Retinotopic Mapping Research Articles

Activation of multiple Eph receptors on neuronal membranes correlates with the onset of optic neuropathy

BackgroundOptic neuropathy is a major cause of irreversible blindness, yet the molecular determinants that contribute to neuronal demise have not been fully elucidated. Several studies have identified ‘ephrin signaling’ as one of the most dysregulated pathways in the early pathophysiology of optic neuropathy with varied etiologies. Developmentally, gradients in ephrin signaling coordinate retinotopic mapping via repulsive modulation of cytoskeletal dynamics in neuronal membranes. Little is known about the role ephrin signaling plays in the post-natal visual system and its correlation with the onset of optic neuropathy.MethodsPostnatal mouse retinas were collected for mass spectrometry analysis for erythropoietin-producing human hepatocellular (Eph) receptors. Optic nerve crush (ONC) model was employed to induce optic neuropathy, and proteomic changes during the acute phase of neuropathic onset were analyzed. Confocal and super-resolution microscopy determined the cellular localization of activated Eph receptors after ONC injury. Eph receptor inhibitors assessed the neuroprotective effect of ephrin signaling modulation.ResultsMass spectrometry revealed expression of seven Eph receptors (EphA2, A4, A5, B1, B2, B3, and B6) in postnatal mouse retinal tissue. Immunoblotting analysis indicated a significant increase in phosphorylation of these Eph receptors 48 h after ONC. Confocal microscopy demonstrated the presence of both subclasses of Eph receptors within the retina. Stochastic optical reconstruction microscopy (STORM) super-resolution imaging combined with optimal transport colocalization analysis revealed a significant co-localization of activated Eph receptors with injured neuronal cells, compared to uninjured neuronal and/or injured glial cells, 48 h post-ONC. Eph receptor inhibitors displayed notable neuroprotective effects for retinal ganglion cells (RGCs) after six days of ONC injury.ConclusionsOur findings demonstrate the functional presence of diverse Eph receptors in the postnatal mammalian retina, capable of modulating multiple biological processes. Pan-Eph receptor activation contributes to the onset of neuropathy in optic neuropathies, with preferential activation of Eph receptors on neuronal processes in the inner retina following optic nerve injury. Notably, Eph receptor activation precedes neuronal loss. We observed a neuroprotective effect on RGCs upon inhibiting Eph receptors. Our study highlights the importance of investigating this repulsive pathway in early optic neuropathies and provides a comprehensive characterization of the receptors present in the developed retina of mice, relevant to both homeostasis and disease processes.

Variability of visual field maps in human early extrastriate cortex challenges the canonical model of organization of V2 and V3.

Visual field maps in human early extrastriate areas (V2 and V3) are traditionally thought to form mirror-image representations which surround the primary visual cortex (V1). According to this scheme, V2 and V3 form nearly symmetrical halves with respect to the calcarine sulcus, with the dorsal halves representing lower contralateral quadrants, and the ventral halves representing upper contralateral quadrants. This arrangement is considered to be consistent across individuals, and thus predictable with reasonable accuracy using templates. However, data that deviate from this expected pattern have been observed, but mainly treated as artifactual. Here, we systematically investigate individual variability in the visual field maps of human early visual cortex using the 7T Human Connectome Project (HCP) retinotopy dataset. Our results demonstrate substantial and principled inter-individual variability. Visual field representation in the dorsal portions of V2 and V3 was more variable than in their ventral counterparts, including substantial departures from the expected mirror-symmetrical patterns. In addition, left hemisphere retinotopic maps were more variable than those in the right hemisphere. Surprisingly, only one-third of individuals had maps that conformed to the expected pattern in the left hemisphere. Visual field sign analysis further revealed that in many individuals the area conventionally identified as dorsal V3 shows a discontinuity in the mirror-image representation of the retina, associated with a Y-shaped lower vertical representation. Our findings challenge the current view that inter-individual variability in early extrastriate cortex is negligible, and that the dorsal portions of V2 and V3 are roughly mirror images of their ventral counterparts.

Retinotopic connectivity maps of human visual cortex with unconstrained eye movements.

Human visual cortex contains topographic visual field maps whose organization can be revealed with retinotopic mapping. Unfortunately, constraints posed by standard mapping hinder its use in patients, atypical subject groups, and individuals at either end of the lifespan. This severely limits the conclusions we can draw about visual processing in such individuals. Here, we present a novel data-driven method to estimate connective fields, resulting in fine-grained maps of the functional connectivity between brain areas. We find that inhibitory connectivity fields accompany, and often surround facilitatory fields. The visual field extent of these inhibitory subfields falls off with cortical magnification. We further show that our method is robust to large eye movements and myopic defocus. Importantly, freed from the controlled stimulus conditions in standard mapping experiments, using entertaining stimuli and unconstrained eye movements our approach can generate retinotopic maps, including the periphery visual field hitherto only possible to map with special stimulus displays. Generally, our results show that the connective field method can gain knowledge about retinotopic architecture of visual cortex in patients and participants where this is at best difficult and confounded, if not impossible, with current methods.

A novel simulation paradigm utilising MRI-derived phosphene maps for cortical prosthetic vision

Objective. We developed a realistic simulation paradigm for cortical prosthetic vision and investigated whether we can improve visual performance using a novel clustering algorithm. Approach. Cortical visual prostheses have been developed to restore sight by stimulating the visual cortex. To investigate the visual experience, previous studies have used uniform phosphene maps, which may not accurately capture generated phosphene map distributions of implant recipients. The current simulation paradigm was based on the Human Connectome Project retinotopy dataset and the placement of implants on the cortices from magnetic resonance imaging scans. Five unique retinotopic maps were derived using this method. To improve performance on these retinotopic maps, we enabled head scanning and a density-based clustering algorithm was then used to relocate centroids of visual stimuli. The impact of these improvements on visual detection performance was tested. Using spatially evenly distributed maps as a control, we recruited ten subjects and evaluated their performance across five sessions on the Berkeley Rudimentary Visual Acuity test and the object recognition task. Main results. Performance on control maps is significantly better than on retinotopic maps in both tasks. Both head scanning and the clustering algorithm showed the potential of improving visual ability across multiple sessions in the object recognition task. Significance. The current paradigm is the first that simulates the experience of cortical prosthetic vision based on brain scans and implant placement, which captures the spatial distribution of phosphenes more realistically. Utilisation of evenly distributed maps may overestimate the performance that visual prosthetics can restore. This simulation paradigm could be used in clinical practice when making plans for where best to implant cortical visual prostheses.

Retinotopic connectivity maps are robust to large eye movements and optical blur

Retinotopic connectivity maps are robust to large eye movements and optical blur

Comparing retinotopic maps of children and adults reveals a late-stage change in how V1 samples the visual field

Comparing retinotopic maps of children and adults reveals a late-stage change in how V1 samples the visual field

Three-dimensional topography of eye-specific domains in the lateral geniculate nucleus of pigmented and albino rats.

We previously revealed the presence of ocular dominance columns (ODCs) in the primary visual cortex (V1) of pigmented rats. On the other hand, previous studies have shown thatthe ipsilateral-eye domains of the dorsal lateral geniculate nucleus (dLGN) are segregated into a handful of patches in pigmented rats. To investigate the three-dimensional (3D) topography of the eye-specific patches of the dLGN and its relationship with ODCs, we injected different tracers into the right and left eyes and examined strain difference, development, and plasticity of the patches. Furthermore, we applied the tissue clearing technique to reveal the 3D morphology of the LGN and were able to observe entire retinotopic map of the rat dLGN at a certain angle. Our results show that the ipsilateral domains of the dLGN appear mesh-like at any angle and are developed at around time of eye-opening. Their development was moderately affected by abnormal visual experience, but the patch formation was not disrupted. In albino Wistar rats, ipsilateral patches were observed in the dLGN, but they were much fewer, especially near the central visual field. These results provide insights into how ipsilateral patches of the dLGN arise, and how the geniculo-cortical arrangement is different between rodents and primates.

Signaling - transcription interactions in mouse retinal ganglion cells early axon pathfinding -a literature review.

Sending an axon out of the eye and into the target brain nuclei is the defining feature of retinal ganglion cells (RGCs). The literature on RGC axon pathfinding is vast, but it focuses mostly on decision making events such as midline crossing at the optic chiasm or retinotopic mapping at the target nuclei. In comparison, the exit of RGC axons out of the eye is much less explored. The first checkpoint on the RGC axons' path is the optic cup - optic stalk junction (OC-OS). OC-OS development and the exit of the RGC pioneer axons out of the eye are coordinated spatially and temporally. By the time the optic nerve head domain is specified, the optic fissure margins are in contact and the fusion process is ongoing, the first RGCs are born in its proximity and send pioneer axons in the optic stalk. RGC differentiation continues in centrifugal waves. Later born RGC axons fasciculate with the more mature axons. Growth cones at the end of the axons respond to guidance cues to adopt a centripetal direction, maintain nerve fiber layer restriction and to leave the optic cup. Although there is extensive information on OC-OS development, we still have important unanswered questions regarding its contribution to the exit of the RGC axons out of the eye. We are still to distinguish the morphogens of the OC-OS from the axon guidance molecules which are expressed in the same place at the same time. The early RGC transcription programs responsible for axon emergence and pathfinding are also unknown. This review summarizes the molecular mechanisms for early RGC axon guidance by contextualizing mouse knock-out studies on OC-OS development with the recent transcriptomic studies on developing RGCs in an attempt to contribute to the understanding of human optic nerve developmental anomalies. The published data summarized here suggests that the developing optic nerve head provides a physical channel (the closing optic fissure) as well as molecular guidance cues for the pioneer RGC axons to exit the eye.

Enriched environment exposure during development positively impacts the structure and function of the visual cortex in mice

Optimal conditions of development have been of interest for decades, since genetics alone cannot fully explain how an individual matures. In the present study, we used optical brain imaging to investigate whether a relatively simple enrichment can positively influence the development of the visual cortex of mice. The enrichment paradigm was composed of larger cages housing multiple mice that contained several toys, hiding places, nesting material and a spinning wheel that were moved or replaced at regular intervals. We compared C57BL/6N adult mice (> P60) that had been raised either in an enriched environment (EE; n = 16) or a standard (ST; n = 12) environment from 1 week before birth to adulthood, encompassing all cortical developmental stages. Here, we report significant beneficial changes on the structure and function of the visual cortex following environmental enrichment throughout the lifespan. More specifically, retinotopic mapping through intrinsic signal optical imaging revealed that the size of the primary visual cortex was greater in mice reared in an EE compared to controls. In addition, the visual field coverage of EE mice was wider. Finally, the organization of the cortical representation of the visual field (as determined by cortical magnification) versus its eccentricity also differed between the two groups. We did not observe any significant differences between females and males within each group. Taken together, these data demonstrate specific benefits of an EE throughout development on the visual cortex, which suggests adaptation to their environmental realities.

Comparing retinotopic maps of children and adults reveals a late-stage change in how V1 samples the visual field

Adult visual performance differs with angular location –it is better for stimuli along the horizontal than vertical, and lower than upper vertical meridian of the visual field. These perceptual asymmetries are paralleled by asymmetries in cortical surface area in primary visual cortex (V1). Children, unlike adults, have similar visual performance at the lower and upper vertical meridian. Do children have similar V1 surface area representing the upper and lower vertical meridian? Using MRI, we measure the surface area of retinotopic maps (V1-V3) in children and adults. Many features of the maps are similar between groups, including greater V1 surface area for the horizontal than vertical meridian. However, unlike adults, children have a similar amount of V1 surface area representing the lower and upper vertical meridian. These data reveal a late-stage change in V1 organization that may relate to the emergence of the visual performance asymmetry along the vertical meridian by adulthood.

Comparison of Stimulus Types for Retinotopic Cortical Mapping of Macular Disease.

Retinotopic maps acquired using functional magnetic resonance imaging (fMRI) provide a valuable adjunct in the assessment of macular function at the level of the visual cortex. The present study quantitatively assessed the performance of different visual stimulation approaches for mapping visual field coverage. Twelve patients with geographic atrophy (GA) secondary to age-related macular degeneration (AMD) were examined using high-resolution ultra-high field fMRI (Siemens Magnetom 7T) and microperimetry (MP; Nidek MP-3). The population receptive field (pRF)-based coverage maps obtained with two different stimulus techniques (moving bars, and rotating wedges and expanding rings) were compared with the results of MP. Correspondence between MP and pRF mapping was quantified by calculating the simple matching coefficient (SMC). Stimulus choice is shown to bias the spatial distribution of pRF centers and eccentricity values with pRF sizes obtained from wedge/ring or bar stimulation showing systematic differences. Wedge/ring stimulation results show a higher number of pRF centers in foveal areas and strongly reduced pRF sizes compared to bar stimulation runs. A statistical comparison shows significantly higher pRF center numbers in the foveal 2.5 degrees region of the visual field for wedge/ring compared to bar stimuli. However, these differences do not significantly influence SMC values when compared to MP (bar <2.5 degrees: 0.88 ± 0.13; bar >2.5 degrees: 0.88 ± 0.11; wedge/ring <2.5 degrees: 0.89 ± 0.12 wedge/ring; >2.5 degrees: 0.86 ± 0.10) for the peripheral visual field. Both visual stimulation designs examined can be applied successfully in patients with GA. Although the two designs show systematic differences in the distribution of pRF center locations, this variability has minimal impact on the SMC when compared to the MP outcome.

White matter BOLD signals at 7 Tesla reveal visual field maps in optic radiation and vertical occipital fasciculus

There is growing evidence that blood-oxygen-level-dependent (BOLD) activity in the white matter (WM) can be detected by functional magnetic resonance imaging (fMRI). However, the functional relevance and significance of WM BOLD signals remain controversial. Here we investigated whether 7T BOLD fMRI can reveal fine-scale functional organizations of a WM bundle. Population receptive field (pRF) analyses of the 7T retinotopy dataset from the Human Connectome Project revealed clear contralateral retinotopic organizations of two visual WM bundles: the optic radiation (OR) and the vertical occipital fasciculus (VOF). The retinotopic maps of OR are highly consistent with post-mortem dissections and diffusion tractographies, while the VOF maps are compatible with the dorsal and ventral visual areas connected by the WM. Similar to the grey matter (GM) visual areas, both WM bundles show over-representations of the central visual field and increasing pRF size with eccentricity. Hemodynamic response functions of visual WM were slower and wider compared with those of GM areas. These findings clearly demonstrate that WM BOLD at 7 Tesla is closely coupled with neural activity related to axons, encoding highly specific information that can be used to characterize fine-scale functional organizations of a WM bundle.

Mapping anatomical connectivity between visual cortex and the pulvinar in human neonates

The primate pulvinar contains extensive, reciprocal connections with visual cortex. Through this connectivity the pulvinar is thought to regulate communication within and between cortical visual areas and facilitate the processing of low- and high-level visual information (Saalmann & Kastner 2011). Here, we explored the development of these connections in humans. We analyzed diffusion imaging data collected in neonates (37-42w gestation) as part of the developing human connectome project (Bastiani et al. 2019). To identify cortical regions for tracking, we aligned an adult retinotopic atlas (Wang et al. 2015) to each neonate’s native brain. Probabilistic tractography (FSL) was performed between individual retinotopic maps and the pulvinar in accordance with prior analyses in adults (Arcaro et al. 2015). Briefly, an initial tractography analysis was performed to identify the most probable thalamic radiations interconnecting cortical areas and the pulvinar. A second analysis restricted to tracking through these radiations identified voxels in the pulvinar with the largest connectivity values with each cortical map. Data were normalized and aligned to a 40-week neonate template for group-level analyses. For occipital cortical maps, the largest connectivity values were within the ventral lateral pulvinar. Medial to this tracking was a region of the ventral pulvinar that was most strongly connected with motion-sensitive maps TO1&2. For posterior parietal maps the largest connectivity values were within the dorsal pulvinar. Tracking from parietal maps extended to the superior colliculus. Overall, pulvino-cortical tracking in neonates followed the known white matter pathways in adults and the spatial organization of connections within the pulvinar was in good correspondence with the organization previously found in adults (Arcaro et al. 2015). These results indicate that the pulvino-cortical pathways that support visual processing in adults are established in utero. We propose that these connections play an important role in shaping postnatal development across visual cortex.

Corrigendum to ‘Quantitative Characterization of the Human Retinotopic Map Based on Quasiconformal Mapping’ [Medical Image Analysis Volume 75 (2022) 102230

Corrigendum to ‘Quantitative Characterization of the Human Retinotopic Map Based on Quasiconformal Mapping’ [Medical Image Analysis Volume 75 (2022) 102230

Contrast polarity-specific mapping improves efficiency of neuronal computation for collision detection.

Neurons receive information through their synaptic inputs, but the functional significance of how those inputs are mapped on to a cell's dendrites remains unclear. We studied this question in a grasshopper visual neuron that tracks approaching objects and triggers escape behavior before an impending collision. In response to black approaching objects, the neuron receives OFF excitatory inputs that form a retinotopic map of the visual field onto compartmentalized, distal dendrites. Subsequent processing of these OFF inputs by active membrane conductances allows the neuron to discriminate the spatial coherence of such stimuli. In contrast, we show that ON excitatory synaptic inputs activated by white approaching objects map in a random manner onto a more proximal dendritic field of the same neuron. The lack of retinotopic synaptic arrangement results in the neuron's inability to discriminate the coherence of white approaching stimuli. Yet, the neuron retains the ability to discriminate stimulus coherence for checkered stimuli of mixed ON/OFF polarity. The coarser mapping and processing of ON stimuli thus has a minimal impact, while reducing the total energetic cost of the circuit. Further, we show that these differences in ON/OFF neuronal processing are behaviorally relevant, being tightly correlated with the animal's escape behavior to light and dark stimuli of variable coherence. Our results show that the synaptic mapping of excitatory inputs affects the fine stimulus discrimination ability of single neurons and document the resulting functional impact on behavior.

Seeing and extrapolating motion trajectories share common informative activation patterns in primary visual cortex.

The natural environment is dynamic and moving objects become constantly occluded, engaging the brain in a challenging completion process to estimate where and when the object might reappear. Although motion extrapolation is critical in daily life-imagine crossing the street while an approaching car is occluded by a larger standing vehicle-its neural underpinnings are still not well understood. While the engagement of low-level visual cortex during dynamic occlusion has been postulated, most of the previous group-level fMRI-studies failed to find evidence for an involvement of low-level visual areas during occlusion. In this fMRI-study, we therefore used individually defined retinotopic maps and multivariate pattern analysis to characterize the neural basis of visible and occluded changes in motion direction in humans. To this end, participants learned velocity-direction change pairings (slow motion-upwards; fast motion-downwards or vice versa) during a training phase without occlusion and judged the change in stimulus direction, based on its velocity, during a following test phase with occlusion. We find that occluded motion direction can be predicted from the activity patterns during visible motion within low-level visual areas, supporting the notion of a mental representation of motion trajectory in these regions during occlusion.

Temporal Dynamics of Neural Responses in Human Visual Cortex.

Neural responses to visual stimuli exhibit complex temporal dynamics, including subadditive temporal summation, response reduction with repeated or sustained stimuli (adaptation), and slower dynamics at low contrast. These phenomena are often studied independently. Here, we demonstrate these phenomena within the same experiment and model the underlying neural computations with a single computational model. We extracted time-varying responses from electrocorticographic recordings from patients presented with stimuli that varied in duration, interstimulus interval (ISI) and contrast. Aggregating data across patients from both sexes yielded 98 electrodes with robust visual responses, covering both earlier (V1-V3) and higher-order (V3a/b, LO, TO, IPS) retinotopic maps. In all regions, the temporal dynamics of neural responses exhibit several nonlinear features. Peak response amplitude saturates with high contrast and longer stimulus durations, the response to a second stimulus is suppressed for short ISIs and recovers for longer ISIs, and response latency decreases with increasing contrast. These features are accurately captured by a computational model composed of a small set of canonical neuronal operations, that is, linear filtering, rectification, exponentiation, and a delayed divisive normalization. We find that an increased normalization term captures both contrast- and adaptation-related response reductions, suggesting potentially shared underlying mechanisms. We additionally demonstrate both changes and invariance in temporal response dynamics between earlier and higher-order visual areas. Together, our results reveal the presence of a wide range of temporal and contrast-dependent neuronal dynamics in the human visual cortex and demonstrate that a simple model captures these dynamics at millisecond resolution.SIGNIFICANCE STATEMENT Sensory inputs and neural responses change continuously over time. It is especially challenging to understand a system that has both dynamic inputs and outputs. Here, we use a computational modeling approach that specifies computations to convert a time-varying input stimulus to a neural response time course, and we use this to predict neural activity measured in the human visual cortex. We show that this computational model predicts a wide variety of complex neural response shapes, which we induced experimentally by manipulating the duration, repetition, and contrast of visual stimuli. By comparing data and model predictions, we uncover systematic properties of temporal dynamics of neural signals, allowing us to better understand how the brain processes dynamic sensory information.

Structural Covariance and Heritability of the Optic Tract and Primary Visual Cortex in Living Human Brains.

Individual differences among human brains exist at many scales, spanning gene expression, white matter tissue properties, and the size and shape of cortical areas. One notable example is an approximately 3-fold range in the size of human primary visual cortex (V1), a much larger range than is found in overall brain size. A previous study (Andrews et al., 1997) reported a correlation between optic tract (OT) cross-section area and V1 size in postmortem human brains, suggesting that there may be a common developmental mechanism for multiple components of the visual pathways. We evaluated the relationship between properties of the OT and V1 in a much larger sample of living human brains by analyzing the Human Connectome Project (HCP) 7 Tesla Retinotopy Dataset (including 107 females and 71 males). This dataset includes retinotopic maps measured with functional MRI (fMRI) and fiber tract data measured with diffusion MRI (dMRI). We found a negative correlation between OT fractional anisotropy (FA) and V1 surface area (r = -0.19). This correlation, although small, was consistent across multiple dMRI datasets differing in acquisition parameters. Further, we found that both V1 surface area and OT properties were correlated among twins, with higher correlations for monozygotic (MZ) than dizygotic (DZ) twins, indicating a high degree of heritability for both properties. Together, these results demonstrate covariation across individuals in properties of the retina (OT) and cortex (V1) and show that each is influenced by genetic factors.SIGNIFICANCE STATEMENT The size of human primary visual cortex (V1) has large interindividual differences. These differences do not scale with overall brain size. A previous postmortem study reported a correlation between the size of the human optic tract (OT) and V1. In this study, we evaluated the relationship between the OT and V1 in living humans by analyzing a neuroimaging dataset that included functional MRI (fMRI) and diffusion MRI (dMRI) data. We found a small, but robust correlation between OT tissue properties and V1 size, supporting the existence of structural covariance between the OT and V1 in living humans. The results suggest that characteristics of retinal ganglion cells (RGCs), reflected in OT measurements, are correlated with individual differences in human V1.

Highly accurate retinotopic maps of the physiological blind spot in human visual cortex.

The physiological blind spot is a naturally occurring scotoma corresponding with the optic disc in the retina of each eye. Even during monocular viewing, observers are usually oblivious to the scotoma, in part because the visual system extrapolates information from the surrounding area. Unfortunately, studying this visual field region with neuroimaging has proven difficult, as it occupies only a small part of retinotopic cortex. Here, we used functional magnetic resonance imaging and a novel data-driven method for mapping the retinotopic organization in and around the blind spot representation in V1. Our approach allowed for highly accurate reconstructions of the extent of an observer's blind spot, and out-performed conventional model-based analyses. This method opens exciting opportunities to study the plasticity of receptive fields after visual field loss, and our data add to evidence suggesting that the neural circuitry responsible for impressions of perceptual completion across the physiological blind spot most likely involves regions of extrastriate cortex-beyond V1.

Training the spatially-coded SSVEP BCI on the fly

BackgroundThe spatially-coded SSVEP BCI employs the retinotopic map in the human visual pathway to infer the gaze direction of the operator relative to a flicker stimulus inducing steady-state visual evoked potentials (SSVEPs) in the brain. It has been shown that with this method, up to 16 channels can be encoded using only a single flicker stimulus. Another advantage over conventional frequency-coded SSVEP BCIs, in which channels are encoded by different combinations of frequencies and phases, is that the operator does not have to gaze directly at flickering lights. This can reduce visual fatigue and improve user comfort. Whereas the frequency of the SSVEP response is well predictable, which has enabled the development of frequency-coded SSVEP BCIs which do not require training data, the spatial distribution of the SSVEP response over the scalp differs much more between different people. This requires collecting a substantial amount of training data before the spatially-coded BCI could be put into operation. New methodIn this study we address this issue by combining the spatially-coded BCI with a feedback channel which the operator uses to flag classification errors, and which allows the system to accumulate valid training data while the BCI is used to solve a spatial navigation task. ResultsStarting from the minimal number of samples required by the classification method, the approach achieved an average accuracy of 69 ± 15 %, corresponding to an ITR of 31 ± 17 bits/min, in solving the task for the first time. This accuracy improved to 87 ± 9 % (ITR: 54 ± 14 bits/min) after completing the task 2 more times. Further we show that participants with a stable SSVEP topography over repeated stimulation enable the BCI to achieve higher accuracies. Comparison with existing methodsCompared to a similar system with separate training and application phases, the time to achieve the same output is reduced by more than 50 %. ConclusionsEvaluating the approach in 17 participants suggests that the performance of the spatially-coded BCI with a minimal set of training samples is sufficient to be operational, and that performance keeps improving in the course of its application.