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Belief updating entails the incorporation of new information about the environment into internal models of the world. Bayesian inference is the statistically optimal strategy for performing belief updating in the presence of uncertainty. An important open question is whether the use of cognitive strategies that implement Bayesian inference is dependent upon motivational state and, if so, how this is reflected in electrophysiological signatures of belief updating in the brain. Here, we recorded the EEG of participants performing a simple reward learning task with both monetary and nonmonetary instructive feedback conditions. Our aim was to distinguish the influence of the rewarding properties of feedback on belief updating from the information content of the feedback itself. A Bayesian updating model allowed us to quantify different aspects of belief updating across trials, including the size of belief updates and the uncertainty of beliefs. Faster learning rates were observed in the monetary feedback condition compared to the instructive feedback condition, while belief updates were generally larger, and belief uncertainty smaller, with monetary compared to instructive feedback. Larger amplitudes in the monetary feedback condition were found for three ERP components: the P3a, the feedback-related negativity, and the late positive potential. These findings suggest that motivational state influences inference strategies in reward learning, and this is reflected in the electrophysiological correlates of belief updating.

Significance Navigating a 3D world requires sensitivity to motion in depth. The human visual system extracts two stereoscopic cues to motion in depth but little is known about the visual pathways that support their computation. Here, we show that achromatic inputs drive a 3D motion signal computed from slow changes in binocular disparity. However, the other 3D motion signal, based on interocular velocity differences, may draw preferentially on information carried by the evolutionarily ancient koniocellular pathway. Paradoxically, this pathway is thought to be far less sensitive to both motion and stereoscopic depth, yet, we find evidence for its involvement in the extraction of motion through depth. Our findings demonstrate an unanticipated role for the koniocellular pathway in a fundamental perceptual mechanism., Motion in depth (MID) can be cued by high-resolution changes in binocular disparity over time (CD), and low-resolution interocular velocity differences (IOVD). Computational differences between these two mechanisms suggest that they may be implemented in visual pathways with different spatial and temporal resolutions. Here, we used fMRI to examine how achromatic and S-cone signals contribute to human MID perception. Both CD and IOVD stimuli evoked responses in a widespread network that included early visual areas, parts of the dorsal and ventral streams, and motion-selective area hMT+. Crucially, however, we measured an interaction between MID type and chromaticity. fMRI CD responses were largely driven by achromatic stimuli, but IOVD responses were better driven by isoluminant S-cone inputs. In our psychophysical experiments, when S-cone and achromatic stimuli were matched for perceived contrast, participants were equally sensitive to the MID in achromatic and S-cone IOVD stimuli. In comparison, they were relatively insensitive to S-cone CD. These findings provide evidence that MID mechanisms asymmetrically draw on information in precortical pathways. An early opponent motion signal optimally conveyed by the S-cone pathway may provide a substantial contribution to the IOVD mechanism.

MH was supported by the European Union’s Horizon 2020 Research and Innovation Program under the Marie Skłodowska-Curie grant agreement no. 641805. AW was supported by United Kingdom Biotechnology and Biological Science Research Council (BBSRC) grant number BB/M002543/1, and JH by BBSRC grant number BB/M001660/1. Two stereoscopic cues that underlie the perception of motion-in-depth (MID) are changes in retinal disparity over time (CD) and interocular velocity differences (IOVD). These cues have independent spatiotemporal sensitivity profiles, depend upon different low-level stimulus properties, and are potentially processed along separate cortical pathways. Here, we ask whether these MID cues code for different motion directions: do they give rise to discriminable patterns of neural signals, and is there evidence for their convergence onto a single “motion-in-depth” pathway? To answer this, we use a decoding algorithm to test whether, and when, patterns of electroencephalogram (EEG) signals measured from across the full scalp, generated in response to CD- and IOVD-isolating stimuli moving toward or away in depth can be distinguished. We find that both MID cue type and 3D-motion direction can be decoded at different points in the EEG timecourse and that direction decoding cannot be accounted for by static disparity information. Remarkably, we find evidence for late processing convergence: IOVD motion direction can be decoded relatively late in the timecourse based on a decoder trained on CD stimuli, and vice versa. We conclude that early CD and IOVD direction decoding performance is dependent upon fundamentally different low-level stimulus features, but that later stages of decoding performance may be driven by a central, shared pathway that is agnostic to these features. Overall, these data are the first to show that neural responses to CD and IOVD cues that move toward and away in depth can be decoded from EEG signals, and that different aspects of MID-cues contribute to decoding performance at different points along the EEG timecourse. Publisher PDF

Two binocular sources of information serve motion-in-depth (MID) perception: changes in disparity over time (CD), and interocular velocity differences (IOVD). While CD requires the computation of small spatial disparities, IOVD could be computed from a much lower-resolution signal. IOVD signals therefore might still be available under conditions of binocular vision impairment (BVI) with limited or no stereopsis, for example, amblyopia.Sensitivity to CD and IOVD was measured in adults who had undergone therapy to correct optical misalignment or amblyopia in childhood (n = 16), as well as normal vision controls with good stereoacuity (n = 8). Observers discriminated the interval containing a smoothly oscillating MID "test" stimulus from a "control" stimulus in a two-interval forced choice paradigm.Of the BVI observers with no static stereoacuity (n = 9), one displayed evidence for sensitivity to IOVD only, while there was otherwise no sensitivity for either CD or IOVD in the group. Generally, BVI observers with measurable stereoacuity (n = 7) displayed a pattern resembling the control group: showing a similar sensitivity for both cues. A neutral density filter placed in front of the fixing eye in a subset of BVI observers did not improve performance.In one BVI observer there was preserved sensitivity to IOVD but not CD, though overall only those BVI observers with at least gross stereopsis were able to detect disparity- or velocity-based cues to MID. The results imply that these logically distinct information sources are somehow coupled, and in some cases BVI observers with no stereopsis may still retain sensitivity to IOVD.

Virtual reality presents a new set of challenges and opportunities for both engineers and neuroscientists. Here we provide an overview of a programme designed by a group of psychologists, neuroscientists and VR specialists to address some of the most outstanding issues in the field ranging from the very low-level (for example, how the brain processed motion-in-depth signals generated by stereoscopic display devices) to the very high level (how virtual environments can lead to a sense of immersion and emotional engagement). We present data from psychophysical, electrophysiological and neuroimaging experiments and explain how different research methodologies can be applied to different problems in the field of VR/AR. We end by describing an open-source, extensible software package for studying issues in VR that can interface to common laboratory measurement equipment and discussing future directions and challenges facing the neuroscience and VR engineering communities.

Belief updating entails the incorporation of new information about the environment into internal models of the world. Bayesian inference is the statistically optimal strategy for performing belief updating in the presence of uncertainty. An important open question is whether the use of cognitive strategies that implement Bayesian inference is dependent upon motivational state and, if so, how this is reflected in electrophysiological signatures of belief updating in the brain. Here we recorded the electroencephalogram of participants performing a simple reward learning task with both monetary and non-monetary instructive feedback conditions. Our aim was to distinguish the influence of the rewarding properties of feedback on belief updating from the information content of the feedback itself. A Bayesian updating model allowed us to quantify different aspects of belief updating across trials, including the size of belief updates and the uncertainty of beliefs. Faster learning rates were observed in the monetary feedback condition compared to the instructive feedback condition, while belief updates were generally larger, and belief uncertainty smaller, with monetary compared to instructive feedback. Larger amplitudes in the monetary feedback condition were found for three event-related potential components: the P3a, the feedback-related negativity (FRN) and the late positive potential (LPP). These findings suggest that motivational state influences inference strategies in reward learning, and this is reflected in the electrophysiological correlates of belief updating.

Anisotropies in the cortical representation of various stimulus parameters can reveal the fundamental mechanisms by which sensory properties are analysed and coded by the brain. One example is the preference for motion radial to the point of fixation (i.e. centripetal or centrifugal) exhibited in mammalian visual cortex. In two experiments, this study used functional magnetic resonance imaging (fMRI) to explore the determinants of these radial biases for motion in functionally-defined areas of human early visual cortex, and in particular their dependence upon eccentricity which has been indicated in recent reports. In one experiment, the cortical response to wide-field random dot kinematograms forming 16 different complex motion patterns (including centrifugal, centripetal, rotational and spiral motion) was measured. The response was analysed according to preferred eccentricity within four different eccentricity ranges. Response anisotropies were characterised by enhanced activity for centripetal or centrifugal patterns that changed systematically with eccentricity in visual areas V1-V3 and hV4 (but not V3A/B or V5/MT+). Responses evolved from a preference for centrifugal over centripetal patterns close to the fovea, to a preference for centripetal over centrifugal at the most peripheral region stimulated, in agreement with previous work. These effects were strongest in V2 and V3. In a second experiment, the stimuli were restricted to within narrow annuli either close to the fovea (0.75-1.88°) or further in the periphery (4.82-6.28°), in a way that preserved the local motion information available in the first experiment. In this configuration a preference for radial motion (centripetal or centrifugal) persisted but the dependence upon eccentricity disappeared. Again this was clearest in V2 and V3. A novel interpretation of the dependence upon eccentricity of motion anisotropies in early visual cortex is offered that takes into account the spatiotemporal "predictability" of the moving pattern. Such stimulus predictability, and its relationship to models of predictive coding, has found considerable support in recent years in accounting for a number of other perceptual and neural phenomena.

Previous research into the effects of viewpoint change on face recognition has typically dealt with rotations around the head's vertical axis (yaw). Another common, although less studied, source of viewpoint variation in faces is rotation around the head's horizontal pitch axis (pitch). In the current study we used both a sequential matching task and an old/new recognition task to examine the effect of viewpoint change following rotation about both pitch and yaw axes on human face recognition. The results of both tasks showed that recognition performance was better for faces rotated about yaw compared to pitch. Further, recognition performance for faces rotated upwards on the pitch axis was better than for faces rotated downwards. Thus, equivalent angular rotations about pitch and yaw do not produce equivalent viewpoint-dependent declines in recognition performance.

Multivariate decoding analyses are widely applied to functional magnetic resonance imaging (fMRI) data, but there is controversy over their interpretation. Orientation decoding in primary visual cortex (V1) reflects coarse-scale biases, including an over-representation of radial orientations. But fMRI responses to clockwise and counter-clockwise spirals can also be decoded. Because these stimuli are matched for radial orientation, while differing in local orientation, it has been argued that fine-scale columnar selectivity for orientation contributes to orientation decoding. We measured fMRI responses in human V1 to both oriented gratings and spirals. Responses to oriented gratings exhibited a complex topography, including a radial bias that was most pronounced in the peripheral representation, and a near-vertical bias that was most pronounced near the foveal representation. Responses to clockwise and counter-clockwise spirals also exhibited coarse-scale organization, at the scale of entire visual quadrants. The preference of each voxel for clockwise or counter-clockwise spirals was predicted from the preferences of that voxel for orientation and spatial position (i.e., within the retinotopic map). Our results demonstrate a bias for local stimulus orientation that has a coarse spatial scale, is robust across stimulus classes (spirals and gratings), and suffices to explain decoding from fMRI responses in V1.

Surface segregation provides an efficient way to parse the visual scene for perceptual analysis. Here, we investigated the segregation of a bivectorial motion display into transparent surfaces through a psychophysical task and fMRI. We found that perceptual transparency correlated with neural activity in the early areas of the visual cortex, suggesting these areas may be involved in the segregation of motion-defined surfaces. Two oppositely rotating, uniquely colored random dot kinematograms (RDKs) were presented either sequentially or in a spatially interleaved manner, displayed at varying alternation frequencies. Participants reported the color and rotation direction pairing of the RDKs in the psychophysical task. The spatially interleaved display generated the percept of motion transparency across the range of frequencies tested, yielding ceiling task performance. At high alternation frequencies, performance on the sequential display also approached ceiling, indicative of perceived transparency. However, transparency broke down in lower alternation frequency sequential displays, producing performance close to chance. A corresponding pattern mirroring the psychophysical data was also evident in univariate and multivariate analyses of the fMRI BOLD activity in visual cortical areas V1, V2, V3, V3AB, hV4, and V5/MT+. Using gray RDKs, we found significant presentation by frequency interactions in most areas; differences in BOLD signal between presentation types were significant only at the lower alternation frequency. Multivariate pattern classification was similarly unable to discriminate between presentation types at the higher frequency. This study provides evidence that early visual cortex may code for motion-defined surface segregation, which in turn may enable perceptual transparency.

Complex patterns of image motion (contracting, expanding, rotating, and spiraling fields) are important in the coordination of visually guided behaviors. Whereas specialized detectors in monkey visual cortex show selectivity for particular patterns of complex motion, their representation in human visual cortex remains unclear. In the present study, functional magnetic resonance imaging (fMRI) was used to investigate the sensitivity of functionally defined regions of human visual cortex to parametrically modulated complex motion trajectories, coupled with complementary psychophysical testing. A unique stimulus design made it possible to disambiguate the neural responses and psychophysical sensitivity to complex motions per se from the distribution of local motions relative to the fovea, which are known to enhance cortical activity when presented radial to fixation. This involved presenting several small, separate motion fields in the periphery in a manner that distinguished them from global optic flow patterns. The patterns were morphed through complex motion space in a systematic time-locked fashion when presented in the scanner. Anisotropies were observed in the fMRI signal, marked by an enhanced response to expanding vs. contracting fields, even in early visual cortex. Anisotropies in the psychophysical sensitivity measures followed a similar pattern that was correlated with activity in areas hV4, V5/MT, and MST. This represents the first systematic examination of complex motion perception at both a behavioral and neural level in human observers. The characteristic processing anisotropy revealed in both data sets can inform models of complex motion processing, particularly with respect to computations performed in early visual cortex.

When two dissimilar colours are displayed to the two eyes at overlapping retinal locations, binocular rivalry typically results: a fluctuating struggle for perceptual dominance of each eye's stimulus. We found instead that isoluminant counter-rotating patterns consisting of coloured and achromatic portions can promote an illusory colour ‘misbinding’, where the colours from both eyes were perceived within a single rotating pattern. The achromatic portion of one rotating pattern thus appeared to take on the colour of the other, oppositely rotating pattern. The results suggest that the neural mechanisms of colour binding can operate even while representations of the same patterns' motions are undergoing rivalry, and support the idea that rivalry can occur in isolation within the motion system.

The crustacean stomatogastric ganglion (STG) is modulated by a large number of amines and neuropeptides that are found in descending pathways from anterior ganglia or reach the STG via the hemolymph. Among these are the allatostatin (AST) B types, also known as myoinhibitory peptides (MIPs). We used mass spectrometry to determine the sequences of nine members of the AST-B family of peptides that were found in the stomatogastric nervous system of the crab Cancer borealis. We raised an antibody against Cancer borealis allatostatin-B1 (CbAST-B1; VPNDWAHFRGSWa) and used it to map the distribution of CbAST-B1-like immunoreactivity (-LI) in the stomatogastric nervous system. CbAST-B1-LI was found in neurons and neuropil in the commissural ganglia (CoGs), in somata in the esophageal ganglion (OG), in fibers in the stomatogastric nerve (stn), and in neuropilar processes in the STG. CbAST-B1-LI was blocked by preincubation with 10(-6) M CbAST-B1 and was partially blocked by lower concentrations. Electrophysiological recordings of the effects of CbAST-B1, CbAST-B2, and CbAST-B3 on the pyloric rhythm of the STG showed that all three peptides inhibited the pyloric rhythm in a state-dependent manner. Specifically, all three peptides at 10(-8) M significantly decreased the frequency of the pyloric rhythm when the initial frequency of the pyloric rhythm was below 0.6 Hz. These data suggest important neuromodulatory roles for the CbAST-B family in the stomatogastric nervous system.

Visual feature binding-the mechanism by which our typically coherent and unified perceptual experience arises from distributed neural representations-is the source of much intrigue in the neuroscience of perception. Surprisingly, feature binding can occur in rapidly alternating displays of color-orientation combinations (e.g., rightward-orange, leftward-blue). However, we found that when the angular separation between orientations is reduced, binding is selectively impaired at temporal alternation frequencies around 5 Hz. To isolate the mechanisms involved, we devised a novel display in which color-orientation conjunction information was distributed temporally over two checkered stimuli and was perceptually discriminable only within an intermediate range of temporal frequencies (7.5-15 Hz). We propose that accurate color-orientation judgments at frequencies exceeding 5 Hz depend on the rapid formation of persistent surface representations that can be accessed by binding mechanisms, circumventing the latter's relatively low temporal resolution.

When two fields of dots with different directions of movement are presented in tandem, the perceived direction of one is biased by the presence of the other. Although this ‘‘direction illusion’’ typically involves repulsion, with an exaggeration of the perceived angular difference in direction between the dot fields, attraction effects, where the perceived difference is reduced, have also been found under certain presentation conditions. Earlier literature has been inconsistent, and there is debate surrounding the nature of the interactions that facilitate the direction illusion, as well as whether they occur at a local or global stage of the motion processing hierarchy. Here we measured the operating characteristics of the direction illusion by parametrically varying inducer contrast and coherence while examining the effects of stimulus speed and dichoptic presentation. It was found that the magnitude and sign of the direction illusion differed substantially from earlier research. Furthermore, there appeared to be significant interindividual variability, with dichoptic presentation producing an attractive rather than repulsive direction illusion in some participants.