Reduced Visual Magnocellular Event-Related Potentials in Developmental Dyslexia

(1) Background—the magnocellular hypothesis proposes that impaired development of the visual timing systems in the brain that are mediated by magnocellular (M-) neurons is a major cause of dyslexia. Their function can now be assessed quite easily by analysing averaged visually evoked event-related potentials (VERPs) in the electroencephalogram (EEG). Such analysis might provide a useful, objective biomarker for diagnosing developmental dyslexia. (2) Methods—in adult dyslexics and normally reading controls, we recorded steady state VERPs, and their frequency content was computed using the fast Fourier transform. The visual stimulus was a black and white checker board whose checks reversed contrast every 100 ms. M- cells respond to this stimulus mainly at 10 Hz, whereas parvocells (P-) do so at 5 Hz. Left and right visual hemifields were stimulated separately in some subjects to see if there were latency differences between the M- inputs to the right vs. left hemispheres, and these were compared with the subjects’ handedness. (3) Results—Controls demonstrated a larger 10 Hz than 5 Hz fundamental peak in the spectra, whereas the dyslexics showed the reverse pattern. The ratio of subjects’ 10/5 Hz amplitudes predicted their reading ability. The latency of the 10 Hz peak was shorter during left than during right hemifield stimulation, and shorter in controls than in dyslexics. The latter correlated weakly with their handedness. (4) Conclusion—Steady state visual ERPs may conveniently be used to identify developmental dyslexia. However, due to the limited numbers of subjects in each sub-study, these results need confirmation.

Unified Visual Working Memory without the Anterior Corpus Callosum

One of the most fundamental, and most studied, human cognitive functions is working memory. Yet, it is currently unknown how working memory is unified. In other words, why does a healthy human brain have one integrated capacity of working memory, rather than one capacity per visual hemifield, for instance. Thus, healthy subjects can memorize roughly as many items, regardless of whether all items are presented in one hemifield, rather than throughout two visual hemifields. In this current research, we investigated two patients in whom either most, or the entire, corpus callosum has been cut to alleviate otherwise untreatable epilepsy. Crucially, in both patients the anterior parts connecting the frontal and most of the parietal cortices, are entirely removed. This is essential, since it is often posited that working memory resides in these areas of the cortex. We found that despite the lack of direct connections between the frontal cortices in these patients, working memory capacity is similar regardless of whether stimuli are all presented in one visual hemifield or across two visual hemifields. This indicates that in the absence of the anterior parts of the corpus callosum working memory remains unified. Moreover, it is important to note that memory performance was not similar across visual fields. In fact, capacity was higher when items appeared in the left visual hemifield than when they appeared in the right visual hemifield. Visual information in the left hemifield is processed by the right hemisphere and vice versa. Therefore, this indicates that visual working memory is not symmetric, with the right hemisphere having a superior visual working memory. Nonetheless, a (subcortical) bottleneck apparently causes visual working memory to be integrated, such that capacity does not increase when items are presented in two, rather than one, visual hemifield.

Interhemispheric transfer of working memories

SummaryVisual working memory (WM) storage is largely independent between the left and right visual hemifields/cerebral hemispheres, yet somehow WM feels seamless. We studied how WM is integrated across hemifields by recording neural activity bilaterally from lateral prefrontal cortex. An instructed saccade during the WM delay shifted the remembered location from one hemifield to the other. Before the shift, spike rates and oscillatory power showed clear signatures of memory laterality. After the shift, the lateralization inverted, consistent with transfer of the memory trace from one hemisphere to the other. Transferred traces initially used different neural ensembles from feedforward-induced ones but they converged at the end of the delay. Around the time of transfer, synchrony between the two prefrontal hemispheres peaked in theta and low-gamma frequencies, with a directionality consistent with memory trace transfer. This illustrates how dynamics between the two cortical hemispheres can stitch together WM traces across visual hemifields.

Gamma coherence mediates interhemispheric integration during multiple object tracking

ABSTRACTOur ability to track the paths of multiple visual objects moving between the hemifields requires effective integration of information between the two cerebral hemispheres. Coherent neural oscillations in the gamma band (35–70 Hz) are hypothesised to drive this information transfer. Here we manipulated the need for interhemispheric integration using a novel multiple object tracking (MOT) task in which stimuli either moved between the two visual hemifields—requiring interhemispheric integration—or moved within separate visual hemifields. We used electroencephalography (EEG) to measure interhemispheric coherence during the task. Human observers (21 female; 20 male) were poorer at tracking objects between-versus within-hemifields, reflecting a cost of interhemispheric integration. Critically, gamma coherence was greater in trials requiring interhemispheric integration, particularly between sensors over parieto-occipital areas. In approximately half of the participants, the observed cost of integration was associated with a failure of the cerebral hemispheres to become coherent in the gamma band. Moreover, individual differences in this integration cost correlated with endogenous gamma coherence at these same sensors, though with generally opposing relationships for the real and imaginary part of coherence. The real part (capturing synchronisation with a near-zero phase-lag) benefited between-hemifield tracking; imaginary coherence was detrimental. Finally, instantaneous phase-coherence over the tracking period uniquely predicted between-hemifield tracking performance, suggesting that effective integration benefits from sustained interhemispheric synchronisation. Our results show that gamma coherence mediates interhemispheric integration during MOT, and add to a growing body of work demonstrating that coherence drives communication across cortically distributed neural networks.

Hemifield-specific control of spatial attention and working memory: Evidence from hemifield crossover costs

Attentional tracking and working memory tasks are often performed better when targets are divided evenly between the left and right visual hemifields, rather than contained within a single hemifield (Alvarez & Cavanagh, 2005; Delvenne, 2005). However, this bilateral field advantage does not provide conclusive evidence of hemifield-specific control of attention and working memory, as it can be explained solely from hemifield-limited spatial interference at early stages of visual processing. If control of attention and working memory is specific to each hemifield, maintaining target information should become more difficult as targets move between the two hemifields. Observers in the present study maintained targets that moved either within or between the left and right hemifields, using either attention (Experiment 1) or working memory (Experiment 2). Maintaining spatial information was more difficult when target items moved between the hemifields compared to when target items moved within their original hemifields, consistent with hemifield-specific control of spatial attention and working memory. However, this pattern was not found for maintaining identity information (e.g., color) in working memory (Experiment 3). Together, these results provide evidence that control of spatial attention and working memory is specific to each hemifield, and that hemifield-specific control is a unique signature of spatial processing.

Population receptive field and connectivity properties of the early visual cortex in human albinism

In albinism, the pathological decussation of the temporal retinal afferents at the optic chiasm leads to superimposed representations of opposing hemifields in the visual cortex. Here, we assessed the equivalence of the two representations and the cortico-cortical connectivity of the early visual areas. Applying fMRI-based population receptive field (pRF)-mapping (both hemifield and bilateral mapping) and connective field (CF)-modeling, we investigated the early visual cortex in 6 albinotic participants and 4 controls. In albinism, superimposed retinotopic representations of the contra- and ipsilateral visual hemifield were observed on the hemisphere contralateral to the stimulated eye. This was confirmed by the observation of bilateral pRFs during bilateral mapping. Hemifield mapping revealed similar pRF-sizes for both hemifield representations throughout V1 to V3. The typical increase of V1-sampling extent for V3 compared to V2 was not found for the albinotic participants. The similarity of the pRF-sizes for opposing visual hemifield representations highlights the equivalence of the two maps in the early visual cortex. The altered V1-sampling extent in V3 indicates the adaptation of cortico-cortical connections to the abnormal input of the visual cortex. These findings thus suggest that conservative developmental mechanisms are complemented by alterations of the extrastriate cortico-cortical connectivity.HighlightspRF mapping confirms cortical overlay of opposing visual hemifields in albinism.Equivalent information processing of both hemifields is indicated by similar pRF sizes.CF modeling indicates changes to the cortico-cortical connections at the level of V3.

Two spatially distinct posterior alpha sources fulfill different functional roles in attention

ABSTRACTDirecting attention helps to extract relevant information and suppress distracters. Alpha brain oscillations (8-12Hz) play a fundamental role in this process, with a power decrease facilitating processing of important information and power increase inhibiting brain regions processing irrelevant information. Evidence for this phenomenon arises from visual attention studies (Worden et al., 2000), however, the effect also exists in other modalities, including the somatosensory system (Haegens et al., 2011) and inter-sensory attention tasks (Foxe and Snyder, 2011). We investigated what happens when attention is divided between two modalities using both a multi- and unimodal attention paradigm while recording EEG over 128 scalp electrodes in two separate experiments. In Experiment 1 participants divided their attention between the visual and somatosensory modality to determine the temporal or spatial frequency of a target stimulus (vibrotactile stimulus or Gabor grating). In Experiment 2, participants divided attention between two visual hemifields to identify the orientation of a target Gabor grating. In both experiments, pre-stimulus alpha power in visual areas decreased linearly with increasing attention to visual stimuli. In contrast, alpha power in parietal areas showed lower pre-stimulus alpha power when attention was divided between modalities, compared to unimodal attention. These results suggest that there are two different alpha sources, where one reflects the ‘visual spotlight of attention’ and the other reflects attentional effort. To our knowledge, this is the first study to show that attention recruits two spatially distinct alpha sources in occipital and parietal brain regions, which act simultaneously but serve different functions in attention.SIGNIFICANCE STATEMENTAttention to one spatial location/sensory modality leads to power changes of alpha oscillations (~10Hz) with decreased power over regions processing relevant information and power increases to actively inhibit areas processing ‘to-be-ignored’ information. Here, we used detailed source modelling to investigate EEG data recorded during separate uni-modal (visual) and multi- (visual and somatosensory) attention tasks. Participants either focused their attention on one modality/spatial location or directed it to both. We show for the first time two distinct alpha sources are active simultaneously but play different roles. A sensory (visual) alpha source was linearly modulated by attention representing the ‘visual spotlight of attention’. In contrast, a parietal alpha source was modulated by attentional effort, showing lowest alpha power when attention was divided.

Online data collection for developmental research

The strategies infants and young children use to understand the world around them provide unique insight into the structure of human cognition. However, developmental research is subject to heavy pragmatic constraints on recruiting large numbers of participants, bringing families back for repeat sessions, and working with special populations or diverse samples. These constraints limit the types of questions that can be addressed in the lab as well as the quality of evidence that can be obtained. In this dissertation, I present a new platform, “Lookit,” that allows researchers to conduct developmental experiments online via asynchronous webcam-recorded sessions, with the aim of expanding the set of questions that we can effectively answer. I first present the results of a series of empirical studies conducted in the laboratory to assess difficulty faced by infants in integrating information across visual hemifields (Chapter 2), as an illustration of the creative workarounds in study design necessary to accommodate the difficulty of participant recruitment. The rest of this work concerns the development of the online platform, from designing the prototype (Chapter 3) and initial proof-of-concept studies (Chapter 4) to the demonstration of an interface for researchers to specify and manage their studies on a collaborative platform (Chapter 5). I show that we are able to reliably collect and code dependent measures including looking times, preferential looking, and verbal responses on Lookit; to work with more representative samples than in the lab; and to flexibly implement a wide variety of study designs of interest to developmental researchers.