Theoretical and Technical Issues Concerning the Measurement of Alpha Frequency and the Application of Signal Detection Theory: Comment on Buergers and Noppeney (2022).

Classical and recent evidence has suggested that alpha oscillations play a critical role in temporally discriminating or binding successively presented items. Challenging this view, Buergers and Noppeney [Buergers, S., & Noppeney, U. The role of alpha oscillations in temporal binding within and across the senses. Nature Human Behaviour, 6, 732-742, 2022] found that by combining EEG, psychophysics, and signal detection theory, neither prestimulus nor resting-state alpha frequency influences perceptual sensitivity and bias in the temporal binding task. We propose the following four points that should be considered when interpreting the role of alpha oscillations, and especially their frequency, on perceptual temporal binding: (1) Multiple alpha components can be contaminated in conventional EEG analysis; (2) the effect of alpha frequency on perception will interact with alpha power; (3) prestimulus and resting-state alpha frequency can be different from poststimulus alpha frequency, which is the frequency during temporal binding and should be more directly related to temporal binding; and (4) when applying signal detection theory under the assumption of equal variance, the assumption is often incomplete and can be problematic (e.g., the magnitude relationships between individuals in parametric sensitivity may change when converted into nonparametric sensitivity). Future directions, including solutions to each of the issues, are discussed.

Macromolecular tissue volume mapping of lateral geniculate nucleus subdivisions in living human brains.

The lateral geniculate nucleus (LGN) is a key thalamic nucleus in the visual system, which has an important function in relaying retinal visual input to the visual cortex. The human LGN is composed mainly of magnocellular (M) and parvocellular (P) subdivisions, each of which has different stimulus selectivity in neural response properties. Previous studies have discussed the potential relationship between LGN subdivisions and visual disorders based on psychophysical data on specific types of visual stimuli. However, these relationships remain speculative because non-invasive measurements of these subdivisions are difficult due to the small size of the LGN. Here we propose a method to identify these subdivisions by combining two structural MR measures: high-resolution proton-density weighted images and macromolecular tissue volume (MTV) maps. We defined the M and P subdivisions based on MTV fraction data and tested the validity of the definition by (1) comparing the data with that from human histological studies, (2) comparing the data with functional magnetic resonance imaging measurements on stimulus selectivity, and (3) analyzing the test-retest reliability. The findings demonstrated that the spatial organization of the M and P subdivisions was consistent across subjects and in line with LGN subdivisions observed in human histological data. Moreover, the difference in stimulus selectivity between the subdivisions identified using MTV was consistent with previous physiology literature. The definition of the subdivisions based on MTV was shown to be robust over measurements taken on different days. These results suggest that MTV mapping is a promising approach for evaluating the tissue properties of LGN subdivisions in living humans. This method potentially will enable neuroscientific and clinical hypotheses about the human LGN subdivisions to be tested.

Frequency- and phase-dependent effects of auditory entrainment on attentional blink.

Attentional blink (AB) is the impaired detection of a second target (T2) after a first target has been identified. In this paper, we investigated the functional roles of alpha and theta oscillations on AB by determining how much preceding rhythmic auditory stimulation affected the performance of AB. Healthy young adults participated in the experiment online. We found that when two targets were embedded in rapid serial visual presentation (RSVP) of distractors at 10 Hz (i.e., alpha frequency), the magnitude of AB increased with auditory stimuli. The increase was limited to the case when the frequency and phase of auditory stimuli matched the following RSVP stream. On the contrary, when only two targets were presented without distractors, auditory stimuli at theta, not alpha, increased the AB magnitude. These results indicate that neural oscillations at two different frequencies, namely, alpha and theta, are involved in attentional blink.

Microstructural properties of the vertical occipital fasciculus explain the variability in human stereoacuity.

Stereopsis is a fundamental visual function that has been studied extensively. However, it is not clear why depth discrimination (stereoacuity) varies more significantly among people than other modalities. Previous studies have reported the involvement of both dorsal and ventral visual areas in stereopsis, implying that not only neural computations in cortical areas but also the anatomical properties of white matter tracts connecting those areas can impact stereopsis. Here, we studied how human stereoacuity relates to white matter properties by combining psychophysics, diffusion MRI (dMRI), and quantitative MRI (qMRI). We performed a psychophysical experiment to measure stereoacuity and, in the same participants, we analyzed the microstructural properties of visual white matter tracts on the basis of two independent measurements, dMRI (fractional anisotropy, FA) and qMRI (macromolecular tissue volume; MTV). Microstructural properties along the right vertical occipital fasciculus (VOF), a major tract connecting dorsal and ventral visual areas, were highly correlated with measures of stereoacuity. This result was consistent for both FA and MTV, suggesting that the behavioral-structural relationship reflects differences in neural tissue density, rather than differences in the morphological configuration of fibers. fMRI confirmed that binocular disparity stimuli activated the dorsal and ventral visual regions near VOF endpoints. No other occipital tracts explained the variance in stereoacuity. In addition, the VOF properties were not associated with differences in performance on a different psychophysical task (contrast detection). These series of experiments suggest that stereoscopic depth discrimination performance is, at least in part, constrained by dorso-ventral communication through the VOF.

Decoded fMRI neurofeedback can induce bidirectional confidence changes within single participants.

Neurofeedback studies using real-time functional magnetic resonance imaging (rt-fMRI) have recently incorporated the multi-voxel pattern decoding approach, allowing for fMRI to serve as a tool to manipulate fine-grained neural activity embedded in voxel patterns. Because of its tremendous potential for clinical applications, certain questions regarding decoded neurofeedback (DecNef) must be addressed. Specifically, can the same participants learn to induce neural patterns in opposite directions in different sessions? If so, how does previous learning affect subsequent induction effectiveness? These questions are critical because neurofeedback effects can last for months, but the short- to mid-term dynamics of such effects are unknown. Here we employed a within-subjects design, where participants underwent two DecNef training sessions to induce behavioural changes of opposing directionality (up or down regulation of perceptual confidence in a visual discrimination task), with the order of training counterbalanced across participants. Behavioral results indicated that the manipulation was strongly influenced by the order and the directionality of neurofeedback training. We applied nonlinear mathematical modeling to parametrize four main consequences of DecNef: main effect of change in confidence, strength of down-regulation of confidence relative to up-regulation, maintenance of learning effects, and anterograde learning interference. Modeling results revealed that DecNef successfully induced bidirectional confidence changes in different sessions within single participants. Furthermore, the effect of up- compared to down-regulation was more prominent, and confidence changes (regardless of the direction) were largely preserved even after a week-long interval. Lastly, the effect of the second session was markedly diminished as compared to the effect of the first session, indicating strong anterograde learning interference. These results are interpreted in the framework of reinforcement learning and provide important implications for its application to basic neuroscience, to occupational and sports training, and to therapy.

Decoding time-resolved neural representations of orientation ensemble perception.

The visual system can compute summary statistics of several visual elements at a glance. Numerous studies have shown that an ensemble of different visual features can be perceived over 50-200 ms; however, the time point at which the visual system forms an accurate ensemble representation associated with an individual's perception remains unclear. This is mainly because most previous studies have not fully addressed time-resolved neural representations that occur during ensemble perception, particularly lacking quantification of the representational strength of ensembles and their correlation with behavior. Here, we conducted orientation ensemble discrimination tasks and electroencephalogram (EEG) recordings to decode orientation representations over time while human observers discriminated an average of multiple orientations. We modeled EEG signals as a linear sum of hypothetical orientation channel responses and inverted this model to quantify the representational strength of orientation ensemble. Our analysis using this inverted encoding model revealed stronger representations of the average orientation over 400-700 ms. We also correlated the orientation representation estimated from EEG signals with the perceived average orientation reported in the ensemble discrimination task with adjustment methods. We found that the estimated orientation at approximately 600-700 ms significantly correlated with the individual differences in perceived average orientation. These results suggest that although ensembles can be quickly and roughly computed, the visual system may gradually compute an orientation ensemble over several hundred milliseconds to achieve a more accurate ensemble representation.

The relationship between alpha power and heart rate variability commonly seen in various mental states.

The extensive exploration of the correlation between electroencephalogram (EEG) and heart rate variability (HRV) has yielded inconsistent outcomes, largely attributable to variations in the tasks employed in the studies. The direct relationship between EEG and HRV is further complicated by alpha power, which is susceptible to influences such as mental fatigue and sleepiness. This research endeavors to examine the brain-heart interplay typically observed during periods of music listening and rest. In an effort to mitigate the indirect effects of mental states on alpha power, subjective fatigue and sleepiness were measured during rest, while emotional valence and arousal were evaluated during music listening. Partial correlation analyses unveiled positive associations between occipital alpha2 power (10-12 Hz) and nHF, an indicator of parasympathetic activity, under both music and rest conditions. These findings underscore brain-heart interactions that persist even after the effects of other variables have been accounted for.

Neural correlates of induced motion perception in the human brain.

A physically stationary stimulus surrounded by a moving stimulus appears to move in the opposite direction. There are similarities between the characteristics of this phenomenon of induced motion and surround suppression of directionally selective neurons in the brain. Here, functional magnetic resonance imaging was used to investigate the link between the subjective perception of induced motion and cortical activity. The visual stimuli consisted of a central drifting sinusoid surrounded by a moving random-dot pattern. The change in cortical activity in response to changes in speed and direction of the central stimulus was measured. The human cortical area hMT+ showed the greatest activation when the central stimulus moved at a fast speed in the direction opposite to that of the surround. More importantly, the activity in this area was the lowest when the central stimulus moved in the same direction as the surround and at a speed such that the central stimulus appeared to be stationary. The results indicate that the activity in hMT+ is related to perceived speed modulated by induced motion rather than to physical speed or a kinetic boundary. Early visual areas (V1, V2, V3, and V3A) showed a similar pattern; however, the relationship to perceived speed was not as clear as that in hMT+. These results suggest that hMT+ may be a neural correlate of induced motion perception and play an important role in contrasting motion signals in relation to their surrounding context and adaptively modulating our motion perception depending on the spatial context.

Neural correlates of impulsive compulsive behaviors in Parkinson's disease: A Japanese retrospective study.

BACKGROUND: Impulsive compulsive behaviors (ICBs) often disturb patients with Parkinson's Disease (PD), of which impulse control disorder (ICD) and dopamine dysregulation syndrome (DDS) are two major subsets. The nucleus accumbens (NAcc) is involved in ICB; however, it remains unclear how the NAcc affects cortical function and defines the different behavioral characteristics of ICD and DDS. OBJECTIVES: To identify the cortico-striatal network primarily involved in ICB and the differences in these networks between patients with ICD and DDS using structural and resting-state functional magnetic resonance imaging. METHODS: Patients with PD were recruited using data from a previous cohort study and divided into those with ICB (ICB group) and without ICB (non-ICB group) using the Japanese version of the Questionnaire for Impulsive Compulsive Disorders in Parkinson's Disease (J-QUIP). From these two groups, we extracted 37 pairs matched for age, sex, disease duration, and levodopa equivalent daily dose of dopamine agonists. Patients with ICB were further classified as having ICD or DDS based on the J-QUIP subscore. General linear models were used to compare gray matter volume and functional connectivity (FC) of the NAcc, caudate, and putamen between the ICB and non-ICB groups and between patients with ICD and those with DDS. RESULTS: We found no significant differences in gray matter volumebetween the ICB and non-ICB groups or between patients with ICD and those with DDS. Compared with the non-ICB group, the FC of the right NAcc in the ICB group was lower in the bilateral ventromedial prefrontal cortex and higher in the left middle occipital gyrus. Furthermore, patients with DDS showed higher FC between the right putamen and left superior temporal gyrus and higher FC between the left caudate and bilateral middle occipital gyrus than patients with ICD. In contrast, patients with ICD exhibited higher FC between the left NAcc and the right posterior cingulate cortex than patients with DDS. CONCLUSIONS: The functionally altered network between the right NAcc and ventromedial prefrontal cortex was associated with ICB in PD. In addition, the surrounding cortico-striatal networks may differentiate the behavioral characteristics of patients with ICD and those with DDS.

Human neural responses involved in spatial pooling of locally ambiguous motion signals.

Early visual motion signals are local and one-dimensional (1-D). For specification of global two-dimensional (2-D) motion vectors, the visual system should appropriately integrate these signals across orientation and space. Previous neurophysiological studies have suggested that this integration process consists of two computational steps (estimation of local 2-D motion vectors, followed by their spatial pooling), both being identified in the area MT. Psychophysical findings, however, suggest that under certain stimulus conditions, the human visual system can also compute mathematically correct global motion vectors from direct pooling of spatially distributed 1-D motion signals. To study the neural mechanisms responsible for this novel 1-D motion pooling, we conducted human magnetoencephalography (MEG) and functional MRI experiments using a global motion stimulus comprising multiple moving Gabors (global-Gabor motion). In the first experiment, we measured MEG and blood oxygen level-dependent responses while changing motion coherence of global-Gabor motion. In the second experiment, we investigated cortical responses correlated with direction-selective adaptation to the global 2-D motion, not to local 1-D motions. We found that human MT complex (hMT+) responses show both coherence dependency and direction selectivity to global motion based on 1-D pooling. The results provide the first evidence that hMT+ is the locus of 1-D motion pooling, as well as that of conventional 2-D motion pooling.

Can enhancement and suppression concurrently guide attention? An assessment at the individual level.

BACKGROUND: Although people can pay attention to targets while ignoring distractors, previous research suggests that target enhancement and distractor suppression work separately and independently. Here, we sought to replicate previous findings and re-establish their independence. METHODS: We employed an internet-based psychological experiment. We presented participants with a visual search task in which they searched for a specified shape with or without a singleton. We replicated the singleton-presence benefit in search performance, but this effect was limited to cases where the target color was fixed across all trials. In a randomly intermixed probe task (30% of all trials), the participants searched for a letter among colored probes; we used this task to assess how far attention was separately allocated toward the target or distractor dimensions. RESULTS: We found a negative correlation between target enhancement and distractor suppression, indicating that the participants who paid closer attention to target features ignored distractor features less effectively and vice versa. Averaged data showed no benefit from target color or cost from distractor color, possibly because of the substantial differences in strategy across participants. CONCLUSIONS: These results suggest that target enhancement and distractor suppression guide attention in mutually dependent ways and that the relative contribution of these components depends on the participants' search strategy.

Microstructural Properties of Human Brain Revealed by Fractional Anisotropy Can Predict the After-Effect of Intermittent Theta Burst Stimulation.

Intermittent theta burst stimulation (iTBS) delivered by transcranial magnetic stimulation (TMS) produces a long-term potentiation-like after-effect useful for investigations of cortical function and of potential therapeutic value. However, the iTBS after-effect over the primary motor cortex (M1) as measured by changes in motor evoked potential (MEP) amplitude exhibits a largely unexplained variability across individuals. Here, we present evidence that individual differences in white matter (WM) and gray matter (GM) microstructural properties revealed by fractional anisotropy (FA) predict the magnitude of the iTBS-induced after-effect over M1. The MEP amplitude change in the early phase (5-10 min post-iTBS) was associated with FA values in WM tracts such as right superior longitudinal fasciculus and corpus callosum. By contrast, the MEP amplitude change in the late phase (15-30 min post-iTBS) was associated with FA in GM, primarily in right frontal cortex. These results suggest that the microstructural properties of regions connected directly or indirectly to the target region (M1) are crucial determinants of the iTBS after-effect. FA values indicative of these microstructural differences can predict the potential effectiveness of repetitive TMS for both investigational use and clinical application.

Building a decoder of perceptual decisions from microsaccades and pupil size.

Many studies have reported neural correlates of visual awareness across several brain regions, including the sensory, parietal, and frontal areas. In most of these studies, participants were instructed to explicitly report their perceptual experience through a button press or verbal report. It is conceivable, however, that explicit reporting itself may trigger specific neural responses that can confound the direct examination of the neural correlates of visual awareness. This suggests the need to assess visual awareness without explicit reporting. One way to achieve this is to develop a technique to predict the visual awareness of participants based on their peripheral responses. Here, we used eye movements and pupil sizes to decode trial-by-trial changes in the awareness of a stimulus whose visibility was deteriorated due to adaptation-induced blindness (AIB). In the experiment, participants judged whether they perceived a target stimulus and rated the confidence they had in their perceptual judgment, while their eye movements and pupil sizes were recorded. We found that not only perceptual decision but also perceptual confidence can be separately decoded from the eye movement and pupil size. We discuss the potential of this technique with regard to assessing visual awareness in future neuroimaging experiments.

The DecNef collection, fMRI data from closed-loop decoded neurofeedback experiments.

Decoded neurofeedback (DecNef) is a form of closed-loop functional magnetic resonance imaging (fMRI) combined with machine learning approaches, which holds some promises for clinical applications. Yet, currently only a few research groups have had the opportunity to run such experiments; furthermore, there is no existing public dataset for scientists to analyse and investigate some of the factors enabling the manipulation of brain dynamics. We release here the data from published DecNef studies, consisting of 5 separate fMRI datasets, each with multiple sessions recorded per participant. For each participant the data consists of a session that was used in the main experiment to train the machine learning decoder, and several (from 3 to 10) closed-loop fMRI neural reinforcement sessions. The large dataset, currently comprising more than 60 participants, will be useful to the fMRI community at large and to researchers trying to understand the mechanisms underlying non-invasive modulation of brain dynamics. Finally, the data collection size will increase over time as data from newly run DecNef studies will be added.

V1 Projection Zone Signals in Human Macular Degeneration Depend on Task Despite Absence of Visual Stimulus.

Identifying the plastic and stable components of the visual cortex after retinal loss is an important topic in visual neuroscience and neuro-ophthalmology.1-5 Humans with juvenile macular degeneration (JMD) show significant blood-oxygen-level-dependent (BOLD) responses in the primary visual area (V1) lesion projection zone (LPZ),6 despite the absence of the feedforward signals from the degenerated retina. Our previous study7 reported that V1 LPZ responds to full-field visual stimuli during the one-back task (OBT), not during passive viewing, suggesting the involvement of task-related feedback signals. Aiming to clarify whether visual inputs to the intact retina are necessary for the LPZ responses, here, we measured BOLD responses to tactile and auditory stimuli for both JMD patients and control participants with and without OBT. Participants were instructed to close their eyes during the experiment for the purpose of eliminating retinal inputs. Without OBT, no V1 responses were detected in both groups of participants. With OBT, to the contrary, both stimuli caused substantial V1 responses in JMD patients, but not controls. Furthermore, we also found that the task-dependent activity in V1 LPZ became less pronounced when JMD patients opened their eyes, suggesting that task-related feedback signals can be partially suppressed by residual feedforward signals. Modality-independent V1 LPZ responses only in the task condition suggest that V1 LPZ responses reflect task-related feedback signals rather than reorganized feedforward visual inputs.

Mapping hV4 and ventral occipital cortex: the venous eclipse.

While the fourth human visual field map (hV4) has been studied for two decades, there remain uncertainties about its spatial organization. In analyzing fMRI measurements designed to resolve these issues, we discovered a significant problem that afflicts measurements from ventral occipital cortex, and particularly measurements near hV4. In most hemispheres the fMRI hV4 data are contaminated by artifacts from the transverse sinus (TS). We created a model of the TS artifact and showed that the model predicts the locations of anomalous fMRI responses to simple large-field on-off stimuli. In many subjects, and particularly the left hemisphere, the TS artifact masks fMRI responses specifically in the region of cortex that distinguishes the two main hV4 models. By selecting subjects with a TS displaced from the lateral edge of hV4, we were able to see around the vein. In these subjects, the visual field coverage extends to the lower meridian, or nearly so, consistent with a model in which hV4 is located on the ventral surface and responds to signals throughout the full contralateral hemifield.

Predicting Neural Response Latency of the Human Early Visual Cortex from MRI-Based Tissue Measurements of the Optic Radiation.

Although the non-invasive measurement of visually evoked responses has been extensively studied, the structural basis of variabilities in latency in healthy humans is not well understood. We investigated how tissue properties of optic radiation could predict interindividual variability in the latency of the initial visually evoked component (C1), which may originate from the primary visual cortex (V1). We collected C1 peak latency data using magnetoencephalography (MEG) and checkerboard stimuli, and multiple structural magnetic resonance imaging (MRI) data from 20 healthy subjects. While we varied the contrast and position of the stimuli, the C1 measurement was most reliable when high-contrast stimuli were presented to the lower visual field (LVF). We then attempted to predict interindividual variability in C1 peak latency in this stimulus condition with a multiple regression model using MRI parameters along the optic radiation. We found that this model could predict >20% of variance in C1 latency, when the data were averaged across the hemispheres. The model using the corticospinal tract did not predict variability in C1 latency, suggesting that there is no evidence for generalization to a non-visual tract. In conclusion, our results suggest that the variability in neural latencies in the early visual cortex in healthy subjects can be partly explained by tissue properties along the optic radiation. We discuss the challenges of predicting neural latency using current structural neuroimaging methods and other factors that may explain interindividual variance in neural latency.

Inter-individual Differences in Occipital Alpha Oscillations Correlate with White Matter Tissue Properties of the Optic Radiation.

Neural oscillations at ∼10 Hz, called alpha oscillations, are one of the most prominent components of neural oscillations in the human brain. In recent years, characteristics (power/frequency/phase) of occipital alpha oscillations have been correlated with various perceptual phenomena. However, the relationship between inter-individual differences in alpha oscillatory characteristics and the properties of the underlying brain structures, such as white matter pathways, is unclear. A possibility is that intrinsic occipital alpha oscillations are mediated by thalamocortical interaction; we hypothesized that the most promising candidate for characterizing the intrinsic alpha oscillation is optic radiation (OR), which is the geniculo-cortical pathway carrying signals between the lateral geniculate nucleus (LGN) and primary visual cortex (V1). We used resting-state magnetoencephalography (MEG) and diffusion-weighted/quantitative magnetic resonance imaging (MRI) (dMRI/qMRI) to correlate the frequency and power of occipital alpha oscillations with the tissue properties of the OR by focusing on the different characteristics across individuals. We found that the peak alpha frequency (PAF) negatively correlated with intracellular volume fraction (ICVF), reflecting diffusion properties in intracellular (axonal) space, whereas the peak alpha power was not correlated with any tissue properties measurements. No significant correlation was found between OR and beta frequency/amplitude or between other white matter tract connecting parietal and inferotemporal cortex and alpha frequency/amplitude. These results support the hypothesis that an interaction between thalamic nuclei and early visual areas is essential for the occipital alpha oscillatory rhythm.

Atypical spatial frequency dependence of visual metacognition among schizophrenia patients.

Although altered early stages of visual processing have been reported among schizophrenia patients, how such atypical visual processing may affect higher-level cognition remains largely unknown. Here we tested the hypothesis that metacognitive performance may be atypically modulated by spatial frequency (SF) of visual stimuli among individuals with schizophrenia, given their altered magnocellular function. To study the effect of SF on metacognitive performance, we asked patients and controls to perform a visual detection task on gratings with different SFs and report confidence, and analyzed the data using the signal detection theoretic measure meta-d'. Control subjects showed better metacognitive performance after yes- (stimulus presence) than after no- (stimulus absence) responses ('yes-response advantage') for high SF (HSF) stimuli but not for low SF (LSF) stimuli. The patients, to the contrary, showed a 'yes-response advantage' not only for HSF but also for LSF stimuli, indicating atypical SF dependency of metacognition. An fMRI experiment using the same task revealed that the dorsolateral prefrontal cortex (DLPFC), known to be crucial for metacognition, shows activity mirroring the behavioral results: decoding accuracy of perceptual confidence in DLPFC was significantly higher for HSF than for LSF stimuli in controls, whereas this decoding accuracy was independent of SF in patients. Additionally, the functional connectivity of DLPFC with parietal and visual areas was modulated by SF and response type (yes/no) in a different manner between controls and patients. While individuals without schizophrenia may flexibly adapt metacognitive computations across SF ranges, patients may employ a different mechanism that is independent of SF. Because visual stimuli of low SF have been linked to predictive top-down processing, this may reflect atypical functioning in these processes in schizophrenia.

Visual field maps, population receptive field sizes, and visual field coverage in the human MT+ complex.

Human neuroimaging experiments typically localize motion-selective cortex (MT+) by contrasting responses to stationary and moving stimuli. It has long been suspected that MT+, located on the lateral surface at the temporal-occipital (TO) boundary, contains several distinct visual field maps, although only one coarse map has been measured. Using a novel functional MRI model-based method we identified two maps-TO-1 and TO-2-and measured population receptive field (pRF) sizes within these maps. The angular representation of the first map, TO-1, has a lower vertical meridian on its posterior side at the boundary with the lateral-occipital cortex (i.e., the LO-2 portion). The angular representation continues through horizontal to the upper vertical meridian at the boundary with the second map, TO-2. The TO-2 angle map reverses from upper to lower visual field at increasingly anterior positions. The TO maps share a parallel eccentricity map in which center-to-periphery is represented in the ventral-to-dorsal direction; both maps have an expanded foveal representation. There is a progressive increase in the pRF size from V1/2/3 to LO-1/2 and TO-1/2, with the largest pRF sizes in TO-2. Further, within each map the pRF size increases as a function of eccentricity. The visual field coverage of both maps extends into the ipsilateral visual field, with larger sensitivity to peripheral ipsilateral stimuli in TO-2 than that in TO-1. The TO maps provide a functional segmentation of human motion-sensitive cortex that enables a more complete characterization of processing in human motion-selective cortex.