Structural covariance and heritability of the optic tract and primary visual cortex in living human brains.

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

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.

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.

Occipital White Matter Tracts in Human and Macaque.

We compare several major white-matter tracts in human and macaque occipital lobe using diffusion magnetic resonance imaging. The comparison suggests similarities but also significant differences in the tracts. There are several apparently homologous tracts in the 2 species, including the vertical occipital fasciculus (VOF), optic radiation, forceps major, and inferior longitudinal fasciculus (ILF). There is one large human tract, the inferior fronto-occipital fasciculus, with no corresponding fasciculus in macaque. We could identify the macaque VOF (mVOF), which has been little studied. Its position is consistent with classical invasive anatomical studies by Wernicke. VOF homology is supported by similarity of the endpoints in V3A and ventral V4 across species. The mVOF fibers intertwine with the dorsal segment of the ILF, but the human VOF appears to be lateral to the ILF. These similarities and differences between the occipital lobe tracts will be useful in establishing which circuitry in the macaque can serve as an accurate model for human visual cortex.

Ensemble Tractography.

Tractography uses diffusion MRI to estimate the trajectory and cortical projection zones of white matter fascicles in the living human brain. There are many different tractography algorithms and each requires the user to set several parameters, such as curvature threshold. Choosing a single algorithm with specific parameters poses two challenges. First, different algorithms and parameter values produce different results. Second, the optimal choice of algorithm and parameter value may differ between different white matter regions or different fascicles, subjects, and acquisition parameters. We propose using ensemble methods to reduce algorithm and parameter dependencies. To do so we separate the processes of fascicle generation and evaluation. Specifically, we analyze the value of creating optimized connectomes by systematically combining candidate streamlines from an ensemble of algorithms (deterministic and probabilistic) and systematically varying parameters (curvature and stopping criterion). The ensemble approach leads to optimized connectomes that provide better cross-validated prediction error of the diffusion MRI data than optimized connectomes generated using a single-algorithm or parameter set. Furthermore, the ensemble approach produces connectomes that contain both short- and long-range fascicles, whereas single-parameter connectomes are biased towards one or the other. In summary, a systematic ensemble tractography approach can produce connectomes that are superior to standard single parameter estimates both for predicting the diffusion measurements and estimating white matter fascicles.

A Major Human White Matter Pathway Between Dorsal and Ventral Visual Cortex.

Human visual cortex comprises many visual field maps organized into clusters. A standard organization separates visual maps into 2 distinct clusters within ventral and dorsal cortex. We combined fMRI, diffusion MRI, and fiber tractography to identify a major white matter pathway, the vertical occipital fasciculus (VOF), connecting maps within the dorsal and ventral visual cortex. We use a model-based method to assess the statistical evidence supporting several aspects of the VOF wiring pattern. There is strong evidence supporting the hypothesis that dorsal and ventral visual maps communicate through the VOF. The cortical projection zones of the VOF suggest that human ventral (hV4/VO-1) and dorsal (V3A/B) maps exchange substantial information. The VOF appears to be crucial for transmitting signals between regions that encode object properties including form, identity, and color and regions that map spatial information.

The visual white matter: The application of diffusion MRI and fiber tractography to vision science.

Visual neuroscience has traditionally focused much of its attention on understanding the response properties of single neurons or neuronal ensembles. The visual white matter and the long-range neuronal connections it supports are fundamental in establishing such neuronal response properties and visual function. This review article provides an introduction to measurements and methods to study the human visual white matter using diffusion MRI. These methods allow us to measure the microstructural and macrostructural properties of the white matter in living human individuals; they allow us to trace long-range connections between neurons in different parts of the visual system and to measure the biophysical properties of these connections. We also review a range of findings from recent studies on connections between different visual field maps, the effects of visual impairment on the white matter, and the properties underlying networks that process visual information supporting visual face recognition. Finally, we discuss a few promising directions for future studies. These include new methods for analysis of MRI data, open datasets that are becoming available to study brain connectivity and white matter properties, and open source software for the analysis of these data.

Tractometry of Human Visual White Matter Pathways in Health and Disease.

Diffusion-weighted MRI (dMRI) provides a unique non-invasive view of human brain tissue properties. The present review article focuses on tractometry analysis methods that use dMRI to assess the properties of brain tissue within the long-range connections comprising brain networks. We focus specifically on the major white matter tracts that convey visual information. These connections are particularly important because vision provides rich information from the environment that supports a large range of daily life activities. Many of the diseases of the visual system are associated with advanced aging, and tractometry of the visual system is particularly important in the modern aging society. We provide an overview of the tractometry analysis pipeline, which includes a primer on dMRI data acquisition, voxelwise model fitting, tractography, recognition of white matter tracts, and calculation of tract tissue property profiles. We then review dMRI-based methods for analyzing visual white matter tracts: the optic nerve, optic tract, optic radiation, forceps major, and vertical occipital fasciculus. For each tract, we review background anatomical knowledge together with recent findings in tractometry studies on these tracts and their properties in relation to visual function and disease. Overall, we find that measurements of the brain's visual white matter are sensitive to a range of disorders and correlate with perceptual abilities. We highlight new and promising analysis methods, as well as some of the current barriers to progress toward integration of these methods into clinical practice. These barriers, such as variability in measurements between protocols and instruments, are targets for future development.

White matter tracts adjacent to the human cingulate sulcus visual area (CSv).

Human cingulate sulcus visual area (CSv) was first identified as an area that responds selectively to visual stimulation indicative of self-motion. It was later shown that the area is also sensitive to vestibular stimulation as well as to bodily motion compatible with locomotion. Understanding the anatomical connections of CSv will shed light on how CSv interacts with other parts of the brain to perform information processing related to self-motion and navigation. A previous neuroimaging study (Smith et al. 2018, Cerebral Cortex, 28, 3685-3596) used diffusion-weighted magnetic resonance imaging (dMRI) to examine the structural connectivity of CSv, and demonstrated connections between CSv and the motor and sensorimotor areas in the anterior and posterior cingulate sulcus. The present study aimed to complement this work by investigating the relationship between CSv and adjacent major white matter tracts, and to map CSv's structural connectivity onto known white matter tracts. By re-analysing the dataset from Smith et al. (2018), we identified bundles of fibres (i.e. streamlines) from the whole-brain tractography that terminate near CSv. We then assessed to which white matter tracts those streamlines may belong based on previously established anatomical prescriptions. We found that a significant number of CSv streamlines can be categorised as part of the dorsalmost branch of the superior longitudinal fasciculus (SLF I) and the cingulum. Given current thinking about the functions of these white matter tracts, our results support the proposition that CSv provides an interface between sensory and motor systems in the context of self-motion.

A prominent vertical occipital white matter fasciculus unique to primate brains.

Vision in humans and other primates enlists parallel processing streams in the dorsal and ventral visual cortex, known to support spatial and object processing, respectively. These streams are bridged, however, by a prominent white matter tract, the vertical occipital fasciculus (VOF), identified in both classical neuroanatomy and recent diffusion-weighted magnetic resonance imaging (dMRI) studies. Understanding the evolution of the VOF may shed light on its origin, function, and role in visually guided behaviors. To this end, we acquired high-resolution dMRI data from the brains of select mammalian species, including anthropoid and strepsirrhine primates, a tree shrew, rodents, and carnivores. In each species, we attempted to delineate the VOF after first locating the optic radiations in the occipital white matter. In all primate species examined, the optic radiation was flanked laterally by a prominent and coherent white matter fasciculus recognizable as the VOF. By contrast, the equivalent analysis applied to four non-primate species from the same superorder as primates (tree shrew, ground squirrel, paca, and rat) failed to reveal white matter tracts in the equivalent location. Clear evidence for a VOF was also absent in two larger carnivore species (ferret and fox). Although we cannot rule out the existence of minor or differently organized homologous fiber pathways in the non-primate species, the results suggest that the VOF has greatly expanded, or possibly emerged, in the primate lineage. This adaptation likely facilitated the evolution of unique visually guided behaviors in primates, with direct impacts on manual object manipulation, social interactions, and arboreal locomotion.

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.

Functional and Structural Properties of Interhemispheric Interaction between Bilateral Precentral Hand Motor Regions in a Top Wheelchair Racing Paralympian.

Long-term motor training can cause functional and structural changes in the human brain. Assessing how the training of specific movements affects specific parts of the neural circuitry is essential to understand better the underlying mechanisms of motor training-induced plasticity in the human brain. We report a single-case neuroimaging study that investigated functional and structural properties in a professional athlete of wheelchair racing. As wheelchair racing requires bilateral synchronization of upper limb movements, we hypothesized that functional and structural properties of interhemispheric interactions in the central motor system might differ between the professional athlete and controls. Functional and diffusion magnetic resonance imaging (fMRI and dMRI) data were obtained from a top Paralympian (P1) in wheelchair racing. With 23 years of wheelchair racing training starting at age eight, she holds an exceptional competitive record. Furthermore, fMRI and dMRI data were collected from three other paraplegic participants (P2-P4) with long-term wheelchair sports training other than wheelchair racing and 37 able-bodied control volunteers. Based on the fMRI data analyses, P1 showed activation in the bilateral precentral hand sections and greater functional connectivity between these sections during a right-hand unimanual task. In contrast, other paraplegic participants and controls showed activation in the contralateral hemisphere and deactivation in the ipsilateral hemisphere. Moreover, dMRI data analysis revealed that P1 exhibited significantly lower mean diffusivity along the transcallosal pathway connecting the bilateral precentral motor regions than control participants, which was not observed in the other paraplegic participants. These results suggest that long-term training with bilaterally synchronized upper-limb movements may promote bilateral recruitment of the precentral hand sections. Such recruitment may affect the structural circuitry involved in the interhemispheric interaction between the bilateral precentral regions. This study provides valuable evidence of the extreme adaptability of the human brain.

Evaluation of simultaneous multi-slice readout-segmented diffusion-weighted MRI acquisition in human optic nerve measurements.

Diffusion-weighted magnetic resonance imaging (dMRI) is the only available method to measure the tissue properties of white matter tracts in living human brains and has opened avenues for neuroscientific and clinical studies on human white matter. However, dMRI using conventional simultaneous multi-slice (SMS) single-shot echo planar imaging (ssEPI) still presents challenges in the analyses of some specific white matter tracts, such as the optic nerve, which are heavily affected by susceptibility-induced artifacts. In this study, we evaluated dMRI data acquired by using SMS readout-segmented EPI (rsEPI), which aims to reduce susceptibility-induced artifacts by dividing the acquisition space into multiple segments along the readout direction to reduce echo spacing. To this end, we acquired dMRI data from 11 healthy volunteers by using SMS ssEPI and SMS rsEPI, and then compared the dMRI data of the human optic nerve between the SMS ssEPI and SMS rsEPI datasets by visual inspection of the datasets and statistical comparisons of fractional anisotropy (FA) values. In comparison with the SMS ssEPI data, the SMS rsEPI data showed smaller susceptibility-induced distortion and exhibited a significantly higher FA along the optic nerve. In summary, this study demonstrates that despite its prolonged acquisition time, SMS rsEPI is a promising approach for measuring the tissue properties of the optic nerve in living humans and will be useful for future neuroscientific and clinical investigations of this pathway.

Hyper-Adaptation in the Human Brain: Functional and Structural Changes in the Foot Section of the Primary Motor Cortex in a Top Wheelchair Racing Paralympian.

The human brain has the capacity to drastically alter its somatotopic representations in response to congenital or acquired limb deficiencies and dysfunctions. The main purpose of the present study was to elucidate such extreme adaptability in the brain of an active top wheelchair racing Paralympian (participant P1) who has congenital paraplegia (dysfunction of bilateral lower limbs). Participant P1 has undergone long-term wheelchair racing training using bilateral upper limbs and has won a total of 19 medals in six consecutive summer Paralympic games as of 2021. We examined the functional and structural changes in the foot section of the primary motor cortex (M1) in participant P1 as compared to able-bodied control participants. We also examined the functional and structural changes in three other individuals (participants P2, P3, and P4) with acquired paraplegia, who also had long-term non-use period of the lower limbs and had undergone long-term training for wheelchair sports (but not top athletes at the level of participant P1). We measured brain activity in all the participants using functional magnetic resonance imaging (MRI) when bimanual wrist extension-flexion movement was performed, and the structural MRI images were collected. Compared to 37 control participants, participant P1 showed significantly greater activity in the M1 foot section during the bimanual task, and significant local GM expansion in this section. Significantly greater activity in the M1 foot section was also observed in participant P4, but not in P2 and P3, and the significant local GM expansion was observed in participant P2, but not in P3 and P4. Thus, functional or structural change was observed in an acquired paraplegic participant, but was not observed in all the paraplegic participants. The functional and structural changes typically observed in participant P1 may represent extreme adaptability of the human brain. We discuss the results in terms of a new idea of hyper-adaptation.

Multi-Contrast Magnetic Resonance Imaging of Visual White Matter Pathways in Patients With Glaucoma.

Purpose: Glaucoma is a disorder that involves visual field loss caused by retinal ganglion cell damage. Previous diffusion magnetic resonance imaging (dMRI) studies have demonstrated that retinal ganglion cell damage affects tissues in the optic tract (OT) and optic radiation (OR). However, because previous studies have used a simple diffusion tensor model to analyze dMRI data, the microstructural interpretation of white matter tissue changes remains uncertain. In this study, we used a multi-contrast MRI approach to further clarify the type of microstructural damage that occurs in patients with glaucoma. Methods: We collected dMRI data from 17 patients with glaucoma and 30 controls using 3-tesla (3T) MRI. Using the dMRI data, we estimated three types of tissue property metrics: intracellular volume fraction (ICVF), orientation dispersion index (ODI), and isotropic volume fraction (IsoV). Quantitative T1 (qT1) data, which may be relatively specific to myelin, were collected from all subjects. Results: In the OT, all four metrics showed significant differences between the glaucoma and control groups. In the OR, only the ICVF showed significant between-group differences. ICVF was significantly correlated with qT1 in the OR of the glaucoma group, although qT1 did not show any abnormality at the group level. Conclusions: Our results suggest that, at the group level, tissue changes in OR caused by glaucoma might be explained by axonal damage, which is reflected in the intracellular diffusion signals, rather than myelin damage. The significant correlation between ICVF and qT1 suggests that myelin damage might also occur in a smaller number of severe cases.

Whether dots moving in two directions appear coherent or transparent depends on directional biases induced by surrounding motion.

When two random-dot patterns moving in different directions are superimposed, motion appears coherent or transparent depending on the directional difference. In addition, when a pattern is surrounded by another pattern that is moving, the perceived motion of the central stimulus is biased away from the direction of the surrounding motion. That phenomenon is known as induced motion. How is the perception of motion coherence and transparency modulated by surrounding motion? It was found that two random-dot horizontal motions surrounded by another stimulus in downward motion appeared to move in two oblique directions: left-up and right-up. Consequently, when motion transparency occurs, each of the two motions interacts independently with the induced motion direction. Furthermore, for a central stimulus consisting of two physical motions in left-up and right-up directions, the presence of the surrounding stimulus in a vertical motion modulated the perceptual solution of motion coherence/transparency such that if interactions with an induced motion signal narrow the apparent directional difference between the two central motions, then motion coherence is preferred over motion transparency. Therefore, whether a moving stimulus is perceived as coherent or transparent is determined based on the internal representation of motion directions, which can be altered by spatial interactions between adjacent regions.

Age dependency and lateralization in the three branches of the human superior longitudinal fasciculus.

The superior longitudinal fascicle/fasciculus (SLF) is a major white matter tract connecting the frontal and parietal cortices in humans. Although the SLF has often been analyzed as a single entity, several studies have reported that the SLF is segregated into three distinct branches (SLF I, II, and III). They have also reported the right lateralization of the SLF III volume and discussed its relationship with lateralized cortical functions in the fronto-parietal network. However, to date, the homogeneity or heterogeneity of the age dependency and lateralization properties of SLF branches have not been fully clarified. Through this study, we aimed to clarify the age dependency and lateralization of SLF I-III by analyzing diffusion-weighted MRI (dMRI) and quantitative R1 (qR1) map datasets collected from a wide range of age groups, mostly comprising right-handed children, adolescents, adults, and seniors (6 to 81 years old). The age dependency in dMRI measurement (fractional anisotropy, FA) was heterogeneous among the three SLF branches, suggesting that these branches are regulated by distinct developmental and aging processes. Lateralization analysis on SLF branches revealed that the right SLF III was larger than the left SLF III in adults, replicating previous reports. FA measurement also suggested that, in addition to SLF III, SLF II was lateralized to the right hemisphere in adolescents and adults. We further found a left lateralization of SLF I in qR1 data, a microstructural measurement sensitive to myelin levels, in adults. These findings suggest that the SLF sub-bundles are distinct entities in terms of age dependency and lateralization.

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.

Neuronal population decoding explains the change in signal detection sensitivity caused by task-irrelevant perceptual bias.

Spatiotemporal context in sensory stimulus has profound effects on neural responses and perception, and it sometimes affects task difficulty. Recently reported experimental data suggest that human detection sensitivity to motion in a target stimulus can be enhanced by adding a slow surrounding motion in an orthogonal direction, even though the illusory motion component caused by the surround is not relevant to the task. It is not computationally clear how the task-irrelevant component of motion modulates the subject's sensitivity to motion detection. In this study, we investigated the effects of encoding biases on detection performance by modeling the stochastic neural population activities. We modeled two types of modulation on the population activity profiles caused by a contextual stimulus: one type is identical to the activity evoked by a physical change in the stimulus, and the other is expressed more simply in terms of response gain modulation. For both encoding schemes, the motion detection performance of the ideal observer is enhanced by a task-irrelevant, additive motion component, replicating the properties observed for real subjects. The success of these models suggests that human detection sensitivity can be characterized by a noisy neural encoding that limits the resolution of information transmission in the cortical visual processing pathway. On the other hand, analyses of the neuronal contributions to the task predict that the effective cell populations differ between the two encoding schemes, posing a question concerning the decoding schemes that the nervous system used during illusory states.