Neural Dynamics during Binocular Rivalry: Indications from Human Lateral Geniculate Nucleus.

When two sufficiently different stimuli are presented to each eye, perception alternates between them. This binocular rivalry is conceived as a competition for representation in the single stream of visual consciousness. The magnocellular (M) and parvocellular (P) pathways, originating in the retina, encode disparate information, but their potentially different contributions to binocular rivalry have not been determined. Here, we used functional magnetic resonance imaging to measure the human lateral geniculate nucleus (LGN), where the M and P neurons are segregated into layers receiving input from a single eye. We had three participants (one male, two females) and used achromatic stimuli to avoid contributions from color opponent neurons that may have confounded previous studies. We observed activity in the eye-specific regions of LGN correlated with perception, with similar magnitudes during rivalry or physical stimuli alternations, also similar in the M and P regions. These results suggest that LGN activity reflects our perceptions during binocular rivalry and is not simply an artifact of color opponency. Further, perception appears to be a global phenomenon in the LGN, not just limited to a single information channel.

A specialized channel for encoding auditory transients in the magnocellular division of the human medial geniculate nucleus.

We test the hypothesis that there exists a generalized magnocellular system in the brain optimized for temporal processing. In the visual system, it is well known that the magnocellular layers in the lateral geniculate nucleus (LGN) are strongly activated by transients and quickly habituate. However, little is known about the perhaps analogous magnocellular division of the medial geniculate nucleus (MGN), the auditory relay in the thalamus. We measured the functional responses of the MGN in 11 subjects who passively listened to sustained and transient nonlinguistic sounds, using functional MRI. We observed that voxels in the ventromedial portion of the MGN, corresponding to the magnocellular division, exhibited a robust preference to transient sounds, consistently across subjects, whereas the remainder of the MGN did not discriminate between sustained and transient sounds. We conclude that the magnocellular neurons in the MGN parallel the magnocellular neurons in its visual counterpart, LGN, and constitute an information stream specialized for encoding auditory dynamics.

A three-response task reveals how attention alters decision criteria but not appearance.

Whether attention alters appearance or just changes decision criteria continues to be controversial. When subjects are forced to choose which of two equal targets, one of which has been pre-cued, has a higher contrast, they tend to choose the cued target. This has been interpreted as attention increasing the apparent contrast of the cued target. However, when subjects must decide whether the two targets have equal or unequal contrast, they respond veridically with no apparent effect of attention. The discrepancy between these comparative and equality judgments is explained by attention altering the decision criteria but not appearance. We supposed that when subjects are forced to choose which of two apparently equal targets has the higher contrast, they tend to proportion their uncertainty in favor of the cued target. To test this hypothesis, we used a three-response task, in which subjects chose which target had the higher contrast but also had the option to report that the targets appeared equal. This task disentangled potential attention effects on appearance from those on the decision criteria. We found that subjects with narrower criteria about what constituted equal contrast were more likely to choose the cued target, supporting the uncertainty stealing hypothesis. Across the population, the effects of the attentional cue are explained as changes in the decision criteria and not changes in appearance.

Distinguishing Hemodynamics from Function in the Human LGN Using a Temporal Response Model.

We developed a temporal population receptive field model to differentiate the neural and hemodynamic response functions (HRF) in the human lateral geniculate nucleus (LGN). The HRF in the human LGN is dominated by the richly vascularized hilum, a structure that serves as a point of entry for blood vessels entering the LGN and supplying the substrates of central vision. The location of the hilum along the ventral surface of the LGN and the resulting gradient in the amplitude of the HRF across the extent of the LGN have made it difficult to segment the human LGN into its more interesting magnocellular and parvocellular regions that represent two distinct visual processing streams. Here, we show that an intrinsic clustering of the LGN responses to a variety of visual inputs reveals the hilum, and further, that this clustering is dominated by the amplitude of the HRF. We introduced a temporal population receptive field model that includes separate sustained and transient temporal impulse response functions that vary on a much short timescale than the HRF. When we account for the HRF amplitude, we demonstrate that this temporal response model is able to functionally segregate the residual responses according to their temporal properties.

Larger Auditory Cortical Area and Broader Frequency Tuning Underlie Absolute Pitch.

Absolute pitch (AP), the ability of some musicians to precisely identify and name musical tones in isolation, is associated with a number of gross morphological changes in the brain, but the fundamental neural mechanisms underlying this ability have not been clear. We presented a series of logarithmic frequency sweeps to age- and sex-matched groups of musicians with or without AP and controls without musical training. We used fMRI and population receptive field (pRF) modeling to measure the responses in the auditory cortex in 61 human subjects. The tuning response of each fMRI voxel was characterized as Gaussian, with independent center frequency and bandwidth parameters. We identified three distinct tonotopic maps, corresponding to primary (A1), rostral (R), and rostral-temporal (RT) regions of auditory cortex. We initially hypothesized that AP abilities might manifest in sharper tuning in the auditory cortex. However, we observed that AP subjects had larger cortical area, with the increased area primarily devoted to broader frequency tuning. We observed anatomically that A1, R and RT were significantly larger in AP musicians than in non-AP musicians or control subjects, which did not differ significantly from each other. The increased cortical area in AP in areas A1 and R were primarily low frequency and broadly tuned, whereas the distribution of responses in area RT did not differ significantly. We conclude that AP abilities are associated with increased early auditory cortical area devoted to broad-frequency tuning and likely exploit increased ensemble encoding.SIGNIFICANCE STATEMENT Absolute pitch (AP), the ability of some musicians to precisely identify and name musical tones in isolation, is associated with a number of gross morphological changes in the brain, but the fundamental neural mechanisms have not been clear. Our study shows that AP musicians have significantly larger volume in early auditory cortex than non-AP musicians and non-musician controls and that this increased volume is primarily devoted to broad-frequency tuning. We conclude that AP musicians are likely able to exploit increased ensemble representations to encode and identify frequency.

The Flash-Lag, Fröhlich and Related Motion Illusions Are Natural Consequences of Discrete Sampling in the Visual System.

The Fröhlich effect and flash-lag effect, in which moving objects appear advanced along their trajectories compared to their actual positions, have defied a simple and consistent explanation. Here, I show that these illusions can be understood as a natural consequence of temporal compression in the human visual system. Discrete sampling at some stage of sensory perception has long been considered, and if it were true, it would necessarily lead to these illusions of motion. I show that the discrete perception hypothesis, with a single free parameter, the perceptual moment or sampling rate, can quantitatively explain all of the scenarios of the Fröhlich and flash-lag effect. I interpret discrete perception as the implementation of data compression in the brain, and our conscious perception as the reconstruction of the compressed input.

Hemispheric asymmetries in the orientation and location of the lateral geniculate nucleus in dyslexia.

Human brain asymmetry reflects normal specialization of functional roles and may derive from evolutionary, hereditary, developmental, experiential, and pathological factors (Toga & Thompson, 2003). Geschwind and Galaburda (1985) suggested that processing difficulties in dyslexia are due to structural differences between hemispheres. Because of its potential significance to the controversial magnocellular theory of dyslexia, we investigated hemispheric differences in the human lateral geniculate nucleus (LGN), the primary visual relay and control nucleus in the thalamus, in subjects with dyslexia compared to normal readers. We acquired and averaged multiple high-resolution proton density (PD) weighted structural magnetic resonance imaging (MRI) volumes to measure in detail the anatomical boundaries of the LGN in each hemisphere. We observed hemispheric asymmetries in the orientation of the nucleus in subjects with dyslexia that were absent in controls. We also found differences in the location of the LGN between hemispheres in controls but not in subjects with dyslexia. Neither the precise anatomical differences in the LGN nor their functional consequences are known, nor is it clear whether the differences might be causes or effects of dyslexia.

Measuring Connectivity in the Primary Visual Pathway in Human Albinism Using Diffusion Tensor Imaging and Tractography.

In albinism, the number of ipsilaterally projecting retinal ganglion cells (RGCs) is significantly reduced. The retina and optic chiasm have been proposed as candidate sites for misrouting. Since a correlation between the number of lateral geniculate nucleus (LGN) relay neurons and LGN size has been shown, and based on previously reported reductions in LGN volumes in human albinism, we suggest that fiber projections from LGN to the primary visual cortex (V1) are also reduced. Studying structural differences in the visual system of albinism can improve the understanding of the mechanism of misrouting and subsequent clinical applications. Diffusion data and tractography are useful for mapping the OR (optic radiation). This manuscript describes two algorithms for OR reconstruction in order to compare brain connectivity in albinism and controls.An MRI scanner with a 32-channel head coil was used to acquire structural scans. A T1-weighted 3D-MPRAGE sequence with 1 mm(3) isotropic voxel size was used to generate high-resolution images for V1 segmentation. Multiple proton density (PD) weighted images were acquired coronally for right and left LGN localization. Diffusion tensor imaging (DTI) scans were acquired with 64 diffusion directions. Both deterministic and probabilistic tracking methods were run and compared, with LGN as the seed mask and V1 as the target mask. Though DTI provides relatively poor spatial resolution, and accurate delineation of OR may be challenging due to its low fiber density, tractography has been shown to be advantageous both in research and clinically. Tract based spatial statistics (TBSS) revealed areas of significantly reduced white matter integrity within the OR in patients with albinism compared to controls. Pairwise comparisons revealed a significant reduction in LGN to V1 connectivity in albinism compared to controls. Comparing both tracking algorithms revealed common findings, strengthening the reliability of the technique.

Population Receptive Field Estimation Reveals New Retinotopic Maps in Human Subcortex.

The human subcortex contains multiple nuclei that govern the transmission of information to and among cortical areas. In the visual domain, these nuclei are organized into retinotopic maps. Because of their small size, these maps have been difficult to precisely measure using phase-encoded functional magnetic resonance imaging, particularly in the eccentricity dimension. Using instead the population receptive field model to estimate the response properties of individual voxels, we were able to resolve two previously unreported retinotopic maps in the thalamic reticular nucleus and the substantia nigra. We measured both the polar angle and eccentricity components, receptive field size and hemodynamic response function delay, in the these nuclei and in the lateral geniculate nucleus, the superior colliculus, and the lateral and intergeniculate pulvinars. The anatomical boundaries of these nuclei were delineated using multiple averaged proton density-weighted images and were used to constrain and confirm the functional activations. Deriving the retinotopic organization of these small, subcortical nuclei is the first step in exploring their response properties and their roles in neural dynamics.

Morphological differences in the lateral geniculate nucleus associated with dyslexia.

Developmental dyslexia is a common learning disability characterized by normal intelligence but difficulty in skills associated with reading, writing and spelling. One of the most prominent, albeit controversial, theories of dyslexia is the magnocellular theory, which suggests that malfunction of the magnocellular system in the brain is responsible for the behavioral deficits. We sought to test the basis of this theory by directly measuring the lateral geniculate nucleus (LGN), the only location in the brain where the magnocellular and parvocellular streams are spatially disjoint. Using high-resolution proton-density weighted MRI scans, we precisely measured the anatomical boundaries of the LGN in 13 subjects with dyslexia (five female) and 13 controls (three female), all 22-26 years old. The left LGN was significantly smaller in volume in subjects with dyslexia and also differed in shape; no differences were observed in the right LGN. The functional significance of this asymmetry is unknown, but these results are consistent with the magnocellular theory and support theories of dyslexia that involve differences in the early visual system.