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.

Investigating the human binocular visual system using multi-modal magnetic resonance imaging.

Having two forward-facing eyes with slightly different viewpoints enables animals, including humans, to discriminate fine differences in depth (disparities), which can facilitate interaction with the world. The binocular visual system starts in the primary visual cortex because that is where information from the eyes is integrated for the first time. Magnetic resonance imaging (MRI) is an ideal tool to non-invasively investigate this system since it can provide a range of detailed measures about structure, function, neurochemistry and connectivity of the human brain. Since binocular disparity is used for both action and object recognition, the binocular visual system is a valuable model system in neuroscience for understanding how basic sensory cues are transformed into behaviourally relevant signals. In this review, we consider how MRI has contributed to the understanding of binocular vision and depth perception in the human brain. Firstly, MRI provides the ability to image the entire brain simultaneously to compare the contribution of specific visual areas to depth perception. A large body of work using functional MRI has led to an understanding of the extensive networks of brain areas involved in depth perception, but also the fine-scale macro-organisation for binocular processing within individual visual areas. Secondly, MRI can uncover mechanistic information underlying binocular combination with the use of MR spectroscopy. This method can quantify neurotransmitters including GABA and glutamate within restricted regions of the brain, and evaluate the role of these inhibitory and excitatory neurochemicals in binocular vision. Thirdly, it is possible to measure the nature and microstructure of pathways underlying depth perception using diffusion MRI. Understanding these pathways provides insight into the importance of the connections between areas implicated in depth perception. Finally, MRI can help to understand changes in the visual system resulting from amblyopia, a neural condition where binocular vision does not develop correctly in childhood.

Heritable functional architecture in human visual cortex.

How much of the functional organization of our visual system is inherited? Here we tested the heritability of retinotopic maps in human visual cortex using functional magnetic resonance imaging. We demonstrate that retinotopic organization shows a closer correspondence in monozygotic (MZ) compared to dizygotic (DZ) twin pairs, suggesting a partial genetic determination. Using population receptive field (pRF) analysis to examine the preferred spatial location and selectivity of these neuronal populations, we estimate a heritability around 10-20% for polar angle preferences and spatial selectivity, as quantified by pRF size, in extrastriate areas V2 and V3. Our findings are consistent with heritability in both the macroscopic arrangement of visual regions and stimulus tuning properties of visual cortex. This could constitute a neural substrate for variations in a range of perceptual effects, which themselves have been found to be at least partially genetically determined. These findings also add convergent evidence for the hypothesis that functional map topology is linked with cortical morphology.

Emerging evidence for cell-autonomous axon guidance.

Current models of axon guidance within the central nervous system (CNS) involve the presentation of environmental cues to navigating growth cones. The surrounding and target tissues present a variety of ligands that either restrict or promote growth, thus providing pathfinding instructions to developing axons. Recent findings show that RGMb, a GPI anchored extracellular protein present on retinal ganglion cells, down-regulates Wnt3a signaling by lowering LRP5 levels at the membrane surface. When RGMb is phosphorylated by the extracellular tyrosine kinase VLK, phosphorylated RGMb (p-RGMb) is internalized and carries LRP5 towards intracellular compartments. In the eye, a dorsal-high ventral-low gradient of VLK generates a dorsal-low ventral-high gradient of LRP5 that modulates Wnt3a signaling. These molecules, which are all expressed by individual RGCs, generate Wnt-signal gradients along the dorso-ventral axis of the retina, resulting in differential axon growth which in turn regulates proper retino-tectal/collicular map formation. This pathway represents a regulatory mechanism whereby extracellular phosphorylation generates what may be the first example of a unique self-guiding mechanism that affects neuronal-target connections independent of paracrine signals from the surrounding target tissue.

Survival of retinal ganglion cells after damage to the occipital lobe in humans is activity dependent.

Damage to the optic radiations or primary visual cortex leads to blindness in all or part of the contralesional visual field. Such damage disconnects the retina from its downstream targets and, over time, leads to trans-synaptic retrograde degeneration of retinal ganglion cells. To date, visual ability is the only predictor of retinal ganglion cell degeneration that has been investigated after geniculostriate damage. Given prior findings that some patients have preserved visual cortex activity for stimuli presented in their blind field, we tested whether that activity explains variability in retinal ganglion cell degeneration over and above visual ability. We prospectively studied 15 patients (four females, mean age = 63.7 years) with homonymous visual field defects secondary to stroke, 10 of whom were tested within the first two months after stroke. Each patient completed automated Humphrey visual field testing, retinotopic mapping with functional magnetic resonance imaging, and spectral-domain optical coherence tomography of the macula. There was a positive relation between ganglion cell complex (GCC) thickness in the blind field and early visual cortex activity for stimuli presented in the blind field. Furthermore, residual visual cortex activity for stimuli presented in the blind field soon after the stroke predicted the degree of retinal GCC thinning six months later. These findings indicate that retinal ganglion cell survival after ischaemic damage to the geniculostriate pathway is activity dependent.

Adaptive smoothing based on Gaussian processes regression increases the sensitivity and specificity of fMRI data.

Temporal and spatial filtering of fMRI data is often used to improve statistical power. However, conventional methods, such as smoothing with fixed-width Gaussian filters, remove fine-scale structure in the data, necessitating a tradeoff between sensitivity and specificity. Specifically, smoothing may increase sensitivity (reduce noise and increase statistical power) but at the cost loss of specificity in that fine-scale structure in neural activity patterns is lost. Here, we propose an alternative smoothing method based on Gaussian processes (GP) regression for single subjects fMRI experiments. This method adapts the level of smoothing on a voxel by voxel basis according to the characteristics of the local neural activity patterns. GP-based fMRI analysis has been heretofore impractical owing to computational demands. Here, we demonstrate a new implementation of GP that makes it possible to handle the massive data dimensionality of the typical fMRI experiment. We demonstrate how GP can be used as a drop-in replacement to conventional preprocessing steps for temporal and spatial smoothing in a standard fMRI pipeline. We present simulated and experimental results that show the increased sensitivity and specificity compared to conventional smoothing strategies. Hum Brain Mapp 38:1438-1459, 2017. © 2016 Wiley Periodicals, Inc.

Downstream mediators of Ten-m3 signalling in the developing visual pathway.

BACKGROUND: The formation of visuotopically-aligned projections in the brain is required for the generation of functional binocular circuits. The mechanisms which underlie this process are unknown. Ten-m3 is expressed in a broad high-ventral to low-dorsal gradient across the retina and in topographically-corresponding gradients in primary visual centres. Deletion of Ten-m3 causes profound disruption of binocular visual alignment and function. Surprisingly, one of the most apparent neuroanatomical changes-dramatic mismapping of ipsilateral, but not contralateral, retinal axons along the representation of the nasotemporal retinal axis-does not correlate well with Ten-m3's expression pattern, raising questions regarding mechanism. The aim of this study was to further our understanding of the molecular interactions which enable the formation of functional binocular visual circuits. METHODS: Anterograde tracing, gene expression studies and protein pull-down experiments were performed. Statistical significance was tested using a Kolmogorov-Smirnov test, pairwise-fixed random reallocation tests and univariate ANOVAs. RESULTS: We show that the ipsilateral retinal axons in Ten-m3 knockout mice are mismapped as a consequence of early axonal guidance defects. The aberrant invasion of the ventral-most region of the dorsal lateral geniculate nucleus by ipsilateral retinal axons in Ten-m3 knockouts suggested changes in the expression of other axonal guidance molecules, particularly members of the EphA-ephrinA family. We identified a consistent down-regulation of EphA7, but none of the other EphA-ephrinA genes tested, as well as an up-regulation of ipsilateral-determinants Zic2 and EphB1 in visual structures. We also found that Zic2 binds specifically to the intracellular domain of Ten-m3 in vitro. CONCLUSION: Our findings suggest that Zic2, EphB1 and EphA7 molecules may work as effectors of Ten-m3 signalling, acting together to enable the wiring of functional binocular visual circuits.

Intersession reliability of population receptive field estimates.

Population receptive field (pRF) analysis is a popular method to infer spatial selectivity of voxels in visual cortex. However, it remains largely untested how stable pRF estimates are over time. Here we measured the intersession reliability of pRF parameter estimates for the central visual field and near periphery, using a combined wedge and ring stimulus containing natural images. Sixteen healthy human participants completed two scanning sessions separated by 10-114 days. Individual participants showed very similar visual field maps for V1-V4 on both sessions. Intersession reliability for eccentricity and polar angle estimates was close to ceiling for most visual field maps (r>.8 for V1-3). PRF size and cortical magnification (CMF) estimates showed strong but lower overall intersession reliability (r≈.4-.6). Group level results for pRF size and CMF were highly similar between sessions. Additional control experiments confirmed that reliability does not depend on the carrier stimulus used and that reliability for pRF size and CMF is high for sessions acquired on the same day (r>.6). Our results demonstrate that pRF mapping is highly reliable across sessions.

Areas activated during naturalistic reading comprehension overlap topological visual, auditory, and somatotomotor maps.

Cortical mapping techniques using fMRI have been instrumental in identifying the boundaries of topological (neighbor-preserving) maps in early sensory areas. The presence of topological maps beyond early sensory areas raises the possibility that they might play a significant role in other cognitive systems, and that topological mapping might help to delineate areas involved in higher cognitive processes. In this study, we combine surface-based visual, auditory, and somatomotor mapping methods with a naturalistic reading comprehension task in the same group of subjects to provide a qualitative and quantitative assessment of the cortical overlap between sensory-motor maps in all major sensory modalities, and reading processing regions. Our results suggest that cortical activation during naturalistic reading comprehension overlaps more extensively with topological sensory-motor maps than has been heretofore appreciated. Reading activation in regions adjacent to occipital lobe and inferior parietal lobe almost completely overlaps visual maps, whereas a significant portion of frontal activation for reading in dorsolateral and ventral prefrontal cortex overlaps both visual and auditory maps. Even classical language regions in superior temporal cortex are partially overlapped by topological visual and auditory maps. By contrast, the main overlap with somatomotor maps is restricted to a small region on the anterior bank of the central sulcus near the border between the face and hand representations of M-I. Hum Brain Mapp 37:2784-2810, 2016. © 2016 The Authors Human Brain Mapping Published by Wiley Periodicals, Inc.

The human cortical areas V6 and V6A.

In macaque, it has long been known since the late nineties that the medial parieto-occipital sulcus (POS) contains two regions, V6 and V6A, important for visual motion and action. While V6 is a retinotopically organized extrastriate area, V6A is a broadly retinotopically organized visuomotor area constituted by a ventral and dorsal subdivision (V6Av and V6Ad), both containing arm movement-related cells active during spatially directed reaching movements. In humans, these areas have been mapped only in recent years thanks to neuroimaging methods. In a series of brain mapping studies, by using a combination of functional magnetic resonance imaging methods such as wide-field retinotopy and task-evoked activity, we mapped human areas V6 (Pitzalis et al., 2006) and V6Av (Pitzalis et al., 2013 d) retinotopically and defined human V6Ad functionally as a pointing-selective region situated anteriorly in the close proximity of V6Av (Tosoni et al., 2014). Like in macaque, human V6 is a motion area (e.g., Pitzalis et al., 2010, 2012, 2013 a, b , c ), while V6Av and V6Ad respond to pointing movements (Tosoni et al., 2014). The retinotopic organization (when present), anatomical position, neighbor relations, and functional properties of these three areas closely resemble those reported for macaque V6 (Galletti et al., 1996, 1999 a), V6Av, and V6Ad (Galletti et al., 1999 b; Gamberini et al., 2011). We suggest that information on objects in depth which are translating in space, because of the self-motion, is processed in V6 and conveyed to V6A for evaluating object distance in a dynamic condition such as that created by self-motion, so to orchestrate the eye and arm movements necessary to reach or avoid static and moving objects in the environment.

Impact of chiasma opticum malformations on the organization of the human ventral visual cortex.

Congenital malformations of the optic chiasm, such as enhanced and reduced crossing of the optic nerve fibers, are evident in albinism and achiasma, respectively. In early visual cortex the resulting additional visual input from the ipsilateral visual hemifield is superimposed onto the normal retinotopic representation of the contralateral visual field, which is likely due to conservative geniculo-striate projections. Counterintuitively, this organization in early visual cortex does not have profound consequences on visual function. Here we ask, whether higher stages of visual processing provide a correction to the abnormal representation allowing for largely normal perception. To this end we assessed the organization patterns of early and ventral visual cortex in five albinotic, one achiasmic, and five control participants. In albinism and achiasma the mirror-symmetrical superposition of the ipsilateral and contalateral visual fields was evident not only in early visual cortex, but also in the higher areas of the ventral processing stream. Specifically, in the visual areas VO1/2 and PHC1/2 no differences in the extent, the degree of superposition, and the magnitude of the responses were evident in comparison to the early visual areas. Consequently, the highly atypical organization of the primary visual cortex was propagated downstream to highly specialized processing stages in an undiminished and unchanged manner. This indicates largely unaltered cortico-cortical connections in both types of misrouting, i.e., enhanced and reduced crossing of the optic nerves. It is concluded that main aspects of visual function are preserved despite sizable representation abnormalities in the ventral visual processing stream.

Using functional magnetic resonance imaging to explore the flashed face distortion effect.

The flashed face distortion (FFD) effect was coined by Tangen, Murphy, and Thompson (2011) in their second-place winner of the 2012 Best Illusion of the Year Contest. The FFD arises when people view various eye-aligned faces that are sequentially flashed in the visual periphery, and gradually the faces appear to be deformed and grotesque. In this functional magnetic resonance imaging (fMRI) study, participants were presented with four conditions: (a) one face pair changing only its illumination; (b) two and (c) three alternating face pairs; and (d) nonrepeated face pairs. Participants rated the magnitude of each illusion immediately after each block. Results showed that the receptive region of the early visual cortex (V1-V4), and category-selective areas such as the fusiform face area (FFA) and occipital face area (OFA), responded proportionally to the participants' rated FFD strength. A random-effects voxelwise analysis further revealed positively correlated areas (including the medial and superolateral frontal areas) and negatively correlated areas (including the precuneus, postcentral gyrus, right insula, and bilateral middle frontal gyri) with respect to participants' ratings. Time series correlations among these nine ROIs (four positive and five negative) indicated that most participants showed a clustering of the two separate ROI types. Exploratory factor analysis (EFA) also demonstrated the segregation of the positive and negative ROIs; additionally, two subsystems were identified within the negative ROIs. These results suggest that the FFD is mediated by at least two networks: one that is likely responsible for perception and another that is likely responsible for subjective feelings and engagement.

qPRF: A system to accelerate population receptive field decoding.

Patterns of BOLD response can be decoded using the population receptive field (PRF) model to reveal how visual input is represented on the cortex (Dumoulin and Wandell, 2008). The time cost of evaluating the PRF model is high, often requiring days to decode BOLD signals for a small cohort of subjects. We introduce the qPRF, an efficient method for decoding that reduced the computation time by a factor of 1436 when compared to another widely available PRF decoder (Kay, Winawer, Mezer and Wandell, 2013) on a benchmark of data from the Human Connectome Project (HCP; Van Essen, Smith, Barch, Behrens, Yacoub and Ugurbil, 2013). With a specially designed data structure and an efficient search algorithm, the qPRF optimizes the five PRF model parameters according to a least-squares criterion. To verify the accuracy of the qPRF solutions, we compared them to those provided by Benson, Jamison, Arcaro, Vu, Glasser, Coalson, Van Essen, Yacoub, Ugurbil, Winawer and Kay (2018). Both hemispheres of the 181 subjects in the HCP data set (a total of 10,753,572 vertices, each with a unique BOLD time series of 1800 frames) were decoded by qPRF in 15.2 hours on an ordinary CPU. The absolute difference in R 2 reported by Benson et al. and achieved by the qPRF was negligible, with a median of 0.39% ( R 2 units being between 0% and 100%). In general, the qPRF yielded a slightly better fitting solution, achieving a greater R 2 on 99.7% of vertices. The qPRF may facilitate the development and computation of more elaborate models based on the PRF framework, as well as the exploration of novel clinical applications.

Measuring spatial visual loss in rats by retinotopic mapping of the superior colliculus using a novel multi-electrode array technique.

BACKGROUND: The retinotopic map property of the superior colliculus (SC) is a reliable indicator of visual functional changes in rodents. Electrophysiological mapping of the SC using a single electrode has been employed for measuring visual function in rat and mouse disease models. Single electrode mapping is highly laborious requiring long-term exposure to the SC surface and prolonged anesthetic conditions that can adversely affect the mapping data. NEW METHOD: To avoid the above-mentioned issues, we fabricated a fifty-six (56) electrode multi-electrode array (MEA) for rapid and reliable visual functional mapping of the SC. Since SC is a dome-shaped structure, the array was made of electrodes with dissimilar tip lengths to enable simultaneous and uniform penetration of the SC. RESULTS: SC mapping using the new MEA was conducted in retinal degenerate (RD) Royal College of Surgeons (RCS) rats and rats with focal retinal damage induced by green diode laser. For SC mapping, the MEA was advanced into the SC surface and the visual activities were recorded during full-filed light stimulation of the eye. Based on the morphological examination, the MEA electrodes covered most of the exposed SC area and penetrated the SC surface at a relatively uniform depth. MEA mapping in RCS rats (n=9) demonstrated progressive development of a scotoma in the SC that corresponded to the degree of photoreceptor loss. MEA mapping in the laser damaged rats demonstrated the presence of a scotoma in the SC area that corresponded to the location of retinal laser injury. COMPARISON WITH EXISTING METHODS AND CONCLUSIONS: The use of MEA for SC mapping is advantageous over single electrode recording by enabling faster recordings and reducing anesthesia time. This study establishes the feasibility of the MEA technique for rapid and efficient SC mapping, particularly advantageous for evaluating therapeutic effects in retinal degenerate rat disease models.

A new method for estimating population receptive field topography in visual cortex.

We introduce a new method for measuring visual population receptive fields (pRF) with functional magnetic resonance imaging (fMRI). The pRF structure is modeled as a set of weights that can be estimated by solving a linear model that predicts the Blood Oxygen Level-Dependent (BOLD) signal using the stimulus protocol and the canonical hemodynamic response function. This method does not make a priori assumptions about the specific pRF shape and is therefore a useful tool for uncovering the underlying pRF structure at different spatial locations in an unbiased way. We show that our method is more accurate than a previously described method (Dumoulin and Wandell, 2008) which directly fits a 2-dimensional isotropic Gaussian pRF model to predict the fMRI time-series. We demonstrate that direct-fit models do not fully capture the actual pRF shape, and can be prone to pRF center mislocalization when the pRF is located near the border of the stimulus space. A quantitative comparison demonstrates that our method outperforms the direct-fit methods in the pRF center modeling by achieving higher explained variance of the BOLD signal. This was true for direct-fit isotropic Gaussian, anisotropic Gaussian, and difference of isotropic Gaussians model. Importantly, our model is also capable of exploring a variety of pRF properties such as surround suppression, receptive field center elongation, orientation, location and size. Additionally, the proposed method is particularly attractive for monitoring pRF properties in the visual areas of subjects with lesions of the visual pathways, where it is difficult to anticipate what shape the reorganized pRF might take. Finally, the method proposed here is more efficient in computation time than direct-fit methods, which need to search for a set of parameters in an extremely large searching space. Instead, this method uses the pRF topography to constrain the space that needs to be searched for the subsequent modeling.

Developmental dissociation of visual dorsal stream parvo and magnocellular representations and the functional impact of negative retinotopic BOLD responses.

Localized neurodevelopmental defects provide an opportunity to study structure-function correlations in the human nervous system. This unique multimodal case report of epileptogenic dysplasia in the visual cortex allowed exploring visual function across distinct pathways in retinotopic regions and the dorsal stream, in relation to fMRI retinotopic mapping and spike triggered BOLD responses. Pre-surgical EEG/video monitoring, MRI/DTI, EEG/fMRI, PET and SPECT were performed to characterize structure/function correlations in this patient with a very early lesion onset. In addition, we included psychophysical methods (assessing parvo/konio and magnocellular pathways) and retinotopic mapping. We could identify dorsal stream impairment (with extended contrast sensitivity deficits within the input magno system contrasting with more confined parvocellular deficits) with disrupted active visual field input representations in regions neighboring the lesion. Simultaneous EEG/fMRI identified perilesional and retinotopic bilaterally symmetric BOLD deactivation triggered by interictal spikes, which matched the contralateral spread of magnocellular dysfunction revealed in the psychophysical tests. Topographic changes in retinotopic organization further suggested long term functional effects of abnormal electrical discharges during brain development. We conclude that fMRI based visual field cortical mapping shows evidence for retinotopic dissociation between magno and parvocellular function well beyond striate cortex, identifiable in high level dorsal visual representations around visual area V3A which is consistent with the effects of epileptic spike triggered negative BOLD.

Minimizing biases in estimating the reorganization of human visual areas with BOLD retinotopic mapping.

There is substantial interest in using functional magnetic resonance imaging (fMRI) retinotopic mapping techniques to examine reorganization of the occipital cortex after vision loss in humans and nonhuman primates. However, previous reports suggest that standard phase encoding and the more recent population Receptive Field (pRF) techniques give biased estimates of retinotopic maps near the boundaries of retinal or cortical scotomas. Here we examine the sources of this bias and show how it can be minimized with a simple modification of the pRF method. In normally sighted subjects, we measured fMRI responses to a stimulus simulating a foveal scotoma; we found that unbiased retinotopic map estimates can be obtained in early visual areas, as long as the pRF fitting algorithm takes the scotoma into account and a randomized "multifocal" stimulus sequence is used.

Evidence of target enhancement and distractor suppression in early visual areas.

Although the mechanisms of target enhancement and distractor suppression have been investigated along the visual processing hierarchy, there remains some unknown as to the role of perceptual load on the competition between different task-related information as attention deployment is manipulated. We present an fMRI spatial cueing paradigm, in which 32 participants had to attend to either a left or a right hemifield location and to indicate the orientation of the target Gabor that was presented simultaneously to a noise patch distractor. Critically, the target could appear at either the cued, valid location or at the uncued, invalid location; in the latter, the noise patch distractor appeared at the cued location. Perceptual load was manipulated by the presence or absence of high-contrast Gabor patches close to the fixation cross, which acted as lateral masks. Behavioural results indicated that participants performed more efficiently in validly cued trials compared to invalidly cued trials and under low compared to high load. Enhancement effects for targets and suppression effects for noise patches were greater in early visual areas at high load, that is in the presence of lateral masks. These results are in line with the hypothesis that attention results in both target enhancement and distractor suppression, and that these effects are most marked under high perceptual load. Theoretical implications of these results for different models of attention are discussed.

A tightly controlled fMRI dataset for receptive field mapping in human visual cortex.

Four right-handed, healthy subjects participated in a visual stimulation experiment. Subjects were viewing a dartboard-shaped flickering checkerboard stimulus, divided into 4 rings and 12 segments, defining 48 sectors in the visual field. Local contrast in each sector was continuously varying across four levels and updated every 3 s. To maintain fixation, subjects had to respond to a stimulus at the center of the visual field. During the entire experiment, in which subjects performed 8 runs, each consisting of 100 trials, brain activity was measured with functional magnetic resonance imaging (MRI). Using a 3-T Siemens Trio MRI scanner, 220 echo-planar images were acquired in each run, with a repetition time of 1.5 s and voxel size of 3 x 3 x 3 mm. The dataset is publicly available from OpenNeuro and additionally includes region of interest maps for visual areas V1 to V4, left and right, obtained from another retinotopic mapping experiment. As such, the dataset allows for accurate mapping of receptive fields and their properties across several stages of human visual cortex.

Training the spatially-coded SSVEP BCI on the fly.

BACKGROUND: The 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 METHOD: In 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. RESULTS: Starting 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 METHODS: Compared to a similar system with separate training and application phases, the time to achieve the same output is reduced by more than 50 %. CONCLUSIONS: Evaluating 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.