A theory for the use of visual orientation information which exploits the columnar structure of striate cortex

A neural model is constructed based on the structure of a visual orientation hypercolumn in mammalian striate cortex. It is then assumed that the perceived orientation of visual contours is determined by the pattern of neuronal activity across orientation columns. Using statistical estimation theory, limits on the precision of orientation estimation and discrimination are calculated. These limits are functions of single unit response properties such as orientation tuning width, response amplitude and response variability, as well as the degree of organization in the neural network. It is shown that a network of modest size, consisting of broadly orientation selective units, can reliably discriminate orientation with a precision equivalent to human performance. Of the various network parameters, the discrimination threshold depends most critically on the number of cells in the hypercolumn. The form of the dependence on cell number correctly predicts the results of psychophysical studies of orientation discrimination. The model system's performance is also consistent with psychophysical data in two situations in which human performance is not optimal. First, interference with orientation discrimination occurs when multiple stimuli activate cells in the same hypercolumn. Second, systematic errors in the estimation of orientation can occur when a stimulus is composed of intersecting lines. The results demonstrate that it is possible to relate neural activity to visual performance by an examination of the pattern of activity across orientation columns. This provides support for the hypothesis that perceived orientation is determined by the distributed pattern of neural activity. The results also encourage the view of neural activity. The results also are determined by the responses of many neurons rather than the sensitivity of individual cells.     

A Laterally Interconnected Neural Architecture in MST Accounts for Psychophysical Discrimination of Complex Motion Patterns

The complex patterns of visual motion formed across the retina during self-motion, often referred to as optic flow, provide a rich source of information describing our dynamic relationship within the environment. Psychophysical studies indicate the existence of specialized detectors for component motion patterns (radial, circular, planar) that are consistent with the visual motion properties of cells in the medial superior temporal area (MST) of nonhuman primates. Here we use computational modeling and psychophysics to investigate the structural and functional role of these specialized detectors in performing a graded motion pattern (GMP) discrimination task. In the psychophysical task perceptual discrimination varied significantly with the type of motion pattern presented, suggesting perceptual correlates to the preferred motion bias reported in MST. Simulated perceptual discrimination in a population of independent MST-like neural responses showed inconsistent psychophysical performance that varied as a function of the visual motion properties within the population code. Robust psychophysical performance was achieved by fully interconnecting neural populations such that they inhibited nonpreferred units. Taken together, these results suggest that robust processing of the complex motion patterns associated with self-motion and optic flow may be mediated by an inhibitory structure of neural interactions in MST.     

蝇视系统的图形—背景分辨和初级模式分辨

本文通过行为实验及计算机模拟进一步证明,蝇视系统的自发模式辨别可以看作是图形—背景分辨的特殊情况.关键在于蝇的模式分辨是由运动检测器实现的.运动检测器不仅对模式速度反应,也对模式的结构特性反应.本文提出,人视系统的模式分辨也可能部分地由运动检测器来实现.     

Visual discrimination of simple geometrical patterns: I. Measurements for multiple element stimuli

We describe a method for the measurement of visual discrimination between simple patterns. The target to be discriminated is embedded in a background consisting of multiple, randomly positioned but identical elements, and is distinguished by a single parameter such as magnification or relative rotation. The positions of the target and background elements are varied randomly between presentations and discrimination for different values of the target parameter is measured in terms of the time taken for detection of the target. Using this method, we have studied discrimination of rotation and of magnification for simple pattern elements such as lines, triangles and squares. The results for rotation discrimination are interpreted as evidence for the activity of two discrimination mechanisms, one sensitive to the orientation of the lines from which the pattern elements are constructed and the other to the orientation of the pattern element relative to the visual field.     

Brain-Stimulation Induced Blindsight: Unconscious Vision or Response Bias?

A dissociation between visual awareness and visual discrimination is referred to as “blindsight”. Blindsight results from loss of function of the primary visual cortex (V1) which can occur due to cerebrovascular accidents (i.e. stroke-related lesions). There are also numerous reports of similar, though reversible, effects on vision induced by transcranial Magnetic Stimulation (TMS) to early visual cortex. These effects point to V1 as the “gate” of visual awareness and have strong implications for understanding the neurological underpinnings of consciousness. It has been argued that evidence for the dissociation between awareness of, and responses to, visual stimuli can be a measurement artifact of the use of a high response criterion under yes-no measures of visual awareness when compared with the criterion free forced-choice responses. This difference between yes-no and forced-choice measures suggests that evidence for a dissociation may actually be normal near-threshold conscious vision. Here we describe three experiments that tested visual performance in normal subjects when their visual awareness was suppressed by applying TMS to the occipital pole. The nature of subjects’ performance whilst undergoing occipital TMS was then verified by use of a psychophysical measure (d'') that is independent of response criteria. This showed that there was no genuine dissociation in visual sensitivity measured by yes-no and forced-choice responses. These results highlight that evidence for visual sensitivity in the absence of awareness must be analysed using a bias-free psychophysical measure, such as d'', In order to confirm whether or not visual performance is truly unconscious.     

Discrete and continuous modes of curved-line discrimination controlled by effective stimulus duration

In previous experiments two extreme modes of visual discrimination performance have been investigated by measuring small differences in pattern shape at points along a continuum of pattern shapes. These two modes, associated with discrete and continuous encoding processes, were obtained by simultaneously manipulating the number of pattern components in the display and the effective duration of the display. In this experiment, discrimination performance was measured for a fixed number of pattern components, namely three, and variable display time course. The stimuli used were curved lines drawn from a continuum with curvature parameter s. There were three stimulus time courses: (1) 2-s stimulus display, (2) 100-ms stimulus display, and (3) 100-ms stimulus display followed by a post-stimulus mask. Discrimination performance declined smoothly and monotonically with s for (1), but varied non-monotonically with s revealing a central peak for (3). Performance for (2) was intermediate between that for (1) and that for (3). A reduction in effective stimulus duration alone was thus sufficient to cause a transition from continuous to discrete modes of discrimination performance, a result which may be compatible with an explanation of variable discrimination modes based on a method of successive internal approximations of the stimulus.     

Performance of African elephants (Loxodonta africana) and California sea lions (Zalophus californianus) on a two-choice object discrimination task

This study documents the ability of African elephants (Loxodonta africana) and California sea lions (Zalophus californianus) to perform a two-choice object discrimination task. Both species were able to perform this task. However, California sea lions took fewer trials overall to reach a criterion of 10 consecutively correct responses than did African elephants. The performance of California sea lions did not change significantly during this study. However, African elephants showed a gradual learning of the task, as exhibited by a gradual decrease in the number of trials needed to reach criterion. This performance difference may reflect differences in either visual abilities or cognitive functioning, which in turn may be influenced by either different evolutionary pressures exerted on herbivores and carnivores, or by the context in which visual information is used in the lives of these animals. © 1994 Wiley-Liss, Inc.     

Motor Cost Influences Perceptual Decisions

Perceptual decision making has been widely studied using tasks in which subjects are asked to discriminate a visual stimulus and instructed to report their decision with a movement. In these studies, performance is measured by assessing the accuracy of the participants’ choices as a function of the ambiguity of the visual stimulus. Typically, the reporting movement is considered as a mere means of reporting the decision with no influence on the decision-making process. However, recent studies have shown that even subtle differences of biomechanical costs between movements may influence how we select between them. Here we investigated whether this purely motor cost could also influence decisions in a perceptual discrimination task in detriment of accuracy. In other words, are perceptual decisions only dependent on the visual stimulus and entirely orthogonal to motor costs? Here we show the results of a psychophysical experiment in which human subjects were presented with a random dot motion discrimination task and asked to report the perceived motion direction using movements of different biomechanical cost. We found that the pattern of decisions exhibited a significant bias towards the movement of lower cost, even when this bias reduced performance accuracy. This strongly suggests that motor costs influence decision making in visual discrimination tasks for which its contribution is neither instructed nor beneficial.     

A detection theoretic explanation of blindsight suggests a link between conscious perception and metacognition

Blindsight refers to the rare ability of V1-damaged patients to perform visual tasks such as forced-choice discrimination, even though these patients claim not to consciously see the relevant stimuli. This striking phenomenon can be described in the formal terms of signal detection theory. (i) Blindsight patients use an unusually conservative criterion to detect targets. (ii) In discrimination tasks, their confidence ratings are low and (iii) such confidence ratings poorly predict task accuracy on a trial-by-trial basis. (iv) Their detection capacity (d') is lower than expected based on their performance in forced-choice tasks. We propose a unifying explanation that accounts for these features: that blindsight is due to a failure to represent and update the statistical information regarding the internal visual neural response, i.e. a failure in metacognition. We provide computational simulation data to demonstrate that this model can qualitatively account for the detection theoretic features of blindsight. Because such metacognitive mechanisms are likely to depend on the prefrontal cortex, this suggests that although blindsight is typically due to damage to the primary visual cortex, distal influence to the prefrontal cortex by such damage may be critical. Recent brain imaging evidence supports this view.     

Breakfast in the nook and dinner in the dining room: time-of-day discrimination in rats

Four studies were conducted which demonstrate that most (63%) male Sprague-Dawley rats can attain criterion, nine correct choices over ten consecutive trials, on a time-of-day discrimination in an elevated T-maze, but that the task is relatively difficult. The discrimination required that the rats go to one goal arm during a morning session and the other in an afternoon session. The sessions always began at the same time and were at least 6 h apart. A larger proportion of rats attained criterion and required fewer trials when the discriminative cue was a maze insert providing visual and tactile stimulation (0.72), orientation and position of the maze in the room (0.88), or the rats were required to always make the same left or right turn (0.94). Also, once criterion was attained, rats trained on time-of-day discrimination only made about 70% correct choices with continued training. Housing the rats with continuous light, all laboratory noises masked with white noise, and a random feeding schedule did not prevent them from acquiring the time-place discrimination. Testing the rats with a random number of trials during morning and afternoon sessions and with added or omitted sessions revealed that the rats did not use response or session alternation strategies to perform the discrimination. Also, the particular experimenter administering the morning or afternoon sessions did not serve as a cue for the discrimination. The relative difficulty of the task suggests that time of day does not normally function as a discriminative stimulus for choices, but probably as a contextual stimulus. Further, performance of the task in the absence of time-of-day cues suggests that the discrimination is based on event memory combined with an internal timing mechanism.     

Modulation of activity in human visual area V1 during memory masking

Neurons in the primary visual cortex, V1, are specialized for the processing of elemental features of the visual stimulus, such as orientation and spatial frequency. Recent fMRI evidence suggest that V1 neurons are also recruited in visual perceptual memory; a number of studies using multi-voxel pattern analysis have successfully decoded stimulus-specific information from V1 activity patterns during the delay phase in memory tasks. However, consistent fMRI signal modulations reflecting the memory process have not yet been demonstrated. Here, we report evidence, from three subjects, that the low V1 BOLD activity during retention of low-level visual features is caused by competing interactions between neural populations coding for different values along the spectrum of the dimension remembered. We applied a memory masking paradigm in which the memory representation of a masker stimulus interferes with a delayed spatial frequency discrimination task when its frequency differs from the discriminanda with ±1 octave and found that impaired behavioral performance due to masking is reflected in weaker V1 BOLD signals. This cross-channel inhibition in V1 only occurs with retinotopic overlap between the masker and the sample stimulus of the discrimination task. The results suggest that memory for spatial frequency is a local process in the retinotopically organized visual cortex.     

The Temporal Dynamics of Early Visual Cortex Involvement in Behavioral Priming

Transcranial magnetic stimulation (TMS) allows for non-invasive interference with ongoing neural processing. Applied in a chronometric design over early visual cortex (EVC), TMS has proved valuable in indicating at which particular time point EVC must remain unperturbed for (conscious) vision to be established. In the current study, we set out to examine the effect of EVC TMS across a broad range of time points, both before (pre-stimulus) and after (post-stimulus) the onset of symbolic visual stimuli. Behavioral priming studies have shown that the behavioral impact of a visual stimulus can be independent from its conscious perception, suggesting two independent neural signatures. To assess whether TMS-induced suppression of visual awareness can be dissociated from behavioral priming in the temporal domain, we thus implemented three different measures of visual processing, namely performance on a standard visual discrimination task, a subjective rating of stimulus visibility, and a visual priming task. To control for non-neural TMS effects, we performed electrooculographical recordings, placebo TMS (sham), and control site TMS (vertex). Our results suggest that, when considering the appropriate control data, the temporal pattern of EVC TMS disruption on visual discrimination, subjective awareness and behavioral priming are not dissociable. Instead, TMS to EVC disrupts visual perception holistically, both when applied before and after the onset of a visual stimulus. The current findings are discussed in light of their implications on models of visual awareness and (subliminal) priming.     

Prosimian learning capacities

A survey of the literature on learning by prosimian primates contradicts a traditional view that they are untrainable. They learn simple operants such as bar pressing readily, work well for the usual rewards, and respond in predictable fashion on schedules of partial reinforcement. Two-dimensional visual discriminations involving pattern and brightness also are learned, as are three-dimensional object discriminations. In the area of complex learning, they not only acquire a variety of difficult tasks, but they also reach a final level of performance that is at or near the level of anthropoids. In each of the following areas, at least one experiment is described in which a prosimian species did as well as one or more anthropoid species: object discrimination learning set, discrimination reversal, delayed response, instrumentation, oddity, and relational problems. The anthropoids to which they are most apt to compare favorably are New World monkeys, a group which they resemble physically, especially with respect to degree of manipulativeness, cortical development, and visual acuity.     

Exposure to Organic Solvents Used in Dry Cleaning Reduces Low and High Level Visual Function

PurposeTo investigate whether exposure to occupational levels of organic solvents in the dry cleaning industry is associated with neurotoxic symptoms and visual deficits in the perception of basic visual features such as luminance contrast and colour, higher level processing of global motion and form (Experiment 1), and cognitive function as measured in a visual search task (Experiment 2).MethodsThe Q16 neurotoxic questionnaire, a commonly used measure of neurotoxicity (by the World Health Organization), was administered to assess the neurotoxic status of a group of 33 dry cleaners exposed to occupational levels of organic solvents (OS) and 35 age-matched non dry-cleaners who had never worked in the dry cleaning industry. In Experiment 1, to assess visual function, contrast sensitivity, colour/hue discrimination (Munsell Hue 100 test), global motion and form thresholds were assessed using computerised psychophysical tests. Sensitivity to global motion or form structure was quantified by varying the pattern coherence of global dot motion (GDM) and Glass pattern (oriented dot pairs) respectively (i.e., the percentage of dots/dot pairs that contribute to the perception of global structure). In Experiment 2, a letter visual-search task was used to measure reaction times (as a function of the number of elements: 4, 8, 16, 32, 64 and 100) in both parallel and serial search conditions.ResultsDry cleaners exposed to organic solvents had significantly higher scores on the Q16 compared to non dry-cleaners indicating that dry cleaners experienced more neurotoxic symptoms on average. The contrast sensitivity function for dry cleaners was significantly lower at all spatial frequencies relative to non dry-cleaners, which is consistent with previous studies. Poorer colour discrimination performance was also noted in dry cleaners than non dry-cleaners, particularly along the blue/yellow axis. In a new finding, we report that global form and motion thresholds for dry cleaners were also significantly higher and almost double than that obtained from non dry-cleaners. However, reaction time performance on both parallel and serial visual search was not different between dry cleaners and non dry-cleaners.ConclusionsExposure to occupational levels of organic solvents is associated with neurotoxicity which is in turn associated with both low level deficits (such as the perception of contrast and discrimination of colour) and high level visual deficits such as the perception of global form and motion, but not visual search performance. The latter finding indicates that the deficits in visual function are unlikely to be due to changes in general cognitive performance.     

Sensitivity variations in the visual system,contrast resolution and eye movements

Attention is drawn to the fact that under normal visual conditions the sensitivity of the receptor units of the visual system are subject to spatial and temporal variations, and that consequently in performing pattern recognition the visual cortex has to discriminate between external luminance structure and internal sensitivity structure. It is suggested that eye movements are the method by which this discrimination is performed. In a simplified model analysis it is shown that eye movements are a suitable mechanism for this discrimination. Implications of this model for detection threshold and stabilized retinal images are discussed. A new interpretation of the adaptation to sine wave grids is given.     

Arginine vasopressin facilitates reversal learning in albino, but not hooded rats

Male Holtzman albino and Long-Evans hooded rats were administered one microgram of arginine vasopressin (AVP) or a placebo each day after the acquisition trials of a visual white-black discrimination. Animals were then trained in the reversal of the discrimination. Performance was assessed by the number of trials to criterion during acquisition and reversal. Treatment with AVP resulted in significantly fewer trials to criterion during reversal learning in Holtzman albino rats, but did not influence reversal learning in Long-Evans hooded rats. These results provide evidence that AVP may have differential actions on memory processes in different strains of rats.     

Discrimination of coloured patterns by honeybees through chromatic and achromatic cues

We investigated pattern discrimination by worker honeybees, Apis mellifera, focusing on the roles of spectral cues and the angular size of patterns. Free-flying bees were trained to discriminate concentric patterns in a Y-maze. The rewarded pattern could be composed of either a cyan and a yellow colour, which presented both different chromatic and achromatic L-receptor contrast, or an orange and a blue colour, which presented different chromatic cues, but the same L-receptor contrast. The non-rewarded alternative was either a single-coloured disc with the colour of the central disc or the surrounding ring of the pattern, a checkerboard pattern with non-resolvable squares, the reversed pattern, or the elements of the training pattern (disc or ring alone). Bees resolved and learned both colour elements in the rewarded patterns and their spatial properties. When the patterns subtended large visual angles, this discrimination used chromatic cues only. Patterns with yellow or orange central discs were generalised toward the yellow and orange colours, respectively. When the patterns subtended a visual angle close to the detection limit and L-receptor contrast was mediating discrimination, pattern perception was reduced: bees perceived only the pattern element with higher contrast.     

The effect of ecologically relevant variations in light level on the performance of Mongolian gerbils on two visual tasks

Tests of rodent visual capacities are typically performed under standard laboratory illumination. However, light level can have subtle and complex effects on behavior in rodents. We tested Mongolian gerbils (Meriones unguiculatus) – a species that shows individual differences in activity pattern – on visual tasks at three ecologically relevant levels of ambient illuminance: approximating moonlight (1 lx), dawn or dusk (10 lx), and low daylight (100 lx). A jumping task and a grating discrimination quantified depth perception and grating acuity, respectively. Illuminance variations had important effects on behavior. Gerbils jumped faster in lower light on the depth discrimination task, but were also less accurate than in bright light. Daytime activity levels (possibly reflecting variations in activity pattern) mediated these effects, with animals that were awake during a lower proportion of daytime hours exhibiting a trend toward jumping faster with lower light level as compared to more day-active gerbils. Moreover, while illuminance did not affect acuity overall, it interacted with activity levels in a complex way: in both the darkest and lowest light levels, animals that were awake during a greater proportion of daytime hours showed higher acuity levels than animals that were less active during the day. These results indicate that gerbils show behavioral profiles in vision tasks that represent an interaction between visual ability and illuminance-sensitive motivational or emotional actors, perhaps including chronotype. Thus, the most ecologically relevant assessments of the visual performance of rodents will likely be achieved by testing at species-specific preferred light levels.     

Color and luminance contrasts attract independent attention

Paying attention can improve vision in many ways, including some very basic functions such as contrast discrimination, a task that probably reflects very early levels of visual processing. Electrophysiological, psychophysical, and imaging studies on humans as well as single recordings in monkey show that attention can modulate the neuronal response at an early stage of visual processing, probably by acting on the response gain. Here, we measure incremental contrast thresholds for luminance and color stimuli to derive the contrast response of early neural mechanisms and their modulation by attention. We show that, for both cases, attention improves contrast discrimination, probably by multiplicatively increasing the gain of the neuronal response to contrast. However, the effects of attention are highly specific to the visual modality: concurrent attention to a competing luminance, but not chromatic pattern, greatly impedes luminance contrast discrimination; and attending to a competing chromatic, but not luminance, task impedes color contrast discrimination. Thus, the effects of attention are highly modality specific, implying separate attentional resources for different fundamental visual attributes at early stages of visual processing.     

Effects of chronic iTBS‐rTMS and enriched environment on visual cortex early critical period and visual pattern discrimination in dark‐reared rats

Early cortical critical period resembles a state of enhanced neuronal plasticity enabling the establishment of specific neuronal connections during first sensory experience. Visual performance with regard to pattern discrimination is impaired if the cortex is deprived from visual input during the critical period. We wondered how unspecific activation of the visual cortex before closure of the critical period using repetitive transcranial magnetic stimulation (rTMS) could affect the critical period and the visual performance of the experimental animals. Would it cause premature closure of the plastic state and thus worsen experience‐dependent visual performance, or would it be able to preserve plasticity? Effects of intermittent theta‐burst stimulation (iTBS) were compared with those of an enriched environment (EE) during dark‐rearing (DR) from birth. Rats dark‐reared in a standard cage showed poor improvement in a visual pattern discrimination task, while rats housed in EE or treated with iTBS showed a performance indistinguishable from rats reared in normal light/dark cycle. The behavioral effects were accompanied by correlated changes in the expression of brain‐derived neurotrophic factor (BDNF) and atypical PKC (PKCζ/PKMζ), two factors controlling stabilization of synaptic potentiation. It appears that not only nonvisual sensory activity and exercise but also cortical activation induced by rTMS has the potential to alleviate the effects of DR on cortical development, most likely due to stimulation of BDNF synthesis and release. As we showed previously, iTBS reduced the expression of parvalbumin in inhibitory cortical interneurons, indicating that modulation of the activity of fast‐spiking interneurons contributes to the observed effects of iTBS. © 2015 Wiley Periodicals, Inc. Develop Neurobiol 76: 19–33, 2016