On identifying magnocellular and parvocellular responses on the basis of contrast-response functions

It has been proposed that magnocellular and parvocellular sensitivity in schizophrenic individuals can be assessed using steady-state visually evoked potentials (VEPs) to either low-contrast stimuli or stimuli whose contrast is modulated around a high contrast "pedestal" (Green MF, Butler PD, Chen Y, et al. Schizophr Bull. 2009;35:163-181). This suggestion faces 2 difficulties: (1) To use low-contrast stimuli to activate the magnocellular system is inconsistent with lesion studies that have shown that under many conditions, the parvocellular system responds to the lowest contrasts and (2) To rely on contrast-response relationships to identify magnocellular and parvocellular responses is difficult because other neurons exist in the visual system that have contrast-response relationships similar to those of magnocellular and parvocellular cells.     

Does primate motion perception depend on the magnocellular pathway?

This study examined the importance of the primate magnocellular retinocortical pathway in the perception of moving stimuli. A portion of the magnocellular pathway was permanently and selectively interrupted by ibotenic acid injections in the LGN of macaque monkeys. We then tested contrast sensitivity for detecting moving stimuli, as well as two indices of motion perception, contrast sensitivity for opposite direction discrimination and speed difference thresholds, in the affected portion of the visual field. Magnocellular lesions greatly reduced detection contrast sensitivity at high temporal and low spatial frequencies and had a similar effect on contrast sensitivity for opposite direction discrimination under these same stimulus conditions. Consequently, opposite direction discriminations could be made at contrast threshold, suggesting that magnocellular lesions reduced the visibility of stimuli used to test direction perception, but did not act directly on direction perception. Magnocellular lesions also elevated speed difference thresholds under some stimulus conditions. However, this deficit was reduced or eliminated by raising the contrast of the test stimulus. Together, these findings suggest that magnocellular lesions reduce the visibility of stimuli used to test motion perception but that they do not appear to alter motion perception otherwise.     

Magno and parvo stimuli affect illusory directional hearing in normal and dyslexic readers

In an experimental paradigm adapted from Hari (1995), forty observers listened via headphones to 8 binaural clicks: 4 left-ear leading followed by 4 right-ear leading with either 38 or 140 ms interstimulus intervals (ISIs). Concurrently, they viewed either foveal or peripheral visual stimuli designed to activate either the parvocellular or magnocellular pathway. They then reported the perceived location of each click-pair. Our results replicated Hari's finding that observers mistake the perceived location of short ISI click-pairs more often than long. That is, when ISIs were short, the sounds seemed to play across the inside of the head in a phenomenon called illusory directional hearing. However, when click-pairs were accompanied by peripheral visual stimuli that activated the magnocellular pathway, observers were more accurate than when there were no visual stimuli. Conversely, parvocellular-activating foveal visual stimuli produced more illusory hearing than when there were no visual stimuli. These findings suggest that activating the slow sustained parvocellular system may result in a longer processing window. Thirty dyslexic observers who repeated the experimental paradigm had an even longer processing window than control observers indicating that dyslexics may have a magnocellular system deficit.     

Visual sensory processing deficits in schizophrenia: is there anything to the magnocellular account?

Visual processing studies have repeatedly shown impairment in patients with schizophrenia compared to healthy controls. Electroencephalography (EEG) and, specifically, visual evoked potential (VEP) studies have identified an early marker of this impairment in the form of a decrement in the P1 component of the VEP in patients and their clinically unaffected first-degree relatives. Much behavioral and neuroimaging research has implicated specific dysfunction of either the subcortical magnocellular pathway or the cortical visual dorsal stream in this impairment. In this study, EEG responses were obtained to the contrast modulation of checkerboard stimuli using the VESPA (Visual Evoked Spread Spectrum Analysis) method. This was done for a high contrast condition and, in order to bias the stimuli towards the magnocellular pathway, a low contrast condition. Standard VEPs were also obtained using high contrast pattern reversing checkerboards. Responses were measured using high-density electrical scalp recordings in 29 individuals meeting DSM-IV criteria for schizophrenia and in 18 control subjects. Replicating previous research, a large (Cohen's d=1.11) reduction in the P1 component of the VEP was seen in patients when compared with controls with no corresponding difference in the VESPA response to high contrast stimuli. In addition, the low-contrast VESPA displayed no difference between patients and controls. Furthermore, no differences were seen between patients and controls for the C1 components of either the VEP or the high-contrast VESPA. Based on the differing acquisition methods between VEP and VESPA, we discuss these results in terms of contrast gain control and the possibility of dysfunction at the cortical level with initial afferent activity into V1 along the magnocellular pathway being intact when processing is biased towards that pathway using low contrast stimuli.     

Dysfunction of early-stage visual processing in schizophrenia.

OBJECTIVE: Schizophrenia is associated with deficits in higher-order processing of visual information. This study evaluated the integrity of early visual processing in order to evaluate the overall pattern of visual dysfunction in schizophrenia. METHOD: Steady-state visual-evoked potential responses were recorded over the occipital cortex in patients with schizophrenia and in age- and sex-matched comparison volunteers. Visual-evoked potentials were obtained for stimuli composed of isolated squares that were modulated sinusoidally in luminance contrast, number of squares, or chromatic contrast in order to emphasize magnocellular or parvocellular visual pathway activity. RESULTS: Responses of patients to magnocellular-biased stimuli were significantly lower than those of comparison volunteers. These lower response levels were observed in conditions using both low luminance contrast and large squares that biased processing toward the magnocellular pathway. In contrast, responses to stimuli that biased processing toward the parvocellular pathway were not significantly different between schizophrenia patients and comparison volunteers. A significant interaction of group and stimulus type was observed in the condition using low luminance contrast. CONCLUSIONS: These findings suggest a dysfunction of lower-level visual pathways, which was more prominent for magnocellular than parvocellular biased stimuli. The magnocellular pathway helps in orienting toward salient stimuli. A magnocellular pathway deficit could contribute to higher-level visual cognitive deficits in schizophrenia.     

Impaired parvocellular pathway in dyslexic children

Recent studies report that some children with dyslexia have impaired visual processing, specifically in the fast-processing magnocellular pathway. The objective was to study the effect of varying luminance and temporal and spatial frequency on the latency and amplitude of the visual evoked potentials (VEPs) in normal and dyslexic Egyptian children who speak Arabic (a right-left reading and writing system). VEPs were recorded in 52 dyslexic and 41 normal children in the fourth grade using a black and white checkerboard pattern with different checkerboard sizes and different rates of stimuli at high- and low-contrast media. The peak of the major positive wave component (P100) of each waveform and the trough of the previous major negative wave component were identified, and the peak-to-trough amplitude was measured. The latency and amplitude of VEPs in response to different experimental conditions showed significant shortening of P100 latency under high-contrast media and under low spatial frequency in children with dyslexia compared with normal readers. Furthermore, dyslexia children showed prolonged P100 latency in response to high spatial frequency stimulation compared with the low spatial frequency (P=0.003) and significantly higher N1-P1 amplitude under high-contrast media compared with low-contrast media (P=0.02), whilst no such changes were observed in normal readers. These results are suggestive of deficiency within the parvocellular pathway rather than the magnocellular pathway. As reading apparently places demands primarily on the ability to discriminate fine details, which is to say, on the parvocellular system, we suggested that deficiency in this system, at least in Arabic speaking children, could be a predisposing factor in dyslexia.     

TMS to V1 spares discrimination of emotive relative to neutral body postures

This study used TMS to examine the role played by striate cortex (V1) in processing the emotional content of visual stimuli. Participants learned to discriminate two sets of body posture images. For half of each set, the posture's emotional significance (threat versus pleasant) provided a redundant cue for the discrimination; the other half were emotionally neutral. An image was briefly presented at a lateral location in the visual field where a TMS pulse produced a phosphene, or at a control location in the opposite hemifield. A TMS pulse 70–140 ms after stimulus presentation at the phosphene location impaired discrimination of neutral stimuli with little effect on discrimination of emotional stimuli; the two classes of stimuli were equally discriminable when presented at the control location. The results are consistent with the proposal that emotionally salient patterns, such as social threat, can be discriminated independently of the geniculo-striate pathway.     

Stimulus Contrast Affects Spatial Integration in the Lateral Geniculate Nucleus of Macaque Monkeys

Gain-control mechanisms adjust neuronal responses to accommodate the wide range of stimulus conditions in the natural environment. Contrast gain control and extraclassical surround suppression are two manifestations of gain control that govern the responses of neurons in the early visual system. Understanding how these two forms of gain control interact has important implications for the detection and discrimination of stimuli across a range of contrast conditions. Here, we report that stimulus contrast affects spatial integration in the lateral geniculate nucleus of alert macaque monkeys (male and female), whereby neurons exhibit a reduction in the strength of extraclassical surround suppression and an expansion in the preferred stimulus size with low-contrast stimuli compared with high-contrast stimuli. Effects were greater for magnocellular neurons than for parvocellular neurons, indicating stream-specific interactions between stimulus contrast and stimulus size. Within the magnocellular pathway, contrast-dependent effects were comparable for ON-center and OFF-center neurons, despite ON neurons having larger receptive fields, less pronounced surround suppression, and more pronounced contrast gain control than OFF neurons. Together, these findings suggest that the parallel streams delivering visual information from retina to primary visual cortex, serve not only to broaden the range of signals delivered to cortex, but also to provide a substrate for differential interactions between stimulus contrast and stimulus size that may serve to improve stimulus detection and stimulus discrimination under pathway-specific lower and higher contrast conditions, respectively.SIGNIFICANCE STATEMENT Stimulus contrast is a salient feature of visual scenes. Here we examine the influence of stimulus contrast on spatial integration in the lateral geniculate nucleus (LGN). Our results demonstrate that increases in contrast generally increase extraclassical suppression and decrease the size of optimal stimuli, indicating a reduction in the extent of visual space from which LGN neurons integrate signals. Differences between magnocellular and parvocellular neurons are noteworthy and further demonstrate that the feedforward parallel pathways to cortex increase the range of information conveyed for downstream cortical processing, a range broadened by diversity in the ON and OFF pathways. These results have important implications for more complex visual processing that underly the detection and discrimination of stimuli under varying natural conditions.     

Interference with vision by TMS over the occipital pole: a fourth period

We investigated the effect of single-pulse transcranial magnetic stimulation (TMS) over the occipital pole on a forced-choice visual letter-identification task. Magnetic stimuli were applied on the midline but with the initial current directed pseudorandomly toward either left or right hemisphere; visual stimuli were presented randomly in either left or right hemifield; magnetic-visual stimulus onset asynchrony varied randomly between 12 values: -500 ms and from -50 ms to +50 ms in 10 ms steps. The data revealed the existence of a hitherto unknown fourth task-interfering TMS effect that was maximal at -10 ms and specific for magnetic stimulus polarity and visual stimulus location. This -10 ms effect cannot be explained by reflex blinking (as the -50 ms effect can) and direct disruption of letter-induced activity (as the +20 ms and +100 ms effects can), but it could be explained by direct disruption of pre-letter activity or indirect disruption of letter-induced activity.     

Unconscious response priming by shape depends on geniculostriate visual projection

It has been suggested that unconscious visual processing of some stimulus features might occur without the contribution of early visual cortex (V1/V2). In the present study, the causal role of V1/V2 in unconscious processing of simple shapes in intact human brain was studied by applying transcranial magnetic stimulation (TMS) on early visual cortex or lateral occipital cortex (LO) while observers performed a metacontrast-masked response priming task with arrow figures as visual stimuli. Magnetic stimulation of V1/V2 impaired masked priming 30-90 ms after the onset of the prime. Stimulation of LO reduced the magnitude of masked priming at 90-120 ms, but this effect occurred only in the early parts of the priming experiment. A control task measuring the visibility of masked primes indicated that the orientation of masked primes could not be consciously discriminated and that TMS did not influence the conscious visibility of the primes indirectly by reducing the effectiveness of the mask in the critical time windows. We conclude that feedforward sweep of processing from V1/V2 (30-90 ms) to LO (90 ms and above) is necessary for unconscious priming of shape, whereas conscious perception requires also the contribution of recurrent (feedback) processing.     

The "special effect" of case mixing on word identification: neuropsychological and transcranial magnetic stimulation studies dissociating case mixing from contrast reduction

We present neuropsychological and transcranial magnetic stimulation (TMS) evidence with normal readers, that the effects of case mixing and contrast reduction on word identification are qualitatively different. Lesions and TMS applied to the right parietal lobe selectively disrupted the identification of mixed relative to single-case stimuli. Bilateral lesions and TMS applied to the occipital cortex selectively disrupted the identification of low-contrast words. These data suggest that different visual distortions (case mixing, contrast reduction) exert different effects on reading, modulated by contrasting brain regions. Case mixing is a "special" distortion and involves the recruitment of processes that are functionally distinct, and dependent on different regions in the brain, from those required to deal with contrast reduction.     

Enhanced visual perception with occipital transcranial magnetic stimulation

Transcranial magnetic stimulation (TMS) over the occipital pole can produce an illusory percept of a light flash (or ‘phosphene’), suggesting an excitatory effect. Whereas previous reported effects produced by single‐pulse occipital pole TMS are typically disruptive, here we report the first demonstration of a location‐specific facilitatory effect on visual perception in humans. Observers performed a spatial cueing orientation discrimination task. An orientation target was presented in one of two peripheral placeholders. A single pulse below the phosphene threshold applied to the occipital pole 150 or 200 ms before stimulus onset was found to facilitate target discrimination in the contralateral compared with the ipsilateral visual field. At the 150‐ms time window contralateral TMS also amplified cueing effects, increasing both facilitation effects for valid cues and interference effects for invalid cues. These results are the first to show location‐specific enhanced visual perception with single‐pulse occipital pole stimulation prior to stimulus presentation, suggesting that occipital stimulation can enhance the excitability of visual cortex to subsequent perception.     

An electrophysiological study on the interaction between emotional content and spatial frequency of visual stimuli

Previous studies suggest that the magnocellular pathway, a visual processing system that rapidly provides low spatial frequency information to fast-responding structures such as the amygdala, is more involved in the processing of emotional facial expressions than the parvocellular pathway (which conveys all spatial frequencies). The present experiment explored the spatio-temporal characteristics of the spatial frequency modulation of affect-related neural processing, as well as its generalizability to non-facial stimuli. To that aim, the event-related potentials (ERPs) elicited by low-pass filtered (i.e., high spatial frequencies are eliminated) and intact non-facial emotional images were recorded from 31 participants using a 60-electrode array. The earliest significant effect of spatial frequency was observed at 135 ms from stimulus onset: N135 component of the ERPs. In line with previous studies, the origin of N135 was localized at secondary visual areas for low-pass filtered stimuli and at primary areas for intact stimuli. Importantly, this component showed an interaction between spatial frequency and emotional content: within low-pass filtered pictures, negative stimuli elicited the highest N135 amplitudes. By contrast, within intact stimuli, neutral pictures were those eliciting the highest amplitudes. These results suggest that high spatial frequencies are not essential for the initial affect-related processing of visual stimuli, which would mainly rely on low spatial frequency visual information. According to present data, high spatial frequencies would come into play later on.     

Retinotopy of emotion: Perception of negatively valenced stimuli presented at different spatial locations as revealed by event‐related potentials

Scarce previous data on how the location where an emotional stimulus appears in the visual scene modulates its perception suggest that, for functional reasons, a perceptual advantage may exist, vertically, for stimuli presented at the lower visual field (LoVF) and, horizontally, for stimuli presented at the left visual field (LeVF). However, this issue has been explored through a limited number of spatial locations, usually in a single spatial dimension (e.g., horizontal) and invariant eccentricities. Event‐related potentials (ERPs) were recorded from 39 participants perceiving brief neutral (wheels) and emotional stimuli (spiders) presented at 17 different locations, one foveal and 16 at different peripheral coordinates. As a secondary scope, we explored the role of the magnocellular (M) and the parvocellular (P) visual pathways by presenting an isoluminant/heterochromatic (P‐biased) and a heteroluminant/isochromatic version (M‐biased) of each stimulus. Emo > Neu effects were observed in PN1 (120 ms) for stimuli located at fovea, and in PN2 (215 ms) for stimuli located both at fovea and diverse peripheral regions. A factorial approach to these effects further revealed that: (a) emotional stimuli presented in the periphery are efficiently perceived, without evident decrease from para‐ to perifovea; (b) peripheral Emo > Neu effects are reflected 95 ms later than foveal Emo > Neu effects in ERPs; (c) LoVF is more involved than UVF in these effects; (d) our data fail to support the LeVF advantage previously reported, and (e) Emo > Neu effects were significant for both M and P stimuli.     

Timing of right parietal and frontal cortex activity in visuo-spatial perception: a TMS study in normal individuals

In a recent study we showed that repetitive transcranial magnetic stimulation (rTMS) with train duration of 400 ms over right frontal and right posterior parietal cortices gives rise to transitory contralateral visuo-spatial neglect in normal subjects. In the present experiment we investigated whether using single-pulse TMS it is possible to obtain information about the timing of cortical activity related to spatial cognition. Nine healthy subjects performed in baseline condition and during TMS a tachistoscopic task, requiring a forced-choice estimation of the length of the two segments of prebisected horizontal lines. Single-pulse TMS was triggered at various time intervals (150 ms, 225 ms, 300 ms) after visual stimulus onset with a focal coil over P6 and F4 (according to 10/20 EEG system). Relative transitory rightward bias was observed only when parietal TMS was delivered 150 ms after visual stimulus presentation. Frontal stimulation induced no effect on visuo-spatial perception with the time intervals explored.     

Facilitation of rhythmic events in progressive myoclonus epilepsy: a transcranial magnetic stimulation study.

Twelve subjects with progressive myoclonus epilepsy (PME) were studied with transcranial magnetic stimulation (TMS), using single and paired magnetic stimuli at different interstimulus intervals (ISIs), and polygraphic recording. Motor threshold (T) and silent period (SP) were normal. Paired TMS showed a loss of inhibition at 100-150 ms ISI and a marked facilitation at 50 ms ISI of conditioned motor evoked potential (MEP). Polygraphic analysis showed 20 Hz oscillatory activity over the sensorimotor area coupled to contralateral myoclonic jerks. These findings suggest a condition of increased supraspinal excitability and support the evidence of a cortical rhythm in the range of 20 Hz. No direct evidence exists that these findings are mediated by the same intracortical pathway. Furthermore, the normal SP and T suggest that the abnormal excitability is not a constant feature but is evident during rhythmic events.     

The neural signature of phosphene perception

Artificial percepts (phosphenes) can be induced by applying transcranial magnetic stimulation (TMS) over human visual cortex. Although phosphenes have been used to study visual awareness, the neural mechanisms generating them have not yet been delineated. We directly tested the two leading hypotheses of how phosphenes arise. These hypotheses correspond to the two competing views of the neural genesis of awareness: the early, feedforward view and the late, recurrent feedback model. We combined online TMS and EEG recordings to investigate whether the electrophysiological correlates of conscious phosphene perception are detectable early after TMS onset as an immediate local effect of TMS, or only at longer latencies, after interactions of TMS‐induced activity with other visual areas. Stimulation was applied at the intensity threshold at which participants saw a phosphene on half of the trials, and brain activity was recorded simultaneously with electroencephalography. Phosphene perception was associated with a differential pattern of TMS‐evoked brain potentials that started 160–200 ms after stimulation and encompassed a wide array of posterior areas. This pattern was differentiated from the TMS‐evoked potential after stimulation of a control site. These findings suggest that conscious phosphene perception is not a local phenomenon, but arises only after extensive recurrent processing. Hum Brain Mapp, 2010. © 2010 Wiley‐Liss, Inc.     

Clinical neurophysiology of visual and auditory processing in dyslexia: A review

Neurophysiological studies on children and adults with dyslexia provide a deeper understanding of how visual and auditory processing in dyslexia might relate to reading deficits. The goal of this review is to provide an overview of research findings in the last two decades on motion related and contrast sensitivity visual evoked potentials and on auditory event related potentials to basic tone and speech sound processing in dyslexia. These results are particularly relevant for three important theories about causality in dyslexia: the magnocellular deficit hypothesis, the temporal processing deficit hypothesis and the phonological deficit hypothesis. Support for magnocellular deficits in dyslexia are primarily provided from evidence for altered visual evoked potentials to rapidly moving stimuli presented at low contrasts. Consistently ERP findings revealed altered neurophysiological processes in individuals with dyslexia to speech stimuli, but evidence for deficits processing certain general acoustic information relevant for speech perception, such as frequency changes and temporal patterns, are also apparent.     

Phosphenes and transient scotomas induced by magnetic stimulation of the occipital lobe: their topographic relationship

Transcranial magnetic stimulation (TMS) of the visual cortex is known to induce phosphenes and is able to suppress visual perception. To address the topographic relationship of phosphenes and transient scotomas, the visual field of 10 normal subjects was investigated using a perimetric approach. The central visual field (diameter: 20 degrees) was tested at 32 sites. Perceptual thresholds were determined by presenting 1 ms test spots flashed with varying intensity in random order. TMS was applied with a focal figure-of-eight coil placed over the inion. All subjects perceived phosphenes, mostly restricted to one of the lower quadrants within the visual field. In 13 out of 15 investigations, a magnetic stimulus triggered 100 ms after the visual target resulted in a relative scotoma with threshold changes of 8 dB or more. In 9 of 13 investigations, scotomas coincided spatially with sketches of phosphenes made by subjects in a separate test. Scotomas covered only a small percentage of the total visual field, which may explain the failure of previous studies to find perceptual suppression with the focal coil. The present result demonstrates that phosphenes evoked during TMS can serve as a guide for optimal visual stimulus alignment in neuropsychological experiments.     

Dyslexia: verbal impairments in the absence of magnocellular impairments

Sensitivity to dynamic visual and auditory stimuli was assessed in dyslexic children (Grade 7) who at school entrance had suffered from the well-established double-deficit of impaired phonological sensitivity and deficient rapid naming performance. A visual magnocellular deficit was assessed by the coherent motion detection task of the Oxford group. An auditory magnocellular deficit was assessed by the illusory sound movement perception task of Hari and Kiesil?. On both tasks our dyslexic subjects' performance was similar or even better than the performance of normally reading controls. Differences in the inclusion of ADHD cases in dyslexic samples is discussed as a potential explanation of differences in results.