Population Receptive Field Size Research Articles

The Role of Population Receptive Field Sizes in Higher-Order Visual Dysfunction.

Population receptive field (pRF) modeling is an fMRI technique used to retinotopically map visual cortex, with pRF size characterizing the degree of spatial integration. In clinical populations, most pRF mapping research has focused on damage to visual system inputs. Herein, we highlight recent work using pRF modeling to study high-level visual dysfunctions. Larger pRF sizes, indicating coarser spatial processing, were observed in homonymous visual field deficits, aging, and autism spectrum disorder. Smaller pRF sizes, indicating finer processing, were observed in Alzheimer's disease and schizophrenia. In posterior cortical atrophy, a unique pattern was found in which pRF size changes depended on eccentricity. Changes to pRF properties were observed in clinical populations, even in high-order impairments, explaining visual behavior. These pRF changes likely stem from altered interactions between brain regions. Furthermore, some studies suggested that pRF sizes change as part of cortical reorganization, and they can point towards future prognosis.

Population receptive field size varies between thin vs. thick stripes in cortical areas V2/V3

Population receptive field size varies between thin vs. thick stripes in cortical areas V2/V3

Isolate or combine: population receptive field size in (un)crowding

Isolate or combine: population receptive field size in (un)crowding

Simultaneous changes in visual acuity, cortical population receptive field size, visual field map size, and retinal thickness in healthy human aging

Healthy human aging is associated with a deterioration of visual acuity, retinal thinning, visual field map shrinkage and increasing population receptive field sizes. Here we ask how these changes are related to each other in a cross-sectional sample of fifty healthy adults aged 20–80 years. We hypothesized that age-related loss of macular retinal ganglion cells may lead to decreased visual field map sizes, and both may lead to increased pRF sizes in the cortical central visual field representation. We measured our participants’ perceptual corrected visual acuity using standard ophthalmological letter charts. We then measured their early visual field map (V1, V2 and V3) functional population receptive field (pRF) sizes and structural surface areas using fMRI, and their retinal structure using high-definition optical coherence tomography. With increasing age visual acuity decreased, pRF sizes increased, visual field maps surface areas (but not whole-brain surface areas) decreased, and retinal thickness decreased. Among these measures, only functional pRF sizes predicted perceptual visual acuity, and Bayesian statistics support a null relationship between visual acuity and cortical or retinal structure. However, pRF sizes were in turn predicted by cortical structure only (visual field map surface areas), which were only predicted by retinal structure (thickness). These results suggest that simultaneous disruptions of neural structure and function throughout the early visual system may underlie the deterioration of perceptual visual acuity in healthy aging.

The organizational principles of de-differentiated topographic maps in somatosensory cortex.

Topographic maps are a fundamental feature of cortex architecture in the mammalian brain. One common theory is that the de-differentiation of topographic maps links to impairments in everyday behavior due to less precise functional map readouts. Here, we tested this theory by characterizing de-differentiated topographic maps in primary somatosensory cortex (SI) of younger and older adults by means of ultra-high resolution functional magnetic resonance imaging together with perceptual finger individuation and hand motor performance. Older adults' SI maps showed similar amplitude and size to younger adults' maps, but presented with less representational similarity between distant fingers. Larger population receptive field sizes in older adults' maps did not correlate with behavior, whereas reduced cortical distances between D2 and D3 related to worse finger individuation but better motor performance. Our data uncover the drawbacks of a simple de-differentiation model of topographic map function, and motivate the introduction of feature-based models of cortical reorganization.

Investigating the Reliability of Population Receptive Field Size Estimates Using fMRI.

In functional MRI (fMRI), population receptive field (pRF) models allow a quantitative description of the response as a function of the features of the stimuli that are relevant for each voxel. The most popular pRF model used in fMRI assumes a Gaussian shape in the features space (e.g., the visual field) reducing the description of the voxel’s pRF to the Gaussian mean (the pRF preferred feature) and standard deviation (the pRF size). The estimation of the pRF mean has been proven to be highly reliable. However, the estimate of the pRF size has been shown not to be consistent within and between subjects. While this issue has been noted experimentally, here we use an optimization theory perspective to describe how the inconsistency in estimating the pRF size is linked to an inherent property of the Gaussian pRF model. When fitting such models, the goodness of fit is less sensitive to variations in the pRF size than to variations in the pRF mean. We also show how the same issue can be considered from a bias-variance perspective. We compare different estimation procedures in terms of the reliability of their estimates using simulated and real fMRI data in the visual (using the Human Connectome Project database) and auditory domain. We show that, the reliability of the estimate of the pRF size can be improved considering a linear combination of those pRF models with similar goodness of fit or a permutation based approach. This increase in reliability of the pRF size estimate does not affect the reliability of the estimate of the pRF mean and the prediction accuracy.

Submillimeter fMRI reveals a layout of dorsal visual cortex in macaques, remarkably similar to New World monkeys

The macaque dorsal occipital cortex is generally thought to contain an elongated third visual area, V3d, extending along most of the rostral border of area V2. In contrast, our submillimeter retinotopic fMRI maps (0.6-mm isotropic voxels, achieved by implanted phased-array receive coils) consistently show three sectors anterior to V2d. The dorsal (mirror image) sector complies with the traditional V3d definition, and the middle (nonmirror image) sector with V3A. The ventral (mirror image) sector bends away from V2d, as does the ventrolateral posterior area (VLP) in marmosets and the dorsolateral posterior area (DLP) in owl monkeys, and represents the entire contralateral hemifield as V3A does. Its population-receptive field size, however, suggests that this ventral sector is another area at the same hierarchical level as V4d. Hence, contrary to prevailing views, the retinotopic organization of cortex rostral to V2d differs substantially from widely accepted models. Instead, it is evolutionarily largely conserved in Old and New World monkeys given its surprisingly similar overall visuotopic organization.

Radial asymmetries in population receptive field size and cortical magnification factor in early visual cortex

Human visual cortex does not represent the whole visual field with the same detail. Changes in receptive field size, population receptive field (pRF) size and cortical magnification factor (CMF) with eccentricity are well established, and associated with changes in visual acuity with eccentricity. Visual acuity also changes across polar angle. However, it remains unclear how RF size, pRF size and CMF change across polar angle. Here, we examine differences in pRF size and CMF across polar angle in V1, V2 and V3 using pRF modeling of human fMRI data. In these visual field maps, we find smaller pRFs and larger CMFs in horizontal (left and right) than vertical (upper and lower) visual field quadrants. Differences increase with eccentricity, approximately in proportion to average pRF size and CMF. Similarly, we find larger CMFs in the lower than upper quadrant, and again differences increase with eccentricity. However, pRF size differences between lower and upper quadrants change direction with eccentricity. Finally, we find slightly smaller pRFs in the left than right quadrants of V2 and V3, though this difference is very small, and we find no differences in V1 and no differences in CMF. Moreover, differences in pRF size and CMF vary gradually with polar angle and are not limited to the meridians or visual field map discontinuities. PRF size and CMF differences do not consistently follow patterns of cortical curvature, despite the link between cortical curvature and polar angle in V1. Thus, the early human visual cortex has a radially asymmetric representation of the visual field. These asymmetries may underlie consistent reports of asymmetries in perceptual abilities.

Radial asymmetries in population receptive field size and cortical magnification factor in early visual cortex

Radial asymmetries in population receptive field size and cortical magnification factor in early visual cortex

A second-order orientation-contrast stimulus for population-receptive-field-based retinotopic mapping

Visual field or retinotopic mapping is one of the most frequently used paradigms in fMRI. It uses activity evoked by position-varying high luminance contrast visual patterns presented throughout the visual field for determining the spatial organization of cortical visual areas. While the advantage of using high luminance contrast is that it tends to drive a wide range of neural populations – thus resulting in high signal-to-noise BOLD responses – this may also be a limitation, especially for approaches that attempt to squeeze more information out of the BOLD response, such as population receptive field (pRF) mapping. In that case, more selective stimulation of a subset of neurons – despite reduced signals – could result in better characterization of pRF properties. Here, we used a second-order stimulus based on local differences in orientation texture – to which we refer as orientation contrast – to perform retinotopic mapping. Participants in our experiment viewed arrays of Gabor patches composed of a foreground (a bar) and a background. These could only be distinguished on the basis of a difference in patch orientation. In our analyses, we compare the pRF properties obtained using this new orientation contrast-based retinotopy (OCR) to those obtained using classic luminance contrast-based retinotopy (LCR). Specifically, in higher order cortical visual areas such as LO, our novel approach resulted in non-trivial reductions in estimated population receptive field size of around 30%. A set of control experiments confirms that the most plausible cause for this reduction is that OCR mainly drives neurons sensitive to orientation contrast. We discuss how OCR – by limiting receptive field scatter and reducing BOLD displacement – may result in more accurate pRF localization as well. Estimation of neuronal properties is crucial for interpreting cortical function. Therefore, we conclude that using our approach, it is possible to selectively target particular neuronal populations, opening the way to use pRF modeling to dissect the response properties of more clearly-defined neuronal populations in different visual areas.

The development of population receptive field size in visual cortex during childhood

The development of population receptive field size in visual cortex during childhood

Is the Cortical Deficit in Amblyopia Due to Reduced Cortical Magnification, Loss of Neural Resolution, or Neural Disorganization?

The neural basis of amblyopia, a visual deficit affecting 3% of the human population, remains a matter of debate. We undertook the first population receptive field functional magnetic resonance imaging analysis in participants with amblyopia and compared the projections from the amblyopic and fellow normal eye in the visual cortex. The projection from the amblyopic eye was found to have a normal cortical magnification factor, enlarged population receptive field sizes, and topographic disorganization in all early visual areas. This is consistent with an explanation of amblyopia as an immature system with a normal complement of cells whose spatial resolution is reduced and whose topographical map is disordered. This bears upon a number of competing theories for the psychophysical defect and affects future treatment therapies.

Cortical magnification factor and population receptive field size in human V1 predict the bottom-up saliency map

Cortical magnification factor and population receptive field size in human V1 predict the bottom-up saliency map

Visual cortex organisation in a macaque monkey with macular degeneration

The visual field is retinotopically represented in early visual areas. It has been suggested that when adult primary visual cortex (V1) is deprived of normal retinal input it is capable of large-scale reorganisation, with neurons inside the lesion projection zone (LPZ) being visually driven by inputs from intact retinal regions. Early functional magnetic resonance imaging (fMRI) studies in humans with macular degeneration (MD) report >1cm spread of activity inside the LPZ border, whereas recent results report no shift of the LPZ border. Here, we used fMRI population receptive field measurements to study, for the first time, the visual cortex organisation of one macaque monkey with MD and to compare it with normal controls. Our results showed that the border of the V1 LPZ remained stable, suggesting that the deafferented area V1 zone of the MD animal has limited capacity for reorganisation. Interestingly, the pRF size of non-deafferented V1 voxels increased slightly (~20% on average), although this effect appears weaker than that in previous single-unit recording reports. Area V2 also showed limited reorganisation. Remarkably, area V5/MT of the MD animal showed extensive activation compared to controls stimulated over the part of the visual field that was spared in the MD animal. Furthermore, population receptive field size distributions differed markedly in area V5/MT of the MD animal. Taken together, these results suggest that V5/MT has a higher potential for reorganisation after MD than earlier visual cortex.

Plasticity and Stability of the Visual System in Human Achiasma

The absence of the optic chiasm is an extraordinary and extreme abnormality in the nervous system. The abnormality produces highly atypical functional responses in the cortex, including overlapping hemifield representations and bilateral population receptive fields in both striate and extrastriate visual cortex. Even in the presence of these large functional abnormalities, the effect on visual perception and daily life is not easily detected. Here, we demonstrate that in two achiasmic humans the gross topography of the geniculostriate and occipital callosal connections remains largely unaltered. We conclude that visual function is preserved by reorganization of intracortical connections instead of large-scale reorganizations of the visual cortex. Thus, developmental mechanisms of local wiring within cortical maps compensate for the improper gross wiring to preserve function in human achiasma.

The Relationship between Cortical Magnification Factor and Population Receptive Field Size in Human Visual Cortex: Constancies in Cortical Architecture

Receptive field (RF) sizes and cortical magnification factor (CMF) are fundamental organization properties of the visual cortex. At increasing visual eccentricity, RF sizes increase and CMF decreases. A relationship between RF size and CMF suggests constancies in cortical architecture, as their product, the cortical representation of an RF (point image), may be constant. Previous animal neurophysiology studies of this question yield conflicting results. Here, we use fMRI to determine the relationship between the population RF (pRF) and CMF in humans. In average and individual data, the product of CMF and pRF size, the population point image, is near constant, decreasing slightly with eccentricity in V1. Interhemisphere and subject variations in CMF, pRF size, and V1 surface area are correlated, and the population point image varies less than these properties. These results suggest a V1 cortical processing architecture of approximately constant size between humans. Up the visual hierarchy, to V2, V3, hV4, and LO1, the population point image decreases with eccentricity, and both the absolute values and rate of change increase. PRF sizes increase between visual areas and with eccentricity, but when expressed in V1 cortical surface area (i.e., corticocortical pRFs), they are constant across eccentricity in V2/V3. Thus, V2/V3, and to some degree hV4, sample from a constant extent of V1. This may explain population point image changes in later areas. Consequently, the constant factor determining pRF size may not be the relationship to the local CMF, but rather pRF sizes and CMFs in visual areas from which the pRF samples.

The effects of spatial attention and population receptive field size estimation on fMRI topographic mapping signals

The effects of spatial attention and population receptive field size estimation on fMRI topographic mapping signals

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.

PROJECTION OF THE RETINA ON TO STRIATE AND PRESTRIATE CORTEX IN THE SQUIRREL MONKEY, SAIMIRI SCIUREUS.

PROJECTION OF THE RETINA ON TO STRIATE AND PRESTRIATE CORTEX IN THE SQUIRREL MONKEY, SAIMIRI SCIUREUS.