Independent component analysis of natural image sequences yields spatio-temporal filters similar to simple cells in primary visual cortex.

Simple cells in the primary visual cortex process incoming visual information with receptive fields localized in space and time, bandpass in spatial and temporal frequency, tuned in orientation, and commonly selective for the direction of movement. It is shown that performing independent component analysis (ICA) on video sequences of natural scenes produces results with qualitatively similar spatio-temporal properties. Whereas the independent components of video resemble moving edges or bars, the independent component filters, i.e. the analogues of receptive fields, resemble moving sinusoids windowed by steady Gaussian envelopes. Contrary to earlier ICA results on static images, which gave only filters at the finest possible spatial scale, the spatio-temporal analysis yields filters at a range of spatial and temporal scales. Filters centred at low spatial frequencies are generally tuned to faster movement than those at high spatial frequencies.     

Do Male and Female Cowbirds See Their World Differently? Implications for Sex Differences in the Sensory System of an Avian Brood Parasite

Background

Male and female avian brood parasites are subject to different selection pressures: males compete for mates but do not provide parental care or territories and only females locate hosts to lay eggs. This sex difference may affect brain architecture in some avian brood parasites, but relatively little is known about their sensory systems and behaviors used to obtain sensory information. Our goal was to study the visual resolution and visual information gathering behavior (i.e., scanning) of brown-headed cowbirds.

Methodology/Principal Findings

We measured the density of single cone photoreceptors, associated with chromatic vision, and double cone photoreceptors, associated with motion detection and achromatic vision. We also measured head movement rates, as indicators of visual information gathering behavior, when exposed to an object. We found that females had significantly lower density of single and double cones than males around the fovea and in the periphery of the retina. Additionally, females had significantly higher head-movement rates than males.

Conclusions

Overall, we suggest that female cowbirds have lower chromatic and achromatic visual resolution than males (without sex differences in visual contrast perception). Females might compensate for the lower visual resolution by gazing alternatively with both foveae in quicker succession than males, increasing their head movement rates. However, other physiological factors may have influenced the behavioral differences observed. Our results bring up relevant questions about the sensory basis of sex differences in behavior. One possibility is that female and male cowbirds differentially allocate costly sensory resources, as a recent study found that females actually have greater auditory resolution than males.     

The Gaussian derivative model for spatial-temporal vision: I. Cortical model

How do we see the motion of objects as well as their shapes? The Gaussian Derivative (GD) spatial model is extended to time to help answer this question. The GD spatio-temporal model requires only two numbers to describe the complete three-dimensional space-time shapes of individual receptive fields in primate visual cortex. These two numbers are the derivative numbers along the respective spatial and temporal principal axes of a given receptive field. Nine transformation parameters allow for a standard geometric association of these intrinsic axes with the extrinsic environment. The GD spatio-temporal model describes in one framework the following properties of primate simple cell fields: motion properties, number of lobes in space-time, spatial orientation. location, and size. A discrete difference-of-offset-Gaussians (DOOG) model provides a plausible physiological mechanism to form GD-like model fields in both space and time. The GD model hypothesizes that receptive fields at the first stage of processing in the visual cortex approximate 'derivative analyzers' that estimate local spatial and temporal derivatives of the intensity profile in the visual environment. The receptive fields as modeled provide operators that can allow later stages of processing in either a biological or machine vision system to estimate the motion as well as the shapes of objects in the environment.     

THE CORTICAL MAGNIFICATION FACTOR AND PHOTOPIC VISION

1. The concept of a topographical representation of sensory surfaces on the mammalian cerebral cortex is now well established. Furthermore, there is claimed an expanded representation for those sensory surfaces associated with behavioural specializations.
2. In the case of the visual system, the field of view associated with the retina is projected on to the striate cortex, with the central fovea or point of most acute vision occupying a disproportionately large area of representation.
3. The term 'cortical magnification factor', denoted by M, has been introduced to indicate, for a given eccentricity and meridian, the linear distance in mm along the primary visual cortex concerned with each degree of visual field.
4. The quantitative accuracy of this function subsequently has been improved and a relationship established between cortical magnification and visual acuity in man.
5. On the basis of spatio-temporal investigations across the visual field under levels of photopic adaptation, visual scientists have utilized retino-cortical magnification to reconcile aspects of structure and function in the human visual system.     

SPATIAL FLOWER PARAMETERS AND INSECT SPATIAL VISION

The present article reviews recent and older literature on the spatial parameters that flowers display, as well as on the capacities of anthophilous insects to perceive and use these parameters for optimizing their foraging success. Although co-evolution of plants and pollinators has frequently been discussed with respect to floral colours and insect colour vision, it has rarely been assessed with respect to insect spatial vision and spatial floral cues, such as shape, pattern, size, contrast, symmetry, spatial frequency, contour density and orientation of contours. This review is an attempt to fill this gap. From experimental findings and observations on both flowers and insects, we arrive at the conclusion that all of the spatial and spatio-temporal parameters that flowers offer are relevant to the foraging task and are tuned to the insect's visual capacities and visually guided behaviour. We try, in addition, to indicate that temporal cues are closely related to spatial cues, and must therefore be included when flower–pollinator interactions are examined. We include results that show that colour vision and spatial vision have diverged over the course of evolution, particularly regarding the processing of spatio-temporal information, but that colour vision plays a role in the processing of spatial cues that are independent of temporal parameters. By presenting this review we hope to contribute to closer collaboration among scientists working in the vast fields of botany, ecology, evolution, ethology and sensory physiology.     

Knowing with which eye we see: utrocular discrimination and eye-specific signals in human visual cortex

Neurophysiological and behavioral reports converge to suggest that monocular neurons in the primary visual cortex are biased toward low spatial frequencies, while binocular neurons favor high spatial frequencies. Here we tested this hypothesis with functional magnetic resonance imaging (fMRI). Human participants viewed flickering gratings at one of two spatial frequencies presented to either the left or the right eye, and judged which of the two eyes was being stimulated (utrocular discrimination). Using multivoxel pattern analysis we found that local spatial patterns of signals in primary visual cortex (V1) allowed successful decoding of the eye-of-origin. Decoding was above chance for low but not high spatial frequencies, confirming the presence of a bias reported by animal studies in human visual cortex. Behaviorally, we found that reliable judgment of the eye-of-origin did not depend on spatial frequency. We further analyzed the mean response in visual cortex to our stimuli and revealed a weak difference between left and right eye stimulation. Our results are thus consistent with the interpretation that participants use overall levels of neural activity in visual cortex, perhaps arising due to local luminance differences, to judge the eye-of-origin. Taken together, we show that it is possible to decode eye-specific voxel pattern information in visual cortex but, at least in healthy participants with normal binocular vision, these patterns are unrelated to awareness of which eye is being stimulated.     

Visual advantage in deaf adults linked to retinal changes

The altered sensory experience of profound early onset deafness provokes sometimes large scale neural reorganisations. In particular, auditory-visual cross-modal plasticity occurs, wherein redundant auditory cortex becomes recruited to vision. However, the effect of human deafness on neural structures involved in visual processing prior to the visual cortex has never been investigated, either in humans or animals. We investigated neural changes at the retina and optic nerve head in profoundly deaf (N = 14) and hearing (N = 15) adults using Optical Coherence Tomography (OCT), an in-vivo light interference method of quantifying retinal micro-structure. We compared retinal changes with behavioural results from the same deaf and hearing adults, measuring sensitivity in the peripheral visual field using Goldmann perimetry. Deaf adults had significantly larger neural rim areas, within the optic nerve head in comparison to hearing controls suggesting greater retinal ganglion cell number. Deaf adults also demonstrated significantly larger visual field areas (indicating greater peripheral sensitivity) than controls. Furthermore, neural rim area was significantly correlated with visual field area in both deaf and hearing adults. Deaf adults also showed a significantly different pattern of retinal nerve fibre layer (RNFL) distribution compared to controls. Significant correlations between the depth of the RNFL at the inferior-nasal peripapillary retina and the corresponding far temporal and superior temporal visual field areas (sensitivity) were found. Our results show that cross-modal plasticity after early onset deafness may not be limited to the sensory cortices, noting specific retinal adaptations in early onset deaf adults which are significantly correlated with peripheral vision sensitivity.     

Spatio-temporal visual properties in the ascending tectofugal system

Our study compares the spatio-temporal visual receptive field properties of different subcortical stages of the ascending tectofugal visual system. Extracellular single-cell recordings were performed in the superficial (SCs) and intermediate (SCi) layers of the superior colliculus (SC), the suprageniculate nucleus (Sg) of the posterior thalamus and the caudate nucleus (CN) of halothane-anesthetized cats. Neuronal responses to drifting gratings of various spatial and temporal frequencies were recorded. The neurons of each structure responded optimally to low spatial and high temporal frequencies and displayed narrow spatial and temporal frequency tuning. The detailed statistical analysis revealed that according to its stimulus preferences the SCs has markedly different spatio-temporal properties from the homogeneous group formed by the SCi, Sg and CN. The SCs neurons preferred higher spatial and lower temporal frequencies and had broader spatial tuning than the other structures. In contrast to the SCs the visually active SCi, as well as the Sg and the CN neurons possessed consequently similar spatio-temporal preferences. These data support our hypothesis that the visually active SCi, Sg and CN neurons form a homogeneous neuronal population given a similar spatio-temporal frequency preference and a common function in processing of dynamic visual information.     

Spatial and temporal dependencies of cross-orientation suppression in human vision

A well-known property of orientation-tuned neurons in the visual cortex is that they are suppressed by the superposition of an orthogonal mask. This phenomenon has been explained in terms of physiological constraints (synaptic depression), engineering solutions for components with poor dynamic range (contrast normalization) and fundamental coding strategies for natural images (redundancy reduction). A common but often tacit assumption is that the suppressive process is equally potent at different spatial and temporal scales of analysis. To determine whether it is so, we measured psychophysical cross-orientation masking (XOM) functions for flickering horizontal Gabor stimuli over wide ranges of spatio-temporal frequency and contrast. We found that orthogonal masks raised contrast detection thresholds substantially at low spatial frequencies and high temporal frequencies (high speeds), and that small and unexpected levels of facilitation were evident elsewhere. The data were well fit by a functional model of contrast gain control, where (i) the weight of suppression increased with the ratio of temporal to spatial frequency and (ii) the weight of facilitatory modulation was the same for all conditions, but outcompeted by suppression at higher contrasts. These results (i) provide new constraints for models of primary visual cortex, (ii) associate XOM and facilitation with the transient magno- and sustained parvostreams, respectively, and (iii) reconcile earlier conflicting psychophysical reports on XOM.     

Aerial and aquatic visual acuity of the grey bichir Polypterus senegalus,as estimated by optokinetic response

The present study assessed the aerial and aquatic visual abilities of juvenile grey bichir Polypterus senegalus, fish capable of terrestrial locomotion, by measuring the optokinetic response to stimuli of varying speed and spatial frequency. In water, fish tracked slow-moving (2° s⁻¹) stimuli moderately well and fast-moving stimuli very poorly. Spatial acuity was very low compared with many other species, with maximum response observed at 0.05–0.075 stimulus cycles per degree of visual arc; however, it should be noted that adult fish, with their larger eyes, are likely to have somewhat improved spatial acuity. Low spatial acuity and limited stimulus tracking ability might be expected in a nocturnal ambush predator such as P. senegalus, where gaze stabilization may be less crucial and other sensory inputs may have greater importance in perception of the environment. In air, spatial and temporal acuity were both poorer by every measure, but some visual ability persisted. As the eye shows no anatomical specialization for aerial vision, poor vision was expected; however, the large decrease in saccade velocity observed in air trials was unexpected. Stimulus parameters typically have little effect on the characteristics of the saccade, so this finding may suggest that the function of the reflex system itself could be compromised in the aerial vision of some fishes capable of terrestrial locomotion.     

Spatial contrast sensitivity of birds

Contrast sensitivity (CS) is the ability of the observer to discriminate between adjacent stimuli on the basis of their differences in relative luminosity (contrast) rather than their absolute luminances. In previous studies, using a narrow range of species, birds have been reported to have low contrast detection thresholds relative to mammals and fishes. This was an unexpected finding because birds had been traditionally reported to have excellent visual acuity and color vision. This study reports CS in six species of birds that represent a range of visual adaptations to varying environments. The species studied were American kestrels (Falco sparverius), barn owls (Tyto alba), Japanese quail (Coturnix coturnix japonica), white Carneaux pigeons (Columba livia), starlings (Sturnus vulgaris), and red-bellied woodpeckers (Melanerpes carolinus). Contrast sensitivity functions (CSFs) were obtained from these birds using the pattern electroretinogram and compared with CSFs from the literature when possible. All of these species exhibited low CS relative to humans and most mammals, which suggests that low CS is a general characteristic of birds. Their low maximum CS may represent a trade-off of contrast detection for some other ecologically vital capacity such as UV detection or other aspects of their unique color vision.     

Prosimian sensory capacities

This paper reviews laboratory research on sensory capacities of prosimians. Most investigations to date have concentrated on visual and auditory sensitivity with relatively little attention directed toward olfactory and gustatory sensitivity. Diurnal prosimians have at least rudimentary color vision and appear deficient to anthropoids in other visual measures such as acuity. The most striking difference between prosimian and anthropoid auditory sensitivity is the increased sensitivity of the former group to high frequencies. No psychophysical data have been published on olfactory sensitivity of prosimians, but the relatively large olfactory areas in their brains and their propensity for scent marking strongly suggest that they have acute olfactory sensitivity. Most data on gustatory thresholds in prosimians does not permit sensitivity estimation since the data are joint functions of sensitivity as well as preferences.     

Interactive coding of visual spatial frequency and auditory amplitude-modulation rate

Spatial frequency is a fundamental visual feature coded in primary visual cortex, relevant for perceiving textures, objects, hierarchical structures, and scenes, as well as for directing attention and eye movements. Temporal amplitude-modulation (AM) rate is a fundamental auditory feature coded in primary auditory cortex, relevant for perceiving auditory objects, scenes, and speech. Spatial frequency and temporal AM rate are thus fundamental building blocks of visual and auditory perception. Recent results suggest that crossmodal interactions are commonplace across the primary sensory cortices and that some of the underlying neural associations develop through consistent multisensory experience such as audio-visually perceiving speech, gender, and objects. We demonstrate that people consistently and absolutely (rather than relatively) match specific auditory AM rates to specific visual spatial frequencies. We further demonstrate that this crossmodal mapping allows amplitude-modulated sounds to guide attention to and modulate awareness of specific visual spatial frequencies. Additional results show that the crossmodal association is approximately linear, based on physical spatial frequency, and generalizes to tactile pulses, suggesting that the association develops through multisensory experience during manual exploration of surfaces.     

Feedback contributions to visual awareness in human occipital cortex

It has traditionally been assumed that processing within the visual system proceeds in a bottom-up, feedforward manner from retina to higher cortical areas. In addition to feedforward processing, it is now clear that there are also important contributions to sensory encoding that rely upon top-down, feedback (reentrant) projections from higher visual areas to lower ones. By utilizing transcranial magnetic stimulation (TMS) in a metacontrast masking paradigm, we addressed whether feedback processes in early visual cortex play a role in visual awareness. We show that TMS of visual cortex, when timed to produce visual suppression of an annulus serving as a metacontrast mask, induces recovery of an otherwise imperceptible disk. In addition to producing disk recovery, TMS suppression of an annulus was greater when a disk preceded it than when an annulus was presented alone. This latter result suggests that there are effects of the disk on the perceptibility of the subsequent mask that are additive and are revealed with TMS of the visual cortex. These results demonstrate spatial and temporal interactions of conscious vision in visual cortex and suggest that a prior visual stimulus can influence subsequent perception at early stages of visual encoding via feedback projections.     

Luminance-dependence of spatial vision in budgerigars (<Emphasis Type="Italic">Melopsittacus undulatus</Emphasis>) and Bourke’s parrots (<Emphasis Type="Italic">Neopsephotus bourkii</Emphasis>)

Budgerigars (Melopsittacus undulatus) and Bourke’s parrots (Neopsephotus bourkii) are closely related birds with different activity patterns. Budgerigars are strictly diurnal while Bourke’s parrots are active in dim twilight. Earlier studies show that the intensity threshold of colour vision is similar in both species while Bourke’s parrots have larger eyes with a higher density of rods than budgerigars. In this study, we investigate whether this could be an adaptation for better spatial vision in dim light. We used two alternative forced-choice experiments to determine the spatial acuity of both species at light intensities ranging from 0.08 to 73 cd/m². We also determined the spatial contrast sensitivity function (CSF) for bright light in Bourke’s parrots and compare it to existing data for budgerigars. The spatial acuity of Bourke’s parrots was found to be similar to that of budgerigars at all light levels. Also the CSF of Bourke’s parrots is similar to that of budgerigars with a sensitivity peak located between 2.1 and 2.6 cycles/degree. Our findings do not support the hypothesis that Bourke’s parrots have superior spatial acuity in dim light compared to budgerigars and the adaptive value of the relatively rod-rich and large eyes of Bourke’s parrots remains unclear.     

Visual acuity, contrast sensitivity and retinal magnification in a marsupial, the tammar wallaby (Macropus eugenii )

The visual acuity of the tammar wallaby was estimated using a behavioural discrimination task. The wallabies were trained to discriminate a high-contrast (86%) square-wave grating from a grey field of equal luminance (1000–6000 cd m⁻²). Visual-evoked cortical potentials were used to measure the complete contrast sensitivity function. The stimulus was a sinusoidal phase reversal of a sinusoidally modulated grating of various spatial frequencies and contrasts with a mean luminance of 40 cd m⁻². The behavioural acuity was estimated to be about 4.8 cycles/deg. The contrast sensitivity peaked at about 0.15 cycles/deg and declined towards both lower and higher spatial frequencies. The cut-off frequency of the contrast sensitivity function is slightly lower than the behaviourally measured acuity at about 2.7 cycles/deg. The retinal magnification factor was estimated anatomically from laser lesions to be about 0.16 mm/deg. Based on the known ganglion cell density and the retinal magnification factor, an anatomical upper limit to visual acuity of about 6 cycles/deg can be calculated. The differences in estimates of visual acuity between the behavioural and anatomical methods on the one side and physiology on the other side are discussed. Accepted: 28 May 1998     

Vision and cortical map development

Functional maps arise in developing visual cortex as response selectivities become organized into columnar patterns of population activity. Recent studies of developing orientation and direction maps indicate that both are sensitive to visual experience, but not to the same degree or duration. Direction maps have a greater dependence on early vision, while orientation maps remain sensitive to experience for a longer period of cortical maturation. There is also a darker side to experience: abnormal vision through closed lids produces severe impairments in neuronal selectivity, rendering these maps nearly undetectable. Thus, the rules that govern their formation and the construction of the underlying neural circuits are modulated-for better or worse-by early vision. Direction maps, and possibly maps of other properties that are dependent upon precise conjunctions of spatial and temporal signals, are most susceptible to the potential benefits and maladaptive consequences of early sensory experience.     

Contrast transfer function of the visual system]

Visually evoked potentials were used to determine the spatial contrast response function of the visual system and the visual acuity of the pigeon. The spatial contrast response describes the relationship between the contrast in a pattern of vertical stripes, whose luminance is a function of position, and the amplitude of the visually evoked response at various spatial frequencies for a given temporal frequency (pattern reversal frequency); it indicates how particular spatial frequencies are attenuated in the visual system. The visually evoked responses were recorded using monopolar stainless steel electrodes inserted into the stratum griseum superficiale of the optic tectum; the depth of penetration was determined on the basis of a stereotactic atlas. The stimulus patterns were generated on a video monitor placed 75 cm in front of the animal's eye perpendicular to the optic axis. The spatial contrast response function measured at 10% contrast and 0.5 Hz reversal frequency shows a peak at a spatial frequency of 0.5 c/deg, corresponding to 1 degree of visual angle, and decreases progressively at higher spatial frequencies. The high-frequency limit (cut-off frequency) for resolution of sinusoidal gratings, estimated from the contrast response function, is 15.5 c/deg, corresponding to a visual acuity of 1.9 min of arc.     

Modality-specific control of strategic spatial attention in parietal cortex

The neural basis of selective spatial attention presents a significant challenge to cognitive neuroscience. Recent neuroimaging studies have suggested that regions of the parietal and temporal cortex constitute a "supramodal" network that mediates goal-directed attention in multiple sensory modalities. Here we used transcranial magnetic stimulation (TMS) to determine which cortical subregions control strategic attention in vision and touch. Healthy observers undertook an orienting task in which a central arrow cue predicted the location of a subsequent visual or somatosensory target. To determine the attentional role of cortical subregions at different stages of processing, TMS was delivered to the right hemisphere during cue or target events. Results indicated a critical role of the inferior parietal cortex in strategic orienting to visual events, but not to somatosensory events. These findings are inconsistent with the existence of a supramodal attentional network and instead provide direct evidence for modality-specific attentional processing in parietal cortex.