al., 2016; Sasaki et al., 2005). Here, we investigated which of these areas are directly involved in perceptual decisions about symmetry, by recording high-density EEG in participants (n = 25) who made rapid judgments about whether an exemplar image contained rotation symmetry or not. Stimulus-locked sensor-level analysis revealed symmetry-specific activity that increased with increasing order of rotation symmetry. Response-locked analysis identified activity occurring between 600 and 200 ms before the button-press, that was directly related to perceptual decision making. We then used fMRI-informed EEG source imaging to characterize the dynamics of symmetry-specific activity within an extended network of areas in visual cortex. The most consistent cortical source of the stimulus-locked activity was VO1, a topographically organized area in ventral visual cortex, that was highly sensitive to symmetry in a previous study (Kohler et al., 2016). Importantly, VO1 activity also contained a strong decision-related component, suggesting that this area plays a crucial role in perceptual decisions about symmetry. Other candidate areas, such as lateral occipital cortex, had weak stimulus-locked symmetry responses and no evidence of correlation with response timing.

    View details for PubMedID 29175495

  • Distinct Representations of Magnitude and Spatial Position within Parietal Cortex during Number-Space Mapping JOURNAL OF COGNITIVE NEUROSCIENCE Kanayet, F. J., Mattarella-Micke, A., Kohler, P. J., Norcia, A. M., McCandliss, B. D., McClelland, J. L. 2018; 30 (2): 200–218


    Mapping numbers onto space is foundational to mathematical cognition. These cognitive operations are often conceptualized in the context of a "mental number line" and involve multiple brain regions in or near the intraparietal sulcus (IPS) that have been implicated both in numeral and spatial cognition. Here we examine possible differentiation of function within these brain areas in relating numbers to spatial positions. By isolating the planning phase of a number line task and introducing spatiotopic mapping tools from fMRI into mental number line task research, we are able to focus our analysis on the neural activity of areas in anterior IPS (aIPS) previously associated with number processing and on spatiotopically organized areas in and around posterior IPS (pIPS), while participants prepare to place a number on a number line. Our results support the view that the nonpositional magnitude of a numerical symbol is coded in aIPS, whereas the position of a number in space is coded in posterior areas of IPS. By focusing on the planning phase, we are able to isolate activation related to the cognitive, rather than the sensory-motor, aspects of the task. Also, to allow the separation of spatial position from magnitude, we tested both a standard positive number line (0 to 100) and a zero-centered mixed number line (-100 to 100). We found evidence of a functional dissociation between aIPS and pIPS: Activity in aIPS was associated with a landmark distance effect not modulated by spatial position, whereas activity in pIPS revealed a contralateral preference effect.

    View details for PubMedID 29040015

  • Development of Relative Disparity Sensitivity in Human Visual Cortex. journal of neuroscience Norcia, A. M., Gerhard, H. E., Meredith, W. J. 2017; 37 (23): 5608-5619


    Stereopsis is the primary cue underlying our ability to make fine depth judgments. In adults, depth discriminations are supported largely by relative rather than absolute binocular disparity, and depth is perceived primarily for horizontal rather than vertical disparities. Although human infants begin to exhibit disparity-specific responses between 3 and 5 months of age, it is not known how relative disparity mechanisms develop. Here we show that the specialization for relative disparity is highly immature in 4- to 6-month-old infants but is adult-like in 4- to 7-year-old children. Disparity-tuning functions for horizontal and vertical disparities were measured using the visual evoked potential. Infant relative disparity thresholds, unlike those of adults, were equal for vertical and horizontal disparities. Their horizontal disparity thresholds were a factor of ∼10 higher than adults, but their vertical disparity thresholds differed by a factor of only ∼4. Horizontal relative disparity thresholds for 4- to 7-year-old children were comparable with those of adults at ∼0.5 arcmin. To test whether infant immaturity was due to spatial limitations or insensitivity to interocular correlation, highly suprathreshold horizontal and vertical disparities were presented in alternate regions of the display, and the interocular correlation of the interdigitated regions was varied from 0% to 100%. This manipulation regulated the availability of coarse-scale relative disparity cues. Adult and infant responses both increased with increasing interocular correlation by similar magnitudes, but adult responses increased much more for horizontal disparities, further evidence for qualitatively immature stereopsis based on relative disparity at 4-6 months of age.SIGNIFICANCE STATEMENT Stereopsis, our ability to sense depth from horizontal image disparity, is among the finest spatial discriminations made by the primate visual system. Fine stereoscopic depth discriminations depend critically on comparisons of disparity relationships in the image that are supported by relative disparity cues rather than the estimation of single, absolute disparities. Very young human and macaque infants are sensitive to absolute disparity, but no previous study has specifically studied the development of relative disparity sensitivity, a hallmark feature of adult stereopsis. Here, using high-density EEG recordings, we show that 4- to 6-month-old infants display both quantitative and qualitative response immaturities for relative disparity information. Relative disparity responses are adult-like no later than 4-7 years of age.

    View details for DOI 10.1523/JNEUROSCI.3570-16.2017

    View details for PubMedID 28473649

  • Evidence for long-range spatiotemporal interactions in infant and adult visual cortex JOURNAL OF VISION Norcia, A. M., Pei, F., Kohler, P. J. 2017; 17 (6): 12


    The development of spatiotemporal interactions giving rise to classical receptive field properties has been well studied in animal models, but little is known about the development of putative nonclassical mechanisms in any species. Here we used visual evoked potentials to study the developmental status of spatiotemporal interactions for stimuli that were biased to engage long-range spatiotemporal integration mechanisms. We compared responses to widely spaced stimuli presented either in temporal succession or at the same time. The former configuration elicits a percept of apparent motion in adults but the latter does not. Component flash responses were summed to make a linear prediction (no spatiotemporal interaction) for comparison with the measured evoked responses to sequential or simultaneous flash conditions. In adults, linear summation of the separate flash responses measured with 40% contrast stimuli predicted sequential flash responses twice as large as those measured, indicating that the response measured under apparent motion conditions is subadditive. Simultaneous-flash responses at the same spatial separation were also subadditive, but substantially less so. The subadditivity in both cases could be modeled as a simple multiplicative gain term across all electrodes and time points. In infants aged 3-8 months, responses to the stimuli used in adults were similar to their linear predictions at 40%, but the responses measured at 80% contrast resembled the subadditive responses of the adults for both sequential and simultaneous flash conditions. We interpret the developmental data as indicating that adult-like long-range spatiotemporal interactions can be demonstrated by 3-8 months, once stimulus contrast is high enough.

    View details for PubMedID 28622700

  • Optimal use of EEG recordings to target active brain areas with transcranial electrical stimulation. NeuroImage Dmochowski, J. P., Koessler, L., Norcia, A. M., Bikson, M., Parra, L. C. 2017


    To demonstrate causal relationships between brain and behavior, investigators would like to guide brain stimulation using measurements of neural activity. Particularly promising in this context are electroencephalography (EEG) and transcranial electrical stimulation (TES), as they are linked by a reciprocity principle which, despite being known for decades, has not led to a formalism for relating EEG recordings to optimal stimulation parameters. Here we derive a closed-form expression for the TES configuration that optimally stimulates (i.e., targets) the sources of recorded EEG, without making assumptions about source location or distribution. We also derive a duality between TES targeting and EEG source localization, and demonstrate that in cases where source localization fails, so does the proposed targeting. Numerical simulations with multiple head models confirm these theoretical predictions and quantify the achieved stimulation in terms of focality and intensity. We show that constraining the stimulation currents automatically selects optimal montages that involve only a few (4-7) electrodes, with only incremental loss in performance when targeting focal activations. The proposed technique allows brain scientists and clinicians to rationally target the sources of observed EEG and thus overcomes a major obstacle to the realization of individualized or closed-loop brain stimulation.

    View details for DOI 10.1016/j.neuroimage.2017.05.059

    View details for PubMedID 28578130

  • Development of contrast normalization mechanisms during childhood and adolescence VISION RESEARCH Pei, F., Baldassi, S., Tsai, J. J., Gerhard, H. E., Norcia, A. M. 2017; 133: 12-20


    Contrast sensitivity is regulated by neural mechanisms that flexibly adjust responsiveness to optimize stimulus encoding across different environments. Here we studied the developmental status of gain control mechanisms in school-age children (5-17years) and adults using a visual masking paradigm. A variable contrast, spatially random 2-D noise test pattern was masked by the presence of a superimposed independent noise pattern presented at 0, 12 and 40% contrast. Frequency-tagged steady state visual evoked potentials were used to separately record responses to the test (5.14Hz) and the mask (7.2Hz). By incrementally increasing the test contrast we measured contrast response functions for each mask contrast. The unmasked contrast response functions were largely similar in shape across age, but peak amplitude was higher in the children. Masking shifted the contrast response function rightward on the contrast axis in both the adults and older children, elevating contrast thresholds by a similar factor across age. However, in younger children, masking resulted in a change in the slope of the contrast response function. These findings suggest that immaturity in the contrast normalization process persists until approximately 11years of age.

    View details for DOI 10.1016/j.visres.2016.03.010

    View details for Web of Science ID 000399865500002

    View details for PubMedID 27826013

  • In-vivo measurements of human brain tissue conductivity using focal electrical current injection through intracerebral multicontact electrodes. Human brain mapping Koessler, L., Colnat-Coulbois, S., Cecchin, T., Hofmanis, J., Dmochowski, J. P., Norcia, A. M., Maillard, L. G. 2017; 38 (2): 974-986


    In-vivo measurements of human brain tissue conductivity at body temperature were conducted using focal electrical currents injected through intracerebral multicontact electrodes. A total of 1,421 measurements in 15 epileptic patients (age: 28 ± 10) using a radiofrequency generator (50 kHz current injection) were analyzed. Each contact pair was classified as being from healthy (gray matter, n = 696; white matter, n = 530) or pathological (epileptogenic zone, n = 195) tissue using neuroimaging analysis of the local tissue environment and intracerebral EEG recordings. Brain tissue conductivities were obtained using numerical simulations based on conductivity estimates that accounted for the current flow in the local brain volume around the contact pairs (a cube with a side length of 13 mm). Conductivity values were 0.26 S/m for gray matter and 0.17 S/m for white matter. Healthy gray and white matter had statistically different median impedances (P < 0.0001). White matter conductivity was found to be homogeneous as normality tests did not find evidence of multiple subgroups. Gray matter had lower conductivity in healthy tissue than in the epileptogenic zone (0.26 vs. 0.29 S/m; P = 0.012), even when the epileptogenic zone was not visible in the magnetic resonance image (MRI) (P = 0.005). The present in-vivo conductivity values could serve to create more accurate volume conduction models and could help to refine the identification of relevant intracerebral contacts, especially when located within the epileptogenic zone of an MRI-invisible lesion. Hum Brain Mapp 38:974-986, 2017. © 2016 Wiley Periodicals, Inc.

    View details for DOI 10.1002/hbm.23431

    View details for PubMedID 27726249

  • Sparse EEG/MEG source estimation via a group lasso. PloS one Lim, M. n., Ales, J. M., Cottereau, B. R., Hastie, T. n., Norcia, A. M. 2017; 12 (6): e0176835


    Non-invasive recordings of human brain activity through electroencephalography (EEG) or magnetoencelphalography (MEG) are of value for both basic science and clinical applications in sensory, cognitive, and affective neuroscience. Here we introduce a new approach to estimating the intra-cranial sources of EEG/MEG activity measured from extra-cranial sensors. The approach is based on the group lasso, a sparse-prior inverse that has been adapted to take advantage of functionally-defined regions of interest for the definition of physiologically meaningful groups within a functionally-based common space. Detailed simulations using realistic source-geometries and data from a human Visual Evoked Potential experiment demonstrate that the group-lasso method has improved performance over traditional ℓ2 minimum-norm methods. In addition, we show that pooling source estimates across subjects over functionally defined regions of interest results in improvements in the accuracy of source estimates for both the group-lasso and minimum-norm approaches.

    View details for PubMedID 28604790

  • Representation of Maximally Regular Textures in Human Visual Cortex JOURNAL OF NEUROSCIENCE Kohler, P. J., Clarke, A., Yakovleva, A., Liu, Y., Norcia, A. M. 2016; 36 (3): 714–29


    Naturalistic textures with an intermediate degree of statistical regularity can capture key structural features of natural images (Freeman and Simoncelli, 2011). V2 and later visual areas are sensitive to these features, while primary visual cortex is not (Freeman et al., 2013). Here we expand on this work by investigating a class of textures that have maximal formal regularity, the 17 crystallographic wallpaper groups (Fedorov, 1891). We used texture stimuli from four of the groups that differ in the maximum order of rotation symmetry they contain, and measured neural responses in human participants using functional MRI and high-density EEG. We found that cortical area V3 has a parametric representation of the rotation symmetries in the textures that is not present in either V1 or V2, the first discovery of a stimulus property that differentiates processing in V3 from that of lower-level areas. Parametric responses were also seen in higher-order ventral stream areas V4, VO1, and lateral occipital complex (LOC), but not in dorsal stream areas. The parametric response pattern was replicated in the EEG data, and source localization indicated that responses in V3 and V4 lead responses in LOC, which is consistent with a feedforward mechanism. Finally, we presented our stimuli to four well developed feedforward models and found that none of them were able to account for our results. Our results highlight structural regularity as an important stimulus dimension for distinguishing the early stages of visual processing, and suggest a previously unrecognized role for V3 in the visual form-processing hierarchy. Significance statement: Hierarchical processing is a fundamental organizing principle in visual neuroscience, with each successive processing stage being sensitive to increasingly complex stimulus properties. Here, we probe the encoding hierarchy in human visual cortex using a class of visual textures--wallpaper patterns--that are maximally regular. Through a combination of fMRI and EEG source imaging, we find specific responses to texture regularity that depend parametrically on the maximum order of rotation symmetry in the textures. These parametric responses are seen in several areas of the ventral visual processing stream, as well as in area V3, but not in V1 or V2. This is the first demonstration of a stimulus property that differentiates processing in V3 from that of lower-level visual areas.

    View details for PubMedID 26791203

  • Temporal Tuning of Word- and Face-selective Cortex. Journal of cognitive neuroscience Yeatman, J. D., Norcia, A. M. 2016; 28 (11): 1820–27


    Sensitivity to temporal change places fundamental limits on object processing in the visual system. An emerging consensus from the behavioral and neuroimaging literature suggests that temporal resolution differs substantially for stimuli of different complexity and for brain areas at different levels of the cortical hierarchy. Here, we used steady-state visually evoked potentials to directly measure three fundamental parameters that characterize the underlying neural response to text and face images: temporal resolution, peak temporal frequency, and response latency. We presented full-screen images of text or a human face, alternated with a scrambled image, at temporal frequencies between 1 and 12 Hz. These images elicited a robust response at the first harmonic that showed differential tuning, scalp topography, and delay for the text and face images. Face-selective responses were maximal at 4 Hz, but text-selective responses, by contrast, were maximal at 1 Hz. The topography of the text image response was strongly left-lateralized at higher stimulation rates, whereas the response to the face image was slightly right-lateralized but nearly bilateral at all frequencies. Both text and face images elicited steady-state activity at more than one apparent latency; we observed early (141-160 msec) and late (>250 msec) text- and face-selective responses. These differences in temporal tuning profiles are likely to reflect differences in the nature of the computations performed by word- and face-selective cortex. Despite the close proximity of word- and face-selective regions on the cortical surface, our measurements demonstrate substantial differences in the temporal dynamics of word- versus face-selective responses.

    View details for PubMedID 27378330

    View details for PubMedCentralID PMC5045815

  • Development of Three-Dimensional Perception in Human Infants. Annual review of vision science Norcia, A. M., Gerhard, H. E. 2015; 1: 569-594


    The play of light on the retina contains multiple sources of information about the three-dimensional (3D) structure of the world. Some of the best information is derived from differencing operations that act on the images that result from the two eyes' laterally displaced vantage points. Other information is available in systematic retinal patterns of local texture and motion cues. This article describes what is currently known about the development of sensitivity to these binocular and monocular cues for depth in human infants, and it places the results in the context of what is known about the underlying neural mechanisms from work in nonhuman primates and human neuroimaging studies.

    View details for DOI 10.1146/annurev-vision-082114-035835

    View details for PubMedID 28532379

  • Cortical Components of Reaction-Time during Perceptual Decisions in Humans PLOS ONE Dmochowski, J. P., Norcia, A. M. 2015; 10 (11)

    View details for DOI 10.1371/journal.pone.0143339

    View details for PubMedID 26599741

  • Relative sensitivity to low- vs. high-level visual properties in face-sensitive regions of the human ventral occipito-temporal cortex: evidence from intra-cerebral recordings. Journal of vision Liu-Shuang, J., Jonas, J., Ales, J., Norcia, A., Maillard, L., Rossion, B. 2015; 15 (12): 751-?


    Neuroimaging studies have identified a network of regions in the human ventral occipito-temporal cortex (VOTC) sensitive to faces. Using intra-cerebral electrophysiological recordings, we compared the relative responsivity of the different regions to face-structure using a periodic visual stimulation paradigm. Recordings were made in 17 epileptic patients implanted with linear-array depth-electrodes (>1000 contacts in the VOTC). In Experiment 1, we presented sequences of phase-scrambled and intact face stimuli (equalised for low-level image properties) alternating at a fixed frequency of 3Hz (6 images/s) while patients completed an orthogonal task. Significant periodic EEG responses (SSVEPs) reflecting high-level face structure processing were found at 3 Hz, while responses reflecting low-level visual processing (e.g. local contrast change) were recorded at 6 Hz. An index of high-level processing (3Hz/(3Hz+6Hz)) showed a posterior-to-anterior gradient, reflecting increased sensitivity to face structure going from the occipital to the temporo-polar cortex, with a right hemisphere lateralisation. In Experiment 2, we further investigated face structure processing with a sweep VEP paradigm (Ales et al., 2012; 4/17 patients). Stimulation was similar to Experiment 1 except the visibility (phase coherence) of the face image was parametrically varied so that a face gradually emerged over the course of a sequence. The patients performed a face detection task during the trials. Responses at 3Hz, objective markers of face detection, were found in all patients on contacts in the lateral fusiform gyrus and inferior occipital gyrus that overlapped with those defined in Experiment 1. All regions showed non-linear voltage vs coherence functions at 3Hz, with thresholds lying around ≈40% coherence. Sites in the lateral fusiform gyrus had a distinct step-like response profile suggestive of categorical face detection. These findings demonstrate that face-sensitive regions are organised with a posterior-to-anterior gradient of increasingly selective high-level visual processing. These regions appear to detect face structure in a non-linear/categorical manner. Meeting abstract presented at VSS 2015.

    View details for DOI 10.1167/15.12.751

    View details for PubMedID 26326439

  • A Representational Similarity Analysis of the Dynamics of Object Processing Using Single-Trial EEG Classification PLOS ONE Kaneshiro, B., Guimaraes, M. P., Kim, H., Norcia, A. M., Suppes, P. 2015; 10 (8)


    The recognition of object categories is effortlessly accomplished in everyday life, yet its neural underpinnings remain not fully understood. In this electroencephalography (EEG) study, we used single-trial classification to perform a Representational Similarity Analysis (RSA) of categorical representation of objects in human visual cortex. Brain responses were recorded while participants viewed a set of 72 photographs of objects with a planned category structure. The Representational Dissimilarity Matrix (RDM) used for RSA was derived from confusions of a linear classifier operating on single EEG trials. In contrast to past studies, which used pairwise correlation or classification to derive the RDM, we used confusion matrices from multi-class classifications, which provided novel self-similarity measures that were used to derive the overall size of the representational space. We additionally performed classifications on subsets of the brain response in order to identify spatial and temporal EEG components that best discriminated object categories and exemplars. Results from category-level classifications revealed that brain responses to images of human faces formed the most distinct category, while responses to images from the two inanimate categories formed a single category cluster. Exemplar-level classifications produced a broadly similar category structure, as well as sub-clusters corresponding to natural language categories. Spatiotemporal components of the brain response that differentiated exemplars within a category were found to differ from those implicated in differentiating between categories. Our results show that a classification approach can be successfully applied to single-trial scalp-recorded EEG to recover fine-grained object category structure, as well as to identify interpretable spatiotemporal components underlying object processing. Finally, object category can be decoded from purely temporal information recorded at single electrodes.

    View details for DOI 10.1371/journal.pone.0135697

    View details for Web of Science ID 000359926900040

    View details for PubMedCentralID PMC4546653

  • The Structural Properties of Major White Matter Tracts in Strabismic Amblyopia. Investigative ophthalmology & visual science Duan, Y., Norcia, A. M., Yeatman, J. D., Mezer, A. 2015; 56 (9): 5152-5160


    In order to better understand whether white matter structural deficits are present in strabismic amblyopia, we performed a survey of the tissue properties of 28 major white matter tracts using diffusion and quantitative magnetic resonance imaging approaches.We used diffusion-based tensor modeling and a new quantitative T1 protocol to measure fractional anisotropy (FA), mean diffusivity (MD), and myelin-sensitive T1 values. We surveyed tracts in the occipital lobe, including the vertical occipital fasciculus (VOF)-a newly rediscovered tract that bridges dorsal and ventral areas of the occipital lobe, as well as tracts across the rest of the brain.Adults with long-standing strabismic amblyopia show tract-specific elevations in MD. We rank-ordered the tracts on the basis of their MD effect-size. The four most affected tracts were the anterior frontal corpus callosum (ACC), the right VOF, the left inferior longitudinal fasciculus (ILF) and the left optic radiation.The results suggest that most white matter tissue properties are relatively robust to the early visual insult caused by strabismus. However, strabismic amblyopia does affect MD, not only in occipital tracts, such as the VOF and optic radiation, but also in long range association tracts connecting visual cortex to the frontal and temporal lobes (ILF) and connecting the two hemispheres (ACC).

    View details for DOI 10.1167/iovs.15-17097

    View details for PubMedID 26241402

    View details for PubMedCentralID PMC4525637

  • How to use fMRI functional localizers to improve EEG/MEG source estimation. Journal of neuroscience methods Cottereau, B. R., Ales, J. M., Norcia, A. M. 2015; 250: 64-73


    EEG and MEG have excellent temporal resolution, but the estimation of the neural sources that generate the signals recorded by the sensors is a difficult, ill-posed problem. The high spatial resolution of functional MRI makes it an ideal tool to improve the localization of the EEG/MEG sources using data fusion. However, the combination of the two techniques remains challenging, as the neural generators of the EEG/MEG and BOLD signals might in some cases be very different. Here we describe a data fusion approach that was developed by our team over the last decade in which fMRI is used to provide source constraints that are based on functional areas defined individually for each subject. This mini-review describes the different steps that are necessary to perform source estimation using this approach. It also provides a list of pitfalls that should be avoided when doing fMRI-informed EEG/MEG source imaging. Finally, it describes the advantages of using a ROI-based approach for group-level analysis and for the study of sensory systems.

    View details for DOI 10.1016/j.jneumeth.2014.07.015

    View details for PubMedID 25088693

    View details for PubMedCentralID PMC4383720

  • Predicting cortical dark/bright asymmetries from natural image statistics and early visual transforms. PLoS computational biology Cooper, E. A., Norcia, A. M. 2015; 11 (5)


    The nervous system has evolved in an environment with structure and predictability. One of the ubiquitous principles of sensory systems is the creation of circuits that capitalize on this predictability. Previous work has identified predictable non-uniformities in the distributions of basic visual features in natural images that are relevant to the encoding tasks of the visual system. Here, we report that the well-established statistical distributions of visual features--such as visual contrast, spatial scale, and depth--differ between bright and dark image components. Following this analysis, we go on to trace how these differences in natural images translate into different patterns of cortical input that arise from the separate bright (ON) and dark (OFF) pathways originating in the retina. We use models of these early visual pathways to transform natural images into statistical patterns of cortical input. The models include the receptive fields and non-linear response properties of the magnocellular (M) and parvocellular (P) pathways, with their ON and OFF pathway divisions. The results indicate that there are regularities in visual cortical input beyond those that have previously been appreciated from the direct analysis of natural images. In particular, several dark/bright asymmetries provide a potential account for recently discovered asymmetries in how the brain processes visual features, such as violations of classic energy-type models. On the basis of our analysis, we expect that the dark/bright dichotomy in natural images plays a key role in the generation of both cortical and perceptual asymmetries.

    View details for DOI 10.1371/journal.pcbi.1004268

    View details for PubMedID 26020624

    View details for PubMedCentralID PMC4447361

  • Predicting Cortical Dark/Bright Asymmetries from Natural Image Statistics and Early Visual Transforms PLOS COMPUTATIONAL BIOLOGY Cooper, E. A., Norcia, A. M. 2015; 11 (5)

    View details for DOI 10.1371/journal.pcbi.1004268

    View details for Web of Science ID 000356700200038

    View details for PubMedID 26020624

  • Maximally reliable spatial filtering of steady state visual evoked potentials. NeuroImage Dmochowski, J. P., Greaves, A. S., Norcia, A. M. 2015; 109: 63-72


    Due to their high signal-to-noise ratio (SNR) and robustness to artifacts, steady state visual evoked potentials (SSVEPs) are a popular technique for studying neural processing in the human visual system. SSVEPs are conventionally analyzed at individual electrodes or linear combinations of electrodes which maximize some variant of the SNR. Here we exploit the fundamental assumption of evoked responses - reproducibility across trials - to develop a technique that extracts a small number of high SNR, maximally reliable SSVEP components. This novel spatial filtering method operates on an array of Fourier coefficients and projects the data into a low-dimensional space in which the trial-to-trial spectral covariance is maximized. When applied to two sample data sets, the resulting technique recovers physiologically plausible components (i.e., the recovered topographies match the lead fields of the underlying sources) while drastically reducing the dimensionality of the data (i.e., more than 90% of the trial-to-trial reliability is captured in the first four components). Moreover, the proposed technique achieves a higher SNR than that of the single-best electrode or the Principal Components. We provide a freely-available MATLAB implementation of the proposed technique, herein termed "Reliable Components Analysis".

    View details for DOI 10.1016/j.neuroimage.2014.12.078

    View details for PubMedID 25579449

  • The effect of contrast polarity reversal on face detection: Evidence of perceptual asymmetry from sweep VEP VISION RESEARCH Liu-Shuang, J., Ales, J. M., Rossion, B., Norcia, A. M. 2015; 108: 8-19


    Contrast polarity inversion (i.e., turning dark regions light and vice versa) impairs face perception. We investigated the perceptual asymmetry between positive and negative polarity faces (matched for overall luminance) using a sweep VEP approach in the context of face detection (Journal of Vision 12 (2012) 1-18). Phase-scrambled face stimuli alternated at a rate of 3 Hz (6 images/s). The phase coherence of every other stimulus was parametrically increased so that a face gradually emerged over a 20-s stimulation sequence, leading to a 3 Hz response reflecting face detection. Contrary to the 6 Hz response, reflecting low-level visual processing, this 3 Hz response was larger and emerged earlier over right occipito-temporal channels for positive than negative polarity faces. Moreover, the 3 Hz response emerged abruptly to positive polarity faces, whereas it increased linearly for negative polarity faces. In another condition, alternating between a positive and a negative polarity face also elicited a strong 3 Hz response, indicating an asymmetrical representation of positive and negative polarity faces even at supra-threshold levels (i.e., when both stimuli were perceived as faces). Overall, these findings demonstrate distinct perceptual representations of positive and negative polarity faces, independently of low-level cues, and suggest qualitatively different detection processes (template-based matching for positive polarity faces vs. linear accumulation of evidence for negative polarity faces).

    View details for DOI 10.1016/j.visres.2015.01.001

    View details for Web of Science ID 000350781100002

    View details for PubMedID 25595857

  • Separable effects of inversion and contrast-reversal on face detection thresholds and response functions: A sweep VEP study JOURNAL OF VISION Liu-Shuang, J., Ales, J., Rossion, B., Norcia, A. M. 2015; 15 (2)


    The human brain rapidly detects faces in the visual environment. We recently presented a sweep visual evoked potential approach to objectively define face detection thresholds as well as suprathreshold response functions (Ales, Farzin, Rossion, & Norcia, 2012). Here we determined these parameters are affected by orientation (upright vs. inverted) and contrast polarity (positive vs. negative), two manipulations that disproportionately disrupt the perception of faces relative to other object categories. Face stimuli parametrically increased in visibility through phase-descrambling while alternating with scrambled images at a fixed presentation rate of 3 Hz (6 images/s). The power spectrum and mean luminance of all stimuli were equalized. As a face gradually emerged during a stimulation sequence, EEG responses at 3 Hz appeared at ≈35% phase coherence over right occipito-temporal channels, replicating previous observations. With inversion and contrast-reversal, the 3-Hz amplitude decreased by ≈20%-50% and the face detection threshold increased by ≈30%-60% coherence. Furthermore, while the 3-Hz response emerged abruptly and saturated quickly for normal faces, suggesting a categorical neural response, the response profile for inverted and negative polarity faces was shallower and more linear, indicating gradual and continuously increasing activation of the underlying neural population. These findings demonstrate that inversion and contrast-reversal increase the threshold and modulate the suprathreshold response function of face detection.

    View details for DOI 10.1167/15.2.11

    View details for Web of Science ID 000350679800011

    View details for PubMedID 25761329

  • Development of Three-Dimensional Perception in Human Infants ANNUAL REVIEW OF VISION SCIENCE, VOL 1 Norcia, A. M., Gerhard, H. E. 2015; 1: 569-594
  • A Representational Similarity Analysis of the Dynamics of Object Processing Using Single-Trial EEG Classification. PloS one Kaneshiro, B., Perreau Guimaraes, M., Kim, H., Norcia, A. M., Suppes, P. 2015; 10 (8)


    The recognition of object categories is effortlessly accomplished in everyday life, yet its neural underpinnings remain not fully understood. In this electroencephalography (EEG) study, we used single-trial classification to perform a Representational Similarity Analysis (RSA) of categorical representation of objects in human visual cortex. Brain responses were recorded while participants viewed a set of 72 photographs of objects with a planned category structure. The Representational Dissimilarity Matrix (RDM) used for RSA was derived from confusions of a linear classifier operating on single EEG trials. In contrast to past studies, which used pairwise correlation or classification to derive the RDM, we used confusion matrices from multi-class classifications, which provided novel self-similarity measures that were used to derive the overall size of the representational space. We additionally performed classifications on subsets of the brain response in order to identify spatial and temporal EEG components that best discriminated object categories and exemplars. Results from category-level classifications revealed that brain responses to images of human faces formed the most distinct category, while responses to images from the two inanimate categories formed a single category cluster. Exemplar-level classifications produced a broadly similar category structure, as well as sub-clusters corresponding to natural language categories. Spatiotemporal components of the brain response that differentiated exemplars within a category were found to differ from those implicated in differentiating between categories. Our results show that a classification approach can be successfully applied to single-trial scalp-recorded EEG to recover fine-grained object category structure, as well as to identify interpretable spatiotemporal components underlying object processing. Finally, object category can be decoded from purely temporal information recorded at single electrodes.

    View details for DOI 10.1371/journal.pone.0135697

    View details for PubMedID 26295970

  • The steady-state visual evoked potential in vision research: A review JOURNAL OF VISION Norcia, A. M., Appelbaum, L. G., Ales, J. M., Cottereau, B. R., Rossion, B. 2015; 15 (6)


    Periodic visual stimulation and analysis of the resulting steady-state visual evoked potentials were first introduced over 80 years ago as a means to study visual sensation and perception. From the first single-channel recording of responses to modulated light to the present use of sophisticated digital displays composed of complex visual stimuli and high-density recording arrays, steady-state methods have been applied in a broad range of scientific and applied settings.The purpose of this article is to describe the fundamental stimulation paradigms for steady-state visual evoked potentials and to illustrate these principles through research findings across a range of applications in vision science.

    View details for DOI 10.1167/15.6.4

    View details for Web of Science ID 000357858600004

    View details for PubMedID 26024451

  • Visuocortical Function in Infants With a History of Neonatal Jaundice INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Hou, C., Norcia, A. M., Madan, A., Good, W. V. 2014; 55 (10): 6443-6449


    High concentrations of unconjugated bilirubin are neurotoxic and cause brain damage in newborn infants. However, the exact level of bilirubin that may be neurotoxic in a given infant is unknown. The aim of this study was to use a quantitative measure of neural activity, the swept parameter visual evoked potential (sVEP) to determine the relationship between neonatal bilirubin levels and visual responsivity several months later.We compared sVEP response functions over a wide range of contrast, spatial frequency, and Vernier offset sizes in 16 full-term infants with high bilirubin levels (>10 mg/dL) and 18 age-matched infants with no visible neonatal jaundice, all enrolled at 14 to 22 weeks of age. The group means of sVEP thresholds and suprathreshold response amplitudes were compared. The correlation between individual sVEP thresholds and bilirubin levels in jaundiced infants was studied.Infants who had a history of neonatal jaundice showed lower response amplitudes (P < 0.05) and worse or immeasurable sVEP thresholds compared with control infants for all three measures (P < 0.05). Swept parameter visual evoked potential thresholds for Vernier offset were correlated with bilirubin level (P < 0.05), but spatial acuity and contrast sensitivity measures in the infants with neonatal jaundice were not (P > 0.05).These results indicate that elevated neonatal bilirubin levels affect measures of visual function in infancy up to at least 14 to 22 weeks of postnatal age.

    View details for DOI 10.1167/iovs.14-14261

    View details for Web of Science ID 000343147100042

    View details for PubMedID 25183760

    View details for PubMedCentralID PMC4197714

  • Dissociation of part-based and integrated neural responses to faces by means of electroencephalographic frequency tagging EUROPEAN JOURNAL OF NEUROSCIENCE Boremanse, A., Norcia, A. M., Rossion, B. 2014; 40 (6): 2987-2997

    View details for DOI 10.1111/ejn.12663

    View details for Web of Science ID 000342739000013

  • Perceived depth in natural images reflects encoding of low-level luminance statistics. journal of neuroscience Cooper, E. A., Norcia, A. M. 2014; 34 (35): 11761-11768


    Sighted animals must survive in an environment that is diverse yet highly structured. Neural-coding models predict that the visual system should allocate its computational resources to exploit regularities in the environment, and that this allocation should facilitate perceptual judgments. Here we use three approaches (natural scenes statistical analysis, a reanalysis of single-unit data from alert behaving macaque, and a behavioral experiment in humans) to address the question of how the visual system maximizes behavioral success by taking advantage of low-level regularities in the environment. An analysis of natural scene statistics reveals that the probability distributions for light increments and decrements are biased in a way that could be exploited by the visual system to estimate depth from relative luminance. A reanalysis of neurophysiology data from Samonds et al. (2012) shows that the previously reported joint tuning of V1 cells for relative luminance and binocular disparity is well matched to a predicted distribution of binocular disparities produced by natural scenes. Finally, we show that a percept of added depth can be elicited in images by exaggerating the correlation between luminance and depth. Together, the results from these three approaches provide further evidence that the visual system allocates its processing resources in a way that is driven by the statistics of the natural environment.

    View details for DOI 10.1523/JNEUROSCI.1336-14.2014

    View details for PubMedID 25164671

    View details for PubMedCentralID PMC4145177

  • Acuity-independent effects of visual deprivation on human visual cortex PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA Hou, C., Pettet, M. W., Norcia, A. M. 2014; 111 (30): E3120-E3128


    Visual development depends on sensory input during an early developmental critical period. Deviation of the pointing direction of the two eyes (strabismus) or chronic optical blur (anisometropia) separately and together can disrupt the formation of normal binocular interactions and the development of spatial processing, leading to a loss of stereopsis and visual acuity known as amblyopia. To shed new light on how these two different forms of visual deprivation affect the development of visual cortex, we used event-related potentials (ERPs) to study the temporal evolution of visual responses in patients who had experienced either strabismus or anisometropia early in life. To make a specific statement about the locus of deprivation effects, we took advantage of a stimulation paradigm in which we could measure deprivation effects that arise either before or after a configuration-specific response to illusory contours (ICs). Extraction of ICs is known to first occur in extrastriate visual areas. Our ERP measurements indicate that deprivation via strabismus affects both the early part of the evoked response that occurs before ICs are formed as well as the later IC-selective response. Importantly, these effects are found in the normal-acuity nonamblyopic eyes of strabismic amblyopes and in both eyes of strabismic patients without amblyopia. The nonamblyopic eyes of anisometropic amblyopes, by contrast, are normal. Our results indicate that beyond the well-known effects of strabismus on the development of normal binocularity, it also affects the early stages of monocular feature processing in an acuity-independent fashion.

    View details for DOI 10.1073/pnas.1404361111

    View details for Web of Science ID 000339500200013

    View details for PubMedID 25024230

    View details for PubMedCentralID PMC4121844

  • The evolution of a disparity decision in human visual cortex. NeuroImage Cottereau, B. R., Ales, J. M., Norcia, A. M. 2014; 92: 193-206


    We used fMRI-informed EEG source-imaging in humans to characterize the dynamics of cortical responses during a disparity-discrimination task. After the onset of a disparity-defined target, decision-related activity was found within an extended cortical network that included several occipital regions of interest (ROIs): V4, V3A, hMT+ and the Lateral Occipital Complex (LOC). By using a response-locked analysis, we were able to determine the timing relationships in this network of ROIs relative to the subject's behavioral response. Choice-related activity appeared first in the V4 ROI almost 200 ms before the button press and then subsequently in the V3A ROI. Modeling of the responses in the V4 ROI suggests that this area provides an early contribution to disparity discrimination. Choice-related responses were also found after the button-press in ROIs V4, V3A, LOC and hMT+. Outside the visual cortex, choice-related activity was found in the frontal and temporal poles before the button-press. By combining the spatial resolution of fMRI-informed EEG source imaging with the ability to sort out neural activity occurring before, during and after the behavioral manifestation of the decision, our study is the first to assign distinct functional roles to the extra-striate ROIs involved in perceptual decisions based on disparity, the primary cue for depth.

    View details for DOI 10.1016/j.neuroimage.2014.01.055

    View details for PubMedID 24513152

  • Dynamics and cortical distribution of neural responses to 2D and 3D motion in human. Journal of neurophysiology Cottereau, B. R., McKee, S. P., Norcia, A. M. 2014; 111 (3): 533-543


    The perception of motion-in-depth is important for avoiding collisions and for the control of vergence eye-movements and other motor actions. Previous psychophysical studies have suggested that sensitivity to motion-in-depth has a lower temporal processing limit than the perception of lateral motion. The present study used functional MRI-informed EEG source-imaging to study the spatiotemporal properties of the responses to lateral motion and motion-in-depth in human visual cortex. Lateral motion and motion-in-depth displays comprised stimuli whose only difference was interocular phase: monocular oscillatory motion was either in-phase in the two eyes (lateral motion) or in antiphase (motion-in-depth). Spectral analysis was used to break the steady-state visually evoked potentials responses down into even and odd harmonic components within five functionally defined regions of interest: V1, V4, lateral occipital complex, V3A, and hMT+. We also characterized the responses within two anatomically defined regions: the inferior and superior parietal cortex. Even harmonic components dominated the evoked responses and were a factor of approximately two larger for lateral motion than motion-in-depth. These responses were slower for motion-in-depth and were largely independent of absolute disparity. In each of our regions of interest, responses at odd-harmonics were relatively small, but were larger for motion-in-depth than lateral motion, especially in parietal cortex, and depended on absolute disparity. Taken together, our results suggest a plausible neural basis for reduced psychophysical sensitivity to rapid motion-in-depth.

    View details for DOI 10.1152/jn.00549.2013

    View details for PubMedID 24198326

  • Flies and humans share a motion estimation strategy that exploits natural scene statistics. Nature neuroscience Clark, D. A., Fitzgerald, J. E., Ales, J. M., Gohl, D. M., Silies, M. A., Norcia, A. M., Clandinin, T. R. 2014; 17 (2): 296-303


    Sighted animals extract motion information from visual scenes by processing spatiotemporal patterns of light falling on the retina. The dominant models for motion estimation exploit intensity correlations only between pairs of points in space and time. Moving natural scenes, however, contain more complex correlations. We found that fly and human visual systems encode the combined direction and contrast polarity of moving edges using triple correlations that enhance motion estimation in natural environments. Both species extracted triple correlations with neural substrates tuned for light or dark edges, and sensitivity to specific triple correlations was retained even as light and dark edge motion signals were combined. Thus, both species separately process light and dark image contrasts to capture motion signatures that can improve estimation accuracy. This convergence argues that statistical structures in natural scenes have greatly affected visual processing, driving a common computational strategy over 500 million years of evolution.

    View details for DOI 10.1038/nn.3600

    View details for PubMedID 24390225

  • An objective index of individual face discrimination in the right occipito-temporal cortex by means of fast periodic oddball stimulation. Neuropsychologia Liu-Shuang, J., Norcia, A. M., Rossion, B. 2014; 52: 57-72


    We introduce an approach based on fast periodic oddball stimulation that provides objective, high signal-to-noise ratio (SNR), and behavior-free measures of the human brain's discriminative response to complex visual patterns. High-density electroencephalogram (EEG) was recorded for human observers presented with 60s sequences containing a base-face (A) sinusoidally contrast-modulated at a frequency of 5.88Hz (F), with face size varying every cycle. Different oddball-faces (B, C, D…) were introduced at fixed intervals (every 4 stimuli=F/5=1.18Hz: AAAABAAAACAAAAD…). Individual face discrimination was indexed by responses at this 1.18Hz oddball frequency. Following only 4min of recording, significant responses emerged at exactly 1.18Hz and its harmonics (e.g., 2F/5=2.35Hz, 3F/5=3.53Hz…), with up to a 300% signal increase over the right occipito-temporal cortex. This response was present in all participants, for both color and greyscale faces, providing a robust implicit neural measure of individual face discrimination. Face inversion or contrast-reversal did not affect the basic 5.88Hz periodic response over medial occipital channels. However, these manipulations substantially reduced the 1.18Hz oddball discrimination response over the right occipito-temporal region, indicating that this response reflects high-level processes that are partly face-specific. These observations indicate that fast periodic oddball stimulation can be used to rapidly and objectively characterize the discrimination of visual patterns and may become invaluable in characterizing this process in typical adult, developmental, and neuropsychological patient populations.

    View details for DOI 10.1016/j.neuropsychologia.2013.10.022

    View details for PubMedID 24200921

  • Electrophysiological measures of low-level vision reveal spatial processing deficits and hemispheric asymmetry in autism spectrum disorder. Journal of vision Pei, F., Baldassi, S., Norcia, A. M. 2014; 14 (11)


    There is accumulating evidence from electrophysiological studies that low-level visual processing is atypical in individuals with autism spectrum disorders (ASDs). Abnormalities in early stages of sensory processing are of interest because they could lead to downstream functional deficits in social or cognitive domains. Using steady-state visual evoked potentials (SSVEPs), we studied how well spatial information is transmitted over a wide range of spatial frequencies (2-30 cycles/deg), including those at the limit of visibility (visual acuity). SSVEPs were recorded over 128 channels in 16 ASD participants between 5 and 17 years old and 17 age-matched, neurotypical (NT) participants. We observed a selective reduction of the amplitude of the SSVEP second harmonic pattern reversal response between 5 and 17 cycles/deg. Responses measured at the fourth harmonic were normal at all spatial frequencies tested, as were responses at the lowest and highest spatial frequencies at the second harmonic. The reduction of second harmonic responses occurred preferentially over right occipital electrodes. Because response abnormalities are restricted to a specific response harmonic and to specific ranges of spatial frequency, we can rule out nonspecific differences between the ASD participants and the NT controls. This particular pattern of loss, combined with the observed exaggeration of the loss over the right hemisphere, suggests that a highly specific neural substrate early in the visual pathway is compromised in ASD.

    View details for DOI 10.1167/14.11.3

    View details for PubMedID 25194015

  • Linking perception to neural activity as measured by visual evoked potentials VISUAL NEUROSCIENCE Norcia, A. M. 2013; 30 (5-6): 223-227


    Linking propositions have played an important role in refining our understanding of the relationship between neural activity and perception. Over the last 40 years, visual evoked potentials (VEPs) have been used in many different ways to address questions of the relationship between neural activity and perception. This review organizes and discusses this research within the linking proposition framework developed by Davida Teller, and her colleagues. A series of examples from the VEP literature illustrates each of the five classes of linking propositions originally proposed by Davida Teller. The related concept of the bridge locus-the site at which neural activity can be said to first be proscriptive of perception-is discussed and a suggestion is made that the concept be expanded to include an evolution over time and cortical area.

    View details for DOI 10.1017/S0952523813000205

    View details for Web of Science ID 000330363700006

    View details for PubMedID 23879990

  • The 6 Hz fundamental stimulation frequency rate for individual face discrimination in the right occipito-temporal cortex NEUROPSYCHOLOGIA Alonso-Prieto, E., Van Belle, G., Liu-Shuang, J., Norcia, A. M., Rossion, B. 2013; 51 (13): 2863-2875


    What is the stimulus presentation rate at which the human brain can discriminate each exemplar of a familiar visual category? We presented faces at 14 frequency rates (1.0-16.66 Hz) to human observers while recording high-density electroencephalogram (EEG). Different face exemplars elicited a larger steady-state visual evoked (ssVEP) response than when the same face was repeated, but only for stimulation frequencies between 4 and 8.33 Hz, with a maximal difference at 5.88 Hz (170 ms cycle). The effect was confined to the exact stimulation frequency and localized over the right occipito-temporal cortex. At high frequency rates (>10 Hz), the response to different and identical exemplars did not differ, suggesting that the fine-grained analysis needed for individual face discrimination cannot be completed before the next face interrupts, or competes, with the processed face. At low rates (<3 Hz), repetition suppression could not be identified at the stimulation frequency, suggesting that the neural response to an individual face is temporally dispersed and distributed over different processes. These observations indicate that at a temporal rate of 170 ms (6 faces/s) the face perception network is able to fully discriminate between each individual face presented, providing information about the temporal bottleneck of individual face discrimination in humans. These results also have important practical implications for optimizing paradigms that rely on repetition suppression, and open an avenue for investigating complex visual processes at an optimal range of stimulation frequency rates.

    View details for DOI 10.1016/j.neuropsychologia.2013.08.018

    View details for Web of Science ID 000328869400040

    View details for PubMedID 24007879

  • The time course of shape discrimination in the human brain NEUROIMAGE Ales, J. M., Appelbaum, L. G., Cottereau, B. R., Norcia, A. M. 2013; 67: 77-88


    The lateral occipital cortex (LOC) activates selectively to images of intact objects versus scrambled controls, is selective for the figure-ground relationship of a scene, and exhibits at least some degree of invariance for size and position. Because of these attributes, it is considered to be a crucial part of the object recognition pathway. Here we show that human LOC is critically involved in perceptual decisions about object shape. High-density EEG was recorded while subjects performed a threshold-level shape discrimination task on texture-defined figures segmented by either phase or orientation cues. The appearance or disappearance of a figure region from a uniform background generated robust visual evoked potentials throughout retinotopic cortex as determined by inverse modeling of the scalp voltage distribution. Contrasting responses from trials containing shape changes that were correctly detected (hits) with trials in which no change occurred (correct rejects) revealed stimulus-locked, target-selective activity in the occipital visual areas LOC and V4 preceding the subject's response. Activity that was locked to the subjects' reaction time was present in the LOC. Response-locked activity in the LOC was determined to be related to shape discrimination for several reasons: shape-selective responses were silenced when subjects viewed identical stimuli but their attention was directed away from the shapes to a demanding letter discrimination task; shape-selectivity was present across four different stimulus configurations used to define the figure; LOC responses correlated with participants' reaction times. These results indicate that decision-related activity is present in the LOC when subjects are engaged in threshold-level shape discriminations.

    View details for DOI 10.1016/j.neuroimage.2012.10.044

    View details for Web of Science ID 000314144600008

    View details for PubMedID 23116814

  • On determining the intracranial sources of visual evoked potentials from scalp topography: A reply to Kelly et al. (this issue) NEUROIMAGE Ales, J. M., Yates, J. L., Norcia, A. M. 2013; 64: 703-711


    The cruciform model posits that if a Visual Evoked Potential component originates in cortical area V1, then stimuli placed in the upper versus lower visual field will generate responses with opposite polarity at the scalp. In our original paper (Ales et al., 2010b) we showed that the cruciform model provides an insufficient criterion for identifying V1 sources. This conclusion was reached on the basis of simulations that used realistic 3D models of early visual areas to simulate scalp topographies expected for stimuli of different sizes and shapes placed in different field locations. The simulations indicated that stimuli placed in the upper and lower visual field produce polarity inverting scalp topographies for activation of areas V2 and V3, but not for area V1. As a consequence of the non-uniqueness of the polarity inversion criterion, we suggested that past studies using the cruciform model had not adequately excluded contributions from sources outside V1. In their comment on our paper, Kelly et al. (this issue) raise several concerns with this suggestion. They claim that our initial results did not use the proper stimulus locations to constitute a valid test of the cruciform model. Kelly et al., also contend that the cortical source of the initial visually evoked component (C1) can be identified based on latency and polarity criteria derived from intracranial recordings in non-human primates. In our reply we show that simulations using the suggested critical stimulus locations are consistent with our original findings and thus do not change our conclusions regarding the use of the polarity inversion criterion. We further show that the anatomical assumptions underlying the putatively optimal locations are not consistent with available V1 anatomical data. We then address the non-human primate data, describing how differences in stimuli across studies and species confound an effective utilization of the non-human primate data for interpreting human evoked potential responses. We also show that, considered more broadly, the non-human primate literature shows that multiple visual areas onset simultaneously with V1. We suggest several directions for future research that will further clarify how to make the best use of scalp data for inferring cortical sources.

    View details for DOI 10.1016/j.neuroimage.2012.09.009

    View details for Web of Science ID 000312504200066

    View details for PubMedID 22982584

  • Assessment of OLED displays for vision research. Journal of vision Cooper, E. A., Jiang, H., Vildavski, V., Farrell, J. E., Norcia, A. M. 2013; 13 (12)


    Vision researchers rely on visual display technology for the presentation of stimuli to human and nonhuman observers. Verifying that the desired and displayed visual patterns match along dimensions such as luminance, spectrum, and spatial and temporal frequency is an essential part of developing controlled experiments. With cathode-ray tubes (CRTs) becoming virtually unavailable on the commercial market, it is useful to determine the characteristics of newly available displays based on organic light emitting diode (OLED) panels to determine how well they may serve to produce visual stimuli. This report describes a series of measurements summarizing the properties of images displayed on two commercially available OLED displays: the Sony Trimaster EL BVM-F250 and PVM-2541. The results show that the OLED displays have large contrast ratios, wide color gamuts, and precise, well-behaved temporal responses. Correct adjustment of the settings on both models produced luminance nonlinearities that were well predicted by a power function ("gamma correction"). Both displays have adjustable pixel independence and can be set to have little to no spatial pixel interactions. OLED displays appear to be a suitable, or even preferable, option for many vision research applications.

    View details for DOI 10.1167/13.12.16

    View details for PubMedID 24155345

    View details for PubMedCentralID PMC3807585

  • An objective signature for visual binding of face parts in the human brain. Journal of vision Boremanse, A., Norcia, A. M., Rossion, B. 2013; 13 (11)


    Whether and how the parts of a visual object are grouped together to form an integrated ("holistic") representation is a central question in cognitive neuroscience. Although the face is considered to be the quintessential example of holistic representation, this issue has been the subject of much debate in face perception research. The implication of holistic processing is that the response to the whole cannot be predicted from the sum of responses to the parts. Here we apply techniques from nonlinear systems analysis to provide an objective measure of the nonlinear integration of parts into a whole, using the left and right halves of a face stimulus as the parts. High-density electroencephalogram (EEG) was recorded in 15 human participants presented with two halves of a face stimulus, flickering at different frequencies (5.88 vs. 7.14 Hz). Besides specific responses at these fundamental frequencies, reflecting part-based responses, we found intermodulation components (e.g., 7.14 - 5.88 = 1.26 Hz) over the right occipito-temporal hemisphere, reflecting nonlinear integration of the face halves. Part-based responses did not depend on the relative alignment of the two face halves, their spatial separation, or whether the face was presented upright or inverted. By contrast, intermodulations were virtually absent when the two halves were spatially misaligned and separated. Inversion of the whole face configuration also reduced specifically the intermodulation components over the right occipito-temporal cortex. These observations indicate that the intermodulation components constitute an objective, configuration-specific signature of an emergent neural representation of the whole face that is distinct from that generated by the parts themselves.

    View details for DOI 10.1167/13.11.6

    View details for PubMedID 24023273

  • Increasing the accuracy of electromagnetic inverses using functional area source correlation constraints HUMAN BRAIN MAPPING Cottereau, B. R., Ales, J. M., Norcia, A. M. 2012; 33 (11): 2694-2713


    Estimating cortical current distributions from electroencephalographic (EEG) or magnetoencephalographic data is a difficult inverse problem whose solution can be improved by the addition of priors on the associated neural responses. In the context of visual activation studies, we propose a new approach that uses a functional area constrained estimator (FACE) to increase the accuracy of the reconstructions. It derives the source correlation matrix from a segmentation of the cortex into areas defined by retinotopic maps of the visual field or by functional localizers obtained independently by fMRI. These areas are computed once for each individual subject and the associated estimators can therefore be reused for any new study on the same participant. The resulting FACE reconstructions emphasize the activity of sources within these areas or enforce their intercorrelations. We used realistic Monte-Carlo simulations to demonstrate that this approach improved our estimates of a diverse set of source configurations. Reconstructions obtained from a real EEG dataset demonstrate that our priors improve the localization of the cortical areas involved in horizontal disparity processing.

    View details for DOI 10.1002/hbm.21394

    View details for Web of Science ID 000310264500015

    View details for PubMedID 21938755

    View details for PubMedCentralID PMC3637966

  • Spatial Contrast Sensitivity Vision Loss in Children with Cortical Visual Impairment INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Good, W. V., Hou, C., Norcia, A. M. 2012; 53 (12): 7730-7734


    Although cortical visual impairment (CVI) is the leading cause of bilateral vision impairment in children in Western countries, little is known about the effects of CVI on visual function. The aim of this study was to compare visual evoked potential measures of contrast sensitivity and grating acuity in children with CVI with those of age-matched typically developing controls.The swept parameter visual evoked potential (sVEP) was used to measure contrast sensitivity and grating acuity in 34 children with CVI at 5 months to 5 years of age and in 16 age-matched control children. Contrast thresholds and spatial frequency thresholds (grating acuities) were derived by extrapolating the tuning functions to zero amplitude. These thresholds and maximal suprathreshold response amplitudes were compared between groups.Among 34 children with CVI, 30 had measurable but reduced contrast sensitivity with a median threshold of 10.8% (range 5.0%-30.0% Michelson), and 32 had measurable but reduced grating acuity with median threshold 0.49 logMAR (9.8 c/deg, range 5-14 c/deg). These thresholds were significantly reduced, compared with age-matched control children. In addition, response amplitudes over the entire sweep range for both measures were significantly diminished in children with CVI compared with those of control children.Our results indicate that spatial contrast sensitivity and response amplitudes are strongly affected by CVI. The substantial degree of loss in contrast sensitivity suggests that contrast is a sensitive measure for evaluating vision deficits in patients with CVI.

    View details for DOI 10.1167/iovs.12-9775

    View details for Web of Science ID 000313053500043

    View details for PubMedID 23060143

  • Visual gain control abnormalities in Autism Spectrum Disorders 16th World Congress of Psychophysiology of the International Organization of Psychophysiology (IOP) Pei, F., Baldassi, S., Norcia, A. ELSEVIER SCIENCE BV. 2012: 298–99
  • Bridging the gap: global disparity processing in the human visual cortex JOURNAL OF NEUROPHYSIOLOGY Cottereau, B. R., McKee, S. P., Norcia, A. M. 2012; 107 (9): 2421-2429


    The human stereoscopic system is remarkable in its ability to utilize widely separated features as references to support fine depth discrimination. In a search for possible neural substrates of this ability, we recorded high-density EEG and used a distributed inverse technique to estimate population-level disparity responses in five regions of interest (ROIs): V1, V3A, hMT+, V4, and lateral occipital complex (LOC). The stimulus was a central modulating disk surrounded by a correlated "reference" annulus presented in the fixation plane. We varied a gap separating the disk from the annulus parametrically from 0 to 5.5° as a test of long-range disparity integration. In the V1, LOC, and hMT+ ROIs, the responses with gaps >0.5° were equal to those obtained in a control condition where the surround was composed of uncorrelated noise (no reference). By contrast, in the V4 and V3A ROIs, responses with gaps as large as 5.5° were still significantly higher than the control. As a test of the spatial distribution of the disparity reference information, we manipulated the properties of the stimulus by placing noise between the center and the surround or throughout the surround. The V3A ROI was particularly sensitive to disparity noise between the center and annulus regions, suggesting an important contribution of disparity edge detectors in this ROI.

    View details for DOI 10.1152/jn.01051.2011

    View details for Web of Science ID 000303600900011

    View details for PubMedID 22323636

    View details for PubMedCentralID PMC3362244

  • The Time Course of Segmentation and Cue-Selectivity in the Human Visual Cortex PLOS ONE Appelbaum, L. G., Ales, J. M., Norcia, A. M. 2012; 7 (3)


    Texture discontinuities are a fundamental cue by which the visual system segments objects from their background. The neural mechanisms supporting texture-based segmentation are therefore critical to visual perception and cognition. In the present experiment we employ an EEG source-imaging approach in order to study the time course of texture-based segmentation in the human brain. Visual Evoked Potentials were recorded to four types of stimuli in which periodic temporal modulation of a central 3° figure region could either support figure-ground segmentation, or have identical local texture modulations but not produce changes in global image segmentation. The image discontinuities were defined either by orientation or phase differences across image regions. Evoked responses to these four stimuli were analyzed both at the scalp and on the cortical surface in retinotopic and functional regions-of-interest (ROIs) defined separately using fMRI on a subject-by-subject basis. Texture segmentation (tsVEP: segmenting versus non-segmenting) and cue-specific (csVEP: orientation versus phase) responses exhibited distinctive patterns of activity. Alternations between uniform and segmented images produced highly asymmetric responses that were larger after transitions from the uniform to the segmented state. Texture modulations that signaled the appearance of a figure evoked a pattern of increased activity starting at ∼143 ms that was larger in V1 and LOC ROIs, relative to identical modulations that didn't signal figure-ground segmentation. This segmentation-related activity occurred after an initial response phase that did not depend on the global segmentation structure of the image. The two cue types evoked similar tsVEPs up to 230 ms when they differed in the V4 and LOC ROIs. The evolution of the response proceeded largely in the feed-forward direction, with only weak evidence for feedback-related activity.

    View details for DOI 10.1371/journal.pone.0034205

    View details for Web of Science ID 000303894900070

    View details for PubMedID 22479566

  • Dynamics of Normalization Underlying Masking in Human Visual Cortex JOURNAL OF NEUROSCIENCE Tsai, J. J., Wade, A. R., Norcia, A. M. 2012; 32 (8): 2783-2789


    Stimulus visibility can be reduced by other stimuli that overlap the same region of visual space, a process known as masking. Here we studied the neural mechanisms of masking in humans using source-imaged steady state visual evoked potentials and frequency-domain analysis over a wide range of relative stimulus strengths of test and mask stimuli. Test and mask stimuli were tagged with distinct temporal frequencies and we quantified spectral response components associated with the individual stimuli (self terms) and responses due to interaction between stimuli (intermodulation terms). In early visual cortex, masking alters the self terms in a manner consistent with a reduction of input contrast. We also identify a novel signature of masking: a robust intermodulation term that peaks when the test and mask stimuli have equal contrast and disappears when they are widely different. We fit all of our data simultaneously with family of a divisive gain control models that differed only in their dynamics. Models with either very short or very long temporal integration constants for the gain pool performed worse than a model with an integration time of ∼30 ms. Finally, the absolute magnitudes of the response were controlled by the ratio of the stimulus contrasts, not their absolute values. This contrast-contrast invariance suggests that many neurons in early visual cortex code relative rather than absolute contrast. Together, these results provide a more complete description of masking within the normalization framework of contrast gain control and suggest that contrast normalization accomplishes multiple functional goals.

    View details for DOI 10.1523/JNEUROSCI.4485-11.2012

    View details for Web of Science ID 000300716600021

    View details for PubMedID 22357861

    View details for PubMedCentralID PMC3337145

  • Disparity-Specific Spatial Interactions: Evidence from EEG Source Imaging JOURNAL OF NEUROSCIENCE Cottereau, B. R., McKee, S. P., Ales, J. M., Norcia, A. M. 2012; 32 (3): 826-840


    Using cortical source estimation techniques based on high-density EEG and fMRI measurements in humans, we measured how a disparity-defined surround influenced the responses to the changing disparity of a central disk within five visual ROIs: V1, V4, lateral occipital complex (LOC), hMT+, and V3A. The responses in the V1 ROI were not consistently affected either by changes in the characteristics of the surround (correlated or uncorrelated) or by its disparity value, consistent with V1 being responsive only to absolute, not relative, disparity. Correlation in the surround increased the responses in the V4, LOC, and hMT+ ROIs over those measured with the uncorrelated surround. Thus, these extrastriate areas contain neurons that are sensitive to disparity differences. However, their evoked responses did not vary systematically with the surround disparity. Responses in the V3A ROI, in contrast, were increased by correlation in the surround and varied with its disparity. We modeled these V3A responses as attributable to a gain modulation of the absolute disparity response, where the gain amplitude is proportional to the center-surround disparity difference. An additional experiment identified a nonlinear center-surround interaction in V3A that facilitates the responses when center and surround are misaligned but suppresses it when they share the same disparity plane.

    View details for DOI 10.1523/JNEUROSCI.2709-11.2012

    View details for Web of Science ID 000299324900008

    View details for PubMedID 22262881

    View details for PubMedCentralID PMC3313655

  • Distinct effects of attention on the neural responses to form and motion processing: A SSVEP source-imaging study JOURNAL OF VISION Palomares, M., Ales, J. M., Wade, A. R., Cottereau, B. R., Norcia, A. M. 2012; 12 (10)


    We measured neural responses to local and global aspects of form and motion stimuli using frequency-tagged, steady-state visual evoked potentials (SSVEPs) combined with magnetic resonance imaging (MRI) data. Random dot stimuli were used to portray either dynamic Glass patterns (Glass, 1969) or coherent motion displays. SSVEPs were used to estimate neural activity in a set of fMRI-defined visual areas in each subject. To compare activity associated with local versus global processing, we analyzed two frequency components of the SSVEP in each visual area: the high temporal frequency at which the local dots were updated (30 Hz) and the much lower frequency corresponding to updates in the global structure (0.83 Hz). Local and global responses were evaluated in the context of two different behavioral tasks--subjects had to either direct their attention toward or away from the global coherence of the stimuli. The data show that the effect of attention on global and local responses is both stimulus and visual area dependent. When attention was directed away from stimulus coherence, both local and global responses were higher in the coherent motion than Glass pattern condition. Directing attention to coherence in Glass patterns enhanced global activity in areas LOC, hMT+, V4, V3a, and V1, while attention to global motion modulated responses by a smaller amount in a smaller set of areas: V4, hMT+, and LOC. In contrast, directing attention towards stimulus coherence weakly increased local responses to both coherent motion and Glass patterns. These results suggest that visual attention differentially modulates the activity of early visual areas at both local and global levels of structural encoding.

    View details for DOI 10.1167/12.10.15

    View details for Web of Science ID 000313886400015

    View details for PubMedID 23019120

  • Effect of Grade I and II Intraventricular Hemorrhage on Visuocortical Function in Very Low Birth Weight Infants SEEING AND PERCEIVING Madan, A., Norcia, A. M., Hou, C., Pettet, M. W., Good, W. V. 2012; 25 (2): 143-154


    The neurological outcome for infants with Grade I/II intraventricular hemorrhage (IVH) is debated. The aim of this study was to determine whether very low birth weight infants (VLBW, <1500 g) with Grade I/II (IVH) have altered visuocortical activity compared with infants with no IVH. We assessed the quantitative swept parameter visual evoked potential (sVEP) responses evoked by three different visual stimuli. Data from 52 VLBW infants were compared with data from 13 infants with Grade I or II IVH, enrolled at 5-7 months corrected age. Acuity thresholds and suprathreshold response amplitudes were compared. Grating acuity (GA), contrast sensitivity (CS) and vernier acuity (VA) were each worse in the Grade I/II IVH compared with the no IVH groups (8.24 cpd in IVH group vs. 13.07 cpd in no IVH group for GA; 1.44% vs. 1.18% for CS and 1.55 arcmin vs. 0.58 arcmin for VA). The slopes of the response amplitude for CS and VA were significantly lower in IVH infants. The spatial frequency tuning function was shifted downward on the spatial frequency axis, without a change in slope. These results indicate that Grade I/II IVH are associated with deleterious effects on cortical vision development and function.

    View details for DOI 10.1163/187847612X626381

    View details for Web of Science ID 000303371100002

    View details for PubMedID 22371027

  • An objective method for measuring face detection thresholds using the sweep steady-state visual evoked response JOURNAL OF VISION Ales, J. M., Farzin, F., Rossion, B., Norcia, A. M. 2012; 12 (10)


    We introduce a sensitive method for measuring face detection thresholds rapidly, objectively, and independently of low-level visual cues. The method is based on the swept parameter steady-state visual evoked potential (ssVEP), in which a stimulus is presented at a specific temporal frequency while parametrically varying ("sweeping") the detectability of the stimulus. Here, the visibility of a face image was increased by progressive derandomization of the phase spectra of the image in a series of equally spaced steps. Alternations between face and fully randomized images at a constant rate (3/s) elicit a robust first harmonic response at 3 Hz specific to the structure of the face. High-density EEG was recorded from 10 human adult participants, who were asked to respond with a button-press as soon as they detected a face. The majority of participants produced an evoked response at the first harmonic (3 Hz) that emerged abruptly between 30% and 35% phase-coherence of the face, which was most prominent on right occipito-temporal sites. Thresholds for face detection were estimated reliably in single participants from 15 trials, or on each of the 15 individual face trials. The ssVEP-derived thresholds correlated with the concurrently measured perceptual face detection thresholds. This first application of the sweep VEP approach to high-level vision provides a sensitive and objective method that could be used to measure and compare visual perception thresholds for various object shapes and levels of categorization in different human populations, including infants and individuals with developmental delay.

    View details for DOI 10.1167/12.10.18

    View details for Web of Science ID 000313886400018

    View details for PubMedID 23024355

  • Piecing it together: Infants' neural responses to face and object structure JOURNAL OF VISION Farzin, F., Hou, C., Norcia, A. M. 2012; 12 (13)


    Integration of local elements into a coherent global form is a fundamental aspect of visual object recognition. How the different hierarchically organized stages of visual analysis develop in order to support object representation in infants remains unknown. The aim of this study was to investigate structural encoding of natural images in 4- to 6-month-old infants and adults. We used the steady-state visual evoked potential (ssVEP) technique to measure cortical responses specific to the global structure present in object and face images, and assessed whether differential responses were present for these image categories. This study is the first to apply the ssVEP method to high-level vision in infants. Infants and adults responded to the structural relations present in both image categories, and topographies of the responses differed based on image category. However, while adult responses to face and object structure were localized over occipitotemporal scalp areas, only infant face responses were distributed over temporal regions. Therefore, both infants and adults show object category specificity in their neural responses. The topography of the infant response distributions indicates that between 4 and 6 months of age, structure encoding of faces occurs at a higher level of processing than that of objects.

    View details for DOI 10.1167/12.13.6

    View details for Web of Science ID 000313888000006

    View details for PubMedID 23220577

    View details for PubMedCentralID PMC3587016

  • Visual Cortical Function in Very Low Birth Weight Infants without Retinal or Cerebral Pathology INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Hou, C., Norcia, A. M., Madan, A., Tith, S., Agarwal, R., Good, W. V. 2011; 52 (12): 9091-9098


    Preterm infants are at high risk of visual and neural developmental deficits. However, the development of visual cortical function in preterm infants with no retinal or neurologic morbidity has not been well defined. To determine whether premature birth itself alters visual cortical function, swept parameter visual evoked potential (sVEP) responses of healthy preterm infants were compared with those of term infants.Fifty-two term infants and 58 very low birth weight (VLBW) infants without significant retinopathy of prematurity or neurologic morbidities were enrolled. Recruited VLBW infants were between 26 and 33 weeks of gestational age, with birth weights of less than 1500 g. Spatial frequency, contrast, and vernier offset sweep VEP tuning functions were measured at 5 to 7 months' corrected age. Acuity and contrast thresholds were derived by extrapolating the tuning functions to 0 amplitude. These thresholds and suprathreshold response amplitudes were compared between groups.Preterm infants showed increased thresholds (indicating decreased sensitivity to visual stimuli) and reductions in amplitudes for all three measures. These changes in cortical responsiveness were larger in the <30 weeks ' gestational age subgroup than in the ≥30 weeks' gestational age subgroup.Preterm infants with VLBW had measurable and significant changes in cortical responsiveness that were correlated with gestational age. These results suggest that premature birth in the absence of identifiable retinal or neurologic abnormalities has a significant effect on visual cortical sensitivity at 5 to 7 months' of corrected age and that gestational age is an important factor in visual development.

    View details for DOI 10.1167/iovs.11-7458

    View details for Web of Science ID 000297631400043

    View details for PubMedID 22025567

  • Distinct cortical responses to 2D figures defined by motion contrast VISION RESEARCH Fesi, J. D., Yannes, M. P., Brinckman, D. D., Norcia, A. M., Ales, J. M., Gilmore, R. O. 2011; 51 (19): 2110-2120


    Motion contrast contributes to the segregation of a two-dimensional figure from its background, yet many questions remain about its neural mechanisms. We measured steady-state visual evoked potential (SSVEP) responses to moving dot displays in which figure regions emerged from and disappeared into the background at a specific temporal frequency (1.2Hz, F1), based on regional differences of dot direction and global direction coherence. The goal was to measure the cortical response function across a range of motion contrast magnitudes. In two experiments using both a low channel count electrode array (Experiment 1) and a high density array (Experiment 2), we observed two distinct phase-locked evoked responses that were similar across motion contrast type. A response at 1.2Hz (1F1) increased in amplitude with increasing magnitudes of direction or coherence contrast. A response at 2.4Hz (2F1) increased in amplitude, but saturated at low levels of direction or coherence contrast. The two components showed different scalp distributions - the 1F1 was strongest along medial occipital channels, while the 2F1 was bilaterally distributed. Taken together, the studies suggest that figures defined by different types of motion contrast are processed by cortical systems with similar dynamics, and that there are separable neural systems devoted to (i) signaling the absolute magnitude of motion contrast and (ii) detecting when a figure defined by motion contrast appears and disappears from view.

    View details for DOI 10.1016/j.visres.2011.07.015

    View details for Web of Science ID 000295489200002

    View details for PubMedID 21820002

  • Contrast Gain Control Abnormalities in Idiopathic Generalized Epilepsy ANNALS OF NEUROLOGY Tsai, J. J., Norcia, A. M., Ales, J. M., Wade, A. R. 2011; 70 (4): 574-582


    The origin of neural hyperexcitability underlying idiopathic generalized epilepsy (IGE) is not known. The objective of this study is to identify evidence of hyperexcitability in precisely measured visual evoked responses and to understand the nature of changes in excitation and inhibition that lead to altered responses in human patients with IGE.Steady-state visual-evoked potentials (VEPs) to contrast reversing gratings were recorded over a wide range of stimulus contrast. VEPs were analyzed at the pattern reversal rate using spectral analysis. Ten patients with IGE and 13 healthy subjects participated. All subjects had normal visual acuity and had no history of photic-induced seizures or photoparoxysmal electroencephalograph (EEG) activity.At a group level, the amplitude of visual responses did not saturate at high stimulus contrast in patients, as it did in the control subjects. This reflects an abnormality in neuronal gain control. The VEPs did not have sufficient power to reliably distinguish patients from controls at an individual level. Parametric modeling using a standard gain control framework showed that the abnormality lay in reduced inhibition from neighboring neurons rather than increased excitatory response to the stimulus.Visual evoked responses reveal changes in a fundamental mechanism regulating neuronal sensitivity. These changes may give rise to hyperexcitability underlying generalized epilepsy.

    View details for DOI 10.1002/ana.22462

    View details for Web of Science ID 000296396700010

    View details for PubMedID 21710621

    View details for PubMedCentralID PMC3183388

  • Orientation tuning in the visual cortex of 3-month-old human infants VISION RESEARCH Baker, T. J., Norcia, A. M., Candy, T. R. 2011; 51 (5): 470-478


    Sensitivity to orientation is critical for making a whole and complete picture of the world. We measured the orientation tuning of mechanisms in the visual cortex of typically developing 3-month-olds and adults using a nonlinear analysis of the two-input steady-state Visually Evoked Potential (VEP). Two gratings, one a fixed test and the other a variable orientation masker were tagged with distinct temporal frequencies and the corresponding evoked responses were measured at the harmonics of the test and masker frequencies and at a frequency equal to the sum of the two stimulus frequencies. The magnitude of the sum frequency component depended strongly on the relative orientation of the test and masker in both infants and adults. The VEP tuning bandwidths of the 3-month-olds measured at the sum frequency were similar to those of adults, suggesting that behavioral immaturities in functions such as orientation discrimination and contour integration may result from other immaturities in long-range lateral projections or feedback mechanisms.

    View details for DOI 10.1016/j.visres.2011.01.003

    View details for Web of Science ID 000288618900002

    View details for PubMedID 21236289

  • Disparity-Tuned Population Responses from Human Visual Cortex JOURNAL OF NEUROSCIENCE Cottereau, B. R., McKee, S. P., Ales, J. M., Norcia, A. M. 2011; 31 (3): 954-965


    We used source imaging of visual evoked potentials to measure neural population responses over a wide range of horizontal disparities (0.5-64 arcmin). The stimulus was a central disk that moved back and forth across the fixation plane at 2 Hz, surrounded either by binocularly uncorrelated dots (disparity noise) or by correlated dots presented in the fixation plane. Both disk and surround were composed of dynamic random dots to remove coherent monocular information. Disparity tuning was measured in five visual regions of interest (ROIs) [V1, human middle temporal area (hMT+), V4, lateral occipital complex (LOC), and V3A], defined in separate functional magnetic resonance imaging scans. The disparity tuning functions peaked between 2 and 16 arcmin for both types of surround in each ROI. Disparity tuning in the V1 ROI was unaffected by the type of surround, but surround correlation altered both the amplitude and phase of the disparity responses in the other ROIs. Response amplitude increased when the disk was in front of the surround in the V3A and LOC ROIs, indicating that these areas encode figure-ground relationships and object convexity. The correlated surround produced a consistent phase lag at the second harmonic in the hMT+ and V4 ROIs without a change in amplitude, while in the V3A ROI, both phase and amplitude effects were observed. Sensitivity to disparity context is thus widespread in visual cortex, but the dynamics of these contextual interactions differ across regions.

    View details for DOI 10.1523/JNEUROSCI.3795-10.2011

    View details for Web of Science ID 000286373700020

    View details for PubMedID 21248120

  • Impaired visual decision-making in individuals with amblyopia JOURNAL OF VISION Farzin, F., Norcia, A. M. 2011; 11 (14)


    This study examined the effects of amblyopia on perceptual decision-making processes to determine the consequences of visual deprivation on the development of higher level cortical networks outside of the visual cortex. A variant of the Eriksen flanker task was used to measure response time and accuracy for decisions made in the presence of response-selection conflict. Performance of adults with amblyopia was compared to that of neurotypical participants of the same age. Additionally, simple and choice reaction time tasks presented in the visual and the auditory modality were used to control for factors such as feature visibility, crowding, and motor execution speed. A selective deficit in response time for visual decisions was found when individuals with amblyopia used either the amblyopic or non-amblyopic (dominant) eye, and this deficit was independent of visual acuity, motor time, and performance accuracy. In trial conditions that provoked response-selection conflict, responses were significantly delayed in amblyopic relative to neurotypical participants and were not subject to standard trial sequence effects. Our results indicate that, beyond the known effects of abnormal visual experience on visual cortex, suboptimal binocular input during a developmental critical period may also impact cortical connections to downstream areas of the brain, including parietal and frontal cortices, that are believed to underlie decision and response-selection processes.

    View details for DOI 10.1167/11.14.6

    View details for Web of Science ID 000298635300006

    View details for PubMedID 22147222

    View details for PubMedCentralID PMC3250232

  • V1 is not uniquely identified by polarity reversals of responses to upper and lower visual field stimuli NEUROIMAGE Ales, J. M., Yates, J. L., Norcia, A. M. 2010; 52 (4): 1401-1409


    The cruciform hypothesis states that if a visual evoked potential component originates in V1, then stimuli placed in the upper versus lower visual fields will generate responses with opposite polarity at the scalp. This diagnostic has been used by many studies as a definitive marker of V1 sources. To provide an empirical test of the validity of the cruciform hypothesis, we generated forward models of cortical areas V1, V2 and V3 that were based on realistic estimates of the 3-D shape of these areas and the shape and conductivity of the brain, skull and scalp. Functional MRI was used to identify the location of early visual areas and anatomical MRI data was used to construct detailed cortical surface reconstructions and to generate boundary element method forward models of the electrical conductivity of each participant's head. These two data sets for each subject were used to generate simulated scalp activity from the dorsal and ventral subdivisions of each visual area that correspond to the lower and upper visual field representations, respectively. The predicted topographies show that sources in V1 do not fully conform to the cruciform sign-reversal. Moreover, contrary to the model, retinotopic visual areas V2 and V3 show polarity reversals for upper and lower field stimuli. The presence of a response polarity inversion for upper versus lower field stimuli is therefore an insufficient criterion for identifying responses as having originated in V1.

    View details for DOI 10.1016/j.neuroimage.2010.05.016

    View details for Web of Science ID 000280695200028

    View details for PubMedID 20488247

  • Configural specificity of the lateral occipital cortex NEUROPSYCHOLOGIA Appelbaum, L. G., Ales, J. M., COTTEREAU, B., Norcia, A. M. 2010; 48 (11): 3323-3328


    While regions of the lateral occipital cortex (LOC) are known to be selective for objects relative to feature-matched controls, it is not known what set of cues or configurations are used to promote this selectivity. Many theories of perceptual organization have emphasized the figure-ground relationship as being especially important in object-level processing. In the present work we studied the role of perceptual organization in eliciting visual evoked potentials from the object selective LOC. To do this, we used two-region stimuli in which the regions were modulated at different temporal frequencies and were comprised of either symmetric or asymmetric arrangements. The asymmetric arrangement produced an unambiguous figure-ground relationship consistent with a smaller figure region surrounded by a larger background, while four different symmetric arrangements resulted in ambiguous figure-ground relationships but still possessed strong kinetic boundaries between the regions. The surrounded figure-ground arrangement evoked greater activity in the LOC relative to first-tier visual areas (V1-V3). Response selectivity in the LOC, however, was not present for the four different types of symmetric stimuli. These results suggest that kinetic texture boundaries alone are not sufficient to trigger selective processing in the LOC, but that the spatial configuration of a figure that is surrounded by a larger background is both necessary and sufficient to selectively activate the LOC.

    View details for DOI 10.1016/j.neuropsychologia.2010.07.016

    View details for Web of Science ID 000281890200018

    View details for PubMedID 20638395

  • Quantitative Fiber Tracking of the Optic Radiation Is Correlated with Visual-Evoked Potential Amplitude in Preterm Infants AMERICAN JOURNAL OF NEURORADIOLOGY Glass, H. C., Berman, J. I., Norcia, A. M., Rogers, E. E., Henry, R. G., Hou, C., Barkovich, A. J., Good, W. V. 2010; 31 (8): 1424-1429


    Children born preterm are at risk for adverse outcome, including visual impairment. We examined the relationship between neonatal DTI and sVEP in children born preterm to determine whether visual outcomes are related to early measurements of brain microstructure.Subjects were born at <34 weeks gestation and imaged before term-equivalent age. DTI fiber tracking was used to delineate the optic radiations and measure tract-specific average FA, D(av), and parallel and transverse diffusivity. Visual-evoked response amplitudes were measured as a function of spatial frequency, contrast, and vernier offset size with sVEP at 6-20 months after birth. The association between DTI and sVEP was assessed by using the Spearman correlation coefficient and linear regression for repeated measures.Nine children with 15 scans were included. The peak response amplitudes for spatial frequency sweeps were associated with increasing FA and decreasing D(av) and transverse diffusivity (P ≤ .006) but not with parallel diffusivity (P = 1). There was only modest association with the swept contrast condition and no detectable association with the vernier offset sweeps.Microstructure of the optic radiations measured shortly after birth is associated with quantitatively measured responses elicited by moderate-to-high contrast spatiotemporal gratings in infancy. These findings are in keeping with studies showing a relationship between brain microstructure and function. While the clinical impact is not known, quantitative neuroimaging of white matter may ultimately be important for predicting outcome in preterm neonates.

    View details for DOI 10.3174/ajnr.A2110

    View details for Web of Science ID 000282804100015

    View details for PubMedID 20448011

  • Connecting the Dots: How Local Structure Affects Global Integration in Infants JOURNAL OF COGNITIVE NEUROSCIENCE Palomares, M., Pettet, M., Vildavski, V., Hou, C., Norcia, A. 2010; 22 (7): 1557-1569


    Glass patterns are moirés created from a sparse random-dot field paired with its spatially shifted copy. Because discrimination of these patterns is not based on local features, they have been used extensively to study global integration processes. Here, we investigated whether 4- to 5.5-month-old infants are sensitive to the global structure of Glass patterns by measuring visual-evoked potentials. Although we found strong responses to the appearance of the constituent dots, we found sensitivity to the global structure of the Glass patterns in the infants only over a very limited range of spatial separation. In contrast, we observed robust responses in the infants when we connected the dot pairs of the Glass pattern with lines. Moreover, both infants and adults showed differential responses to exchanges between line patterns portraying different global structures. A control study varying luminance contrast in adults suggests that infant sensitivity to global structure is not primarily limited by reduced element visibility. Together our results suggest that the insensitivity to structure in conventional Glass patterns is due to inefficiencies in extracting the local orientation cues generated by the dot pairs. Once the local orientations are made unambiguous or when the interpolation span is small, infants can integrate these signals over the image.

    View details for Web of Science ID 000279057500015

    View details for PubMedID 19642888

  • Human neutrophil alloantigen-1a,-1b,-2,-3a and-4a frequencies in Brazilians TISSUE ANTIGENS Norcia, A. M., Sugano, E. Y., Chiba, A. K., Moritz, E., Guirao, F. P., Yamamoto, M., Bordin, J. O. 2009; 74 (5): 404-407


    Human neutrophil reactive antibodies may cause clinical disorders such as transfusion-related acute lung injury, febrile transfusion reactions, alloimmune neonatal neutropenia, immune neutropenia after stem cell transplantation, refractoriness to granulocyte transfusion, drug-induced neutropenia and autoimmune neutropenia. Using the granulocyte immunofluorescence test by flow cytometry, the phenotypic frequencies of the human neutrophil alloantigens (HNA)-1a, -1b, -2, -3a and -4a were determined in 100 healthy Brazilian persons. Neutrophils were separated from blood samples by sedimentation, centrifugated and incubated with HNA-specific alloantibody plus fluorescein isothiocyanate-labeled F(ab')(2) fragments of anti-human IgG. The results showed that the phenotype frequencies of HNA-1a, -1b, -2a, -3a and -4a were 65%, 83%, 97%, 95% and 94%, respectively. We detected that neutrophils from 17% of Brazilians typed positive only with anti-HNA-1a (HNA-1a/a), 35% only with anti-HNA-1b (HNA-1b/b) and 48% reacted with both antibodies (HNA-1a/b). The frequencies found for HNA-1a and -1b were quite similar to that reported among Africans and American-Africans, but different from those found in Japanese and Chinese. In addition, our data showed that the frequencies of HNA-2, -3a and -4a in Brazilians were comparable with those observed in Caucasians. The determination of HNAs frequencies among populations with distinct racial backgrounds is important not only for anthropological reasons, but also for neonatal typing in suspected cases of alloimmune neutropenia or when patients are severely neutropenic.

    View details for DOI 10.1111/j.1399-0039.2009.01357.x

    View details for Web of Science ID 000270833200005

    View details for PubMedID 19737365

  • Spatio-temporal tuning of coherent motion evoked responses in 4-6 month old infants and adults VISION RESEARCH Hou, C., Gilmore, R. O., Pettet, M. W., Norcia, A. M. 2009; 49 (20): 2509-2517


    Motion cues provide a rich source of information about translations of the observer through the environment as well as the movements of objects and surfaces. While the direction of motion can be extracted locally these local measurements are, in general, insufficient for determining object and surface motions. To study the development of local and global motion processing mechanisms, we recorded Visual Evoked Potentials (VEPs) in response to dynamic random dot displays that alternated between coherent rotational motion and random motion at 0.8 Hz. We compared the spatio-temporal tuning of the evoked response in 4-6 months old infants to that of adults by recording over a range of dot displacements and temporal update rates. Responses recorded at the frequency of the coherent motion modulation were tuned for displacement at the occipital midline in both adults in infants. Responses at lateral electrodes were tuned for speed in adults, but not in infants. Infant responses were maximal at a larger range of spatial displacement than that of adults. In contrast, responses recorded at the dot-update rate showed a more similar parametric displacement tuning and scalp topography in infants and adults. Taken together, our results suggest that while local motion processing is relatively mature at 4-6 months, global integration mechanisms exhibit significant immaturities at this age.

    View details for DOI 10.1016/j.visres.2009.08.007

    View details for Web of Science ID 000270861900008

    View details for PubMedID 19679146

  • Backward masking suppresses collinear facilitation in the visual cortex VISION RESEARCH Sterkin, A., Yehezkel, O., Bonneh, Y. S., Norcia, A., Polat, U. 2009; 49 (14): 1784-1794


    Perceptual facilitation in detecting low-contrast Gabor patches (GPs) is induced by collinearly oriented high-contrast flankers. Our recent Visual Evoked Potentials (VEPs) study provided new physiological evidence for these collinear interactions, reflected by nonlinear modulation of multiple waveform components and frequencies [Sterkin, A., Yehezkel, O., Bonneh, Y. S., Norcia, A., & Polat, U. (2008). Multi-component correlate for lateral collinear interactions in the human visual cortex. Vision Research, 48(15), 1641-1647]. Here we used VEPs to study the temporal structure of this process. Low-contrast, foveal target GP (T) was simultaneously flanked by two collinear high-contrast GPs with a spatial separation that induces facilitation of T (lateral masking, LM). Another mask, identical to LM, was presented at different time-intervals (ISIs) after LM (backward masking, BM-on-LM). The responses were compared to separate waveforms evoked by T-alone and mask-alone at different ISIs. BM canceled the physiological markers of facilitation at an ISI of 50 ms, in agreement with earlier psychophysical findings, whereas no BM effect on T-alone was observed. This ISI coincides with the active time-window of lateral interactions, confirming our working model. The waveform amplitude of the negative N1 peak of LM was modulated toward the linear prediction of no interactions and the spectrum was shifted toward suppression, with no evidence of facilitation. Moreover, the P1 peak amplitude of BM was decreased at the same ISI, indicating that there is a mutual interference in cortical representation of both events. Waveform subtraction between BM-on-LM and LM suggests a mechanism of extended persistence of the target representation underlying facilitation in LM. We suggest an explanation for the role of improved detection of collinear stimuli in grouping of contours.

    View details for DOI 10.1016/j.visres.2009.04.013

    View details for Web of Science ID 000267504800008

    View details for PubMedID 19383510

  • Evidence for Visual Compromise in Preverbal Children with Orbital Vascular Birthmarks AMERICAN JOURNAL OF OPHTHALMOLOGY Good, W. V., Hou, C., Frieden, I. J., Norcia, A. M. 2009; 147 (4): 679-682


    To learn whether electrophysiological changes indicating amblyopia occur even in the absence of clinically recognizable amblyopia.Prospective study.Four consecutive infants between 7 and 19 months of age with unilateral periocular vascular lesions that intermittently obstructed vision in the affected eye and no clinical evidence of amblyopia were evaluated. No child had anisometropia greater than 0.50 diopter in the greatest meridian or strabismus. Sweep visual evoked potential vernier acuity was measured under monocular viewing conditions with the fellow eye tested as the control.Response amplitudes and acuity thresholds were significantly diminished in the affected eyes. A phase analysis showed slowing of the response in the affected eyes compared with the control eyes.An amblyopia-like effect on vernier acuity occurred in infants with unilateral periocular vascular birthmarks when the lesion caused intermittent occlusion of the eye. Whether long-term effects will occur is unknown, but children with no clinically apparent amblyopia in the setting of a vascular mark or other cause of intermittent occlusion of the visual axis should be followed, since these electrophysiology findings suggest amblyopia may be present.

    View details for DOI 10.1016/j.ajo.2008.11.007

    View details for Web of Science ID 000264566600018

    View details for PubMedID 19195640

  • Disparity Tuning of Binocular Facilitation and Suppression after Normal versus Abnormal Visual Development INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Norcia, A. M., Hale, J., Pettet, M. W., McKee, S. P., Harrad, R. A. 2009; 50 (3): 1168-1175


    To study the pattern of facilitatory and suppressive binocular interactions in stereodeficient patients with strabismus and in healthy controls.Visual evoked potentials were recorded in response to a Vernier onset/offset pattern presented to one eye, either monocularly or paired dichoptically with a straight vertical square-wave grating, which, when fused with the target in the other eye, gave rise to a percept of a series of bands appearing in depth from an otherwise uniform plane or with a grating that contained offsets that produced a standing disparity and the appearance of a constantly segmented image, portions of which moved in depth.Participants with normal stereopsis showed facilitative and suppressive binocular interactions that depended on which dichoptic target was presented. Patients with longstanding, constant strabismus lacked normal facilitative binocular interactions. The response to a normally facilitative stimulus was reduced below the monocular level when it was presented to the dominant eye of patients without anisometropia, consistent with classical strabismic suppression of the nondominant eye. The dominant eye of strabismic patients without anisometropia retained suppressive input from crossed but not uncrossed disparity stimuli presented to the nondominant eye.Abnormal disparity processing can be detected with the dichoptic VEP method we describe. Our results suggest that suppression in stereoblind, nonamblyopic observers is determined by a binocular mechanism responsive to disparity. In some cases, the sign of the disparity is important, and this suggests a mechanism that can explain diplopia in patients made exotropic after surgery for esotropia.

    View details for DOI 10.1167/iovs.08-2281

    View details for Web of Science ID 000263665000026

    View details for PubMedID 19098323

  • Assessing direction-specific adaptation using the steady-state visual evoked potential: Results from EEG source imaging JOURNAL OF VISION Ales, J. M., Norcia, A. M. 2009; 9 (7)


    Studying directional selectivity using neuroimaging in humans is difficult because the resolution is insufficient to directly access directionally selective activity. Here we used motion adaptation of the steady-state visual evoked potential (SSVEP) and source imaging in the frequency domain to detect brain areas that contain direction-selective cells. This study uses a definitive electrophysiological marker for direction-specific adaptation in the SSVEP to localize cortical areas that are direction selective. It has been shown previously that an oscillating stimulus produces an SSVEP response that is dominated by even harmonics of the stimulus frequency. This pattern of response is consistent with equal population responses to each direction of motion. Prolonged exposure to unidirectional motion induces an asymmetry in the population response that is consistent with adaptation of direction-selective cells. This asymmetry manifests itself in the presence of odd harmonic components after adaptation Critically, the feature that indicates the direction used for adaptation is the phase of the odd-harmonic responses. We recorded this signature of direction selectivity in a group of observers whose retinotopic visual areas had been defined from fMRI mapping. We find direction-specific responses throughout retinotopic cortex, with the largest effect in areas V1 (occipital pole) and V3/V3a (dorsal).

    View details for DOI 10.1167/9.7.8

    View details for Web of Science ID 000269824300008

    View details for PubMedID 19761323

  • Attentive and pre-attentive aspects of figural processing JOURNAL OF VISION Appelbaum, L. G., Norcia, A. M. 2009; 9 (11)


    Here we use the steady-state visual evoked potential (SSVEP) to study attentive versus non-attentive processing of simple texture-defined shapes. By "tagging" the figure and background regions with different temporal frequencies, the method isolates response components associated with the figure region, the background region, and with non-linear spatio-temporal interactions between regions. Each of these response classes has a distinct scalp topography that is preserved under differing attentional task demands. In one task, attention was directed to discrimination of shape changes in the figure region. In the other task, a difficult letter discrimination was used to divert attentive processing resources away from the texture-defined form. Larger task-dependent effects were observed for figure responses and for the figure/background interaction than for the background responses. The figure region responses were delayed in occipital areas in the shape versus letter task conditions, while the region interactions were enhanced, especially in frontal areas. While a basic differentiation of figure from background processing occurred independent of task, attentive processing of elementary shapes recruited later occipital activity for figure processing and sustained non-linear figure/background interaction in frontal areas. Collectively, these results indicate that basic aspects of scene segmentation proceed pre-attentively, but that directed attention to the object shape engages a widely distributed network of brain areas including frontal and occipital regions.

    View details for DOI 10.1167/9.11.18

    View details for Web of Science ID 000273280700018

    View details for PubMedID 20053081

  • Multi-component correlate for lateral collinear interactions in the human visual cortex VISION RESEARCH Sterkin, A., Yehezkel, O., Bonneh, Y. S., Norcia, A., Polat, U. 2008; 48 (15): 1641-1647


    Perceptual facilitation, a decrease in detection threshold for low-contrast Gabor patches (GPs) occurs when the GP is flanked by collinearly oriented high-contrast patches. There is earlier evidence suggesting a spatial architecture of excitatory and inhibitory interactions. Here we used Visual Evoked Potentials (VEPs) to study the temporal structure of this process. We measured VEPs elicited by a foveal near-threshold target GP presented in isolation (T), T in the presence of two flanking collinear high-contrast GPs (lateral masking, LM), or the flankers alone (F). Stimuli were presented for 50 ms every 1000 ms. The choice of the set parameters elicited behavioral facilitation of T detection. Significant modulation of peak amplitudes in LM compared with linearly summed waveforms elicited by T and F was found for five alternating polarity components, ranging from 65 to 290 ms after stimulus onset. In the frequency domain, suppression at lower frequencies (up to 0.8 log units) was followed by facilitation at higher frequencies (4-6 Hz, up to 0.8 log units). Although no differences in the latencies were found, lateral interactions were reflected by non-linear waveform modulation of multiple components and frequencies, including components as early as 65-75 ms. Spectrum analysis suggests that both suppression and facilitation may be found for the same configuration of stimuli, simultaneously, distributed at different temporal frequencies and/or sources. The physiological correlates of lateral interactions may thus originate at multiple sources, only some of which are explicitly facilitatory. The final perceptual outcome of this complex spatio-temporal representation is determined by combining sensory and cognitive factors.

    View details for DOI 10.1016/j.visres.2008.04.018

    View details for Web of Science ID 000257581300007

    View details for PubMedID 18538813

  • Figure-ground interaction in the human visual cortex JOURNAL OF VISION Appelbaum, L. G., Wade, A. R., Pettet, M. W., Vildavski, V. Y., Norcia, A. M. 2008; 8 (9)


    Discontinuities in feature maps serve as important cues for the location of object boundaries. Here we used multi-input nonlinear analysis methods and EEG source imaging to assess the role of several different boundary cues in visual scene segmentation. Synthetic figure/ground displays portraying a circular figure region were defined solely by differences in the temporal frequency of the figure and background regions in the limiting case and by the addition of orientation or relative alignment cues in other cases. The use of distinct temporal frequencies made it possible to separately record responses arising from each region and to characterize the nature of nonlinear interactions between the two regions as measured in a set of retinotopically and functionally defined cortical areas. Figure/background interactions were prominent in retinotopic areas, and in an extra-striate region lying dorsal and anterior to area MT+. Figure/background interaction was greatly diminished by the elimination of orientation cues, the introduction of small gaps between the two regions, or by the presence of a constant second-order border between regions. Nonlinear figure/background interactions therefore carry spatially precise, time-locked information about the continuity/discontinuity of oriented texture fields. This information is widely distributed throughout occipital areas, including areas that do not display strong retinotopy.

    View details for DOI 10.1167/8.9.8

    View details for Web of Science ID 000258709300008

    View details for PubMedID 18831644

  • Neural correlates of transformational apparent motion PERCEPTION Mirabella, G., Norcia, A. M. 2008; 37 (9): 1368-1379


    Transformational apparent motion (TAM) arises when a shape that is abruptly flashed on and off next to a static shape of similar color or texture appears as a protrusion that extends and retracts smoothly from the static object. Here we report that the strength of the TAM percept can be predicted from the waveform of visual evoked potentials (VEPs) measured while observers rated their percepts. The VEPs at pattern onset and offset are maximally symmetric when the static inducer and the flashing patches of the display are of the same contrast. VEP symmetry is affected by how the two patches can be matched as a single surface and may reflect the relative contribution of different motion and object detection systems in visual cortex.

    View details for DOI 10.1068/p5972

    View details for Web of Science ID 000260949900005

    View details for PubMedID 18986063

  • Abnormalities of coherent motion processing in strabismic amblyopia: Visual-evoked potential measurements JOURNAL OF VISION Hou, C., Pettet, M. W., Norcia, A. M. 2008; 8 (4)


    Coherent motion responses of patients with mild to moderate strabismic amblyopia were compared to those of normals using visual-evoked potentials (VEPs). Responses were elicited by dynamic random-dot kinematograms that alternated at 0.83 Hz between globally coherent (left-right) and incoherent (random) motion states. Tuning curves were measured at the first harmonic of the global motion update rate (0.83 Hz) and at the first harmonic of the dot update rate (20 Hz) for spatial displacements 3.1 to 27.9 arcmin (1.6 to 9.3 deg/s). Responses locked to the changes in the global organization of the local direction vectors were an inverted U-shaped function of displacement/speed in the normal-vision observers and in the fellow eyes of the strabismus patients while the tuning function of the amblyopic eyes was shifted to larger displacements/higher speeds. Responses at the dot update rate were reduced in amplitude and altered in timing in both eyes of the patients. The results are consistent with both local and global deficits in motion processing in strabismic amblyopia.

    View details for DOI 10.1167/8.4.2

    View details for Web of Science ID 000255976000002

    View details for PubMedID 18484841

  • Development of cortical responses to optic flow VISUAL NEUROSCIENCE Gilmore, R. O., Hou, C., Pettet, M. W., Norcia, A. M. 2007; 24 (6): 845-856


    Humans discriminate approaching objects from receding ones shortly after birth, and optic flow associated with self-motion may activate distinctive brain networks, including the human MT+ complex. We sought evidence for evoked brain activity that distinguished radial motion from other optic flow patterns, such as translation or rotation by recording steady-state visual evoked potentials (ssVEPs), in both adults and 4-6 month-old infants to direction-reversing optic flow patterns. In adults, radial flow evoked distinctive brain responses in both the time and frequency domains. Differences between expansion/contraction and both translation and rotation were especially strong in lateral channels (PO7 and PO8), and there was an asymmetry between responses to expansion and contraction. In contrast, infants' evoked response waveforms to all flow types were equivalent, and showed no evidence of the expansion/contraction asymmetry. Infants' responses were largest and most reliable for the translation patterns in which all dots moved in the same direction. This pattern of response is consistent with an account in which motion processing systems detecting locally uniform motion develop earlier than do systems specializing in complex, globally non-uniform patterns of motion, and with evidence suggesting that motion processing undergoes prolonged postnatal development.

    View details for DOI 10.1017/S0952523807070769

    View details for Web of Science ID 000251977300008

    View details for PubMedID 18093371

  • Validation study of VEP vernier acuity in normal-vision and amblyopic adults INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Hou, C., Good, W. V., Norcia, A. M. 2007; 48 (9): 4070-4078


    Vernier displacement thresholds can be measured with swept-parameter visual evoked potentials (sVEPs) and may therefore be useful in pre- or nonverbal subjects. This study was conducted to test whether sVEP vernier thresholds are valid measures of the visibility of vernier offsets in two different settings.Vernier acuity thresholds were measured psychophysically and electrophysiologically using square-wave gratings containing vernier displacements modulated at 3.76 Hz. The detectability of the vernier alignment cue was degraded by introducing either gaps or standing offsets in the stimulus. These manipulations were performed in normal-vision observers. In a second experiment, psychophysical and sVEP vernier acuity were measured in amblyopic observers.sVEP thresholds and overall amplitudes in normal observers were strongly affected by the introduction of gaps or standing offsets, as were psychophysical thresholds. Psychophysical and sVEP vernier offset thresholds were significantly correlated in the amblyopic eyes, as were sVEP and optotype interocular threshold differences. sVEP amplitudes of patients with strabismus were lower than those of patients with anisometropic amblyopia, even though optotype acuities were the same in the two groups.Vernier acuity thresholds derived from the sVEP tap mechanisms that are specific for the relative position of stimulus elements, and they correlate with perceptual visibility in normal and amblyopic observers. Because of this correlation and because sVEP thresholds can be measured without the need for instruction or behavioral responses, they may be useful in assessing visual function in pre- and nonverbal patients.

    View details for DOI 10.1167/iovs.06-1368

    View details for Web of Science ID 000249061900024

    View details for PubMedID 17724189

  • Sensitivity and configuration-specificity of orientation-defined texture processing in infants and adults VISION RESEARCH Pei, F., Pettet, M. W., Norcia, A. M. 2007; 47 (3): 338-348


    Here we use textures made up of widely spaced Gabor patches to compare infant and adult sensitivity to the global organization of the elements comprising the textures. Visual Evoked Potentials (VEPs) were recorded to alterations between random images and images containing varying proportions of patches that were of the same orientation. The patches were placed on rectangular, hexagonal or random lattices. Texture-specific responses were robust in adults and their VEP threshold was reached when 1-17% of the patches had the same orientation in the structured image. Infant thresholds were approximately 20-60%. While infants are capable of detecting the global structure of our textures, their sensitivity is low. In adults we found, unexpectedly, that sensitivity and response gain were higher for horizontal compared to vertical global orientations. Infant sensitivity was the same for the two orientations. Comparable orientation anisotropies have not been previously reported for gratings, suggesting that the Gabor-defined textures are tapping different mechanisms. There were small, but measurable effects of the lattice type in adults, with the rectangular lattice producing the largest responses.

    View details for DOI 10.1016/j.visres.2006.10.005

    View details for Web of Science ID 000244016000006

    View details for PubMedID 17188321

  • Cue-invariant networks for figure and background processing in human visual cortex JOURNAL OF NEUROSCIENCE Appelbaum, L. G., Wade, A. R., Vildavski, V. Y., Pettet, M. W., Norcia, A. M. 2006; 26 (45): 11695-11708


    Lateral occipital cortical areas are involved in the perception of objects, but it is not clear how these areas interact with first tier visual areas. Using synthetic images portraying a simple texture-defined figure and an electrophysiological paradigm that allows us to monitor cortical responses to figure and background regions separately, we found distinct neuronal networks responsible for the processing of each region. The figure region of our displays was tagged with one temporal frequency (3.0 Hz) and the background region with another (3.6 Hz). Spectral analysis was used to separate the responses to the two regions during their simultaneous presentation. Distributed source reconstructions were made by using the minimum norm method, and cortical current density was measured in a set of visual areas defined on retinotopic and functional criteria with the use of functional magnetic resonance imaging. The results of the main experiments, combined with a set of control experiments, indicate that the figure region, but not the background, was routed preferentially to lateral cortex. A separate network extending from first tier through more dorsal areas responded preferentially to the background region. The figure-related responses were mostly invariant with respect to the texture types used to define the figure, did not depend on its spatial location or size, and mostly were unaffected by attentional instructions. Because of the emergent nature of a segmented figure in our displays, feedback from higher cortical areas is a likely candidate for the selection mechanism by which the figure region is routed to lateral occipital cortex.

    View details for DOI 10.1523/JNEUROSCI.2741-06.2006

    View details for Web of Science ID 000241892700021

    View details for PubMedID 17093091

  • Visual development in very low birth weight infants PEDIATRIC RESEARCH Mirabella, G., Kjaer, P. K., Norcia, A. M., Good, W. V., Madan, A. 2006; 60 (4): 435-439


    Extremely preterm infants are at risk for neurodevelopmental problems and the visual system is particularly vulnerable. However, development of visual function in preterm infants with little or no retinal or neurologic injury has not been well defined. This study compared development of visual function in preterm infants without severe retinopathy of prematurity (ROP), intraventricular hemorrhage (IVH) or periventricular leukomalacia (PVL) to that of term infants at 5-7 mo corrected age. Twenty-one very low birth weight (VLBW) preterm infants (24-32 wk gestational age, weighing < 1500 g), and 22 healthy term infants were tested at 5-7 mo corrected age. Infants with any IVH/PVL and > Stage II ROP or Plus disease were excluded. Contrast sensitivity, grating acuity, and vernier acuity were measured using swept-parameter visual evoked potentials. Thresholds and maximum amplitudes were compared between groups. VLBW and term infants showed no differences in sensitivity for contrast (67.5 versus 63.8), grating resolution (12.4 versus 12.5 cpd) or vernier acuity (1.2 versus 1.0 arcmin). However, the amplitudes for swept contrast (p < 0.03) and swept vernier offset (p < 0.04) stimuli were higher in VLBW infants. Visual thresholds in VLBW infants without serious retinal or neurologic abnormalities were not significantly different from those of term infants, suggesting that increased visual experience does not influence visual sensitivity. The higher amplitudes in VLBW infants, suggests that visual experience may affect responses to suprathreshold stimuli.

    View details for DOI 10.1203/01.pdr.0000238249.44088.2c

    View details for Web of Science ID 000240707500012

    View details for PubMedID 16940247

  • Neural correlates of shape-from-shading VISION RESEARCH Hou, C., Pettet, M. W., Vildavski, V. Y., Norcia, A. M. 2006; 46 (6-7): 1080-1090


    Visual evoked potentials were recorded during presentation of a single stimulus that generated bi-stable perceptual alternation between two different three-dimensional percepts. One interpretation (asymmetric) changed depth structure from flat to corrugated in depth and the other (symmetric) had the appearance of a flat surface translating laterally behind a set of apertures. Responses during perception of the asymmetric three-dimensional structure contained larger negative components than did responses during perception of the symmetric three-dimensional structure. Control experiments suggest that the interpretation of depth structure is selected after junction information caused by the interplay between shading and object shape is extracted.

    View details for DOI 10.1016/j.visres.2005.10.017

    View details for Web of Science ID 000236073400032

    View details for PubMedID 16321419

  • The repeatability of best corrected acuity in normal and amblyopic children 4 to 12 years of age INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Chen, S. I., Chandna, A., Norcia, A. M., Pettet, M., Stone, D. 2006; 47 (2): 614-619


    The main purpose of this work was to measure repeatability of line-by-line logMAR (logarithm of the minimum angle of resolution) acuity in normal and amblyopic children, while adequately controlling for optical defocus.The Lea Symbols Chart is a constant-crowding, equal-logMAR increment chart similar in design to the Early Treatment Diabetes Retinopathy Study [ETDRS] chart. LogMAR visual acuity was tested twice in each eye of 32 amblyopic and 11 normal children. Each test commenced with screening in which one of the three central symbols was chosen for identification starting with the 1.0- or 0.9-logMAR line, progressing to every second line until incorrect identification occurred. Symbol-by-symbol presentation then commenced at the logMAR line containing the last correctly identified symbol. The threshold was recorded as the last logMAR line where four of four or four of five correct responses occurred (i.e., line-by-line scoring). Retesting by the same examiner was identical and occurred within the same session.There was no significant difference in repeatability among normal, fellow, or amblyopic eyes. The difference between test and retest thresholds lay between +/-0.10 logMAR in 93% of eyes. The 95% limits of agreement for the difference was +/-0.18 logMAR. Repeatability in eyes tested first did not differ from that in those tested second in either the normal or amblyopic groups.In the age-group tested, the line-by-line method of threshold scoring compares favorably with previous reports of both line-by-line and interpolated threshold scoring. There was no clinically meaningful difference in repeatability between the normal and amblyopic children tested.

    View details for DOI 10.1167/iovs.05-0610

    View details for Web of Science ID 000234922100022

    View details for PubMedID 16431958

  • Measurement of position acuity in strabismus and amblyopia: Specificity of the vernier VEP paradigm INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Chen, S. I., Norcia, A. M., Pettet, M. W., Chandna, A. 2005; 46 (12): 4563-4570


    An objective measure of positional acuity is desirable in the nonverbal clinical population. This study was conducted to investigate the specificity of the vernier VEP as a measure of positional acuity, evaluating the potential confound of asymmetric motion responses that may be present in some groups of patients. These motion responses could masquerade as position-specific responses, since they occur at the same response frequency as the vernier-related response.Twelve observers with early-onset esotropia (EOE), 30 children with untreated amblyopia, and 15 control children underwent swept vernier VEP acuity testing accompanied by a swept motion control stimulus. The control condition was used to detect the presence of artifactual responses not related to position sensitivity. The patients with EOE were selected for high levels of motion asymmetry as documented with oscillating gratings presented monocularly. As a measure of motion confound (penetration), the proportion of first-harmonic responses recorded in the control condition was determined.The penetration rate in the vernier condition in each study group (EOE: 0.93%; amblyopes: 4.26%; normal subjects: 2.40%) and the entire group (2.85%) was acceptably low. The level of penetration was not significantly influenced by the presence of amblyopia.The vernier VEP paradigm, when applied in the manner described, can be interpreted as a measure of position sensitivity. The presence of motion asymmetry or untreated amblyopia does not affect the validity of vernier measurements made.

    View details for DOI 10.1167/iovs.05-0792

    View details for Web of Science ID 000233578600031

    View details for PubMedID 16303949

  • Event-related potentials show configural specificity of global form processing NEUROREPORT Pei, F., Pettet, M. W., Vildavski, V. Y., Norcia, A. M. 2005; 16 (13): 1427-1430


    Glass patterns are a type of moiré created when a random-dot field is overlaid with a rotated, translated or dilated copy. The overall form of the moiré cannot be detected using local processing mechanisms, and because of this, Glass patterns are useful probes of global form processing. Here, we use event-related potentials to show that certain global organizations (concentric structure created by rotation and radial structure produced by dilation) produce much larger brain responses than others (linear structure created by translation). The results are consistent with the existence of specialized form processing mechanisms in the extrastriate cortex.

    View details for Web of Science ID 000232456900004

    View details for PubMedID 16110264

  • A VEP measure of the binocular fusion of horizontal and vertical disparities INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Hale, J., Harrad, R. A., MCKEE, S. P., Pettet, M. W., Norcia, A. M. 2005; 46 (5): 1786-1790


    Because of the lateral separation of the orbits, the retinal images differ in the two eyes. These differences are reconciled into a single image through sensory and motor fusional mechanisms. This study demonstrates electrophysiologically the effects that normal horizontal and vertical fusional processes have on the processing of monocular position signals.VEPs were recorded in 16 healthy adults in response to a vernier onset-offset target presented to one eye. The vernier offsets appeared and disappeared at 2 Hz and were introduced into bar targets that were oriented either vertically (horizontal offsets) or horizontally (vertical offsets). The magnitude of the offsets was varied over the range of 0.5 to 10 arc min. VEP amplitude was measured as a function of the size of the dynamic offset under monocular viewing conditions and in the presence of two different static targets presented to the other eye. One of the static targets matched the dynamic test, except that it had no vernier offsets. The other static target, the static pedestal, matched the dynamic test, but contained a set of static vernier offsets in locations corresponding to the locations of the dynamic offsets presented to the other eye.VEP amplitude was a monotonically increasing function of vernier offset size under monocular viewing conditions. The addition of the static target without offsets in the other eye resulted in an increased amplitude VEP response. The addition of the static target with vernier offsets resulted in a decrease in VEP amplitude for both horizontal and vertical disparities.The normal process of fusion results in a single visual direction. To obtain a single visual direction, the visual system must synthesize a binocular visual direction that differs from the monocular components. One of the conditions (the static pedestal with offsets) produces binocular visual direction shifts that degrade the appearance of vernier onset-offset, and reduce VEP amplitude for both horizontal and vertical disparities. This characteristic evoked response marker is a promising tool for measuring binocular fusion objectively in patients with strabismus.

    View details for DOI 10.1167/iovs.04-0954

    View details for Web of Science ID 000228708000037

    View details for PubMedID 15851583

  • Development of sensitivity to texture and contour information in the human infant JOURNAL OF COGNITIVE NEUROSCIENCE Norcia, A. M., Pei, F., Bonneh, Y., Hou, C., Sampath, V., Pettet, M. W. 2005; 17 (4): 569-579


    Texture discrimination and bounding contour extraction are essential parts of the object segmentation and shape discrimination process. As such, successful texture and contour processing are key components underlying the development of the perception of both objects and surfaces. By recording visual-evoked potentials, we investigate whether young infants can detect orientation-defined textures and contours. We measured responses to an organized texture comprised of many Gabor patches of the same orientation, alternated with images containing the same number of patches, but all of random orientation. These responses were compared with a control condition consisting of the alternation between two independently random configurations. Significant difference potentials were found as early as 2-5 months, as were significant odd harmonics in the test conditions. Responses were also measured to Gabor patches organized either as circles (all patches tangent to an imaginary circular path) alternated with pinwheels (all patches having a fixed orientation offset from the path). Infants between 6 and 13 months also showed sensitivity to the global organization of the elements along contours. Differential responses to our texture and contour stimuli and their controls could only have been generated by mechanisms that are capable of comparing the relative orientation of 2 or more patches, as no local information at a single patch distinguished the random and organized textures or the circle and pinwheel configurations.

    View details for Web of Science ID 000228360600003

    View details for PubMedID 15829078

  • Suppression of monocular visual direction under fused binocular stimulation: Evoked potential measurements JOURNAL OF VISION Norcia, A. M., MCKEE, S. P., Bonneh, Y., Pettet, M. W. 2005; 5 (1): 34-44


    Visual evoked potentials (VEPs) were recorded in response to a vernier onset/offset target presented to one eye that was combined with matching static targets in the other eye. The monocular response was dominated by a negative peak at 160 ms that occurred after a set of offsets was introduced into a one-dimensional random bar pattern. The static targets produced no discernible VEP response by themselves, but when fused binocularly with the oscillating vernier target, they produced shifts in perceived visual direction that influenced the VEP response. A vernier target fused with static vertical bars was perceived to alternate in depth between a flat surface and one broken into two interleaved surfaces. The response to this "surface-breaking" was as large or larger than the response to the monocular vernier offset. This response was much reduced when the oscillating vernier was fused with a static offset vernier (5' offset) that produced a percept of segregated regions moving in depth. Apparently, the VEP is strongly driven by shifts in visual direction that alter surface, texture, or contour contiguity.

    View details for DOI 10.1167/5.1.4

    View details for Web of Science ID 000227037600004

    View details for PubMedID 15831065

  • Experience-expectant development of contour integration mechanisms in human visual cortex JOURNAL OF VISION Norcia, A. M., Sampath, V., Hou, C., Pettet, M. W. 2005; 5 (2): 116-130


    Extended contours are a common feature of natural images. Most previous studies have considered contour integration as a two-dimensional process of linking like-oriented elements along their common orientation axis. Yet contours exist in a three-dimensional world, and one might therefore ask about the relationship between contour integration and binocular vision. Using an event-related potential assay of contour integration, we demonstrate that patients with strabismic amblyopia show a relative insensitivity to Gabor-defined contours in their dominant eyes, all of which had normal acuity. These deficits were not seen in the dominant eyes of patients with anisometropic amblyopia without strabismus, but were present in the amblyopic eyes of patients with either strabismus or anisometropia. Deficits were also found in both eyes of strabismus patients who had normal visual acuity in each eye, but who had strongly reduced or absent stereopsis. These results suggest that the maturation of contour detection mechanisms depends at least in part on the presence of normal binocular interaction during a developmental critical period.

    View details for DOI 10.1167/5.2.3

    View details for Web of Science ID 000227249100003

    View details for PubMedID 15831072

  • Recovery of contour integration in relation to LogMAR visual acuity during treatment of amblyopia in children INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Chandna, A., Gonzalez-Martin, J. A., Norcia, A. M. 2004; 45 (11): 4016-4022


    In several studies, researchers have found that integration of orientation information along contours defined by Gabor patches is abnormal in patients with strabismus and in untreated patients with anisometropic amblyopia. In this study, the rate and degree of recovery of contour-integration deficits were compared with the recovery of logMAR (logarithm of the minimum angle of resolution) visual acuity deficits in patients newly diagnosed with amblyopia secondary to anisometropia, strabismus, or both.Contour-detection thresholds and optotype acuity were measured in 17 newly diagnosed anisometropic amblyopes, in 6 patients with strabismic amblyopia, and in 4 patients with combined anisometropic and strabismic amblyopia. Contour-detection thresholds were measured with a card-based procedure. Treatment comprised full refractive correction and full-time total occlusion therapy, when necessary. Visual function was measured at monthly visits during the course of treatment, with an average follow-up period of 16 weeks (12-24 weeks) for the entire group. Complete data were obtained from 23 patients through 8 weeks of follow-up.Significant interocular differences in contour-detection thresholds were present in 16 of the 27 patients at the first visit after initial refractive correction. Interocular differences in contour-detection thresholds declined to normal levels in most of the patients within 8 weeks of the initiation of treatment. Interocular acuity differences remained significant in many of the patients (19/23) at 8 weeks of follow-up and continued to decline, but did not fully normalize, over the remainder of the follow-up period.Refractive correction alone or in combination with occlusion therapy produces a normalization of contour-integration thresholds in amblyopia that is more rapid and complete than that achieved for visual acuity.

    View details for DOI 10.1167/iovs.03-0795

    View details for Web of Science ID 000224678200024

    View details for PubMedID 15505051

  • Development of the spatial organization and dynamics of lateral interactions in the human visual system JOURNAL OF NEUROSCIENCE Hou, C., Pettet, M. W., Sampath, V., Candy, T. R., Norcia, A. M. 2003; 23 (25): 8630-8640


    Psychophysical thresholds and neuronal responses for isolated stimuli are strongly modified by nearby stimuli in the visual field. We studied the orientation and position specificity of these contextual interactions using a dual-frequency visual-evoked potential technique in developing human infants and adults. One set of small, oriented stimulus elements (targets) was tagged with a temporal frequency f1 of 4.52 Hz. The addition of an abutting second set of similar patches (flankers) tagged at f2 = 2.58 Hz had three effects: (1) The flankers reduced the second and fourth harmonic responses to the targets. This reduction was independent of flanker orientation or position and age. (2) The response to the combination of targets and flankers also contained nonlinear interaction terms (1f1 +/- 1f2) that were tuned for flanker orientation and position in adults, but only for flanker orientation in infants 8-31 weeks of age. (3) Nonlinear interaction terms recorded at 2f1 +/- 2f2 were large and untuned for flanker orientation and position in adults but were nearly absent in the youngest infants. The three forms of nonlinear interaction, thus, have differences in sensitivity to flanker orientation and position and differential growth trends, indicating that they are generated by different mechanisms. These three forms of interaction could serve different functional roles. The first process provides a nonselective gain control that is fully functional in early infancy. The second process, which develops slowly, is selective for the specific form of the stimuli. The third process, which is also immature, pools across orientation.

    View details for Web of Science ID 000185534500002

    View details for PubMedID 14507962

  • Late maturation of visual hyperacuity PSYCHOLOGICAL SCIENCE Skoczenski, A. M., Norcia, A. M. 2002; 13 (6): 537-541


    We used a visual evoked-potential measure to study the development of two components of pattern vision, vernier acuity and grating acuity, in humans from early infancy through adolescence. These two visual functions develop at similar rates and have nearly the same absolute values between 1 month and 6 years of age. After age 6, grating acuity is constant at the adult level, but vernier acuity continues to improve, becoming a hyperacuity. Vernier acuity reaches asymptotic levels around age 14 years. These results suggest that adultlike vernier hyperacuity is not limited by spatial resolution or sensitivity of small receptive fields, but rather that the limitation is imposed by higher-level processing. Sensitivity, connections in visual cortical areas, or both therefore retain plasticity throughout childhood and into adolescence.

    View details for Web of Science ID 000178993800008

    View details for PubMedID 12430838

  • Temporal dynamics of the human response to symmetry JOURNAL OF VISION Norcia, A. M., Candy, T. R., Pettet, M. W., Vildavski, V. Y., Tyler, C. W. 2002; 2 (2): 132-139


    Symmetry is a highly salient feature of animals, plants, and the constructed environment. Although the perceptual phenomenology of symmetry processing is well understood, little is known about the underlying neural mechanisms. Here we use visual evoked potentials to measure the time course of neural events associated with the extraction of symmetry in random dot fields. We presented sparse random dot patterns that were symmetric about both the vertical and horizontal axes. Symmetric patterns were alternated with random patterns of the same density every 500 msec, using new exemplars of symmetric and random patterns on each image update. Random/random exchanges were used as a control. The response to updates of random patterns was multiphasic, consisting of P65, N90, P110, N140 and P220 peaks. The response to symmetric/random sequences was indistinguishable from that for random/random sequences up to about 220 msec, after which the response to symmetric patterns became relatively more negative. Symmetry in random dot patterns thus appears to be extracted after an initial response phase that is indifferent to configuration. These results are consistent with the hypothesis (Lee, Mumford, Romero, & Lamme, 1998; Tyler & Baseler, 1998) that the symmetry property is extracted by processing in extrastriate cortex.

    View details for DOI 10.1167/2.2.1

    View details for Web of Science ID 000206907600001

    View details for PubMedID 12678588

  • Extending the modelfest image/threshold database into the spatio-temporal domain. HUMAN VISION AND ELECTRONIC IMAGING VII Carney, T., Klein, S. A., BEUTTER, B., Norcia, A., Chen, C. C., Tyler, C. W., Makous, W., Watson, A., Cropper, S. J., Popple, A. V., Robertson, K., Manahilov, V., Simpson, B., Wenzel, K. 2002; 4662: 138-148
  • Neural correlates of object-based attention JOURNAL OF VISION Pei, F., Pettet, M. W., Norcia, A. M. 2002; 2 (9): 588-596


    Much research has been directed toward disentangling the "units" of attention: Is attention directed to locations in space, visual objects, or to individual features of an object? Moreover, there is considerable interest in whether attention increases the gain of neural mechanisms (signal enhancement) or acts by other means, such as reducing noise or narrowing channel tuning. To address these questions, we used a direct measure of signal strength: the amplitude of visual evoked potentials and a task in which selection could be based on a depth order cue but not on location. Attended and nonattended stimuli were presented at different temporal frequencies, and, thus, responses to the two stimuli could be analyzed separately even though they were presented simultaneously. Attention increased the amplitude of the second harmonic component of the response, but not the fourth harmonic. In addition, responses measured at the second harmonic, but not at the fourth harmonic, were larger for stimuli seen as behind. The results are consistent with the fourth harmonic being generated at a stage of processing that is not accessible to attention and where depth order has not been extracted. The second harmonic, on the other hand, is modifiable by attention and shows evidence for differential encoding of depth order.

    View details for DOI 10.1167/2.9.1

    View details for Web of Science ID 000206908200001

    View details for PubMedID 12678630

  • Normalization models applied to orientation masking in the human infant JOURNAL OF NEUROSCIENCE Candy, T. R., Skoczenski, A. M., Norcia, A. M. 2001; 21 (12): 4530-4541


    Human infants can discriminate the orientation of lines within the first week after birth (Atkinson et al., 1988; Slater et al., 1988) but have immature orientation-selective pattern masking until after 6 months of age (Morrone and Burr, 1986). Here the development of orientation processing is further examined using a visual-evoked potential paradigm and normalization models of pattern masking. Contrast response functions were measured for 1 cycle per degree (cpd) gratings, counterphase-reversed in contrast at either 3.3 or 5.5 Hz. A second 1 cpd, 20% contrast, 8.3 Hz grating of either the same or orthogonal orientation was added as a mask. Evoked responses associated with the test grating, the mask, and intermodulation between the two were individually extracted using spectral analysis of the scalp-recorded EEG. Adults exhibited orientation selectivity in the masking of their test component responses and in nonlinear intermodulation between the test and mask stimuli. Infants <5 months old, however, demonstrated nonselective masking or a reversed selectivity in their responses to the test component, with adult-like orientation selectivity in their intermodulation responses. Within the context of a normalization model of pattern masking, the results are consistent with the existence of oriented filters early in life the responses of which are normalized immaturely until approximately 5 months of age.

    View details for Web of Science ID 000169279900047

    View details for PubMedID 11404441

  • Colinear facilitation promotes reliability of single-cell responses in cat striate cortex EXPERIMENTAL BRAIN RESEARCH Kasamatsu, T., Polat, U., Pettet, M. W., Norcia, A. M. 2001; 138 (2): 163-172


    Behavior is controlled by neural activity in the brain. The final outcome of this neural control may critically depend on the firing reliability of individual neurons. A nearly constant, proportional relationship is usually found between the response mean and response variance. Here we asked whether lateral interactions within striate cortex that modulate response magnitude also proportionately modify the response variance of cortical neurons. In many cases, response variability depended on stimulus organization: discrete flankers colinearly placed well outside the neuron's receptive field increased response magnitude without a proportional increase in variance, thus improving the neuron's response reliability. Since colinear flanker facilitation is often seen near the neuron's firing threshold, increased response reliability for weak stimuli may contribute to enhancing perceptual saliency.

    View details for Web of Science ID 000168951000003

    View details for PubMedID 11417457

  • Contrast response characteristics of long-range lateral interactions in cat striate cortex 28th Annual Meeting of the Society-for-Neuroscience Chen, C. C., Kasamatsu, T., Polat, U., Norcia, A. M. LIPPINCOTT WILLIAMS & WILKINS. 2001: 655–61


    Single-cell responses in visual cortex to a target falling within their receptive field can be modified by collinear flanking stimuli concurrently presented outside the receptive field. Here, we report the presence of four types of contrast-dependent lateral effects: (1) facilitation at low target contrasts and suppression at high contrasts, (2) facilitation that increases with contrast, (3) suppression that increases with contrast, and (4) suppression at low contrasts with facilitation at high contrasts. We propose a sensitivity modulation model that accounts for all the four types of lateral effects by changes in two parameters. In this model, activation of neighboring neurons changes the sensitivities of the target neuron to both the direct feedforward input and inhibitory, divisive feedback from neighboring neurons.

    View details for Web of Science ID 000167504700007

    View details for PubMedID 11277558

  • Contour integration deficits in anisometropic amblyopia INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Chandna, A., Pennefather, P. M., Kovacs, I., Norcia, A. M. 2001; 42 (3): 875-878


    Previous retrospective studies have found that integration of orientation information along contours defined by Gabor patches is abnormal in strabismic, but not in anisometropic, amblyopia. This study was conducted to reexamine the question of whether anisometropic amblyopes have contour integration deficits prospectively in an untreated sample, to isolate the effects of the disease from the effects of prior treatment-factors that may have confounded the results in previous retrospective studies.Contour detection thresholds, optotype acuity, and stereoacuity were measured in a group of 19 newly diagnosed anisometropic amblyopes before initiation of occlusion therapy. Contour detection thresholds were measured using a card-based procedure.Significant interocular differences in contour detection thresholds were present in 14 of the 19 patients with anisometropic amblyopia.Contour integration deficits are a common, but not universal, finding in untreated anisometropic amblyopia. Differences in the prevalence of contour integration deficits between the present study and that of another study may lie in differences in treatment history and/or in the sensitivity of the two different contour integration tasks.

    View details for Web of Science ID 000167206600047

    View details for PubMedID 11222553

  • Changes in cortical activity during suppression in stereoblindness NEUROREPORT Norcia, A. M., Harrad, R. A., Brown, R. J. 2000; 11 (5): 1007-1012


    Patients with strabismus or anisometropic amblyopia fixate and attend with one eye and suppress the image from the other eye. Here we use a visual evoked potential technique to show that patients who lack normal stereopsis retain suppressive binocular interactions but lack a characteristic form of non-linear binocular interaction that is present in normal observers. Oscillating grating targets presented at different temporal frequencies in the two eyes evoke a strong response in normal observers at a frequency equal to the sum of the two input frequencies for fusable targets but not for rivalrous ones. However increasing contrast in one eye reduces the response amplitude from the other eye under either fusable (dichoptic masking) or rivalrous conditions. Stereo-deficient observers lack the sum-frequency response, but retain dichoptic masking interactions. Dichoptic masking is stronger when the masker is presented to the patients' dominant rather than non-dominant eyes, suggesting that a subset of preserved binocular inhibitory interactions form the basis of clinical suppression.

    View details for Web of Science ID 000086385300026

    View details for PubMedID 10790873

  • A new test of contour integration deficits in patients with a history of disrupted binocular experience during visual development VISION RESEARCH Kovacs, I., Polat, U., Pennefather, P. M., Chandna, A., Norcia, A. M. 2000; 40 (13): 1775-1783


    Previous studies have suggested that the integration of orientation information across space is impaired in amblyopia. We developed a method for quantifying orientation-domain processing using a test format that is suitable for clinical application. The test comprises a graded series of cards where each card includes a closed path (contour) of high contrast Gabor signals embedded in a random background of Gabor signals. Contour visibility in both normals and patients with histories of abnormal binocular vision depends jointly on the spacing of elements on the contour as well as background element density. Strabismic amblyopes show significant degradation of performance compared to normals. Small but significant losses in sensitivity were also observed in a group of non-amblyopic strabismus patients. Threshold measurements made with contrast reducing diffusers indicated that the amblyopic loss is not due to the reduced contrast sensitivity of the amblyopic eye. An abnormal pattern of long-range connectivity between spatial filters or a loss of such connectivity appears to be the primary source of contour integration deficits in amblyopia and strabismus.

    View details for Web of Science ID 000087362500012

    View details for PubMedID 10814762

  • Development of rivalry and dichoptic masking in human infants Annual Meeting of the Association-for-Research-in-Vision-and-Ophthalmology (ARVO) Brown, R. J., Candy, T. R., Norcia, A. M. ASSOC RESEARCH VISION OPHTHALMOLOGY INC. 1999: 3324–33


    To examine the development of rivalry, dichoptic masking, and binocular interactions in infants more than 5 months of age using the visual evoked potential (VEP).VEPs were recorded in 35 infants between 5 and 15 months of age and 23 adults between 13 and 59 years of age. Counterphasing, sinusoidal, 1 cycle/deg gratings were presented dichoptically. Responses from each eye were isolated by "tagging" each half-image with a different temporal frequency (5 or 7.5 Hz). Observers were presented with fixed 80% contrast gratings in each eye in experiment 1. Rivalry was detected on the basis of a negative correlation between the simultaneously measured response amplitudes at the second harmonics of the two eye-tagging frequencies. In a second analysis of the same data, response amplitudes recorded under dichoptic viewing conditions were compared to those obtained in a monocular control condition (dichoptic masking). In experiment 2, a 40% fixed-contrast grating was presented to one eye, whereas the other eye viewed a grating that was swept in contrast from 1% to 67%. Dichoptic masking was measured as the reduction in the fixed-grating response caused by the variable contrast grating.Experiment 1: although adults showed evidence of VEP amplitude alternations between the eyes for cross-oriented half-images (physiological rivalry), infants did not. This immature response to rivalrous stimuli occurred despite the presence of responses at nonlinear combination frequencies recorded with gratings of the same orientation in each eye, a definitive indication of binocular interaction. In addition, both iso- and cross-oriented half-images produced less dichoptic masking in infants than in adults in this experiment. Experiment 2: dichoptic masking in the infants was equivalent to that seen in adults with parallel gratings in the two eyes; however, masking with cross-oriented configurations was approximately five times weaker in the infants relative to the adults.The authors have identified a set of stimulus conditions under which infants between 5 and 15 months of age fail to demonstrate physiological rivalry despite the presence of binocular interactions. The observed lack of binocular rivalry may be the result of a specific immaturity in dichoptic, cross-orientation suppression.

    View details for Web of Science ID 000083939500036

    View details for PubMedID 10586959

  • Electrophysiological correlates of vernier and relative motion mechanisms in human visual cortex VISUAL NEUROSCIENCE Norcia, A. M., Wesemann, W., Manny, R. E. 1999; 16 (6): 1123-1131


    Vernier onset/offset thresholds were measured both psychophysically and with the steady-state VEP by introducing a series of horizontal breaks in a vertical square-wave luminance grating. Several diagnostic tests indicated that the first harmonic component of the evoked response generated by periodic modulation of offset gratings taps mechanisms that encode the relative position of spatial features. In the first test, a first harmonic component was only found with targets that contained transitions between collinear and noncollinear states. VEP vernier onset/offset thresholds obtained with foveal viewing were in the range of 15-22 arc sec. Control experiments with transitions between symmetrical, noncollinear patterns (relative motion) did not produce first harmonic components, nor did full-field motion of a collinear grating. A second series of experiments showed that VEP thresholds based on the first harmonic component of the vernier onset/offset response had an eccentricity dependence that was very similar to that found in a psychophysical discrimination task that required a left/right position judgment (vernier acuity). Other recordings showed that the first harmonic of the vernier onset/offset VEP was degraded by the introduction of a gap between stimulus elements, as is the displacement threshold. The vernier onset/offset target also produced a second harmonic component that was virtually identical to the one produced by a relative motion stimulus. Displacement thresholds based on these second harmonic components showed a more gradual decline with retinal eccentricity than did the first harmonic component elicited by vernier offsets. The second harmonic of the vernier onset/offset VEP was relatively unaffected by the introduction of gaps between the stimulus elements. The first and second harmonic components of the vernier onset/offset VEP thus tap different mechanisms, both of which support displacement thresholds that are finer than the resolution limits set by the spacing of the photoreceptors (hyperacuity).

    View details for Web of Science ID 000084409800012

    View details for PubMedID 10614592

  • Development of VEP Vernier acuity and grating acuity in human infants INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Skoczenski, A. M., Norcia, A. M. 1999; 40 (10): 2411-2417


    To compare the developmental sequences of two basic measures of pattern vision, Vernier acuity and grating acuity, using steady state visual-evoked potentials (VEPs) and an analysis designed to isolate pattern-specific responses from those due to motion in the Vernier stimulus.The authors recorded VEPs from 57 healthy full-term infants and 4 adults. The grating acuity stimulus was a sinusoidal grating, temporally modulated (appearance-disappearance) at a rate of 3 Hz, with spatial frequency decreasing in linear steps during each 10-second trial. The Vernier acuity stimulus was a vertical square-wave grating with portions of each bar temporally modulated to make offsets appear and disappear at a rate of 3 Hz. Vernier offset size changed in log steps from small to large offsets. The authors recorded each observer's electroencephalogram (EEG) during multiple presentations of each stimulus type, and the EEG was digitized and filtered to obtain the amplitude and phase of the response at the first two harmonics of the stimulus temporal frequency. Thresholds were estimated with an extrapolation technique that took into account the signal-to-noise ratio and phase of the response.VEP Vernier acuity and grating acuity develop at different rates, with grating acuity approaching adult levels earlier than Vernier acuity. The within-subject relationship between VEP Vernier acuity and grating acuity follows the same developmental trajectory established by previous psychophysical studies of humans and monkeys.This VEP technique provides a rapid estimate of Vernier acuity in infants. VEP Vernier acuity remains strikingly immature throughout the first year of life, similar to behavioral Vernier acuity. Because Vernier acuity is a sensitive measure of amblyopia, this VEP test may be useful in the future to identify amblyopia and to follow its treatment progress in pediatric patients.

    View details for Web of Science ID 000082239800032

    View details for PubMedID 10476810

  • Asymmetric responses in cortical visually evoked potentials to motion are not derived from eye movements INVESTIGATIVE OPHTHALMOLOGY & VISUAL SCIENCE Wilson, J. R., Noyd, W. W., Aiyer, A. D., Norcia, A. M., Mustari, M. J., Boothe, R. G. 1999; 40 (10): 2435-2439


    Normal neonates and many adults after abnormal visual development have directional preferences for visual stimulus motions; i.e., they give better responses for optokinetic nystagmus (OKN) and visually evoked potentials (VEPs) in one direction than to those in the opposite direction. The authors tested whether the VEP responses were asymmetrical because of abnormal eye movements.VEPs were recorded from the visual cortices of five macaque monkeys: one normal, one neonate, and three reared with alternating monocular occlusion (AMO). They were lightly anesthetized, followed by paralysis to prevent eye movements. They then had "jittered" vertical grating patterns presented in their visual fields. The steady state VEPs were analyzed with discrete Fourier transforms to obtain the amplitudes and phases of the asymmetries.The normal, control monkey had small, insignificant amplitudes of its asymmetrical Fourier component and random phases that were not 180 degrees out of phase across the left and right eyes. The neonatal monkey and the AMO monkeys all had large, significant asymmetries that were approximately 180 degrees out of phase between the left and right eyes.The neonate and abnormally reared monkeys continued to have asymmetrical responses even after their eyes were paralyzed. Therefore, eye movements cannot be the source of the asymmetrical amplitudes of the VEPs, and the visual cortex is at least one source responsible for asymmetries observed in neonates and adults reared under abnormal visual inputs.

    View details for Web of Science ID 000082239800037

    View details for PubMedID 10476815

  • Oscillatory motion but not pattern reversal elicits monocular motion VEP biases in infantile esotropia VISION RESEARCH Shea, S. J., Chandna, A., Norcia, A. M. 1999; 39 (10): 1803-1811


    Patients with early disruptions of binocularity show cortical directional asymmetries in their steady state monocular VEP response to oscillatory motion. The VEP directional asymmetry is characterized by significant first harmonic components that show a 180 degrees difference in the response phase between the two eyes. By contrast, the normal response is dominated by even-order response harmonics, although some normal observers also have measurable responses at the first harmonic. Experiments and simulations were conducted to determine if the first harmonic in patients could reasonably be attributed to direction selective mechanisms. A secondary goal was to determine whether the first harmonic response of normals was also due to imbalances in direction selective mechanisms. Monocular steady state VEPs were elicited by oscillating 3 c/deg gratings presented at 6 and 10 Hz in normal observers and observers with infantile esotropia. Responses were also obtained to phase-reversing gratings of the same spatial and temporal frequencies. Phase reversal eliminated the majority of first harmonic responses which were recorded for normal observers to oscillatory motion. However, phase reversal did not elicit the cortical motion asymmetry in infantile esotropia. Modeling results suggest that the first harmonic response to oscillatory motion arises due to non-linearities in both direction selective and non-direction-selective mechanisms, with the latter being dominant in patients with early onset strabismus.

    View details for Web of Science ID 000079298900007

    View details for PubMedID 10343872

  • Contour detection threshold: repeatability and learning with 'contour cards' SPATIAL VISION Pennefather, P. M., Chandna, A., Kovacs, I., Polat, U., Norcia, A. M. 1999; 12 (3): 257-266


    Human observers are able to locate contours that are defined solely on the basis of long-range, orientation-domain correlations. The integrity of the mechanisms responsible for second-order contour detection is disrupted by amblyopia (Kovacs et al., 1996; Hess et al., 1997) and it is therefore of interest to develop methods for assessing pediatric patients undergoing treatment for amblyopia. In this study, we have determined the inter-observer and test-retest reliability of a card-based test of second-order contour integration. The magnitude of practice effects was also assessed in both adult and pediatric patient groups. Contour detection thresholds were measured for a closed contour, defined by Gabor patches, embedded in a randomly oriented Gabor-patch background. The visibility of the contour was controlled by varying the density of the background elements. Thresholds, defined in terms of the ratio of contour element spacing to average background spacing were measured with a clinical staircase procedure. Thresholds measured by two observers differed on average by 0.023 +/- 0.075 or about one half the increment between cards. Children and adults showed only small practice effects (0.022 +/- 0.051 vs 0.053 +/- 0.077, respectively) and average unsigned differences between repeated measures were equivalent to approximately 1 card across groups. A card-based test of second-order contour integration produces reliable estimates of contour integration performance in normal and amblyopic observers, including children.

    View details for Web of Science ID 000081631900001

    View details for PubMedID 10442513

  • Elongated physiological summation pools in the human visual cortex VISION RESEARCH Polat, U., Norcia, A. M. 1998; 38 (23): 3735-3741


    The visibility of gratings improves with increasing stimulus area. This effect is usually interpreted as being due to probability summation between the outputs of linear, independent spatial filters, although non-linear spatial summation can have similar effects [1]. In order to distinguish between probabilistic and physiological summation models, we measured contrast thresholds using the Visual Evoked Potential (VEP). Our previous work [2] suggests that spatial summation in the VEP is nonlinear and that it occurs preferentially for collinear configurations. Traditional probability summation models predict that areal summation will improve threshold independent of stimulus configuration. Contrast thresholds were derived from VEP contrast response functions for either circular or elongated Gabor patches with aspect ratios up to 6:1. The carrier orientation was either the same as the patch envelope orientation (collinear) or orthogonal to it. Response amplitudes were larger and contrast sensitivity was higher for collinear configurations. The results are consistent with nonlinear, configuration dependent summation that is more extensive along the axis of orientation.

    View details for Web of Science ID 000075693300007

    View details for PubMedID 9893803

  • OKN, perceptual and VEP direction biases in strabismus VISION RESEARCH Brosnahan, D., Norcia, A. M., Schor, C. M., Taylor, D. G. 1998; 38 (18): 2833-2840


    The present study quantified nasalward/temporalward biases in monocular optokinetic nystagmus (MOKN) and perceived velocity in patients with either early onset esotropia, late onset esotropia and in normals. MOKN was measured with low spatial frequency, small-field gratings drifting at 9.4 degrees/s. MOKN bias was quantified as the ratio of nasalward slow-phase velocity divided by the sum of temporalward and nasalward slow-phase velocities (N/(N + T)). Observers also rated the perceived velocity of gratings moving in nasalward and temporalward directions (3 or 9.4 degrees/s) using a two interval forced choice task. MOKN and perceived velocity biases were correlated negatively in both early onset and late onset groups in the perceptual task--nasalward moving targets were rated as slower than temporalward targets, but in the MOKN task, slow-phase gain was higher for nasalward than for temporalward targets. Oscillatory-motion, visual evoked potentials (VEPs), were recorded in response to 1 c/deg gratings undergoing apparent motion at 10 Hz in a subset of the observers. VEP direction biases were quantified by calculating the ratio of first harmonic response amplitudes to the sum of first and second harmonic amplitudes. Significant correlations were found between the direction biases obtained on all three measures. Perceived velocity and MOKN bias measures were also correlated negatively. Patients with early onset esotropia (infantile esotropia) had larger biases than late onset esotropes or normals on each measure and the biases were more frequently bilateral in the early onset patients. The pattern of result is consistent with early critical periods for the mechanism(s) underlying MOKN, perceived velocity and cortical responsiveness. A single site model for all three asymmetries is unlikely, at least in simple form, because of the negative correlation between MOKN and perceived velocity biases and because of the differences in relative magnitude between the perceptual and MOKN biases.

    View details for Web of Science ID 000074821900011

    View details for PubMedID 9775329

  • Development of directional motion symmetry in the monocular visually evoked potential of infant monkeys VISION RESEARCH Brown, R. J., Wilson, J. R., Norcia, A. M., Boothe, R. G. 1998; 38 (9): 1253-1263


    Motion processing in humans and monkeys exhibit a directional asymmetry during infancy which is not present in adults except following abnormal visual rearing conditions. To characterize the time course for maturation of a symmetric response, we measured the monocular visually evoked potential (MVEP) response to 0.26 c/deg gratings oscillating horizontally at 6 Hz in 13 infant rhesus monkeys between 1 and 52 weeks of age. An asymmetric (F1) and a symmetric (F2) frequency component were extracted from the MVEP using Fourier analysis. At early ages the asymmetric F1 component measured from the two eyes exhibited a 180 deg interocular phase shift, demonstrating that there was a directional bias in opposite directions between the left and right eyes. Although our methods could not determine whether the bias was in the nasal or temporal direction, our results would be consistent with a nasal bias, as has been observed in previous motion studies. Magnitude of the asymmetry was quantified in the form of an asymmetry index, F1/(F1 + F2). Based on developmental changes in the asymmetry index, and phase and amplitudes of F1 and F2, we conclude that the MVEP loses its directional asymmetry at 6 weeks of age. The development of directional motion symmetry observed in monkeys over the first 6 weeks is similar to that observed in humans over the first 5 months.

    View details for Web of Science ID 000073529000007

    View details for PubMedID 9666993

  • Neural noise limitations on infant visual sensitivity NATURE Skoczenski, A. M., Norcia, A. M. 1998; 391 (6668): 697-700


    Visual contrast sensitivity is poor in newborn human infants, but improves rapidly to approach adult levels by 8 months of age. During this period, infant sensitivity can be limited by physical factors affecting photon capture, such as eye size and photoreceptor density. Here we show that infant visual sensitivity is also limited by high levels of noise in the neural transduction process. Using a non-invasive electrophysiological measurement and a visual noise titration technique, we have found that intrinsic neural noise in neonates is approximately nine times higher than in adults. As intrinsic neural noise decreases during infancy, contrast sensitivity improves proportionally, suggesting that neural noise places critical limits on contrast sensitivity throughout development. Moreover, contrast gain control, an inhibitory process that adjusts visual responses to changing stimulation, is in place and operating in infants as young as 6 weeks of age, in spite of high levels of neural noise and significant immaturities in contrast sensitivity. The contrast gain control that we observed in human neonates may serve as a building block for more complex forms of visual inhibition, which develop later in infancy.

    View details for Web of Science ID 000071982500053

    View details for PubMedID 9490413

  • Collinear stimuli regulate visual responses depending on cell's contrast threshold NATURE Polat, U., Mizobe, K., Pettet, M. W., Kasamatsu, T., Norcia, A. M. 1998; 391 (6667): 580-584


    Neurons in the primary visual cortex are selective for the size, orientation and direction of motion of patterns falling within a restricted region of visual space known as the receptive field. The response to stimuli presented within the receptive field can be facilitated or suppressed by other stimuli falling outside the receptive field which, when presented in isolation, fail to activate the cell. Whether this interaction is facilitative or suppressive depends on the relative orientation of pattern elements inside and outside the receptive field. Here we show that neuronal facilitation preferentially occurs when a near-threshold stimulus inside the receptive field is flanked by higher-contrast, collinear elements located in surrounding regions of visual space. Collinear flanks and orthogonally oriented flanks, however, both act to reduce the response to high-contrast stimuli presented within the receptive field. The observed pattern of facilitation and suppression may be the cellular basis for the observation in humans that the detectability of an oriented pattern is enhanced by collinear flanking elements. Modulation of neuronal responses by stimuli falling outside their receptive fields may thus represent an early neural mechanism for encoding objects and enhancing their perceptual saliency.

    View details for Web of Science ID 000071842300050

    View details for PubMedID 9468134

  • Lack of lateral inhibitory interactions in visual cortex of monocularly deprived cats VISION RESEARCH Kasamatsu, T., Kitano, M., Sutter, E. E., Norcia, A. M. 1998; 38 (1): 1-12


    To gain new insight into the effects of monocular deprivation, we studied the visual cortex of adult cats deprived of vision in one eye. Local field potentials were recorded in response to contrast reversal of square-wave gratings modulated in time either by pseudorandom, m-sequences or periodically. We have found that: (1) stimulation of the retinotopic locus of the recording site elicits responses with abnormal waveforms and long latencies from the deprived eye; (2) stimulation of a remote, non-retinotopic locus elicits responses from the non-deprived eye but not from the deprived eye; (3) the monocularly deprived cortex lacks lateral inhibitory interactions which are characteristic of the normal cortex; and (4) steady-state responses showed little difference in spatial-frequency tuning and contrast sensitivity between the deprived and non-deprived eye, mostly conforming to earlier field-potential data in monocular deprivation. Functional lateral interactions appear to be greatly reduced in monocularly deprived cortex.

    View details for Web of Science ID A1998YH85300001

    View details for PubMedID 9474370

  • Spatial frequency masking with the sweep-VEP VISION RESEARCH Peterzell, D. H., Norcia, A. M. 1997; 37 (17): 2349-2359


    A suprathreshold sinewave grating can change the amplitude of the steady-state visual evoked potential (VEP) in response to a test grating if the two are close in spatial frequency (SF). The change in amplitude provides clues to underlying pattern analyzers. Masking was measured in 12 observers using the steady-state VEP. As a test grating reversed at 7 Hz, a masker of similar temporal frequency (9 Hz) but of variable SF was superimposed on it. Test gratings were 1, 3 and 8 c/deg (20% contrast). Within a 10 sec trial, the mask (20 or 40% contrast) was fixed at one of nine SFs or was swept across 19 SFs (5 octaves). The amplitude of the test response (at 14 Hz) was measured as a function of the SF of the masker. Group masking functions were broad (2-3 octaves) and revealed multiple minima. Functions for 1 and 3 c/deg tests each revealed minima near 1 and 3 c/deg. Functions for 8 c/deg tests revealed minima at 3 and 8 c/deg. Doubling the contrast of the mask from 20 to 40% increased masking but in a nonlinear fashion that enlarged the off-peak minima. Swept masks caused slightly more masking than fixed masks, and caused masked amplitudes to exceed unmasked amplitudes (i.e., enhancement) in one condition (3 c/deg.test, 20% contrast mask). The data suggest that each VEP masking function reflects the outputs of multiple spatial analyzers, that a discrete set of analyzers may underlie the data, and that the efficient sweep-VEP can measure SF tuning.

    View details for Web of Science ID A1997XP66800004

    View details for PubMedID 9381671

  • A method for investigating binocular rivalry in real-time with the steady-state VEP VISION RESEARCH Brown, R. J., Norcia, A. M. 1997; 37 (17): 2401-2408


    Under conditions in which the visual system cannot reconcile dissimilar images from the two eyes, perception typically alternates between the two half-images-a process known as binocular rivalry. We report a real-time, steady-state VEP method that is a sensitive detector of the continuous alternations in perceptual dominance across the eyes. This method works by labelling each half-image with a slightly different temporal frequency so that the record generated by each can be recovered from the EEG by spectrum analysis. In this way, one can track the "waxing" and "waning" of the VEP amplitudes for each eye simultaneously during spontaneous rivalry, permitting an analysis of the relative physiological dominance of each eye in real-time. Such alternations were clearly observed in the VEP amplitudes generated by each half-image during rivalry (the amplitudes for the two eyes correlated negatively). In contrast, VEP amplitudes for the two eyes varied either synchronously or randomly when the half-images were allowed to fuse. The instances of physiological dominance of each eye as evidenced by the VEP correlated well with the subjects' report of perceptual dominance. This purely electrophysiological method appears to be suitable for measuring rivalry in non-verbal human or animal subjects, as it does not require active participation from them.

    View details for Web of Science ID A1997XP66800008

    View details for PubMedID 9381675

  • Abnormal long-range spatial interactions in amblyopia VISION RESEARCH Polat, U., Sagi, D., Norcia, A. M. 1997; 37 (6): 737-744


    Neural interactions between widely separated stimuli were explored with psychophysical and visual evoked potential (VEP) measures in normal and amblyopic observers. Contrast detection thresholds were measured psychophysically for small foveally viewed Gabor patches presented in isolation and in the presence of similar, but laterally displaced flanks. The amplitude and phase of VEPs elicited by similar targets were also measured. The presence of neural interaction between the target and flank responses was assessed by comparing the unflanked threshold to the flanked threshold in the psychophysical experiments and by comparing the response predicted by the algebraic sum of test and flank responses to that measured when test and flanks were presented simultaneously. In normal observers simultaneous presentation of test and flank targets produces a VEP response that is up to a factor of two larger than the linear prediction (facilitation). Psychophysical threshold is also facilitated by a comparable factor. Facilitation was found mainly for configurations in which local (carrier) and global (patch) orientations resulted in collinearity, independent of global orientation (meridian). Amblyopic observers showed several deviations from the normal pattern. The facilitation for the collinear configurations was either markedly lower than normal or was replaced by inhibition. The normal pattern of spatial interaction may facilitate the grouping of collinear line segments into smooth curves. In contrast, abnormal long-range spatial interactions may underlie the grouping disorders and perceptual distortions found in amblyopia.

    View details for Web of Science ID A1997WM12200009

    View details for PubMedID 9156218

  • Neurophysiological evidence for contrast dependent long-range facilitation and suppression in the human visual cortex VISION RESEARCH Polat, U., Norcia, A. M. 1996; 36 (14): 2099-2109


    Long-range spatial interactions in human visual cortex were explored using a lateral masking paradigm. Visual evoked potentials (VEPs) elicited by a Gabor signal presented in isolation or in the presence of two flanking high-contrast Gabor signals (masks) were measured. Response amplitude and phase were recorded for a vertically oriented test, for horizontal and vertical masks and for combinations of vertical tests and vertical or horizontal masks. The amplitudes and phases of the test alone and mask alone responses were added coherently to predict the amplitude for collinear and orthogonal lateral masking conditions. Additivity failures were taken as evidence for neural interactions. At a target-to-mask distance of 2 deg, VEP amplitude exceeded the linear prediction for test contrasts in the range of 8-16% for the collinear, co-axial target/mask combination. Measured response phase also led predicted response phase over the same range of contrast. The VEP amplitudes were less than the linear prediction in the orthogonal target/mask combination and measured response phase lagged the predicted phase. Significant facilitation occurred with collinear test/mask combinations up to at least 3 deg of separation (nine wavelengths). Co-oriented, but non-collinear test/mask combinations (oblique test and mask, horizontal test and mask) did not produce facilitation. Contrast gain thus appears to be set over considerable distances in a configuration-specific fashion.

    View details for Web of Science ID A1996UY57500009

    View details for PubMedID 8776476

  • Development of grating acuity and contrast sensitivity in the central and peripheral visual field of the human infant VISION RESEARCH Allen, D., Tyler, C. W., Norcia, A. M. 1996; 36 (13): 1945-1953


    Central and peripheral visual functions were measured simultaneously in 39 infants from 10 to 39 weeks old using a dual-frequency VEP technique. Central acuity and contrast sensitivity over a 4 deg circular field were measured at 6 or 8 Hz. Peripheral acuity and contrast sensitivity were measured simultaneously at the other rate with a semi-circular stimulus extending from 8 to 16 deg. The EEG was analyzed at 12 and 16 Hz to determine the separate responses for the central and peripheral fields. Both central and peripheral VEP acuity developed over the age range tested. Central acuity improved by about a factor of 2.6 over the age range tested, while peripheral acuity improved by about a factor of 2.2. Central acuity was always higher by an average factor of about 2.3. Contrast sensitivity showed similar development for the central and peripheral fields with an early rapid rise in sensitivity.

    View details for Web of Science ID A1996UV00700010

    View details for PubMedID 8759434

  • Abnormal motion processing and binocularity: Infantile esotropia as a model system for effects of early interruptions of binocularity EYE Norcia, A. M. 1996; 10: 259-265


    Infantile esotropia, a common form of strabismus with onset prior to 6 months of age, occurs at a time of rapid visual development. While monocular visual acuity is relatively unaffected in these patients, the majority of them fail to achieve fully normal stereopsis. In addition, these patients show a spectrum of abnormalities in their ocular following responses, visual perception and visual evoked potentials (VEPs) that suggest a failure to develop a normal complement of motion processing mechanisms. While abnormalities of of stereopsis have been studied for many years, motion processing in strabismus is a rapidly evolving area of current research. Motion mechanisms are normally binocular and may form a distinct binocular sub-system. This review summarises which is known about sensory and motor abnormalities in infantile esotropia, with special emphasis on recent motion VEP recordings. The monocular motion VEP shows directional biases early in infancy that are consistent with a nasalward/temporalward response bias. Patients with infantile esotropia maintain their neonatal biases beyond the age at which they normally disappear. The motion VEP biases persist into visual maturity in patients whose strabismus is treated after about 2 years of age. Treatment prior to age 2 can lessen the magnitude of the motion VEP asymmetry and these improvements can be maintained into visual maturity. A recording from the striate cortex of a visually deprived macaque monkey indicates that the motion VEP asymmetry arises early in the visual pathway.

    View details for Web of Science ID A1996UK09600017

    View details for PubMedID 8776457

  • Measurement of vernier and motion sensitivity with the rapid-sweep-VEP 5th Meeting of the Bielschowsky-Society-for-Research-of-Squint Wesemann, W., Norcia, A. M., Manny, R. E. FERDINAND ENKE VERLAG. 1996: 11–17


    The visual abilities of infants and small children, who are unable to communicate verbally, can be determined with "objective" visual acuity tests as, e.g., preferential looking (PL) or visual evoked potentials (VEP). Both methods provide an estimate of grating visual acuity, but are unable to determine optotype visual acuity. Grating acuity, however, is not an optimum indicator for visual performance, as it is less affected than optotype acuity by pathological changes. The diagnostic relevance of objective methods may be improved by testing visual functions that are more strongly degraded by a malfunction of the fovea than is grating acuity. Visual functions such as vernier acuity and relative motion sensitivity are potential candidates.Characteristic properties of vernier- and motion-VEP have been determined on adult subjects with a rapid-sweep-technique at various eccentricities.Both vernier and motion stimuli elicited VEP-responses at the 2nd harmonic (F2) of the stimulus frequency. A VEP-response at the fundamental frequency (F1) could be recorded with the vernier stimulus only. Foveal VEP-thresholds were very similar to psychophysical thresholds lying in the range from 11 to 25 arcsecs. The eccentricity dependence of the VEP-threshold recorded with vernier- and motion stimuli shows different cortical magnifications and supports the notion that the F1 response is related to the detection of the vernier offset, whereas the F2 response is generated by motion components of the stimuli.Vernier and relative motion-related VEP thresholds can be recorded with the steady-state VEP. They may provide more sensitive tests of foveal vision loss than grating visual acuity tests.

    View details for Web of Science ID A1996TV31000003

    View details for PubMedID 8839339



    Monocular oscillatory-motion visual evoked potentials (VEPs) were measured in prospective and retrospective groups of infantile esotropia patients who had been aligned surgically at different ages. A nasalward-temporal response bias that is present prior to surgery was reduced below pre-surgery levels in the prospective group. Patients in the retrospective group who had been aligned before 2 yr of age showed lower levels of response asymmetry than those who were aligned after age 2. The data imply that binocular motion processing mechanisms in infantile esotropia patients are capable of some degree of recovery, and that this plasticity is restricted to a critical period of visual development.

    View details for Web of Science ID A1995TE46000009

    View details for PubMedID 8560799



    A 2-weight adaptive filter that determines the amplitude and phase of steady-state evoked potentials is presented. Reference signals are derived from the visual stimulator that are related to corresponding harmonics of the response and the filter weights are adjusted so as to minimize the squared estimation error between the reference and the recorded signal using the recursive least squares (RLS) method. The filter, which acts as an adaptive bandpass filter, is followed by a detector based on the T2circ statistic. The performance of the RLS adaptive filter was compared to that of the conventional Discrete Fourier Transform (DFT) and the filtered DFT of Tang and Norcia in a series of simulations with known sinusoids buried in Gaussian noise and in EEG noise. In the simulations, the RLS adaptive filter detected signals at about 3-4 times lower signal to noise ratios than did the DFT. The RLS filter also outperformed the filtered DFT. Qualitatively similar results were obtained with human visual evoked potential recordings. The adaptive RLS filter significantly outperforms both the DFT and filtered DFT and is much simpler to implement than the filtered DFT method of Tang and Norcia.

    View details for Web of Science ID A1995QZ16100008

    View details for PubMedID 7750452



    Field potentials were recorded from cat striate cortex, either between an epidural screw electrode and a cannula-electrode inserted deep in the gray matter (transcortical recording) or with a pair of metal microelectrodes. Electrodes were placed bilaterally near the cortical projection of the area centralis. The horizontal separation of the recording tips was approximately 2 mm and approximately 300 microns, respectively. The area of the visual field providing input to the recording site (receptive field) was determined by measuring the field potentials generated by contrast reversal of high-contrast, achromatic bar gratings. Five-degree-diameter grating patches were presented individually over a large area of the visual field. The gratings were contrast-reversed at 4, 6 or 10 Hz, while also being swept in spatial frequency between 0.56 and 5.24 c/deg. The receptive fields were approximately 20 deg across or more, substantially larger than expected on the basis of cortical retinotopy. Responses were also elicited by stimulation of the hemi-field contralateral to that contributing to the classical receptive field, implicating the presence of a callosal projection. The large, spatially distributed receptive fields consisted of patches of high and low sensitivity. Continuous cortical infusion of either 100 microM tetrodotoxin or 10 mM muscimol at the recording site totally suppressed the transcortically recorded field potentials, proving that the local field potentials were generated postsynaptically. The present findings suggest that a cluster of cortical cells near the projection site of the area centralis receives input from remote cortical regions to an extent that is comparable with that of anatomically demonstrated long-range lateral connections.

    View details for Web of Science ID A1995RC03200012

    View details for PubMedID 7672022



    A new method to detect steady-state evoked potentials (EPs) is presented. The technique is based on a two-weight recursive least squares (RLS) adaptive filter and the Tcirc2 statistic. Simulations with known sinusoids buried in Gaussian noise and in EEG noise indicate that the adaptive filter can detect signals at 3 or 4 times lower signal-to-noise ratios that the discrete Fourier transform (DFT). Qualitatively similar results were obtained with human visual evoked potential recordings.

    View details for Web of Science ID A1995RE99400005

    View details for PubMedID 7666685

  • RETINOTOPIC AND NONRETINOTOPIC FIELD POTENTIALS IN CAT VISUAL-CORTEX VISUAL NEUROSCIENCE Kitano, M., Niiyama, K., Kasamatsu, T., Sutter, E. E., Norcia, A. M. 1994; 11 (5): 953-977


    Two types of field potentials were identified in cat visual cortex using contrast reversal of oriented bar gratings: a short-latency fast-local component with a retinotopic organization similar to that seen with single-unit discharges at the same cortical site, and a slow, nonretinotopic component with a longer peak latency. The slow-distributed component had an extensive receptive field mapped by measuring the amplitude of binary kernels and showed strong inhibitory interactions within the receptive field. The peak latency of the slow-local component increased with distance from the retinotopic center, suggesting a possible conduction delay. Both components showed some orientation bias depending on the laminar location, but the bias could be independent of the orientation preferred by single units in the immediate vicinity. The present findings indicate that locally generated field potentials reflect cortical mechanisms for nonlinear integration over wide areas of the visual field.

    View details for Web of Science ID A1994PC70800011

    View details for PubMedID 7947408



    Using the sweep visual evoked potential (VEP), we have measured oscillatory displacement thresholds (OMTs) in 49 infants ranging in age from 7 to 54 weeks of age. The stimuli were high-contrast (80%), sine-wave gratings (1 c/deg) undergoing oscillatory displacements at 6 Hz. In addition to the motion thresholds, contrast thresholds for phase-reversing (6 Hz), 1 c/deg gratings were measured in the same session for 26 infants. In the main experiment, responses were recorded at the second harmonic (F2) of the stimulus frequency (12 Hz) under binocular viewing conditions. Our main finding is that, over the age range during which infants' peak contrast sensitivity (CS) first develops to within a factor of 2 of adult CS (9-12 weeks), infants' sensitivity to grating displacement is a factor of approximately 10 less than adults'. Moreover, infants' sensitivity to oscillatory motion undergoes relatively little development over the period between 2 and 15 months postnatal, gradually achieving a factor of 4.5 below adult values by 1 yr of age. Averaged over the entire age range tested, infants' OMTs were 167 sec arc, a factor of 6.4 times higher than the average OMT (26 sec arc) for 13 adults tested under identical conditions. In contrast, the infants' average CS for reversing gratings averaged only a factor of 2.5 less than the adults' average CS. In a second experiment, we took advantage of a developmental asymmetry in the monocular oscillatory motion VEP which allows for unambiguous identification of direction selective responses from very young infants. Monocular motion VEPs were measured in five infants (8-14 weeks) and their data analyzed at the fundamental frequency (F1). Responses at F1 were present in the monocular motion VEP from each infant and were 180 deg out of phase between the two eyes, identifying them as directional cortical responses with a nasalward/temporalward bias. These directional thresholds were equal to or lower than the symmetric (F2) thresholds. The presence of directional asymmetry in the motion VEP and the similarity of the monocular F1 and F2 OMTs support the notion that the OMTs measured in the main experiment were, in fact, derived from the responses of directionally selective cells in visual cortex. These data also imply that the OMTs are not derived from local contrast-reversal responses. Other models to explain infants' relative insensitivity to oscillatory motion are discussed.

    View details for Web of Science ID A1994PB29700005

    View details for PubMedID 7975278



    We have examined the effects of preoperative, full-time alternate occlusion on the development of visual motion processing mechanisms. Motion visual evoked potentials (MVEPs) were recorded longitudinally in 14 infantile esotropia patients during the course of standard preoperative occlusion therapy. The MVEP in these patients was initially asymmetric in a fashion consistent with a nasalward/temporalward response bias, with a motion asymmetry significantly higher than that of age-matched normals. The magnitude of the developmental motion asymmetry declined significantly after an average of 24 weeks of alternate occlusion. This result implies that the binocular motion-sensitive cells underlying the MVEP retain some degree of plasticity up to at least 1 year of age. Our data suggest further that the persistence of motion asymmetries in untreated infantile esotropia patients is maintained by an active process that can be disrupted by alternate occlusion. Alternate occlusion apparently eliminates a form of abnormal binocular interaction that supports the persistence of the motion asymmetry. We propose that one of the necessary pre-conditions for symmetricization of motion processing in infantile esotropia is the absence of abnormal competitive binocular interactions.

    View details for Web of Science ID A1994NB81500001

    View details for PubMedID 8195968



    Chromatic sensitivity is very low in humans during the first few months of life. We examined whether low chromatic sensitivity reflects a deficiency among chromatic mechanisms or whether it is simply a manifestation of poor visual sensitivity in general. The sweep VEP was used to measure contrast sensitivity to gratings varying in the mixture of red and green components. For infants from 2 to 8 weeks of age, sensitivity to all mixtures was lower than color-normal adults' sensitivity, but infant and adult ratios of luminance/chromatic sensitivity were similar. This finding is consistent with the hypothesis that infants have functional MWS and LWS cones and the requisite post-receptor chromatic mechanisms to compare their signals.

    View details for Web of Science ID A1993LZ83700018

    View details for PubMedID 8249334



    Two related procedures for estimating the parameters of steady-state evoked potentials (SSEPs) are introduced. The first procedure involves an initial stage of digital bandpass filtering followed by a Discrete Fourier Transform analysis. In the second method, a high resolution method based on parametric modelling is applied to the filtered data. The digital pre-filter consists of a non-phase shifting Chebychev bandpass filter. The parametric modelling method considers the evoked-response-plus-noise distribution to consist of a set of exponentially damped sinusoids. The frequency, amplitude, phase and damping factors of these components are estimated by calculating the mean of the forward and backward prediction filters and linear regression. We compared the signal-to-noise ratio (SNR) of the new procedures to the conventional Discrete Fourier Transform method for Monte Carlo simulations utilizing known sinusoids buried in white noise, known sinusoids buried in human EEG noise and for a sample of visual evoked potential data. Both of the new methods produce substantially more accurate and less variable estimates of test sinusoid amplitude. For VEP recording, the EEG background noise level is reduced by 5-6 dB over that obtained with the DFT. The new methods also provide approximately 5 dB better SNR than the DFT for detection of sinusoids based on the Rayleigh statistic. The parametric modelling approach is particularly suited for the analysis of very short data records including cycle-by-cycle analysis of the SSEP.

    View details for Web of Science ID A1993LR33000009

    View details for PubMedID 7688287



    Observer sensitivity to oscillatory step displacements of sine-wave gratings was investigated at various loci in the visual field (0-30 degrees) as a function of contrast. Detection thresholds at 10 Hz and high grating contrasts were approximately 11-15 arcsec in the fovea and 37-47 arcsec at 30 degrees eccentricity. At any given contrast, threshold displacement increases linearly with eccentricity. The data provide evidence against an interpretation based on cortical magnification, because the slope and the scale-free x intercept of the eccentricity function vary strongly with contrast. While foveal thresholds for high-contrast gratings are in the range of the hyperacuities, the oscillatory motion threshold falls off an order of magnitude more slowly than the traditional hyperacuities. Rather than conceiving of the oscillatory motion threshold as a spatial acuity limited by cortical magnification, we suggest an alternative approach that is based on a form of contrast discrimination. Oscillatory motion can be decomposed into the sum of a modulating counterphase grating and a static masking grating, both of which are in spatial quadrature (i.e., 90 degrees out of phase). At low grating contrast, oscillatory motion can be detected when the counterphase component exceeds a constant contrast value. Above a critical contrast value of the static component Cscrit, threshold rises as a power function of contrast with a slope near 1.0. The critical contrast value Cscrit increases linearly with eccentricity, indicating that oscillating gratings observed with the peripheral visual field are less easily masked compared with foveally fixated gratings.

    View details for Web of Science ID A1992JQ42200002

    View details for PubMedID 1403240



    We have compared the performance of an off-axis (knife-edge) photorefractor with that of an on-axis (isotropic) system. Normal infants and children between the ages of 8 and 208 weeks were photographed with each camera both with and without cycloplegia. Refractive errors were estimated for each technique based on equations derived from ray-tracing. These refractions were compared to the results of retinoscopy under cycloplegia. Sensitivity and specificity of the two photorefraction systems were evaluated as a function of the magnitude of meridional hyperopia defined by retinoscopy. We also examined the effect of varying the photorefraction screening criterion. Thirteen percent of the infants in the screening sample presented with +3.50 diopters or more of meridional hyperopia. Using this level of ametropia as a referral criterion, the sensitivity and specificity of the off-axis system for infants without cycloplegia were 83% and 72%, respectively. For the on-axis system, sensitivity and specificity values were 85% and 53%. The use of cycloplegics did not significantly improve the performance of either system, but rather their use degraded the specificity of the on-axis system in the presence of moderate refractive errors. The results of the present study indicate that both on- and off-axis systems are effective in identifying highly ametropic infants, but that the off-axis system results in significantly fewer false positives. Moreover, the off-axis system has the advantages of an inherently greater dynamic range for a fixed camera design, and also more easily interpreted photographs.

    View details for Web of Science ID A1992JK32800009

    View details for PubMedID 1512665



    An optical analysis of eccentric photorefraction (photoretinoscopy) of astigmatic eyes is presented. The size and the angular tilt of the dark crescent appearing in the subject's pupil are derived as a function of five variables: the ametropia of the eye (Dsph, Dcyl, axis), the eccentricity of the flash, e, and the distance of the camera from the subject's eye, dc. A simplified solution and a solution of the inverse problem, which enable one to calculate the degree of ametropia from the size and the tilt of the crescent, are also presented. If the crescent is smaller than the pupil, both the size and the tilt of the dark crescent are independent of the pupil size. The angular tilt of the crescent is also independent of the eccentricity. Characteristic changes of the crescent as a function of the cylinder axis are illustrated for compound and mixed astigmatisms. The validity of the theoretical predictions was experimentally verified on a model eye.

    View details for Web of Science ID A1991GX03200026

    View details for PubMedID 1783948



    Two forms of visual deficit were induced in 12 pairs of kittens (4-6 weeks of age) by monocular lid suture combined with either an opaque soft contact lens ('occlusion' amblyopia) or a clear lens ('diffusion' amblyopia) which had been kept behind the sutured eyelids for 5-9 weeks. The kittens were then reverse-sutured at the age of 9-15 weeks, and the previously open eye was occluded with an opaque lens for the next 5-7 weeks. We compared across the two groups the proportion of binocularly driven cells (group 2-6) and cells predominantly activated by stimulation of the initially deprived eye (groups 1-3). Both values were significantly higher in the 'occlusion' kitten, though the difference was small, compared to the 'diffusion' kitten. In addition, we measured visual acuity and peak contrast sensitivity for the two eyes in 4 of the 12 pairs, using the sweep VEP method applied under anesthesia and paralysis. The 'occlusion' kitten showed consistently better acuity and higher peak contrast sensitivity than the 'diffusion' kitten, when the initially deprived eye was tested. Taken together, the present results suggest that there is a difference in the depth of amblyopia caused by monocular 'occlusion' and monocular 'diffusion'. The cortical effects of the latter are more difficult to reverse than those of the former.

    View details for Web of Science ID A1991FP17000010

    View details for PubMedID 1651147



    Visual evoked potentials (VEPs) were recorded monocularly in response to vertical gratings that underwent oscillatory apparent motion at a temporal frequency of 10 Hz. In normal infants 6 months or younger and in patients with a history of constant strabismus onset before 6 months of age, the oscillatory motion VEP contains a prominent first harmonic component that is temporally 180 degrees out of phase in the two eyes. This pattern is not seen in normal adults and is consistent with the presence of a nasalward/temporalward asymmetry of cortical responsiveness in infants and in patients with early onset strabismus.

    View details for Web of Science ID A1991EX76600027

    View details for PubMedID 1993597

  • DEVELOPMENT OF CONTRAST SENSITIVITY IN THE HUMAN INFANT VISION RESEARCH Norcia, A. M., Tyler, C. W., Hamer, R. D. 1990; 30 (10): 1475-1486


    Contrast sensitivity and grating acuity were measured using the sweep VEP method in a group of 48 infants from 2 to 40 weeks of age and in a group of 10 adults. Sinusoidal gratings were reversed in contrast at 12 alternations per sec at a space-average luminance of 220 cd/m2. During 10 sec trials, either the contrast or the spatial frequency was increased in a series of 19 steps. Thresholds were estimated by extrapolation of the VEP response functions to zero amplitude. The contrast threshold at low spatial frequencies developed rapidly from 7% contrast at 2-3 weeks to an asymptote of 0.5% at 9 weeks. For adults, maximum sensitivity at low spatial frequencies was 0.32-0.22%. The sweep VEP estimate of grating acuity showed a gradual increase in spatial frequency with age, starting at 5 c/deg during the first month and reaching 16.3 c/deg at 8 months. The mean adult acuity was 31.9 c/deg. There appeared to be two phases in the development of contrast sensitivity and acuity. Between 4 and 9 weeks overall contrast sensitivity increased by a factor of 4-5 at all spatial frequencies. Beyond 9 weeks, contrast sensitivity at low spatial frequencies remained constant, while sensitivity increased systematically at higher spatial frequencies.

    View details for Web of Science ID A1990DX33300009

    View details for PubMedID 2247957

  • Polaroid photorefractive screening of infants. Journal of pediatric ophthalmology & strabismus Hsu-Winges, C., Hamer, R. D., Norcia, A. M., Wesemann, H., Chan, C. 1989; 26 (5): 254-260


    We modified a Polaroid SE camera for use as a photoretinoscope. A total of 187 infants between 2 and 18 months of age were photographed using this device. About half of these infants (97) participated in a double blind study in which the results of photorefraction were compared with those of standard cycloplegic retinoscopy. Eighty-three infants were photographed without cycloplegia. Thirty-four infants were photographed while cyclopleged. Photographs were evaluated for significant refractive errors and other ocular abnormalities. The effectiveness of the camera system to screen for significant refractive errors without the use of cycloplegia was assessed. Infants were identified to be at risk by photorefraction if, in any photograph, a hyperopic bright crescent calculated to be greater than or equal to +1.25 D was present in the pupil. Clinically significant refractive errors were defined by the results of cycloplegic retinoscopy: "at-risk" infants had either 3.5 D or more hyperopia in either eye, or astigmatism in either eye greater than or equal to 2.5 D, or anisometropia greater than or equal to 1.5 D. With these clinical criteria and the above photographic screening criterion, the camera's sensitivity and specificity were 83% and 69%, respectively. The present system compares favorably with earlier, more sophisticated units in alerting practitioners to potentially significant refractive errors in infants. Additionally, as a screening tool, this device offers the benefits of being inexpensive and easy to use, and of providing immediate feedback.

    View details for PubMedID 2795416



    The development of monocular and binocular grating acuity was measured in 87 infants, 2-52 weeks of age, using the sweep VEP technique. Average monocular and binocular acuity growth functions were nearly identical, with a small (less than 0.2 octaves) binocular acuity superiority occurring only under 6 months. Interocular acuity differences were small (averaging less than 1/4 octave, unsigned, with a 95% confidence interval of less than +/- 0.6 octaves) and were not significant at any age. These characteristics make the sweep VEP technique a potentially sensitive tool for the detection of monocular visual losses in the early stages of amblyopia.

    View details for Web of Science ID A1989T971200004

    View details for PubMedID 2781730



    Contrast response functions (CRFs) for the VEP were obtained with a Discrete Fourier Transform (DFT) technique employing swept contrast gratings. VEP CRFs in infants were found to have a form similar to those observed in adults, being linear functions of log contrast over a range of near-threshold contrasts. CRFs with low and high contrast lobes were present in infants, as they are in adults. Contrast thresholds were estimated by extrapolation of the CRF to zero microvolts. The effects of additive EEG noise and of the DFT data window on the shape of the measured CRF are considered. For large signals, the measured CRF is nearly independent of the additive noise, but at small signal values additive noise introduces a small bias towards larger amplitudes. The VEP signal-plus-noise distribution was modeled as a family of Rice distributions in order to evaluate the effects of bias on the estimates of threshold. The amount of bias depends inversely upon the slope of the CRF. The amount of bias introduced by a smoothing window also depends upon slope of the CRF as well as the sweep rate. The combined effects of additive noise and window bias were such that the total bias was nearly independent of CRF slope. Sweep VEP contrast thresholds were shown empirically to be unaffected by changes in the range of contrast swept.

    View details for Web of Science ID A1989T971300012

    View details for PubMedID 2603399



    Monocular and binocular grating acuities were measured using a swept spatial frequency visual evoked potential (VEP) technique in a group of fifteen infants with esotropia and alternating fixation. Both monocular and binocular acuity measures fell significantly below the mean for age-matched normals. Infants with esotropia and alternating fixation did not have significant interocular acuity differences.

    View details for Web of Science ID A1988M122000023

    View details for PubMedID 3338891



    The visual evoked potential (VEP) was used to estimate photopic contrast sensitivity of 10-week-old infants over a wide range of spatial frequencies including the acuity limit. Adult and infant VEP contrast sensitivity was compared for sinusoidal luminance gratings reversed in contrast at 6 Hz. Space-average luminance was 220 cd/m2. Grating contrast was swept from well below the measured thresholds to well above them in 10 sec trials. Contrast thresholds were defined as the zero voltage intercept of the initial rising portion of the VEP amplitude versus contrast function. The VEP contrast sensitivity of 10-week-old infants was close to that of the adults for spatial frequencies below about 1 cycle (c)/deg.

    View details for Web of Science ID A1988L896700006

    View details for PubMedID 3335433

  • Visual acuity development in normal and abnormal preterm human infants. Journal of pediatric ophthalmology & strabismus Norcia, A. M., Tyler, C. W., Piecuch, R., Clyman, R., GROBSTEIN, J. 1987; 24 (2): 70-74


    The grating acuity of preterm infants was determined by measurements of the visual evoked potential (VEP) produced by phase alternation of sinusoidal luminance gratings. The development of visual acuity in healthy preterm infants appears to be accelerated when compared with full term infants of the same post-conceptual age. Cortical insults and the more advanced stages of retinopathy of prematurity may adversely affect acuity development as indexed by the VEP. Preterm twins also appear to have lower acuity than preterm monoparous infants.

    View details for PubMedID 3585654



    Kaakinen (1979) presented a method of measuring refractive error and ocular alignment by simultaneous photography of corneal and fundus reflexes. As presented, the technique was unable to detect refractive errors of less than 2-3 diopters (Howland 1980; Kaakinen 1979). We demonstrate that the use of a catadioptric lens and a long working distance can improve sensitivity to less than 1.0 D. Refractions of a model eye, an accommodating eye and ametropic eyes indicate that refractive errors greater than 0.75 D are readily detectable. The improvement in sensitivity achieved by the present system is attributable to a reduction of the angle between the flash source and the entrance pupil of the photorefractor.

    View details for Web of Science ID A1986E199000003

    View details for PubMedID 3776500



    We measured the human contrast sensitivity function (CSF) both electrophysiologically with the steady-state visual evoked potential (VEP) and psychophysically using a method of ascending limits. VEP contrast thresholds were determined using a rapid recording technique in which the contrast of a counterphase modulated sine wave grating was swept logarithmically from 0.5 to 40% over a period of 10 s. For this pattern reversal stimulus the amplitude and phase of the second harmonic response as a function of contrast were measured using a discrete Fourier transform (DFT). Psychophysical thresholds were determined on the same trials used to record the VEP. Near threshold the VEP amplitude vs. contrast function was approximately linear and VEP contrast thresholds were estimated by a linear extrapolation to zero amplitude. The contrast thresholds obtained by the two methods correlated at 0.914 for 5 observers, with a mean discrepancy of only 12%. At higher contrasts, the VEP amplitude vs. contrast function often became nonmonotonic, sometimes showing two amplitude peaks.

    View details for Web of Science ID A1986C735000008

    View details for PubMedID 3728639



    Two prominent frequency components designated f1 and f2 have been identified in the visual evoked response to the transient presentation of sinusoidal luminance gratings in the range of 0.5-8 c/deg. The components occur at temporal frequencies below the alpha band, with the f1 frequency being roughly half that of the f2 frequency. The f1 component is largest at low spatial frequencies with f2 becoming progressively dominant as spatial frequency is increased. The frequency and amplitude of f1 and f2 change substantially over the time course of the response. This has been studied by calculating the temporal frequency spectrum of the transient evoked potential over successive short-time epochs running through the response. Using this technique, the response is shown to consist of narrow-band frequency peaks or 'formants' emerging at different times after stimulus onset. These formants occur at frequencies other than those of the spontaneous EEG and undergo changes in frequency and amplitude over the time course of the response. Two spectrum analysis techniques were employed: the Discrete Fourier Transform and Linear Predictive Coding. Frequency components were successfully identified in single-trial responses using the LPC technique.

    View details for Web of Science ID A1986C198900007

    View details for PubMedID 2420574



    Contrast sensitivity functions (CSF's) were measured for a group of 6-month-old infants using the visual evoked potential (VEP). Sine-wave luminance gratings were counterphase modulated at 12 contrast reversals per s and simultaneously swept in contrast. Each contrast sweep lasted 10 s, spanning a range of 0.5 to 40% contrast in 19 equal logarithmic steps. The amplitude and phase of the response at the second harmonic were determined by a discrete Fourier transform. Contrast thresholds were estimated from a linear extrapolation to zero-amplitude of the VEP amplitude vs. log-contrast function. Contrast sensitivity was found to be nearly adult-like at 1 c/deg, but was not yet mature at higher spatial frequencies.

    View details for Web of Science ID A1986AXY7900003

    View details for PubMedID 3942183



    Refractive errors were determined photographically in a group of infants and children and were compared to conventional cycloplegic retinoscopy. The refractor consisted of a mirror telephoto lens and strobe flash designed to mimic the action of a retinoscope. Significant amblyogenic conditions such as anisometropia and high isoametropia were detectable. Strabismus and media opacities were also recordable. The technique provides a potential mass suited for infants.

    View details for Web of Science ID A1986AYV8900006

    View details for PubMedID 3951813



    Visual sensitivity to stereoscopic disparity changes was measured both psychophysically and by means of evoked potentials. The binocular disparity of a dynamic random-dot stereogram portraying a single flat plane alternated between two values symmetrical about the plane of fixation. The threshold for disparity alternation of the stereoscopic plane was determined at alternation rates between 4 and 12 depth reversals per second (rps). Evoked potential and forced-choice psychophysical estimates of stereoscopic threshold at each reversal frequency agreed, with a mean discrepancy of only +/- 0.1 log units. Evoked potential amplitude was a linear function of log disparity up to about 15 arc min peak to peak disparity. For larger disparities, the evoked potential amplitude versus log disparity function was found to be nonmonotonic with a dip occurring at approximately 26 arc min disparity. Responses to fine disparities of less than 20 arc min lay close to one temporal phase while those evoked by coarse disparities greater than 40 arc min lay near a different phase. The data suggest that disparity processing mechanisms either undergo dynamic changes as disparity increases or that processing shifts between at least two independent mechanisms.

    View details for Web of Science ID A1985AVZ8700008

    View details for PubMedID 3832583



    The reliability of visual evoked potential (VEP) measurements of acuity was determined by estimating acuity for sinewave luminance gratings which were counterphase modulated at either 12 or 20 reversals/sec (rps). Gratings were swept in spatial frequency beyond the acuity limit and acuity was estimated from an extrapolation based on the last peak in the VEP amplitude versus spatial frequency function. Twenty-five infants ranging in age from 17 to 25 weeks were studied. Individual 10 sec sweeps resulted in records with a criterion response in 65-75% of trials. The reliability of acuities obtained from individual 10 sec sweeps was +/- 0.54 octaves at 95% confidence across 12 and 20 rps recording conditions (RMS error of +/- 0.27 octaves). The best acuity attained by each infant on either a single sweep, or on their vector average, was reliable to +/- 0.38 octaves at 95% confidence (RMS error of +/- 0.19 octaves) compared to a range of individual acuities of about 2 octaves. Much of the variability of sweep VEP acuity in cross-sectional samples of infants is therefore attributable to reliable individual differences rather than to measurement error. In testing individual infants our analysis indicates that choice of temporal frequency accounts for only 14% of the variation in acuity estimates within subjects.

    View details for Web of Science ID A1985AUC7000004

    View details for PubMedID 2412787



    The grating acuity of 197 infants from 1 week to 53 weeks of age was measured using the visual evoked potential (VEP) in response to counterphase grating stimulation. The gratings were presented as a 10 sec spatial frequency sweep which spanned the acuity limit. The amplitude and phase of the second harmonic response were extracted by discrete Fourier analysis. The VEP amplitude versus spatial frequency function showed narrow spatial frequency tuning with amplitude peaks at one or more spatial frequencies. The phase of the response at medium to high spatial frequencies was generally constant at a spatial frequency peak, followed by a progressive phase lag with increasing spatial frequency. Grating acuity was estimated by linear extrapolation to zero microvolts of the highest spatial frequency peak in the VEP amplitude versus spatial frequency function. This visual acuity estimate increased from a mean of 4.5 c/deg during the first month to about 20 c/deg at 8-13 months of age. The VEP acuities at 1 month are a factor of three to five higher than previously reported for pattern reversal or pattern appearance stimuli. By 8 months VEP grating resolution was not reliably different from adult levels in the same apparatus.

    View details for Web of Science ID A1985AUP2000005

    View details for PubMedID 4090273


    View details for Web of Science ID A1985AVV5700003

    View details for PubMedID 19493780



    Temporal processing limits were determined for two types of stereoscopic percept associated with square wave disparity alternation: apparent depth motion and depth pulsation. The stimuli were dynamic random dot stereograms containing no monocular cues for either target motion or disparity change. The percept of a single noise plane undergoing apparent depth motion coincided with the extent of a large peak in the low frequency portion of the evoked potential amplitude spectrum. The limit for apparent depth motion was approximately 6 Hz. Above this frequency two pulsating depth planes were seen simultaneously. Depth pulsations wer visible up to 14 Hz and an evoked potential occurred in synchrony with each disparity change (up to 28 depth reversals/sec). Above 14 Hz two transparent planes were perceived without depth pulsation and no stereoscopic evoked potential could be recorded. The results indicate a higher temporal resolution for stereoscopic position change than has been reported in previous studies of apparent depth motion.

    View details for Web of Science ID A1984SQ13800001

    View details for PubMedID 6740960



    Event-related brain potentials (ERPs) in response to tachistoscopically presented photographs of 2 human faces were recorded for 4--7-month-old infants. For each infant 1 face was chosen to be presented frequently (p = .88, a low-information event) and the other infrequently (p = .12, a high-information event). Both types of events elicited in our infants a long-latency negative ERP wave (ca. 700 msec), termed Nc, and a long-latency positive wave (ca. 1,360 msec), termed Pc. We found that the discrepant, infrequently presented face elicited Nc waves which were higher in amplitude and longer in latency than those elicited by the frequent face. These differences suggest that our infants were able to remember the frequently presented face from trial to trial and to discriminate it from the discrepant face. The discrepant event elicited Pc waves which were insignificantly higher in amplitude than those elicited by frequent events. In adults and children, discrepant events have been found by numerous researchers to elicit positive P3 waves (latency ca. 300--800 msec). In our study, however, such waves could not be discerned. So, of all of the ERP waves which have been related to cognitive processes, the wave which is maturationally the earliest to appear is the Nc wave, which has been related to the perception of attention-getting events or events of interest to the subject. Our findings suggest that ERP responses could provide a sensitive means for investigating infant cognitive development since they do not depend upon an integrated motor-response system.

    View details for Web of Science ID A1981MH02200004

    View details for PubMedID 7285651