Functional measurements of human ventral occipital cortex: retinotopy and colour

Human colour vision originates in the cone photoreceptors, whose spatial density peaks in the fovea and declines rapidly into the periphery. For this reason, one expects to find a large representation of the cone-rich fovea in those cortical locations that support colour perception. Human occipital cortex contains several distinct foveal representations including at least two that extend onto the ventral surface: a region thought to be critical for colour vision. To learn more about these ventral signals, we used functional magnetic resonance imaging to identify visual field maps and colour responsivity on the ventral surface. We found a visual map of the complete contralateral hemifield in a 4 cm(2) region adjacent to ventral V3; the foveal representation of this map is confluent with that of areas V1/2/3. Additionally, a distinct foveal representation is present on the ventral surface situated 3-5 cm anterior from the confluent V1/2/3 foveal representations. This organization is not consistent with the definition of area V8, which assumes the presence of a quarter field representation adjacent to V3v. Comparisons of responses to luminance-matched coloured and achromatic patterns show increased activity to the coloured stimuli beginning in area V1 and extending through the new hemifield representation and further anterior in the ventral occipital lobe.     

The two axes of the human eye and inversion of the retinal layers: The basis for the interpretation of the retina as a phase grating optical,cellular 3D chip

The question of why the human eye has two axes, a photopic visual axis, and an eye axis, is just as justified as the one of why the fovea is not on the eye axis, but instead is on the visual axis. An optical engineer would have omitted the second axis and placed the fovea on the eye axis. The answer to the question of why the design of the real eye differs from the logic of the engineer is found in its prenatal development. The biaxial structure was the only possible consequence of the decision to invert the retinal layers. Accordingly, this is of considerable importance. It, in turn, forms the basis of the interpretation of the retina as a cellular 3D phase grating, and can provide a grating-optical interpretation of adaptive effects (Purkinje shift) and aperture phenomena (Stiles-Crawford effects I and II, Bezold-Brücke phenomenon) and visual acuity data in photopic and scotopic vision.     

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.     

Multiple spotlights of attentional selection in human visual cortex

Spatially directed attention strongly enhances visual perceptual processing. The metaphor of the "spotlight" has long been used to describe spatial attention; however, there has been considerable debate as to whether spatial attention must be unitary or may be split between discrete regions of space. This question was addressed here through functional MR imaging of human subjects as they performed a task that required simultaneous attention to two briefly displayed and masked targets at locations separated by distractor stimuli. These data reveal retinotopically specific enhanced activation in striate and extrastriate visual cortical representations of the two attended stimuli and no enhancement at the intervening representation of distractor stimuli. This finding of two spotlights was obtained within a single cortical hemisphere and across the two hemispheres. This provides direct evidence that spatial attention can select, in parallel, multiple low-level perceptual representations.     

The number and distribution of Meynert cells in area 17 of the macaque monkey

A study has been made of the number and distribution of the Meynert cells in area 17 of the visual cortex in the macaque monkey. There are between 30 000 and 40 000 Meynert cells in one hemisphere and these are randomly distributed with an average uniform area density of about 25 cells per square millimetre. The area density of Meynert cells is similar in the representation of the fovea and the perifoveal cortex. These findings are discussed in relation to the basic uniformity of the functional architecture of the visual cortex.     

Features of the Retinotopic Representation in the Visual Wulst of a Laterally Eyed Bird,the Zebra Finch (Taeniopygia guttata)

The visual wulst of the zebra finch comprises at least two retinotopic maps of the contralateral eye. As yet, it is not known how much of the visual field is represented in the wulst neuronal maps, how the organization of the maps is related to the retinal architecture, and how information from the ipsilateral eye is involved in the activation of the wulst. Here, we have used autofluorescent flavoprotein imaging and classical anatomical methods to investigate such characteristics of the most posterior map of the multiple retinotopic representations. We found that the visual wulst can be activated by visual stimuli from a large part of the visual field of the contralateral eye. Horizontally, the visual field representation extended from -5° beyond the beak tip up to +125° laterally. Vertically, a small strip from -10° below to about +25° above the horizon activated the visual wulst. Although retinal ganglion cells had a much higher density around the fovea and along a strip extending from the fovea towards the beak tip, these areas were not overrepresented in the wulst map. The wulst area activated from the foveal region of the ipsilateral eye, overlapped substantially with the middle of the three contralaterally activated regions in the visual wulst, and partially with the other two. Visual wulst activity evoked by stimulation of the frontal visual field was stronger with contralateral than with binocular stimulation. This confirms earlier electrophysiological studies indicating an inhibitory influence of the activation of the ipsilateral eye on wulst activity elicited by stimulating the contralateral eye. The lack of a foveal overrepresentation suggests that identification of objects may not be the primary task of the zebra finch visual wulst. Instead, this brain area may be involved in the processing of visual information necessary for spatial orientation.     

The sense of touch in the star-nosed mole: from mechanoreceptors to the brain

Star-nosed moles are somatosensory specialists that explore their environment with 22 appendages that ring their nostrils. The appendages are covered with sensory domes called Eimer's organs. Each organ is associated with a Merkel cell-neurite complex, a lamellated corpuscle, and a series of 5-10 free nerve endings that form a circle of terminal swellings. Anatomy and electrophysiological recordings suggest that Eimer's organs detect small shapes and textures. There are parallels between the organization of the mole's somatosensory system and visual systems of other mammals. The centre of the star is a tactile fovea used for detailed exploration of objects and prey items. The tactile fovea is over-represented in the neocortex, and this is evident in the modular, anatomically visible representation of the star. Multiple maps of the star are visible in flattened cortical preparations processed for cytochrome oxidase or NADPH-diaphorase. Star-nosed moles are the fastest known foragers among mammals, able to identify and consume a small prey item in 120 ms. Together these behavioural and nervous system specializations have made star-nosed moles an intriguing model system for examining general and specialized aspects of mammalian touch.     

Fovea-Periphery Axis Symmetry of Surround Modulation in the Human Visual System

A visual stimulus activates different sized cortical area depending on eccentricity of the stimulus. Here, our aim is to understand whether the visual field size of a stimulus or cortical size of the corresponding representation determines how strongly it interacts with other stimuli. We measured surround modulation of blood-oxygenation-level-dependent signal and perceived contrast with surrounds that extended either towards the periphery or the fovea from a center stimulus, centered at 6° eccentricity. This design compares the effects of two surrounds which are identical in visual field size, but differ in the sizes of their cortical representations. The surrounds produced equally strong suppression, which suggests that visual field size of the surround determines suppression strength. A modeled population of neuronal responses, in which all the parameters were experimentally fixed, captured the pattern of results both in psychophysics and functional magnetic resonance imaging. Although the fovea-periphery anisotropy affects nearly all aspects of spatial vision, our results suggest that in surround modulation the visual system compensates for it.     

Representation of three-dimensional visual space in the cerebral cortex

This article reviews two issues relevant to the topic of how three-dimensional space is represented in the cerebral cortex. The first is the question of how individual neurons encode information that might contribute to stereoscopic estimation of visual depth. Particular attention is given to the current understanding of the neural representation of motion through three-dimensional space and to the complexities that arise in interpreting neuronal responses to this complex stimulus parameter. The second issue considered is the disorderlines that exists in the retinotopic mapping of the visual field in some cortical visual areas. Several extrastriate areas have been found to contain maps of the contralateral visual hemifield that are disorderly in the sense that the representation of various parts of the visual field are often misplaced or grossly over-or under-represented. It is suggested that this disorderlines may in some cases represent adaptations to facilitate certain types of visual functions.     

Stochastic re-calibration: contextual effects on perceived tilt

The human visual system exaggerates the difference between the tilts of adjacent lines or grating patches. In addition to this tilt illusion, we found that oblique flanks reduced acuity for small changes of tilt in the centre of the visual field. However, no flanks--regardless of their tilts--decreased sensitivity to contrast. Thus, the foveal tilt illusion should not be attributed to orientation-selective lateral inhibition. Nor is it similar to conventional crowding, which typically does not impair letter recognition in the fovea. Our observers behaved as though the reference orientation (horizontal) had a small tilt in the direction of the flanks. We suggest that the extent of this re-calibration varies randomly over trials, and we demonstrate that this stochastic re-calibration can explain flank-induced acuity loss in the fovea.     

Deployment of Spatial Attention towards Locations in Memory Representations. An EEG Study

Recalling information from visual short-term memory (VSTM) involves the same neural mechanisms as attending to an actually perceived scene. In particular, retrieval from VSTM has been associated with orienting of visual attention towards a location within a spatially-organized memory representation. However, an open question concerns whether spatial attention is also recruited during VSTM retrieval even when performing the task does not require access to spatial coordinates of items in the memorized scene. The present study combined a visual search task with a modified, delayed central probe protocol, together with EEG analysis, to answer this question. We found a temporal contralateral negativity (TCN) elicited by a centrally presented go-signal which was spatially uninformative and featurally unrelated to the search target and informed participants only about a response key that they had to press to indicate a prepared target-present vs. -absent decision. This lateralization during VSTM retrieval (TCN) provides strong evidence of a shift of attention towards the target location in the memory representation, which occurred despite the fact that the present task required no spatial (or featural) information from the search to be encoded, maintained, and retrieved to produce the correct response and that the go-signal did not itself specify any information relating to the location and defining feature of the target.     

A nose that looks like a hand and acts like an eye: the unusual mechanosensory system of the star-nosed mole

The star-nosed mole (Condylura cristata) has a snout surrounded by 22 fleshy and mobile appendages. This unusual structure is not an olfactory organ, as might be assumed from its location, nor is it used to manipulate objects as might be guessed from its appearance. Rather, the star is devoted to the sense of touch, and for this purpose the appendages are covered with thousands of small mechanoreceptive Eimer's organs. Recent behavioral studies find that the star acts much like a tactile eye, having a small behavioral focus, or “fovea” at the center – used for detailed explorations of objects of interest. The peripheral and central nervous systems of the mole reflect these behavioral specializations, such that the small behavioral focus on the nose is more densely innervated in the periphery, and has a greatly enlarged representation in the somatosensory cortex. This somatosensory representation of the tactile fovea is not correlated with anatomical parameters (innervation density) as found in other species, but rather is highly correlated with patterns of behavior. The many surprising parallels between the somatosensory system of the mole, and the visual systems of other mammals, suggest a convergent and perhaps common organization for highly developed sensory systems. Accepted: 31 May 1999     

A star in the brainstem reveals the first step of cortical magnification

A fundamental question in the neurosciences is how central nervous system (CNS) space is allocated to different sensory inputs. Yet it is difficult to measure innervation density and corresponding representational areas in the CNS of most species. These measurements can be made in star-nosed moles (Condylura cristata) because the cortical representation of nasal rays is visible in flattened sections and afferents from each ray can be counted. Here we used electrophysiological recordings combined with sections of the brainstem to identify a large, visible star representation in the principal sensory nucleus (PrV). PrV was greatly expanded and bulged out of the brainstem rostrally to partially invade the trigeminal nerve. The star representation was a distinct PrV subnucleus containing 11 modules, each representing one of the nasal rays. The 11 PrV ray representations were reconstructed to obtain volumes and the largest module corresponded to ray 11, the mole's tactile fovea. These measures were compared to fiber counts and primary cortical areas from a previous investigation. PrV ray volumes were closely correlated with the number of afferents from each ray, but afferents from the behaviorally most important, 11(th) ray were preferentially over-represented. This over-representation at the brainstem level was much less than at the cortical level. Our results indicate that PrV provides the first step in magnifying CNS representations of important afferents, but additional magnification occurs at higher levels. The early development of the 11(th), foveal appendage could provide a mechanism for the most important afferents to capture the most CNS space.     

A novel interhemispheric interaction: modulation of neuronal cooperativity in the visual areas

Background

The cortical representation of the visual field is split along the vertical midline, with the left and the right hemi-fields projecting to separate hemispheres. Connections between the visual areas of the two hemispheres are abundant near the representation of the visual midline. It was suggested that they re-establish the functional continuity of the visual field by controlling the dynamics of the responses in the two hemispheres.

Methods/Principal Findings

To understand if and how the interactions between the two hemispheres participate in processing visual stimuli, the synchronization of responses to identical or different moving gratings in the two hemi-fields were studied in anesthetized ferrets. The responses were recorded by multiple electrodes in the primary visual areas and the synchronization of local field potentials across the electrodes were analyzed with a recent method derived from dynamical system theory. Inactivating the visual areas of one hemisphere modulated the synchronization of the stimulus-driven activity in the other hemisphere. The modulation was stimulus-specific and was consistent with the fine morphology of callosal axons in particular with the spatio-temporal pattern of activity that axonal geometry can generate.

Conclusions/Significance

These findings describe a new kind of interaction between the cerebral hemispheres and highlight the role of axonal geometry in modulating aspects of cortical dynamics responsible for stimulus detection and/or categorization.     

Whisker Movements Reveal Spatial Attention: A Unified Computational Model of Active Sensing Control in the Rat

Spatial attention is most often investigated in the visual modality through measurement of eye movements, with primates, including humans, a widely-studied model. Its study in laboratory rodents, such as mice and rats, requires different techniques, owing to the lack of a visual fovea and the particular ethological relevance of orienting movements of the snout and the whiskers in these animals. In recent years, several reliable relationships have been observed between environmental and behavioural variables and movements of the whiskers, but the function of these responses, as well as how they integrate, remains unclear. Here, we propose a unifying abstract model of whisker movement control that has as its key variable the region of space that is the animal''s current focus of attention, and demonstrate, using computer-simulated behavioral experiments, that the model is consistent with a broad range of experimental observations. A core hypothesis is that the rat explicitly decodes the location in space of whisker contacts and that this representation is used to regulate whisker drive signals. This proposition stands in contrast to earlier proposals that the modulation of whisker movement during exploration is mediated primarily by reflex loops. We go on to argue that the superior colliculus is a candidate neural substrate for the siting of a head-centred map guiding whisker movement, in analogy to current models of visual attention. The proposed model has the potential to offer a more complete understanding of whisker control as well as to highlight the potential of the rodent and its whiskers as a tool for the study of mammalian attention.     

Simulated bipolar cells in fovea of human retina

This static bipolar cell (BC) model of the human fovea is based on a number of reasonable assumptions. The human fovea is directly responsible for visual acuity and color vision. The fovea can be considered as having two parts; a central fovea with only red- and green-sensitive cones and a parafovea with blue-sensitive cones added to the other two. A cone mosaic can be precisely organized spatially into unit hexagons that specify inputs to horizontal cells (HC) and BCs. The retina up to and including BCs is piece-wise linear, i.e. at a given steady-state adapting light intensity BC outputs are linear functions of the physical image. BC centers receive inputs directly from weighted cones, while antagonistic surrounds receive inverted inputs from HCs. Appropriate optical and chromatic filtering due to the eye that are taken from human data are incorporated into the model. Chromatic aberrations are simulated by three separate point spread functions that also are taken from human data. Automatic gain control of cones is a function of intensity and wavelength of the steady adapting light.The major part of this work was done while the author was a Senior Research Associate of the National Research Council, USA     

Foveation period and waveforms of congenital ocular nystagmus

Normal visual acuity requires a stationary retinal image on the fovea. If fixation instabilities cause movement of the retinal image across the fovea for a few degrees, visual acuity is diminished. Nystagmus as the fixation instability, consequently, may impair vision. Period of foveation is the area in the wave form, i.e. a brief period of time when the eye is still and is pointed at the object of regard. At this period eye velocity is at a minimum and visual acuity is the best. In the children with congenital ocular nystagmus, using usual clinical equipment (TC 1.0 and TC 0.3 s), was performed electronystagmography (ENG) and analysis of the obtained nystagmus waveforms. In the some patients visual acuity was also examined. The ENG records were classified according to Dell'Osso criteria for waveforms. The findings of jerk nystagmus with extended foveation (J(EF)) and of bidirectional jerk nystagmus (BDJ) were singled out. Foveation time, measured in these weveforms was compared with the visual acuity. Visual acuity was better in the jerk nystafmus weveforms with extended foveation period (J(EF)) than in bidirectional jerk nystagmus with shorter foveation time.     

Auditory fovea and Doppler shift compensation: adaptations for flutter detection in echolocating bats using CF-FM signals

Rhythmical modulations in insect echoes caused by the moving wings of fluttering insects are behaviourally relevant information for bats emitting CF-FM signals with a high duty cycle. Transmitter and receiver of the echolocation system in flutter detecting foragers are especially adapted for the processing of flutter information. The adaptations of the transmitter are indicated by a flutter induced increase in duty cycle, and by Doppler shift compensation (DSC) that keeps the carrier frequency of the insect echoes near a reference frequency. An adaptation of the receiver is the auditory fovea on the basilar membrane, a highly expanded frequency representation centred to the reference frequency. The afferent projections from the fovea lead to foveal areas with an overrepresentation of sharply tuned neurons with best frequencies near the reference frequency throughout the entire auditory pathway. These foveal neurons are very sensitive to stimuli with natural and simulated flutter information. The frequency range of the foveal areas with their flutter processing neurons overlaps exactly with the frequency range where DS compensating bats most likely receive echoes from fluttering insects. This tight match indicates that auditory fovea and DSC are adaptations for the detection and evaluation of insects flying in clutter.     

One picture is worth at least a million neurons

How many neurons participate in the representation of a single visual image? Answering this question is critical for constraining biologically inspired models of object recognition, which vary greatly in their assumptions from few "grandmother cells" to numerous neurons in widely distributed networks. Functional imaging techniques, such as fMRI, provide an opportunity to explore this issue, since they allow the simultaneous detection of the entire neuronal population responding to each stimulus. Several studies have shown that fMRI BOLD signal is approximately proportional to neuronal activity. However, since it provides an indirect measure of this activity, obtaining a realistic estimate of the number of activated neurons requires several intervening steps. Here, we used the extensive knowledge of primate V1 to yield a conservative estimate of the ratio between hemodynamic response and neuronal firing. This ratio was then used, in addition to several cautious assumptions, to assess the number of neurons responding to a single-object image in the entire visual cortex and particularly in object-related areas. Our results show that at least a million neurons in object-related cortex and about two hundred million neurons in the entire visual cortex are involved in the representation of a single-object image.     

The effect of a moving foveal target on the subjective sensation of motion

In this paper, we report on two experiments concerning the effect of the visual field of fovea on the subjective estimation of angular velocity. Experiment 1 investigates the effect of a slow moving target on the perception of self motion. The result of this experiment can be summarized as follows: a slow moving target seen in the visual field of fovea by a stationary person generates in this person a sensation of self rotation in the same direction as the motion of the target. This phenomenon will be called foveal induced ego motion. Experiment 2 investigates the latency for the detection of a self angular acceleration when the person focusses his fovea on a slowly moving target. From the results of this experiment we conclude that the latency for detection of a small self angular acceleration is shorter if the person sees a small foveal target moving with respect to the person in the direction of self rotation than if that small foveal target is moving (with respect to the person) in the opposite direction. The results of these experiments help us in refining existing models of visual-vestibular interaction, by providing a model which accounts for the phenomenon of oculogyral illusion.This research was conducted while serving as a Visiting Professor at the Man Vehicle Laboratory, Department of Aeronautics and Astronautics, Massachusetts Institute of Technology, Cambridge, MA 02139, USA