Various background pattern-effect on saccadic suppression.

It has been proved that the saccadic suppression is a phenomenon closely related to the presence of contours and structures in the visual field. Experiments were performed to clarify whether the structured background influences the pattern of attention distribution (making the stimulus detection more difficult) or whether the elevation of visual threshold is due to the "masking' effect of the moving background image over the retina. Two types of backgrounds were used therefore: those with symbolic meaning in the processing of which "psychological' mechanisms are presumably involved like picture reproductions of famous painters and photographs of nudes, and those lacking semantic significance like computer figures composed of randomly distributed black and white squares with different grain expressed as the entropy of the pattern. The results show that saccadic suppression is primarily a consequence of peripheral mechanisms, probably of lateral inhibition in the visual field occurring in the presence of moving edges over the retina. Psychological factors have to be excluded as being fundamental for saccadic suppression.     

Mechanisms for allocating auditory attention: an auditory saliency map

Our nervous system is confronted with a barrage of sensory stimuli, but neural resources are limited and not all stimuli can be processed to the same extent. Mechanisms exist to bias attention toward the particularly salient events, thereby providing a weighted representation of our environment. Our understanding of these mechanisms is still limited, but theoretical models can replicate such a weighting of sensory inputs and provide a basis for understanding the underlying principles. Here, we describe such a model for the auditory system-an auditory saliency map. We experimentally validate the model on natural acoustical scenarios, demonstrating that it reproduces human judgments of auditory saliency and predicts the detectability of salient sounds embedded in noisy backgrounds. In addition, it also predicts the natural orienting behavior of naive macaque monkeys to the same salient stimuli. The structure of the suggested model is identical to that of successfully used visual saliency maps. Hence, we conclude that saliency is determined either by implementing similar mechanisms in different unisensory pathways or by the same mechanism in multisensory areas. In any case, our results demonstrate that different primate sensory systems rely on common principles for extracting relevant sensory events.     

A model of the saccade-generating system that accounts for trajectory variations produced by competing visual stimuli

Variable saccade trajectories are produced in visual search paradigms in which multiple potential target stimuli are present. These variable trajectories provide a rich source of information that may lead to a deeper understanding of the basic control mechanisms of the saccadic system. We have used published behavioral observations and neural recordings in the superior colliculus (SC), gathered in monkeys performing visual search paradigms, to guide the construction of a new distributed model of the saccadic system. The new model can account for many of the variations in saccade trajectory produced by the appearance of multiple visual stimuli in a search paradigm. The model uses distributed feedback about current eye motion from the brainstem to the SC to reduce activity there at physiologically realistic rates during saccades. The long-range lateral inhibitory connections between SC cells used in previous models have been eliminated to match recent physiological evidence. The model features interactions between visually activated multiple populations of cells in the SC and distributed and topologically organized inhibitory input to the SC from the SNr to produce some of the types of variable saccadic trajectories, including slightly curved and averaging saccades, observed in visual search tasks. The distributed perisaccadic disinhibition of SC from the substantia nigra (SNr) is assumed to have broad spatial tuning. In order to produce the strongly curved saccades occasionally recorded in visual search, the existence of a parallel input to the saccadic burst generators in addition to that provided by the distributed input from the SC is required. The spatiotemporal form of this additional parallel input is computed based on the assumption that the input from the model SC is realistic. In accordance with other recent models, it is assumed that the parallel input comes from the cerebellum, but our model predicts that the parallel input is delayed during highly curved saccadic trajectories.     

Cortical Contributions to Saccadic Suppression

The stability of visual perception is partly maintained by saccadic suppression: the selective reduction of visual sensitivity that accompanies rapid eye movements. The neural mechanisms responsible for this reduced perisaccadic visibility remain unknown, but the Lateral Geniculate Nucleus (LGN) has been proposed as a likely site. Our data show, however, that the saccadic suppression of a target flashed in the right visual hemifield increased with an increase in background luminance in the left visual hemifield. Because each LGN only receives retinal input from a single hemifield, this hemifield interaction cannot be explained solely on the basis of neural mechanisms operating in the LGN. Instead, this suggests that saccadic suppression must involve processing in higher level cortical areas that have access to a considerable part of the ipsilateral hemifield.     

Saccadic eye movements and the detection of fast-moving gratings

Experiments are presented in which the effect of saccadic eye movements on the visibility of sinusoidal gratings drifting with velocities between 2 deg/s and 400 deg/s is investigated. The results demonstrate that saccades are highly useful for detecting this class of stimuli. Due to a saccade, otherwise subthreshold stimuli become visible as short, distinct flashes of the seemingly statinoary pattern. The paper analyzes in detail the dependence of the amount of facilitation on saccade size and relative direction and isolates the additional effect of saccadic suppression. A simple model is proposed which predicts the experimental findings.     

Sensory and behavioral properties of neurons in posterior parietal cortex of the awake, trained monkey.

Neurons in posterior parietal cortex of the awake, trained monkey respond to passive visual and/or somatosensory stimuli. In general, the receptive fields of these cells are large and nonspecific. When these neurons are studied during visually guided hand movements and eye movements, most of their activity can be accounted for by passive sensory stimulation. However, for some visual cells, the response to a stimulus is enhanced when it is to be the target for a saccadic eye movement. This enhancement is selective for eye movements into the visual receptive field since it does not occur with eye movements to other parts of the visual field. Cells that discharge in association with a visual fixation task have foveal receptive fields and respond to the spots of light used as fixation targets. Cells discharging selectively in association with different directions of tracking eye movements have directionally selective responses to moving visual stimuli. Every cell in our sample discharging in association with movement could be driven by passive sensory stimuli. We conclude that the activity of neurons in posterior parietal cortex is dependent on and indicative of external stimuli but not predictive of movement.     

Eye movements modulate visual receptive fields of V4 neurons

The receptive field, defined as the spatiotemporal selectivity of neurons to sensory stimuli, is central to our understanding of the neuronal mechanisms of perception. However, despite the fact that eye movements are critical during normal vision, the influence of eye movements on the structure of receptive fields has never been characterized. Here, we map the receptive fields of macaque area V4 neurons during saccadic eye movements and find that receptive fields are remarkably dynamic. Specifically, before the initiation of a saccadic eye movement, receptive fields shrink and shift towards the saccade target. These spatiotemporal dynamics may enhance information processing of relevant stimuli during the scanning of a visual scene, thereby assisting the selection of saccade targets and accelerating the analysis of the visual scene during free viewing.     

Adaptive body patterning,three‐dimensional skin morphology and camouflage measures of the slender filefish Monacanthus tuckeri on a Caribbean coral reef

The slender filefish is a master of adaptive camouflage and can change its appearance within 1–3 s. Videos and photographs of this animal's cryptic body patterning and behavior were collected in situ under natural light on a Caribbean coral reef. We present an ethogram of body patterning components that includes large‐ and small‐scale spots, stripes and bars that confer a variety of cryptic patterns amidst a range of complex backgrounds. Field images were analyzed to investigate two aspects of camouflage effectiveness: (1) the degree of colour resemblance between animals and their nearby visual stimuli; and (2) the visibility of each fish's actual body outline vs. its illusory outline. Most animals more closely matched the colour of nearby visual stimuli than that of the surrounding background. Three‐dimensional dermal flaps complement the melanophore skin patterns by enhancing the complexity of the fish's physical skin texture to disguise its actual body shape, and the morphology of these structures was studied. The results suggest that the body patterns, skin texture, postures and swimming orientations putatively hinder both the detection and recognition of the fish by potential visual predators. Overall, the rapid speed of change of multiple patterns, colour blending with nearby backgrounds, and the physically complicated edge produced by dermal flaps effectively camouflage this animal among soft corals and macroalgae in the Caribbean Sea.     

Role of statistical symmetries in sensory coding: an optimal scale invariant code for vision

The visual system is the most studied sensory pathway, which is partly because visual stimuli have rather intuitive properties. There are reasons to think that the underlying principle ruling coding, however, is the same for vision and any other type of sensory signal, namely the code has to satisfy some notion of optimality--understood as minimum redundancy or as maximum transmitted information. Given the huge variability of natural stimuli, it would seem that attaining an optimal code is almost impossible; however, regularities and symmetries in the stimuli can be used to simplify the task: symmetries allow predicting one part of a stimulus from another, that is, they imply a structured type of redundancy. Optimal coding can only be achieved once the intrinsic symmetries of natural scenes are understood and used to the best performance of the neural encoder. In this paper, we review the concepts of optimal coding and discuss the known redundancies and symmetries that visual scenes have. We discuss in depth the only approach which implements the three of them known so far: translational invariance, scale invariance and multiscaling. Not surprisingly, the resulting code possesses features observed in real visual systems in mammals.     

Gaze control: role of the parietal cortex

Eye movements constitute one of the most basic means of interacting with our environment, allowing to orient to, localize and scrutinize the variety of potentially interesting objects that surround us. In this review we discuss the role of the parietal cortex in the control of saccadic and smooth pursuit eye movements, whose purpose is to rapidly displace the line of gaze and to maintain a moving object on the central retina, respectively. From single cell recording studies in monkey we know that distinct sub-regions of the parietal lobe are implicated in these two kinds of movement. The middle temporal (MT) and medial superior temporal (MST) areas show neuronal activities related to moving visual stimuli and to ocular pursuit. The lateral intraparietal (LIP) area exhibits visual and saccadic neuronal responses. Electrophysiology, which in essence is a correlation method, cannot entirely solve the question of the functional implication of these areas: are they primarily involved in sensory processing, in motor processing, or in some intermediate function? Lesion approaches (reversible or permanent) in the monkey can provide important information in this respect. Lesions of MT or MST produce deficits in the perception of visual motion, which would argue for their possible role in sensory guidance of ocular pursuit rather than in directing motor commands to the eye muscle. Lesions of LIP do not produce specific visual impairments and cause only subtle saccadic deficits. However, recent results have shown the presence of severe deficits in spatial attention tasks. LIP could thus be implicated in the selection of relevant objects in the visual scene and provide a signal for directing the eyes toward these objects. Functional imaging studies in humans confirm the role of the parietal cortex in pursuit, saccadic, and attentional networks, and show a high degree of overlap with monkey data. Parietal lobe lesions in humans also result in behavioral deficits very similar to those that are observed in the monkey. Altogether, these different sources of data consistently point to the involvement of the parietal cortex in the representation of space, at an intermediate stage between vision and action.     

Neural correlates of saccadic suppression in humans

When you look into a mirror and move your eyes left to right, you will see that you cannot observe your own eye movements. This demonstrates the phenomenon of saccadic suppression: during saccadic eye movements, visual sensitivity is much reduced. Given that humans make more than 100,000 eye movements each day, it is clear why suppression is needed: without it, the motion on the retina would prevent us from seeing anything at all. Psychophysical data show that suppression is stimulus selective: it is strongest for the kind of stimuli that preferentially activate magnocellular thalamic neurons. This has led to the hypothesis that saccadic suppression selectively targets the magnocellular stream. We used fMRI to find brain areas with a stimulus-selective suppression of the BOLD signal that matches the psychophysical data. We found such a neural correlate of saccadic suppression in the dorsal stream (hMT+, V7) and in ventral area V4. These areas receive magnocellular input; hence our findings are consistent with the magnocellular hypothesis. The range of effects in our data and in single cell data, however, argues against a single thalamic mechanism that suppresses all cortical input. Instead, we speculate that saccadic suppression relies on multiple mechanisms operating in different cortical areas.     

The influence of structured visual backgrounds on smooth-pursuit initiation,steady-state pursuit and smooth-pursuit termination

 Smooth-pursuit eye movements were recorded in two rhesus monkeys in order to compare the influence of structured visual backgrounds on smooth-pursuit initiation, steady-state pursuit and pursuit termination. Different target trajectories were used in order to study smooth-pursuit initiation and termination. The influence of visual backgrounds on pursuit initiation was characterized by recording ocular responses elicited by step-ramp target displacements starting from straight ahead. Pursuit termination was characterized by analysing the transition from steady-state smooth-pursuit to fixation when a centripetally directed target ramp was terminated by a small target step in the direction of the ramp as soon as the target had come close to the straightahead position. The quantification of steady-state pursuit was based on ocular responses elicited by either paradigm. In accordance with previous work, we found that the onset of smooth-pursuit eye movements was delayed and initial eye acceleration reduced in the presence of a structured visual background. Likewise, mean eye velocity during steady-state pursuit was reduced by structured visual backgrounds. However, neither the latency nor the time course of smooth-pursuit termination was altered when the homogeneous background was replaced by a structured visual background. The lack of sensitivity of pursuit termination to the presence of visual structured backgrounds supports a previous contention that pursuit termination is mediated by a process which is different from the ones mediating smooth-pursuit initiation and steady-state pursuit. The absence of any noticeable effect of structured backgrounds on pursuit termination suggests that at least the fast component of the optokinetic reflex is suppressed during pursuit termination. Received: 24 October 1994/Accepted in revised form: 16 December 1994     

Turbidity Hampers Mate Choice in a Pipefish

European coastal waters have in recent years become more turbid as algal growth has increased, probably due to eutrophication, global warming and changes in fish communities. Turbidity reduces visibility, and such changes may in turn affect animal behaviour as well as evolutionary processes that are dependent on visual stimuli. In this study we experimentally manipulated water visibility and olfactory cues to investigate mate choice using the sex role‐reversed broad‐nosed pipefish Syngnathus typhle as our study organism. We show that males spent significantly longer time assessing females when they had access to full visual cues, compared to when visibility was reduced. Presence or absence of olfactory cues from females did not affect mate choice, suggesting that the possible use of smell could not make up for a reduction in visibility. This implies that mate choice is environmentally dependent and that an increased turbidity may affect processes of sexual selection through an impaired possibility for visually based mate choice.     

Mechanisms of neural integration in the parietal lobe for visual attention.

The impulse discharges of neurons in the inferior parietal association cortex (area 7) were studied in the alert, behaving rhesus monkey, trained to fixate and follow visual targets. Four classes of cells related to visual or visuomotor function were found. Cells of one of these are sensitive to visual stimuli and have large, contralateral receptive fields with maximal sensitivity in the far temporal quadrants. Cells of the other three classes are related to visuomotor functions: visual fixation, tracking, and saccades. They are neither sensory nor motor in the usual sense for they are activated only by interested fixation of gaze or tracking, or before visually evoked saccadic eye movements. They are not activated during the spontaneous saccades and fixations that the monkey makes while casually exploring his environment. It is hypothesized that the light-sensitive neurons provide the visual input to the visuomotor cells that, in turn, produce a command signal for the direction of visual attention and for shifting the focus of attention from one target to another.     

The role of attentional priority and saliency in determining capacity limits in enumeration and visual working memory

Many common tasks require us to individuate in parallel two or more objects out of a complex scene. Although the mechanisms underlying our abilities to count the number of items, remember the visual properties of objects and to make saccadic eye movements towards targets have been studied separately, each of these tasks require selection of individual objects and shows a capacity limit. Here we show that a common factor--salience--determines the capacity limit in the various tasks. We manipulated bottom-up salience (visual contrast) and top-down salience (task relevance) in enumeration and visual memory tasks. As one item became increasingly salient, the subitizing range was reduced and memory performance for all other less-salient items was decreased. Overall, the pattern of results suggests that our abilities to enumerate and remember small groups of stimuli are grounded in an attentional priority or salience map which represents the location of important items.     

Separate visual representations for perception and action revealed by saccadic eye movements

Some 30 years ago, Trevarthen [1] introduced the idea of two separate visual systems, a focal system for fine motor acts and an ambient system for gross body movements such as ambulation. More recent developments indicating anatomically and physiologically separate pathways in primate vision [2] have led to a different idea of separate visual systems, one for conscious perception and one for action [3]. It has received empirical support from several studies showing that pointing, reaching, and grasping can remain accurate while the perceived position or size of objects is subject to illusory distortion [4-6]. However, much of this evidence has been challenged on the grounds of methodological flaws, particularly failure to match perfectly the conditions for verbal and motor tasks and failure to replicate results [7-10]. Here we take advantage of the strong compression of perceived position that occurs around the time of saccadic eye movements [11, 12]. Under normal lighting conditions, stimuli flashed briefly over a wide range of spatial positions just before saccadic onset are neither seen nor reached for in their veridical positions, but are compressed toward the saccadic target. We validate the idea of separate systems by showing that, in the dark, subjects are able to point accurately to the correct target position, even though their verbal reports are still subject to compression.     

Latency of visually evoked saccadic eye movements

The paper deals with the initiation of visually guided saccades, in order to break down the saccadic reaction time into functionally different periods of time. It takes into account that spatial processing of information is so basic that modelling of saccadic control properties should include spatio-temporal arrangements. The output signal of the saccadic system was measured in response to visual stimuli in which the time between the appearance of a visual stimulus in the peripheral field and the disappearance of the central fixation point was varied. The variation of the mean saccadic latency time, measured with respect to the onset of the peripheral stimulus, as a function of stimulus asynchrony was highly significant. This variation may be represented by a so-called gap-overlap curve, which is characterized here by means of five parameters. A facilitation model is introduced to fit the results of the gap-overlap experiments. The facilitation model for the initiation of visually evoked saccades incorporates a mechanism which governs the efficiency of processing of signals that arise from a stimulus presented at a particular position in space. It explains how visual information may be affected by other sensory information before it is used to command further saccades. It allows determination of saccadic system parameters, such as the peripheral and the foveal afferent processing time, the central processing time for a saccade and the degree of facilitation. These quantities were found to be characteristic for the given test subjects, and where these data could be compared with neurophysiological data, the agreement was within the experimental error.     

Face Orientation and Motion Differently Affect the Deployment of Visual Attention in Newborns and 4-Month-Old Infants

Orienting visual attention allows us to properly select relevant visual information from a noisy environment. Despite extensive investigation of the orienting of visual attention in infancy, it is unknown whether and how stimulus characteristics modulate the deployment of attention from birth to 4 months of age, a period in which the efficiency in orienting of attention improves dramatically. The aim of the present study was to compare 4-month-old infants’ and newborns’ ability to orient attention from central to peripheral stimuli that have the same or different attributes. In Experiment 1, all the stimuli were dynamic and the only attribute of the central and peripheral stimuli to be manipulated was face orientation. In Experiment 2, both face orientation and motion of the central and peripheral stimuli were contrasted. The number of valid trials and saccadic latency were measured at both ages. Our results demonstrated that the deployment of attention is mainly influenced by motion at birth, while it is also influenced by face orientation at 4-month of age. These findings provide insight into the development of the orienting visual attention in the first few months of life and suggest that maturation may be not the only factor that determines the developmental change in orienting visual attention from birth to 4 months.     

Function of a fly motion-sensitive neuron matches eye movements during free flight

Sensing is often implicitly assumed to be the passive acquisition of information. However, part of the sensory information is generated actively when animals move. For instance, humans shift their gaze actively in a sequence of saccades towards interesting locations in a scene. Likewise, many insects shift their gaze by saccadic turns of body and head, keeping their gaze fixed between saccades. Here we employ a novel panoramic virtual reality stimulator and show that motion computation in a blowfly visual interneuron is tuned to make efficient use of the characteristic dynamics of retinal image flow. The neuron is able to extract information about the spatial layout of the environment by utilizing intervals of stable vision resulting from the saccadic viewing strategy. The extraction is possible because the retinal image flow evoked by translation, containing information about object distances, is confined to low frequencies. This flow component can be derived from the total optic flow between saccades because the residual intersaccadic head rotations are small and encoded at higher frequencies. Information about the spatial layout of the environment can thus be extracted by the neuron in a computationally parsimonious way. These results on neuronal function based on naturalistic, behaviourally generated optic flow are in stark contrast to conclusions based on conventional visual stimuli that the neuron primarily represents a detector for yaw rotations of the animal.     

Cortical slow negative potentials during eye fixation and preparing of visually triggered saccades in a man

In 10 right-handed healthy subjects EEGs preceding saccades with mean latent periods were selectively averaged. Two standard schemes of visual stimulation were used: with immediate presentation of a peripheral target stimuli after the central fixation stimulus (a single step paradigm) and with the interval between the stimuli in 200 ms (GAP paradigm). Two waves of slow premotor negativity (early PMN1 and late PMN2) that appeared 930 +/- 79 and 609 +/- 82 ms, respectively, prior to a saccade onset were observed. The PMN2 was followed by the negative potentials N-3, N-2, and N-1 (saccadic initiation potential). It was found that in GAP stimulation condition the PMN1 was less pronounces and N-1 was increased as compared to the single step. These findings suggest that disengage of attention from the central point during the GAP period clears the saccadic system for decision making and initiation of a saccade. Under such conditions, the expectation of a visual target does not require a high level of nonspecific activation and motor attention.