Visual, saccade-related, and cognitive activation of single neurons in monkey extrastriate area V3A

Area V3A is an extrastriate visual area that provides a major input to parietal cortex. To identify the sensory, saccade-related, and cognitive signals carried by V3A neurons, we recorded from single units in alert monkeys during performance of fixation and memory guided saccade tasks. We found that visual responses to stationary stimuli in area V3A were affected by the behavioral relevance of the stimulus. The amplitude of the visual response differed between the memory-guided saccade task, in which the monkey had to use the information provided by the stimulus to guide its behavior, and the fixation task. For 18% (29/163) of V3A neurons, the response was significantly enhanced in the memory-guided saccade task as compared with that in the fixation task. For 8% (13/163) of V3A neurons, the amplitude of response in the memory-guided saccade task was significantly suppressed. We also observed task-related modulation of activity prior to stimulus onset. Among the V3A neurons (37/163) that showed significant differences between tasks in prestimulus activity, the majority (89%; 33/37) showed greater prestimulus activity in the memory-guided saccade task. Task-related increases in prestimulus activity in the memory-guided saccade task were not always matched by increases in the sensory response, indicating that visual responses and prestimulus activity can be modulated independently. Activity in the memory period was suppressed compared with prestimulus activity for 83% (49/59) of the V3A neurons that showed a significant difference in activity (59/197) between these two epochs. For some neurons, memory-period activity dropped even below the baseline level in the fixation task, indicating that there may be an active suppression mechanism. Many V3A neurons (75%, 148/197) also had activity in the saccade epoch. This activity was most prominent immediately after the saccade. Postsaccadic activity was observed even when testing was carried out in total darkness, indicating that this activity reflects, at least in part, extraretinal signals and is not simply a response to visual reafference. These results indicate that several kinds of signals are carried by single neurons in extrastriate area V3A. Moreover, activity in V3A is subject to modulation by extraretinal factors, including attention, anticipation, memory, and saccadic eye movements.     

Saccadic eye movements to visual and auditory targets

 Recent neurophysiological studies of the saccadic ocular motor system have lent support to the hypothesis that this system uses a motor error signal in retinotopic coordinates to direct saccades to both visual and auditory targets. With visual targets, the coordinates of the sensory and motor error signals will be identical unless the eyes move between the time of target presentation and the time of saccade onset. However, targets from other modalities must undergo different sensory-motor transformations to access the same motor error map. Because auditory targets are initially localized in head-centered coordinates, analyzing the metrics of saccades from different starting positions allows a determination of whether the coordinates of the motor signals are those of the sensory system. We studied six human subjects who made saccades to visual or auditory targets from a central fixation point or from one at 10° to the right or left of the midline of the head. Although the latencies of saccades to visual targets increased as stimulus eccentricity increased, the latencies of saccades to auditory targets decreased as stimulus eccentricity increased. The longest auditory latencies were for the smallest values of motor error (the difference between target position and fixation eye position) or desired saccade size, regardless of the position of the auditory target relative to the head or the amplitude of the executed saccade. Similarly, differences in initial eye position did not affect the accuracy of saccades of the same desired size. When saccadic error was plotted as a function of motor error, the curves obtained at the different fixation positions overlapped completely. Thus, saccadic programs in the central nervous system compensated for eye position regardless of the modality of the saccade target, supporting the hypothesis that the saccadic ocular motor system uses motor error signals to direct saccades to auditory targets. Received: 8 September 1995 / Accepted: 22 November 1996     

Perisaccadic mislocalization without saccadic eye movements

Despite frequent saccadic gaze shifts we perceive the surrounding visual world as stable. It has been proposed that the brain uses extraretinal eye position signals to cancel out saccade-induced retinal image motion. Nevertheless, stimuli flashed briefly around the onset of a saccade are grossly mislocalized, resulting in a shift and, under certain conditions, an additional compression of visual space. Perisaccadic mislocalization has been related to a spatio-temporal misalignment of an extraretinal eye position signal with the corresponding saccade. Here, we investigated perceptual mislocalization of human observers both in saccade and fixation conditions. In the latter conditions, the retinal stimulation during saccadic eye movements was simulated by a fast saccade-like shift of the stimulus display. We show that the spatio-temporal pattern of both the shift and compression components of perceptual mislocalization can be surprisingly similar before real and simulated saccades. Our findings suggest that the full pattern of perisaccadic mislocalization can also occur in conditions which are unlikely to involve changes of an extraretinal eye position signal. Instead, we suggest that, under the conditions of our experiments, the arising difficulty to establish a stable percept of a briefly flashed stimulus within a given visual reference frame yields mislocalizations before fast retinal image motion. The availability of visual references appears to exert a major influence on the relative contributions of shift and compression components to mislocalization across the visual field.     

Control of saccade initiation in a countermanding task using visual and auditory stop signals

We examined inhibitory control in an oculomotor countermanding task, where the primary task required a saccadic eye movement be made to a target and a less-frequent secondary task required that the movement be halted. Previous studies have used a visual stimulus presented centrally on the fovea as the signal to stop or countermand a saccade. In these previous studies, there are at least two possible sources of saccadic inhibition: (1) sensory stimulation at the fovea can elicit a bottom-up mechanism, where a visual transient signal can delay or inhibit the developing saccade command; and (2) information based on the task instruction can be used to initiate a top-down mechanism to halt the movement. In the present study, we used both visual and auditory stop signals to test the hypothesis that the bottom-up mechanism is activated only after presentation of a foveal visual stop signal. Subjects were instructed first to look at a central spot and then to look to an eccentric visual target that appeared randomly to the left or right of center. On about one-third of the trials, a stop signal was presented. Three types of stop signals were used with equal probability: a broad-band noise burst (auditory), a central fixation spot (visual), and a combination of the auditory and visual stimuli (combined). Saccadic reaction time and stop-signal accuracy were used to calculate stop signal reaction time (SSRT), an estimate of the time required to inhibit the eye movement. Mean SSRT was longer for the auditory stop signals (201 ms) than for the signals with a foveal visual component (visual 113 ms; combined 91 ms). We conclude that a foveal visual stop signal in an oculomotor countermanding task changes the measure of inhibitory control to reflect not only inhibitory processes but also the sensory information afforded by stimulation at the fovea.     

The time course of perisaccadic receptive field shifts in the lateral intraparietal area of the monkey

Neurons in the lateral intraparietal area of the monkey (LIP) have visual receptive fields in retinotopic coordinates when studied in a fixation task. However, in the period immediately surrounding a saccade these receptive fields often shift, so that a briefly flashed stimulus outside the receptive field will drive the neurons if the eye movement will bring the spatial location of that vanished stimulus into the receptive field. This is equivalent to a transient shift of the retinal receptive field. The process enables the monkey brain to process a stimulus in a spatially accurate manner after a saccade, even though the stimulus appeared only before the saccade. We studied the time course of this receptive field shift by flashing a task-irrelevant stimulus for 100 ms before, during, or after a saccade. The stimulus could appear in receptive field as defined by the fixation before the saccade (the current receptive field) or the receptive field as defined by the fixation after the saccade (the future receptive field). We recorded the activity of 48 visually responsive neurons in LIP of three hemispheres of two rhesus monkeys. We studied 45 neurons in the current receptive field task, in which the saccade removed the stimulus from the receptive field. Of these neurons 29/45 (64%) showed a significant decrement of response when the stimulus appeared 250 ms or less before the saccade, as compared with their activity during fixation. The average response decrement was 38% for those cells showing a significant (P < 0.05 by t-test) decrement. We studied 39 neurons in the future receptive field task, in which the saccade brought the spatial location of a recently vanished stimulus into the receptive field. Of these 32/39 (82%) had a significant response to stimuli flashed for 100 ms in the future receptive field, even 400 ms before the saccade. Neurons never responded to stimuli moved by the saccade from a point outside the receptive field to another point outside the receptive field. Neurons did not necessarily show any saccadic suppression for stimuli moved from one part of the receptive field to another by the saccade. Stimuli flashed <250 ms before the saccade-evoked responses in both the presaccadic and the postsaccadic receptive fields, resulting in an increase in the effective receptive field size, an effect that we suggest is responsible for perisaccadic perceptual inaccuracies.     

Discharge patterns of neurons in the rostral superior colliculus of cat: activity related to fixation of visual and auditory targets

Neurons in the rostral superior colliculus (SC) of alert cats exhibit quasi-sustained discharge patterns related to the fixation of visual targets. Because some SC neurons also respond to auditory stimuli, we investigated whether there is a population of neurons in the rostral SC which is active in relation to fixation of both auditory and visual targets. We identified cells which were active with visual fixation and which continued to discharge if the fixation stimulus was briefly extinguished. The population of neurons exhibited similar discharge characteristics when the fixation stimulus was auditory. Few neurons were significantly more active during fixation of visual targets than during fixation of auditory targets. Most fixation neurons showed a diminished discharge rate during spontaneous (self-generated) saccadic eye movements away from a visual fixation stimulus, regardless of the direction of the saccade. this diminished discharge rate (or pause) typically began, on average, 12.2 ms before saccade onset and the duration of the pause was Ionger than the duration of the saccade. These observations are consistent with the hypothesis that increased discharge of these neurons is related to active fixation and that reductions in their activity are important for the generation of saccades. However, the lack of a precise relationship between pause duration and saccade duration implies that these neurons would be unlikely to project directly to the saccadic burst generator. The mean interval from the beginning of the pauses of fixation neurons to be beginning of the saccades away from fixation targets is also shorter than has been found in brainstem omnipause neurons. By analogy with the concept of a receptive field, agaze position error field depicts the range of gaze position error for which a cell is active. Although fixation neurons appear to encode the magnitude and direction of the error between visual targets and the visual axis, visual error fields at the end of fixating eye movements were significantly larger than those at stimulus onset. For auditory stimuli, this difference was not significant. These observations are compatible with a number of recent experiments indicating that neural signals of eye position are damped or delayed with respect to current eye position.     

Collicular involvement in a saccadic colour discrimination task

Summary We have recorded the neural activity of single superior colliculus (SC) neurons in monkeys engaged in a saccadic target/nontarget discrimination task based on a colour cue. Since correct execution of this task probably depends on cortical signal processing, our experiments are of interest for getting a better insight in the problem of how cortical and subcortical signals, relevant for the visual guidance of saccades, are combined. The experiments were designed to distinguish between two extreme possibilities: 1) The crucial cortical signal affects the saccadic system at or above the level of the SC movement-related cells (serial hypothesis); 2) The colour-based target information bypasses the motor colliculus and affects the saccadic system at a level more downstream (bypass hypothesis). Under conditions where the saccadic system had to select a green target stimulus and to ignore the red nontarget spot, the saccade-related activity in SC visuomotor neurons remained as tightly coupled to the metrics of the saccade as it was in a simple spot-detection task. Since the saccade-related activity of these cells appeared to be based on colour information, we conclude that our data corroborate the serial hypothesis. The initial activity after stimulus onset appeared to be colour nonopponent in all neurons. In some cells the neural activity was quantitatively slightly different for the green target and the red nontarget. Since these minor differences were colour rather than motor response dependent, they were probably not part of the target-selection process. These data suggest the possibility that the decision as to which saccade should be made was largely imposed upon the SC visuomotor cells by an external source. We discuss various possibilities for the origin of the putative intervening signal which orders a saccade by causing a burst in the appropriate SC visuomotor neurons.     

Long-term voltage-sensitive dye imaging reveals cortical dynamics in behaving monkeys

A novel method of chronic optical imaging based on new voltage-sensitive dyes (VSDs) was developed to facilitate the explorations of the spatial and temporal patterns underlying higher cognitive functions in the neocortex of behaving monkeys. Using this system, we were able to explore cortical dynamics, with high spatial and temporal resolution, over period of 相似文献   

Saccade target selection in the superior colliculus during a visual search task

Because real-world scenes typically contain many different potential objects of interest, selecting one goal from many is clearly a fundamental problem faced by the saccadic system. We recorded from visual, movement, and visuo-movement (VM) neurons in the superior colliculus (SC) of monkeys performing a reaction-time visual-search task requiring them to make saccades to an odd-colored target presented with distractors. First, we compared the responses of SC neurons in search with their responses when a single target was presented without distractors (single-stimulus task). Consistent with earlier reports, initial visual activity was smaller in search than in the single-stimulus task, while movement-related activity in the two tasks was comparable. Further experiments showed that much of the reduction in the initial visual response during search was due to lateral inhibition, although a top-down task-related component was also evident. Although the initial visual activity did not discriminate the target from the distractors, some neurons showed a biphasic pattern of visual activity. In VM burst neurons, the second phase of this activity was significantly larger when the target, rather than a distractor, was in the response field. We traced the time course of target/distractor discrimination using receiver operating characteristic (ROC) analysis and found that VM burst neurons, VM prelude neurons, and pure movement neurons discriminated the target from distractors before saccade onset but that phasic and tonic pure visual neurons did not. We also examined the relationship between target/distractor discrimination time and saccade latency. Discrimination in VM burst neurons having a biphasic pattern of visual activity and in many VM prelude neurons occurred after a consistent delay that did not depend on saccade latency, suggesting that these neurons are involved in target selection as well as movement initiation. In contrast, VM burst neurons lacking a biphasic pattern of visual activity, pure movement neurons, and a subset of VM prelude neurons discriminated the target at a time that was well correlated with saccade latency, suggesting that this latter group of neurons is involved in triggering movement execution but not in target selection. Thus a mix of signals likely related to target selection and movement initiation co-exists in different groups of SC neurons. This suggests that certain types of SC neurons participate in the target selection process and that the SC as a whole represents a gateway for target selection signals to be converted into a saccadic command.     

Temporal processing of saccade targets in parietal cortex area LIP during visual search

We studied whether the lateral intraparietal (LIP) area-a subdivision of parietal cortex anatomically interposed between visual cortical areas and saccade executive centers-contains neurons with activity patterns sufficient to contribute to the active process of selecting saccade targets in visual search. Visually responsive neurons were recorded while monkeys searched for a color-different target presented concurrently with seven distractors evenly distributed in a circular search array. We found that LIP neurons initially responded indiscriminately to the presentation of a visual stimulus in their response fields, regardless of its feature and identity. Their activation nevertheless evolved to signal the search target before saccade initiation: an ideal observer could reliably discriminate the target from the individual activation of 60% of neurons, on average, 138 ms after stimulus presentation and 26 ms before saccade initiation. Importantly, the timing of LIP neuronal discrimination varied proportionally with reaction times. These findings suggest that LIP activity reflects the selection of both the search target and the targeting saccade during active visual search.     

The effects of illusory line motion on incongruent saccades: implications for saccadic eye movements and visual attention

A complex neural problem must be solved before a voluntary eye movement is triggered away from a stimulus (antisaccade). The location code activated by a stimulus must be internally translated into an appropriate signal to direct the eyes into the opposite visual field, while the reflexive tendency to look directly at the stimulus must be suppressed. No doubt these extra processes contribute to the ubiquitous slowing of antisaccades. However, there is no consensus on the cognitive mechanisms that contribute to the antisaccade programme. Visual attention is closely associated with the generation of saccadic eye movements and it has been shown that attention will track an illusion of line motion. A series of experiments combined this illusion with a saccadic eye movement that was congruent (i.e. directed towards), or incongruent with (i.e. direct away from), a peripheral target. Experiment 1 showed that congruent saccades had faster reaction times than incongruent saccades. In contrast, Experiments 2 and 3 demonstrated that, with illusory line motion, incongruent saccades now had faster reaction times than congruent saccades. These findings demonstrate that an illusory phenomenon can accelerate the processing of an incongruent relative to a congruent saccade.     

Target selection for saccadic eye movements: direction-selective visual responses in the superior colliculus.

We investigated the role of the superior colliculus (SC) in saccade target selection in rhesus monkeys who were trained to perform a direction-discrimination task. In this task, the monkey discriminated between opposed directions of visual motion and indicated its judgment by making a saccadic eye movement to one of two visual targets that were spatially aligned with the two possible directions of motion in the display. Thus the neural circuits that implement target selection in this task are likely to receive directionally selective visual inputs and be closely linked to the saccadic system. We therefore studied prelude neurons in the intermediate and deep layers of the SC that can discharge up to several seconds before an impending saccade, indicating a relatively high-level role in saccade planning. We used the direction-discrimination task to identify neurons whose prelude activity "predicted" the impending perceptual report several seconds before the animal actually executed the operant eye movement; these "choice predicting" cells comprised approximately 30% of the neurons we encountered in the intermediate and deep layers of the SC. Surprisingly, about half of these prelude cells yielded direction-selective responses to our motion stimulus during a passive fixation task. In general, these neurons responded to motion stimuli in many locations around the visual field including the center of gaze where the visual discriminanda were positioned during the direction-discrimination task. Preferred directions generally pointed toward the location of the movement field of the SC neuron in accordance with the sensorimotor demands of the discrimination task. Control experiments indicate that the directional responses do not simply reflect covertly planned saccades. Our results indicate that a small population of SC prelude neurons exhibits properties appropriate for linking stimulus cues to saccade target selection in the context of a visual discrimination task.     

Direction-selective visual responses in macaque superior colliculus induced by behavioral training

In a previous report, we described a heretofore undetected population of neurons in the intermediate and deep layers of the monkey superior colliculus (SC) that yielded directionally selective visual responses to stimuli presented within the central 4 degrees of the visual field. We observed these neurons in three monkeys that had been extensively trained to perform a visual direction discrimination task in this region of the visual field. The task required the monkeys to report the perceived direction of motion by making a saccadic eye movement to one of two targets aligned with the two possible directions of motion. We hypothesized that these neurons reflect a learned association between visual motion direction and saccade direction formed through extensive training on the direction discrimination task. We tested this hypothesis by searching for direction-selective visual responses in two monkeys that had been trained to perform a similar motion discrimination task in which the direction of stimulus motion was dissociated from the direction of the operant saccade. Strongly directional visual responses were absent in these monkeys, consistent with the notion that extensive training can induce highly specific visual responses in a subpopulation of SC neurons.     

Saccadic impairments in Huntington's disease

Huntington’s disease (HD), a progressive neurological disorder involving degeneration in basal ganglia structures, leads to abnormal control of saccadic eye movements. We investigated whether saccadic impairments in HD (N = 9) correlated with clinical disease severity to determine the relationship between saccadic control and basal ganglia pathology. HD patients and age/sex-matched controls performed various eye movement tasks that required the execution or suppression of automatic or voluntary saccades. In the “immediate” saccade tasks, subjects were instructed to look either toward (pro-saccade) or away from (anti-saccade) a peripheral stimulus. In the “delayed” saccade tasks (pro-/anti-saccades; delayed memory-guided sequential saccades), subjects were instructed to wait for a central fixation point to disappear before initiating saccades towards or away from a peripheral stimulus that had appeared previously. In all tasks, mean saccadic reaction time was longer and more variable amongst the HD patients. On immediate anti-saccade trials, the occurrence of direction errors (pro-saccades initiated toward stimulus) was higher in the HD patients. In the delayed tasks, timing errors (eye movements made prior to the go signal) were also greater in the HD patients. The increased variability in saccadic reaction times and occurrence of errors (both timing and direction errors) were highly correlated with disease severity, as assessed with the Unified Huntington’s Disease Rating Scale, suggesting that saccadic impairments worsen as the disease progresses. Thus, performance on voluntary saccade paradigms provides a sensitive indicator of disease progression in HD. A. Peltsch and A. Hoffman contributed equally.     

Consider the context: Blocked versus interleaved presentation of antisaccade trials

Conclusions about the cognitive and neural requirements of saccade control may differ as a result of stimulus presentation method. This issue was examined in the current study by evaluating behavioral differences in pro- and antisaccade responses among 12 healthy young adults as a function of task presentation method, length of cue-to-target interval, and previous trial type. A 1-s cue-to-target interval fostered goal neglect, indicated by an increase in uncorrected errors and reaction times for "error" saccades. There was also a strong relationship between speed of visual orienting (prosaccade latencies) and failed inhibition (antisaccade errors) for the simultaneous condition. Interestingly, only the simultaneous condition produced task switch costs (on saccade latencies and error response percentages). The saccadic task presentation method, therefore, can influence conclusions about the cognitive operations supporting successful performance.     

Combined auditory and visual stimuli facilitate head saccades in the barn owl (Tyto alba)

The barn owl naturally responds to an auditory or visual stimulus in its environment with a quick head turn toward the source. We measured these head saccades evoked by auditory, visual, and simultaneous, co-localized audiovisual stimuli to quantify multisensory interactions in the barn owl. Stimulus levels ranged from near to well above saccadic threshold. In accordance with previous human psychophysical findings, the owl's saccade reaction times (SRTs) and errors to unisensory stimuli were inversely related to stimulus strength. Auditory saccades characteristically had shorter reaction times but were less accurate than visual saccades. Audiovisual trials, over a large range of tested stimulus combinations, had auditory-like SRTs and visual-like errors, suggesting that barn owls are able to use both auditory and visual cues to produce saccades with the shortest possible SRT and greatest accuracy. These results support a model of sensory integration in which the faster modality initiates the saccade and the slower modality remains available to refine saccade trajectory.     

Correlates of motor planning and postsaccadic fixation in the macaque monkey lateral geniculate nucleus

There is significant controversy regarding the ability of the primate visual system to construct stable percepts from a never-ending stream of brief fixations and rapid saccadic eye movements. In this study, we examined the timing and occurrence of perisaccadic modulation of LGN single-unit activity in awake-behaving macaque monkeys while they made spontaneous saccades in the dark and made visually guided saccades to discrete stimuli located outside the receptive field. Our hypothesis was that the activity of LGN cells is modulated by efference copies of motor plans to produce saccadic eye movements and that this modulation depends neither on the presence of feedforward visual information nor on a corollary discharge of signals directing saccadic eye movements. On average, 25% of LGN cells demonstrated significant perisaccadic modulation. This modulation consisted of a moderate suppression of activity that began more than 100 ms prior to the initiation of a saccadic eye movement and continued beyond the termination of the saccadic eye movement. This suppression was followed by a large enhancement of activity after the eyes arrived at the next fixation. Although members of all three LGN relay cell classes (magnocellular, parvocellular, and koniocellular) demonstrated significant saccade-related suppression and enhancement of activity, more cells demonstrated postsaccadic enhancement (25%) than perisaccadic suppression (17%). In no case did the timing of the modulation coincide directly with saccade duration. The degree of modulation observed did not vary with LGN cell class, LGN receptive field center location, center sign (ON-center or OFF-center), or saccade latency or velocity. The time course of modulation did, however, vary with saccade size such that suppression was longer for longer saccades. The fact that activity from a percentage of LGN cells from all cell classes was modulated in relationship to saccadic eye movements in the absence of direct visual stimulation suggests that this modulation is a general phenomenon not tied to specific types of visual stimuli. Similarly, because the onset of the modulation preceded eye movements by more than 100 ms, it is likely that this modulation reflects higher order motor-planning rather than a corollary of mechanisms in direct control of eye movements themselves. Finally, the fact that the largest modulation is a postsaccadic enhancement of activity may suggest that perisaccadic modulations are designed more for the facilitation of visual information processing once the eyes land at a new location than for filtering unwanted visual stimuli.     

Symbolic cue-driven activity in superior colliculus neurons in a peripheral visual choice task

Recent evidence implicates the superior colliculus (SC) in cognitive processes, such as target selection and control of spatial attention, in addition to the execution of saccadic eye movements. We report here the presence of a cognitive response in some cells in the SC in a task that requires the long-term association of spatial location with an arbitrary color. In this study, using a visual choice response task, we demonstrate that visuomotor neurons in the SC were activated by the appearance of a central symbolic cue delivered outside of the visual response fields of the recorded neurons. This procedure ensures that cognitively generated activity in these SC cells is not confounded with modulation of activity from previous visual stimuli that appeared in the response field of the neurons. The experiments suggest that cognitive signals can activate SC cells by themselves instead of only being able to modulate activities already evoked by visual events. Furthermore, a substantial fraction of these cells accurately reflected cue-aligned target selection in advance of saccade initiation. Our results add further support to other studies that have demonstrated that internally generated signals exist in SC cells.     

The influence of motivational salience on saccade latencies

Eye movements provide a direct link to study the allocation of overt attention to stimuli in the visual field. The initiation of saccades towards visual stimuli is known to be influenced by the bottom-up salience of stimuli as well as the motivational context of the task. Here, we asked whether the initiation of saccades is also influenced by the intrinsic motivational salience of a stimulus. Face stimuli were first associated with positive or negative motivational salience through instrumental learning. The same faces served as target stimuli in a subsequent saccade task, in which their motivational salience was no longer task-relevant. Participants performed either voluntary saccades, which required the selection of the saccade target out of two simultaneously presented stimuli (experiment 1), or reactive saccades, where only the target stimulus was presented (experiment 2). We found a specific effect of learned positive stimulus value on the latencies of voluntary saccades: For faces with high versus low positive motivational salience, saccadic latencies were significantly reduced. No such difference was observed for previously punished faces. In contrast, reactive saccades to both previously rewarded and punished faces were unaffected by learned stimulus value. Our findings show for the first time that saccadic preparation is susceptible to the acquired intrinsic motivational salience of visual stimuli. Based on the observation that only voluntary saccades but not reactive saccades were modulated, we conclude that the recruitment of neural processes for target identification is required to allow for an influence of motivational stimulus salience on saccadic preparation.     

Role of striate cortex and superior colliculus in visual guidance of saccadic eye movements in monkeys.

1. We studied the effect of lesions placed in striate cortex or superior colliculus on the detection of visual stimuli and the accuracy of saccadic eye movements. The monkeys (Macaca mulatta) first learned to respond to a 0.25 degrees spot of light flashed for 150-200 ms in one part of the visual field while they were fixating in order to determine if they could detect the light. The monkeys also learned in a different task to make a saccade to the spot of light when the fixation point went out, and the accuracy of the saccades was measured. 2. Following a unilateral partial ablation of the striate cortex in two monkeys they could not detect the spot of light in the resulting scotoma or saccade to it. The deficit was only relative; if we increased the brightness of the stimulus from the usual 11 cd/m2 to 1,700 cd/m2 against a background of 1 cd/m2 the monkeys were able to detect and to make a saccade to the spot of light. 3. Following about 1 mo of practice on the detection and saccade tasks, the monkeys recovered the ability to detect the spots of light and to make saccades to them without gross errors (saccades made beyond an area of +/-3 average standard deviations). Lowering the stimulus intensity reinstated both the detection and saccadic errors...