Functional evidence for cone-specific connectivity in the human retina

NoPhysiological studies of colour vision have not yet resolved the controversial issue of how chromatic opponency is constructed at a neuronal level. Two competing theories, the cone-selective hypothesis and the random-wiring hypothesis, are currently equivocal to the architecture of the primate retina. In central vision, both schemes are capable of producing colour opponency due to the fact that receptive field centres receive input from a single bipolar cell ¿ the so called `private line arrangement¿. However, in peripheral vision this single-cone input to the receptive field centre is lost, so that any random cone connectivity would result in a predictable reduction in the quality of colour vision. Behavioural studies thus far have indeed suggested a selective loss of chromatic sensitivity in peripheral vision. We investigated chromatic sensitivity as a function of eccentricity for the cardinal chromatic (L/M and S/(L + M)) and achromatic (L + M) pathways, adopting stimulus size as the critical variable. Results show that performance can be equated across the visual field simply by a change of scale (size). In other words, there exists no qualitative loss of chromatic sensitivity across the visual field. Critically, however, the quantitative nature of size dependency for each of the cardinal chromatic and achromatic mechanisms is very specific, reinforcing their independence in terms of anatomy and genetics. Our data provide clear evidence for a physiological model of primate colour vision that retains chromatic quality in peripheral vision, thus supporting the cone-selective hypothesis

Global shape aftereffects in composite radial frequency patterns

YesIndividual radial frequency (RF) patterns are generated by modulating a circle's radius as a sinusoidal function of polar angle and have been shown to tap into global shape processing mechanisms. Composite RF patterns can reproduce the complex outlines of natural shapes and examining these stimuli may allow us to interrogate global shape mechanisms that are recruited in biologically relevant tasks. We present evidence for a global shape aftereffect in a composite RF pattern stimulus comprising two RF components. Manipulations of the shape, location, size and spatial frequency of the stimuli revealed that this aftereffect could only be explained by the attenuation of intermediate-level global shape mechanisms. The tuning of the aftereffect to test stimulus size also revealed two mechanisms underlying the aftereffect; one that was tuned to size and one that was invariant. Finally, we show that these shape mechanisms may encode some RF information. However, the RF encoding we found was not capable of explaining the full extent of the aftereffect, indicating that encoding of other shape features such as curvature are also important in global shape processing.This research was supported by a Biotechnology and Biological Sciences Research Council (BBSRC) grant #BB/L007770/1

Rod Electroretinograms Elicited by Silent Substitution Stimuli from the Light-Adapted Human Eye

YesPurpose: To demonstrate that silent substitution stimuli can be used to generate electroretinograms (ERGs) that effectively isolate rod photoreceptor function in humans without the need for dark adaptation, and that this approach constitutes a viable alternative to current clinical standard testing protocols. Methods: Rod-isolating and non-isolating sinusoidal flicker stimuli were generated on a 4 primary light-emitting diode (LED) Ganzfeld stimulator to elicit ERGs from participants with normal and compromised rod function who had not undergone dark-adaptation. Responses were subjected to Fourier analysis, and the amplitude and phase of the fundamental were used to examine temporal frequency and retinal illuminance response characteristics. Results: Electroretinograms elicited by rod-isolating silent substitution stimuli exhibit low-pass temporal frequency response characteristics with an upper response limit of 30 Hz. Responses are optimal between 5 and 8 Hz and between 10 and 100 photopic trolands (Td). There is a significant correlation between the response amplitudes obtained with the silent substitution method and current standard clinical protocols. Analysis of signal-to-noise ratios reveals significant differences between subjects with normal and compromised rod function. Conclusions: Silent substitution provides an effective method for the isolation of human rod photoreceptor function in subjects with normal as well as compromised rod function when stimuli are used within appropriate parameter ranges. Translational Relevance: This method of generating rod-mediated ERGs can be achieved without time-consuming periods of dark adaptation, provides improved isolation of rod- from cone-based activity, and will lead to the development of faster clinical electrophysiologic testing protocols with improved selectivity

A Direct Demonstration of Functional Differences between Subdivisions of Human V5/MT

Two subdivisions of human V5/MT+: one located posteriorly (MT/TO-1) and the other more anteriorly (MST/TO-2) were identified in human participants using functional magnetic resonance imaging on the basis of their representations of the ipsilateral versus contralateral visual field. These subdivisions were then targeted for disruption by the application of repetitive transcranial magnetic stimulation (rTMS). The rTMS was delivered to cortical areas while participants performed direction discrimination tasks involving 3 different types of moving stimuli defined by the translational, radial, or rotational motion of dot patterns. For translational motion, performance was significantly reduced relative to baseline when rTMS was applied to both MT/TO-1 and MST/TO-2. For radial motion, there was a differential effect between MT/TO-1 and MST/TO-2, with only disruption of the latter area affecting performance. The rTMS failed to reveal a complete dissociation between MT/TO-1 and MST/TO-2 in terms of their contribution to the perception of rotational motion. On the basis of these results, MT/TO-1 and MST/TO-2 appear to be functionally distinct subdivisions of hV5/MT+. While both areas appear to be implicated in the processing of translational motion, only the anterior region (MST/TO-2) makes a causal contribution to the perception of radial motion

An enhanced role for right hV5/MT+ in the analysis of motion in the contra- and ipsi-lateral visual hemi-fields

Previous experiments have demonstrated that transcranial magnetic stimulation (TMS) of human V5/MT+, in either the left or right cerebral hemisphere, can induce deficits in visual motion perception in their respective contra- and ipsi-lateral visual hemi-fields. However, motion deficits in the ipsi-lateral hemi-field are greater when TMS is applied to V5/MT + in the right hemisphere relative to the left hemisphere. One possible explanation for this asymmetry might lie in differential stimulation of sub-divisions within V5/MT + across the two hemispheres. V5/MT + has two major sub-divisions; MT/TO-1 and MST/TO-2, the latter area contains neurons with large receptive fields (RFs) that extend up to 15° further into the ipsi-lateral hemi-field than the former. We wanted to examine whether applying TMS to MT/TO-1 and MST/TO-2 separately could explain the previously reported functional asymmetries for ipsi-lateral motion processing in V5/MT + across right and left cerebral hemispheres. MT/TO-1 and MST/TO-2 were identified in seven subjects using fMRI localisers. In psychophysical experiments subjects identified the translational direction (up/down) of coherently moving dots presented in either the left or right visual field whilst repetitive TMS (25 Hz; 70%) was applied synchronously with stimulus presentation. Application of TMS to MT/TO-1 and MST/TO-2 in the right hemisphere affected translational direction discrimination in both contra-lateral and ipsi-lateral visual fields. In contrast, deficits of motion perception following application of TMS to MT/TO-1 and MST/TO-2 in the left hemisphere were restricted to the contra-lateral visual field. This result suggests an enhanced role for the right hemisphere in processing translational motion across the full visual field

The Spatial Properties of L- and M-Cone Inputs to Electroretinograms That Reflect Different Types of Post-Receptoral Processing

yesWe studied the spatial arrangement of L- and M-cone driven electroretinograms (ERGs) reflecting the activity of magno- and parvocellular pathways. L- and M-cone isolating sine wave stimuli were created with a four primary LED stimulator using triple silent substitution paradigms. Temporal frequencies were 8 and 12 Hz, to reflect cone opponent activity, and 30, 36 and 48 Hz to reflect luminance activity. The responses were measured for full-field stimuli and for different circular and annular stimuli. The ERG data confirm the presence of two different mechanisms at intermediate and high temporal frequencies. The responses measured at high temporal frequencies strongly depended upon spatial stimulus configuration. In the full-field conditions, the L-cone driven responses were substantially larger than the full-field M-cone driven responses and also than the L-cone driven responses with smaller stimuli. The M-cone driven responses at full-field and with 70° diameter stimuli displayed similar amplitudes. The L- and M-cone driven responses measured at 8 and 12 Hz were of similar amplitude and approximately in counter-phase. The amplitudes were constant for most stimulus configurations. The results indicate that, when the ERG reflects luminance activity, it is positively correlated with stimulus size. Beyond 35° retinal eccentricity, the retina mainly contains L-cones. Small stimuli are sufficient to obtain maximal ERGs at low temporal frequencies where the ERGs are also sensitive to cone-opponent processin

Differential processing of the direction and focus of expansion of optic flow stimuli in areas MST and V3A of the human visual cortex

Human neuropsychological and neuroimaging studies have raised the possibility that different attributes of optic flow stimuli, namely radial direction and the position of the focus of expansion (FOE), are processed within separate cortical areas. In the human brain, visual areas V5/MT+ and V3A have been proposed as integral to the analysis of these different attributes of optic flow stimuli. To establish direct causal relationships between neural activity in human (h)V5/MT+ and V3A and the perception of radial motion direction and FOE position, we used transcranial magnetic stimulation (TMS) to disrupt cortical activity in these areas while participants performed behavioral tasks dependent on these different aspects of optic flow stimuli. The cortical regions of interest were identified in seven human participants using standard functional MRI retinotopic mapping techniques and functional localizers. TMS to area V3A was found to disrupt FOE positional judgments but not radial direction discrimination, whereas the application of TMS to an anterior subdivision of hV5/MT+, MST/TO-2 produced the reverse effects, disrupting radial direction discrimination but eliciting no effect on the FOE positional judgment task. This double dissociation demonstrates that FOE position and radial direction of optic flow stimuli are signaled independently by neural activity in areas hV5/MT+ and V3A.NEW & NOTEWORTHY Optic flow constitutes a biologically relevant visual cue as we move through any environment. With the use of neuroimaging and brain-stimulation techniques, this study demonstrates that separate human brain areas are involved in the analysis of the direction of radial motion and the focus of expansion in optic flow. This dissociation reveals the existence of separate processing pathways for the analysis of different attributes of optic flow that are important for the guidance of self-locomotion and object avoidance

Simple reaction times in colour space: the influence of chromaticity, contrast and cone opponency.

NoPURPOSE. This study examined the influence of stimulus chromaticity on simple reaction times (RTs) to determine the stage of chromatic processing that is most influential in their generation. METHODS. Simple RTs were measured in response to the cosinusoidally ramped onset of small, equiluminant, colored Gaussian spots. The chromaticity of these stimuli was varied, to modulate along a series of vectors in color space that included red-green (L-M) and blue-yellow (S-[L+M]) opponent axes. RESULTS. RTs are highly sensitive to small departures from subjective equiluminance. They are also dependent on stimulus chromaticity. The longest RTs are generated in response to equiluminant stimuli that isolate S-cone activity, whereas the shortest are generated by stimuli that modulate the L-M opponent axis. However, temporal processing differences are highly dependent on how the chromatic stimuli are scaled in relation to one another. The differences are reduced when scaling is based on detection threshold. The relationship between chromatic contrast and RT can be described by the modified Piéron equation RT = RT0 + k · C-1. CONCLUSIONS. Simple RTs generated in this study conform to the idea that they are largely determined by cone-opponency mechanisms. The use of cone contrast as a metric for scaling chromatic stimuli exaggerates differences between the temporal responsiveness of L-M and S-(L+M) opponency mechanisms

Specificity and selectivity of chromatic visual evoked potentials

AbstractA paper by Rabin et al. (1994)Vision Research, 34, 2657–2671, claimed that spatially extensive grating stimuli could be used to generate chromatic-specific visual evoked potentials from subjects assumed to have standard spectral sensitivity and tritanopic confusion lines. Here we demonstrate that such spatially extensive stimuli may generate responses which are contaminated by luminance-contrast intrusions. Such intrusions are mainly due to chromatic aberrations and are compounded by the abovementioned assumptions. Claims regarding the chromatic selectivity of VEPs must, therefore, be substantiated by establishing correlations with the known properties of the chromatic system. Coprright © 1996 Elsevier Science Ltd