Primate pupillary light reflex: receptive field characteristics of pretectal luminance neurons

This study examined the response properties of luminance neurons found within the pretectal olivary nucleus (PON), which is the pretectal nucleus that mediates the primate pupillary light reflex. We recorded the activity of 121 single units in alert, behaving rhesus monkeys trained to fixate a back-projected laser spot while a luminance stimulus was presented. The change in the firing rate of luminance neurons was measured as a function of changes in the size, retinal illuminance, and position of the stimulus. We found that these neurons possessed large receptive fields, which were sufficiently distinct that they could be placed into three classes. Approximately 40% of the PON luminance neurons responded well to stimuli presented in either the contralateral or ipsilateral hemifield. These neurons were classified as "bilateral" neurons. In the primate, retinal projections to the pretectum and other retinorecipient nuclei are organized such that direct retinal input can only account for the contralateral hemifield responses of these neurons. Thus the representation of the ipsilateral hemifield in "bilateral" PON cells must result from input from a nonretinal source. Approximately 30% of PON neurons responded only to stimuli presented in the contralateral hemifield. These neurons were classified as "contralateral" neurons. Finally, approximately 30% of PON neurons responded to stimuli presented at or near the animal's fixation point. These neurons were classified as "macular" neurons. The mean firing rates of all classes of neurons increased with increases in stimulus size and luminance within their receptive fields. The thresholds and magnitude of these responses closely matched those that would be appropriate for mediating the pupillary light reflex. In summary, these results suggest that all three classes of PON neurons contribute to the behaviorally observed pupillomotor field characteristics in which stimuli at the macular produce substantially larger pupillary responses than more peripheral stimuli. The contributions of "bilateral" and "contralateral" cells account for pupillary responses evoked by peripheral changes in luminance, whereas the contributions of all three cell classes account for the larger pupillary responses evoked by stimuli in the central visual field.     

Prepulse effects on the photic eyeblink reflex: Evidence for startle-dazzle theory

Weak visual prestimulation effects on the early (R50, 50–80 ms) and late (R80, 80–200 ms) components of the eyeblink response to bright light flashes were studied in 16 normal individuals. At a lead time of 120 ms, R50 was inhibited relative to no-prepulse control trials, whereas R80 was facilitated. According to the proposed startle-dazzle theory, luminance onset transients trigger an initial response, R50, that is functionally related to startle. Sustained stimulation then activates prolonged eyelid (R80) and pupil responses, which serve to minimize retinal bleaching. Although the sensitivity of the photic blink reflex to attention is controversial, R50 latency showed a pattern suggestive of inhibition of return (Posner & Cohen, 1984, Attention and performance, Vol. X, pp. 531–556, Hillsdale, NJ: Erlbaum). Analyses in a patient with unilateral occipital lobe damage supported previous evidence that inhibition by a visual prepulse requires neocortex, but facilitation does not.     

Intermittent contralateral and ipsilateral hemiretinal stimulation and its effect on the phasic stretch reflex

The effect of visual stimulation on the phasic stretch reflex, measured isometrically by way of EMG recordings, was investigated with normal human subjects. Three experimental treatments were applied, namely 3 flashes of light presented as hemiretinal contralateral stimulation prior to tendon taps; similar hemiretinal stimulation presented ipsilaterally; and no flashes of light as a control condition. The analysis of variance showed that the MAP means and standard deviations for both of the lights conditions were significantly greater than for the control condition. Eleven out of 12 subjects showed a larger reflex response to stimulation of the brain ipsilateral to the tendon stimulated. The findings are discussed in terms of visual projections differentially affecting alpha and gamma motoneurons.     

Increases in alpha oscillatory power reflect an active retinotopic mechanism for distracter suppression during sustained visuospatial attention

Human electrophysiological (EEG) studies have demonstrated the involvement of alpha band (8- to 14-Hz) oscillations in the anticipatory biasing of attention. In the context of visual spatial attention within bilateral stimulus arrays, alpha has exhibited greater amplitude over parietooccipital cortex contralateral to the hemifield required to be ignored, relative to that measured when the same hemifield is to be attended. Whether this differential effect arises solely from alpha desynchronization (decreases) over the "attending" hemisphere, from synchronization (increases) over the "ignoring" hemisphere, or both, has not been fully resolved. This is because of the confounding effect of externally evoked desynchronization that occurs involuntarily in response to visual cues. Here, bilateral flickering stimuli were presented simultaneously and continuously over entire trial blocks, such that externally evoked alpha desynchronization is equated in precue baseline and postcue intervals. Equivalent random letter sequences were superimposed on the left and right flicker stimuli. Subjects were required to count the presentations of the target letter "X" at the cued hemifield over an 8-s period and ignore the sequence in the opposite hemifield. The data showed significant increases in alpha power over the ignoring hemisphere relative to the precue baseline, observable for both cue directions. A strong attentional bias necessitated by the subjective difficulty in gating the distracting letter sequence is reflected in a large effect size of 2.1 (eta2 = 0.82), measured from the attention x hemisphere interaction. This strongly suggests that alpha synchronization reflects an active attentional suppression mechanism, rather than a passive one reflecting "idling" circuits.     

Topographic and directional organization of visual motion inputs for the initiation of horizontal and vertical smooth-pursuit eye movements in monkeys

1. The goal of our study was to determine the properties of the visual inputs for pursuit eye movements. In a previous study we presented horizontal target motion along the horizontal meridian and showed that targets were more effective if they moved across the center of the visual field. We have now analyzed the topographic weighting of the inputs for pursuit in greater detail, using targets that moved in all directions and across a wide area of the visual field. 2. Monkeys were rewarded for tracking targets that started at 48 positions in the visual field. The initial positions were spaced equally around 4 circles that were centered at the position of fixation and had radii of 3, 6, 9, and 12 degrees. Targets moved horizontally or vertically at 30 degrees/s. We measured the smooth eye acceleration in the first 80 ms after the initiation of pursuit, before there had been time for visual feedback to affect the position or velocity of the retinal images from the target. 3. For both horizontal and vertical target motion, there were major differences between the early and late intervals in the first 80 ms of pursuit. In the first 20 ms eye acceleration was largely independent of initial target position. In later intervals eye acceleration decreased sharply as a function of initial target eccentricity. The later intervals also showed a pronounced toward/away asymmetry such that the initiation of pursuit was more vigorous for target motion toward than for motion away from the horizontal or vertical meridian. 4. Comparison of the topographic organization of the middle temporal visual area (MT) with our data on pursuit suggests that the topography of cortical maps is smoothed when the visual signals are transmitted to the pursuit system. For example, the superior visual hemifield is underrepresented in cortical motion processing areas, but target motion in the superior and inferior visual hemifields is equally effective for the initiation of pursuit. 5. We investigated the directional organization of the visual inputs for pursuit by presenting targets that started at 6 degrees eccentric and moved in 16 different directions. Horizontal target motion always evoked larger eye accelerations than did vertical target motion. Target motion in oblique directions evoked intermediate values of eye acceleration. 6. Our data show two classes of variation in pursuit performance. First, some subjects showed ideosyncratic variations that were restricted to one hemifield or one direction of target motion. We attribute these variations to differences among subjects in the physiology of visual pathways.(ABSTRACT TRUNCATED AT 400 WORDS)     

Evolution of average evoked potentials in cats during conditioning before and after tegmental lesions.

Sensory-specific and modality nonspecific average evoked potentials (AEPs) were recorded from the cortical areas and subcortical structures in two groups of cats: (1) normal cats which after conditioning were subjected to brainstem lesions and reconditioning; (2) cats with brainstem lesions and subsequent conditioning. A new waveshape of the visually evoked potenials developed in the visual cortex in the course of conditioning to light flashes (LF) in both groups of cats. In normal cats, a new component, with a peak latency from 80-100 msec and with reversed polarity, was observed in place of the late, longlasting, component of the preconditioning AEP. The latency of the new component is longer in cats with brainstem lesions. The first signs of waveform modification occurred relatively early in the process of learning, well before the animal learned to react consistently to the conditioned stimulus. Also, there seemed to be no clear relationship between the modified waveshapes of the AEPs and performance level during a particular session of conditioning. The modifications did not depend on habituation to the long-lasting exposure to light flashes presented alone. The new waveshape was preserved to a variable degree after brainstem lesions, that is, reconditioning never started with a potential characteristic for a naive animal. AEPs to licks in the auditory cortex, also changed during conditioning, although this modification was not so evident as in visual responses. In contrast to evident modifications of evoked responses in sensory specific structures during the process of conditiong, there were very small if any, changes in modality nonspecific structures, including brainstem reticular formation. Modality nonspecific responses were obtained from the brainstem reticular formation and motor cortex to light flashes and clicks, from the visual cortex to auditory stimuli and from the auditory cortex to light flashes. Only poorly developed evoked responses could be detected in the motor-sensory cortex during conditioning to light flashes although rhythmic EEG activity related to presentation of the conditioned stimulus (CS) was observed from this arena - thus indicating that they were not the same phenomena.     

Cortical activation in hemianopia after stroke

Changes in neuronal activity of the visual cortex have been described in patients with hemianopia. The anatomical areas that are involved in neuroplastic changes have not been studied in a larger group of stroke patients with a homogenous structural pathology of the visual cortex. Brain activation was measured in 13 patients with a single ischemic lesion of the striate cortex and partially recovered hemianopia and in 13 age-matched control subjects using blood oxygen level dependent (BOLD) functional magnetic resonance imaging (fMRI). Differences in activation between rest and visual hemifield stimulation were assessed with statistical parametric mapping using group and multi-group studies. In normal subjects, the most significant activation was found in the contralateral primary visual cortex (area 17) and bilaterally in the extrastriate cortex (areas 18 and 19). In patients, these areas were also activated when the intact hemifield was stimulated. During stimulation of the hemianopic side, bilateral activation was seen within the extrastriate cortex, stronger in the ipsilateral (contralesional) hemisphere. Stimulation of the hemianopic visual field is associated with ipsilateral activation of the extrastriate visual cortex. This pattern of activation suggests extensive neuronal plasticity within the visual cortex after postgeniculate ischemic lesions and may have implications for therapeutic interventions.     

Study of target and non-target interplay in spatial attention task

Selective visual attention is the ability to selectively pay attention to the targets while inhibiting the distractors. This paper aims to study the targets and non-targets interplay in spatial attention task while subject attends to the target object present in one visual hemifield and ignores the distractor present in another visual hemifield. This paper performs the averaged evoked response potential (ERP) analysis and time-frequency analysis. ERP analysis agrees to the left hemisphere superiority over late potentials for the targets present in right visual hemifield. Time-frequency analysis performed suggests two parameters i.e. event-related spectral perturbation (ERSP) and inter-trial coherence (ITC). These parameters show the same properties for the target present in either of the visual hemifields but show the difference while comparing the activity corresponding to the targets and non-targets. In this way, this study helps to visualise the difference between targets present in the left and right visual hemifields and, also the targets and non-targets present in the left and right visual hemifields. These results could be utilised to monitor subjects’ performance in brain–computer interface (BCI) and neurorehabilitation.     

Both striate cortex and superior colliculus contribute to visual properties of neurons in superior temporal polysensory area of macaque monkey

Although the tectofugal system projects to the primate cerebral cortex by way of the pulvinar, previous studies have failed to find any physiological evidence that the superior colliculus influences visual activity in the cortex. We studied the relative contributions of the tectofugal and geniculostriate systems to the visual properties of neurons in the superior temporal polysensory area (STP) by comparing the effects of unilateral removal of striate cortex, the superior colliculus, or of both structures. In the intact monkey, STP neurons have large, bilateral receptive fields. Complete unilateral removal of striate cortex did not eliminate visual responses of STP neurons in the contralateral visual hemifield; rather, nearly half the cells still responded to visual stimuli in the hemifield contralateral to the lesion. Thus the visual properties of STP neurons are not completely dependent on the geniculostriate system. Unilateral striate lesions did affect the response properties of STP neurons in three ways. Whereas most STP neurons in the intact monkey respond similarly to stimuli in the two visual hemifields, responses to stimuli in the hemifield contralateral to the striate lesion were usually weaker than responses in the ipsilateral hemifield. Whereas the responses of many STP neurons in the intact monkey were selective for the direction of stimulus motion or for stimulus form, responses in the hemifield contralateral to the striate lesion were not selective for either motion or form. Whereas the median receptive field in the intact monkey extended 80 degrees into the contralateral visual field, the receptive fields of cells with responses in the contralateral field that survived the striate lesions had a median border that extended only 50 degrees into the contralateral visual field. Removal of both striate cortex and the superior colliculus in the same hemisphere abolished the responses of STP neurons to visual stimuli in the hemifield contralateral to the combined lesion. Nearly 80% of the cells still responded to visual stimuli in the hemifield ipsilateral to the lesion. Unilateral removal of the superior colliculus alone had only small effects on visual responses in STP. Receptive-field size and visual response strength were slightly reduced in the hemifield contralateral to the collicular lesion. As in the intact monkey, selectivity for stimulus motion or form were similar in the two visual hemifields. We conclude that both striate cortex and the superior colliculus contribute to the visual responses of STP neurons. Striate cortex is crucial for the movement and stimulus specificity of neurons in STP.(ABSTRACT TRUNCATED AT 400 WORDS)     

Life-span alterations in visually evoked potentials and inhibitory function

Visually evoked potentials (VEPs) elicited by patterned and unpatterned flashes were recorded from 211 healthy males aged 4–90 years. A measure of similarity between the two kinds of VEPs was obtained by correlating the digital values comprising the two waveforms. Across the life-span, correlations followed a U-shaped curve; patterned and unpatterned flash VEPs were most alike for the youngest and oldest subjects. This age effect, localized to scalp areas overlying visual cortex, is compatible with a concept of reduced inhibitory functioning within the visual systems of the young and the old. At central scalp, patterned and unpatterned flash VEP waveforms were more effectively differentiated by the right hemisphere. This observation agrees with reports that the right hemisphere specializes in analyses of spatial material.     

Sustained division of spatial attention to multiple locations within one hemifield

Attending to a location in space significantly improves stimulus perception at that location. Everyday experience requires the deployment of attention to multiple objects at different locations. Recent empirical evidence suggests that the "beam" of attention can be divided between non-contiguous areas of the visual field. Whether this is only possible when stimuli are presented in different hemifields and harder, if not impossible, when stimuli are in the same hemifield is an ongoing debate. Here we use an electrophysiological measure of sustained attentional resource allocation (the steady-state visual evoked potential, SSVEP) to address this question. In combination with behavioural data we demonstrate that splitting the attentional "beam" is in principle possible within one hemifield. However, results showed that task performance was in general lower for same-hemifield presentation as opposed to our previous study with different-hemifield presentation [M.M. Müller, P. Malinowski, T. Gruber, S.A. Hillyard, Sustained division of the attentional spotlight, Nature 424 (2003) 309-312]. SSVEP amplitude showed a mixed pattern of results for stimuli presented in the upper versus lower quadrant of the left visual hemifield under conditions of attending to two separated locations. Results are discussed in the light of the bilateral distribution advantage hypothesis and differences in stimulus salience between the upper and lower visual field.     

Life-span changes in visually evoked potentials at central scalp

Three intensities of patterned flash were used to elicit visually evoked potentials (VEPs) from central scalp of 220 healthy males aged 4–90 years. Intensities were based on visual thresholds. VEP components from children and oldsters were generally larger than those of subjects of intermediate ages and, in response to increased flash intensity, demonstrated greater amplitude enhancement. This pattern of age changes is thought to reflect decreased cortical inhibition in the young and the old with inhibitory capacity being maximal during late adolescent and early adult years. Hemispheric asymmetries were observed. VEP amplitudes from the right hemisphere were larger than those from the left and demonstrated a greater amplitude augmentation to brighter flashes.     

Life-span changes in visually evoked potentials at central scalp

Three intensities of patterned flash were used to elicit visually evoked potentials (VEPs) from central scalp of 220 healthy males aged 4–90 years. Intensities were based on visual thresholds. VEP components from children and oldsters were generally larger than those of subjects of intermediate ages and, in response to increased flash intensity, demonstrated greater amplitude enhancement. This pattern of age changes is thought to reflect decreased cortical inhibition in the young and the old with inhibitory capacity being maximal during late adolescent and early adult years. Hemispheric asymmetries were observed. VEP amplitudes from the right hemisphere were larger than those from the left and demonstrated a greater amplitude augmentation to brighter flashes.     

Illusory conjunctions in visual short‐term memory: Individual differences in corpus callosum connectivity and splitting attention between the two hemifields

Overloading the capacity of visual attention can result in mistakenly combining the various features of an object, that is, illusory conjunctions. We hypothesize that if the two hemispheres separately process visual information by splitting attention, connectivity of corpus callosum—a brain structure integrating the two hemispheres—would predict the degree of illusory conjunctions. In the current study, we assessed two types of illusory conjunctions using a memory‐scanning paradigm; the features were either presented across the two opposite hemifields or within the same hemifield. Four objects, each with two visual features, were briefly presented together followed by a probe‐recognition and a confidence rating for the recognition accuracy. MRI scans were also obtained. Results indicated that successful recollection during probe recognition was better for across hemifields conjunctions compared to within hemifield conjunctions, lending support to the bilateral advantage of the two hemispheres in visual short‐term memory. Age‐related differences regarding the underlying mechanisms of the bilateral advantage indicated greater reliance on recollection‐based processing in young and on familiarity‐based processing in old. Moreover, the integrity of the posterior corpus callosum was more predictive of opposite hemifield illusory conjunctions compared to within hemifield illusory conjunctions, even after controlling for age. That is, individuals with lesser posterior corpus callosum connectivity had better recognition for objects when their features were recombined from the opposite hemifields than from the same hemifield. This study is the first to investigate the role of the corpus callosum in splitting attention between versus within hemifields.     

Flashed pattern-induced activity in the visual system: I. The short latency evoked response recorded from the cat visual cortex

Previous methods for estimating visual acuity have used the visual evoked response conventionally the late visual evoked response components or the steady-state potential. The present experiments were undertaken to evaluate the possible use of short latency flashed pattern evoked responses in estimating pattern dependent activity in the cat visual system. Recordings were made from the skull bone and the dura above the primary visual cortex and intracortically. The visual evoked responses to patterned (checks) and non-patterned light flashes of high intensity and short duration were recorded. The visual evoked response activity recorded from the cortical surface had an onset latency of 14–15 ms. The initial positive-negative potential sequences of the responses were similar for patterned and non-patterned stimuli, however a difference was recorded from 35–40 ms after stimulus. The smallest check size which separated a pattern from a non-pattern VER was in the order of 10 min of arc. The results indicate that the short-latency cortical VER may be used to estimate visual resolution.     

Visual working memory capacity and stimulus categories: a behavioral and electrophysiological investigation

It has recently been suggested that visual working memory capacity may vary depending on the type of material that has to be memorized. Here, we use a delayed match-to-sample paradigm and event-related potentials (ERP) to investigate the neural correlates that are linked to these changes in capacity. A variable number of stimuli (1–4) were presented in each visual hemifield. Participants were required to selectively memorize the stimuli presented in one hemifield. Following memorization, a test stimulus was presented that had to be matched against the memorized item(s). Two types of stimuli were used: one set consisting of discretely different objects (discrete stimuli) and one set consisting of more continuous variations along a single dimension (continuous stimuli). Behavioral results indicate that memory capacity was much larger for the discrete stimuli, when compared with the continuous stimuli. This behavioral effect correlated with an increase in a contralateral negative slow wave ERP component that is known to be involved in memorization. We therefore conclude that the larger working memory capacity for discrete stimuli can be directly related to an increase in activity in visual areas and propose that this increase in visual activity is due to interactions with other, non-visual representations.     

Position selectivity in scene- and object-responsive occipitotemporal regions

Complex visual scenes preferentially activate several areas of the human brain, including the parahippocampal place area (PPA), the retrosplenial complex (RSC), and the transverse occipital sulcus (TOS). The sensitivity of neurons in these regions to the retinal position of stimuli is unknown, but could provide insight into their roles in scene perception and navigation. To address this issue, we used functional magnetic resonance imaging (fMRI) to measure neural responses evoked by sequences of scenes and objects confined to either the left or right visual hemifields. We also measured the level of adaptation produced when stimuli were either presented first in one hemifield and then repeated in the opposite hemifield or repeated in the same hemifield. Although overall responses in the PPA, RSC, and TOS tended to be higher for contralateral stimuli than for ipsilateral stimuli, all three regions exhibited position-invariant adaptation, insofar as the magnitude of adaptation did not depend on whether stimuli were repeated in the same or opposite hemifields. In contrast, object-selective regions showed significantly greater adaptation when objects were repeated in the same hemifield. These results suggest that neuronal receptive fields (RFs) in scene-selective regions span the vertical meridian, whereas RFs in object-selective regions do not. The PPA, RSC, and TOS may support scene perception and navigation by maintaining stable representations of large-scale features of the visual environment that are insensitive to the shifts in retinal stimulation that occur frequently during natural vision.     

Visually triggered K-complexes: a study in New Zealand rabbits

K-complexes are the EEG elements recorded during the state of developing sleep and during slow wave sleep. They are the only EEG components which can be elicited by sensory stimulation during sleep. The peculiarity of New Zealand rabbits to sleep with their eyes open allows the use of visual stimuli to elicit K-complexes. Experiments were performed with three rabbits. For visual stimulation, an elongated screen illuminated by LED flashes was attached to an implant on the animal’s skull. The screen covered 20–120° of the visual field of one eye, and moved with the head during animal motion. One-millisecond flashes (15-s interval) were used during daytime in an illuminated room. Flashes elicited evoked responses, which, during the first stages of sleep, were often accompanied by K-complexes. The induced K-complexes were recorded from electrodes located both above visual and somatosensory areas. Evoked responses to visual stimuli were also recorded from both pairs of electrodes, although they were generated exclusively in the visual cortex. Correlation analysis showed that visual evoked responses and K-complexes induced by this stimulation were generated in visual cortex, and passively spread to the electrodes above the somatosensory area. Investigation of the latencies of induced K-complexes revealed two time windows when these complexes could be seen. Within each window there was no correlation between latency and amplitude of K-complexes. There was also no correlation between amplitudes of the visual evoked responses and K-complexes elicited by these responses. We propose that visual stimulation in light sleep temporarily opens a gate for some independent external signals, which evoke activation of the visual cortex, reflected in K-complexes.     

Cross-Modal Selective Attention Effects on Retinal, Myogenic, Brainstem, and Cerebral Evoked Potentials

Short latency evoked potentials were recorded during a cross-modal selective attention task to evaluate recent proposals that sensory transmission in the peripheral auditory and visual pathways can be modified selectively by centrifugal mechanisms in humans. Twenty young adult subjects attended in turn to either left-ear tones or right-field flashes presented in a randomized sequence, in order to detect infrequent, lower-intensity targets. Attention-related enhancement of longer-latency components, including the visual P105 and the auditory N1/Nd waves and T-complex, showed that subjects were able to adopt a selective sensory set toward either modality. Neither the auditory evoked brainstem potentials nor the early visual components (electroretinogram, occipito-temporal N40, P50, N70 waves) were significantly affected by attention. Measures of retinal B-waves were significantly reduced in amplitude when attention was directed to the flashes, but concurrent recordings of eyelid electromyographic activity and the electro-oculogram indicated that this effect may have resulted from contamination of the retinal recordings by blink microreflex activity. A trend toward greater positivity in the 15-50 ms latency range for auditory evoked potentials to attended tones was observed. These results provide further evidence that the earliest levels of sensory transmission are unaffected by cross-modal selective attention, but that longer latency exogenous and endogenous potentials are enhanced to stimuli in the attended modality.     

Prefrontal modulation of visual processing in humans

Single neuron, evoked potential and metabolic techniques show that attention influences visual processing in extrastriate cortex. We provide anatomical, electrophysiological and behavioral evidence that prefrontal cortex regulates neuronal activity in extrastriate cortex during visual discrimination. Event-related potentials (ERPs) were recorded during a visual detection task in patients with damage in dorsolateral prefrontal cortex. Prefrontal damage reduced neuronal activity in extrastriate cortex of the lesioned hemisphere. These electrophysiological abnormalities, beginning 125 ms after stimulation and lasting for another 500 ms, were accompanied by behavioral deficits in detection ability in the contralesional hemifield. The results provide evidence for intrahemispheric prefrontal modulation of visual processing.