Lucia Riggio

Reorienting attention across the horizontal and vertical meridians: Evidence in favor of a premotor theory of attention

Stimuli presented in a non-attended location are responded to much slower than stimuli presented in an attended one. The hypotheses proposed to explain this effect make reference to covert movement of attention, hemifield inhibition, or attentional gradients. The experiment reported here was aimed at discriminating among these hypotheses. Subjects were cued to attend to one of four possible stimulus locations, which were arranged either horizontally or vertically, above, below, to the right or left of a fixation point. The instructions were to respond manually as fast as possible to the occurrence of a visual stimulus, regardless of whether it occurred in a cued or in a non-cued location. In 70% of the cued trials the stimulus was presented in the cued location and in 30% in one of the non-cued locations. In addition there were trials in which a non-directional cue instructed the subject to pay attention to all four locations. The results showed that the correct orienting of attention yielded a small but significant benefit; the incorrect orienting of attention yielded a large and significant cost; the cost tended to increase as a function of the distance between the attended location and the location that was actually stimulated; and an additional cost was incurred when the stimulated and attended locations were on opposite sides of the vertical or horizontal meridian. We concluded that neither the hypothesis postulating hemifield inhibition nor that postulating movement of attention with a constant time can explain the data. The hypothesis of an attention gradient and that of attention movements with a constant speed are tenable in principle, but they fail to account for the effect of crossing the horizontal and vertical meridians. A hypothesis is proposed that postulates a strict link between covert orienting of attention and programming explicit ocular movements. Attention is oriented to a given point when the oculomotor programme for moving the eyes to this point is ready to be executed. Attentional cost is the time required to erase one ocular program and prepare the next one.

Orienting of attention and eye movements

According to the premotor theory of attention, the mechanisms responsible for spatial attention and the mechanisms involved in programming ocular saccades are basically the same. The aim of the present experiments was to test this claim. In experiment 1 subjects were presented with a visual display consisting of a fixation point and four boxes arranged horizontally and located above the fixation cross. Two of the boxes were in the left visual hemifield, two in the right. A fifth box was located on the vertical meridian below the fixation cross. Digit cues indicated in which of the upper boxes the imperative stimulus was most likely to appear. Subjects were instructed to direct attention to the cued box and to perform a saccadic eye movement to the lower box on presentation of the imperative stimulus. The trajectory of the saccades deviated contralateral to the hemifield in which the imperative stimulus was presented. This deviation was larger when the hemifield where the imperative stimulus was presented was the cued one. In experiment 2, the visual display consisted of five boxes forming a cross. The central box served as a fixation point. The cue was a small line, linked to the central box, pointing to different directions and indicating where the visual imperative stimulus would appear. In 50% of trials, the imperative stimulus was a visual stimulus presented either in one of the lateral boxes or in the central one. In the remaining 50% of trials, the imperative stimulus was a non-lateralised sound. Half the subjects were instructed to make a saccade to the upper box at the presentation of the visual imperative stimulus and to the lower box at the presentation of the acoustic stimulus. Half the subjects received the opposite instructions. The results confirmed that the saccades deviate contralateral to the hemifield of stimulus presentation in the case of visual imperative stimuli. Most importantly, they showed that the saccades deviate contralateral to the cued hemifield, also in the case of acoustic imperative stimuli. Experiment 3 was similar to experiment 2. It confirmed the results of that experiment and showed that slow ocular drifts, which are observed in the time interval between cue and imperative stimulus presentation, cannot explain the ocular deviations. Taken together, the experiments demonstrate that spatial attention allocation leads to an activation of oculomotor circuits, in spite of eye immobility.

The mirror neuron system and action recognition

Mirror neurons, first described in the rostral part of monkey ventral premotor cortex (area F5), discharge both when the animal performs a goal-directed hand action and when it observes another individual performing the same or a similar action. More recently, in the same area mirror neurons responding to the observation of mouth actions have been also found. In humans, through an fMRI study, it has been shown that the observation of actions performed with the hand, the mouth and the foot leads to the activation of different sectors of Brocas area and premotor cortex, according to the effector involved in the observed action, following a somatotopic pattern which resembles the classical motor cortex homunculus. These results strongly support the existence of an execution-observation matching system (mirror neuron system). It has been proposed that this system is involved in action recognition. Experimental evidence in favor of this hypothesis both in the monkey and humans are shortly reviewed.

Processing abstract language modulates motor system activity

Embodiment theory proposes that neural systems for perception and action are also engaged during language comprehension. Previous neuroimaging and neurophysiological studies have only been able to demonstrate modulation of action systems during comprehension of concrete language. We provide neurophysiological evidence for modulation of motor system activity during the comprehension of both concrete and abstract language. In Experiment 1, when the described direction of object transfer or information transfer (e.g., away from the reader to another) matched the literal direction of a hand movement used to make a response, speed of responding was faster than when the two directions mismatched (an action–sentence compatibility effect). In Experiment 2, we used single-pulse transcranial magnetic stimulation to study changes in the corticospinal motor pathways to hand muscles while reading the same sentences. Relative to sentences that do not describe transfer, there is greater modulation of activity in the hand muscles when reading sentences describing transfer of both concrete objects and abstract information. These findings are discussed in relation to the human mirror neuron system.

Spatial attention and eye movements.

We previously showed that when attention is allocated to the right or left of the fixation point, saccades directed to targets located above or below the fixation point deviate contralateral to the attention locus. In the present study, we examined how general this phenomenon is and whether the amount of saccade deviation depends on the location of attention with respect to that of the saccade target. Three experiments were carried out. In experiment 1 the location of the imperative stimulus was uncued. Its presentation exogenously directed attention to its location. In experiment 2 the location of the imperative stimulus was cued by a central cognitive cue. In this experiment attention was endogenously directed to the imperative stimulus location before its presentation (expectancy paradigm). In experiment 3 all stimulus boxes contained a possible imperative stimulus at the display presentation. A central cue, presented subsequently, indicated which of them had to be used for the saccade. In this experiment attention was endogenously directed to the imperative stimulus, but after its presentation (no-expectancy paradigm). The results showed that, regardless of how attention was directed to the imperative stimulus, the vertical saccades deviated contralateral to the attention location. The deviation was larger when attention was in the upper field and the saccade was directed upward (“same hemifield” condition) than when attention was in the upper field and the saccade was directed downward (“opposite hemifield” condition). The same relationship between the “same hemifield” condition and “opposite hemifield” condition was found when attention was in the lower field. Saccadic reaction times (SRTs) were shortest in experiment 2 and longest in experiment 3. In experiment 2, SRTs of the “same hemifield” condition were significantly longer than those of the “opposite hemifield” condition. Taken altogether, these results strongly support the notion that attention allocation in space leads to an activation of oculomotor circuits, in spite of eye immobility. The possible mechanisms responsible for saccade deviations and for greater saccade deviations when attention is in the same hemifield as the programmed ocular saccade are discussed.

Movements of attention in the three spatial dimensions and the meaning of "neutral" cues.

Six experiments were conducted to examine the effect of various attentional manipulations on reaction time to visual stimuli. The first three experiments compared the responses to stimuli presented in the depth (Experiment 1), along the horizontal (Experiment 2), and vertical (Experiment 3) meridians in a valid condition (stimulus presented in the cued position), an invalid condition (stimulus presented in the alternative position to the cued position) and a neutral condition (no information on stimulus position). The most interesting result was the demonstration that attention can be moved along the sagittal plane in the absence of vergence eye movements and that when attention is focused on a certain point, unattended points between this point and the observer (i.e. near points) are responded faster than unattended points beyond it (i.e. far points). In the frontal plane no asymmetry was found between the responses to unattended points above or below the fixation, whereas a certain, albeit non-constant, advantage was present for unattended stimuli on the right of the fixation point in respect to those on the left of it. The second series of experiments was similar to the first one, except that a new situation was introduced in which the fixation point was cued and stimuli could appear either in correspondence to it or in a peripheral position (invalid condition with attention at the fixation point). The results showed that in this new situation the responses to unattended stimuli are much longer than they are under neutral conditions, and as long as they are under conventional invalid condition. It is suggested that the so called neutral condition is a condition of diffuse attention and an attempt is made to explain it in terms of a premotor theory of attention.

WHAT IS CROSSED IN CROSSED-HAND EFFECTS? *

Abstract In choice RT tasks, when the subject crosses the hands two effects are observed: the direction of spatial compatibility reverses and response latency becomes overall longer. It has been proposed that the location of the response is compared to both its anatomical status (i.e., the hand that emits the response) and the location of the stimulus. When the hands are crossed, the former comparison yields the lengthening of RT while the latter yields the reversal in the direction of spatial compatibility. The present study aimed at testing the relative importance of two locational aspects of the response (i.e., the position of the effector and the position of the response goal) which were confounded in all previous studies. In both experiments the hands were always in anatomical uncrossed position but the position of the response goal might or might not be the same as that of the effector. In experiment 1 the responses were emitted by the index fingers, which were either uncrossed or crossed, whereas in experiment 2 the responses were emitted through two sticks, which again were either uncrossed or crossed. The results replicated both the reversal of spatial compatibility and the lengthening of RT. It was concluded that the effects of crossing the hands are due to the crossing of the response goals.

Differential effects of central and peripheral cues on the reorienting of spatial attention

Abstract Observen can be induced to direct attention to a peripheral location by the use of central and peripheral cues. On the basis of a previous experiment in which central cues were used to orient attention, we proposed a premotor hypothesis of attention (Rizzolatti et al., 1987). which postulates a strict Link between covert orienting of attention and the programming of ocular movements. In the present paper, we tried to extend the soope of the premotor hypothesis by using both central and peripheral cues. The stimulus display consisted of a small fixation cross and four boxes for stimulus presentation. The boxes were arranged horizontally above the lixation point. The subjects were asked to attend to one box or all boxes and to respond manually as fast as possible to the occurrence of a visual stimulus, regardless of its location. When only one box was cued, the imperative stimulus appeared in it 70% of the time, whereas it appeared in one of the non-cued boxes in the remaining cases (10% for each b...

Effects of spatial attention on directional manual and ocular responses

Abstract The aim of the present study was to investigate how spatial attention influences directional manual and saccadic reaction times. Two experiments were carried out. In experiment 1 subjects were instructed to perform pointing responses toward targets that were located either in the same or the opposite hemifield with respect to the hemifield in which an imperative stimulus was presented. In experiment 2, they were instructed to make saccadic or pointing responses. The direction of the responses was indicated by the shape of the imperative stimulus. Reaction time (RT), movement time, and, in experiment 2, saccadic trajectory were measured. The imperative stimulus location was either cued (endogenous attention) or uncued. In the latter case the imperative stimulus presentation attracted attention (exogenous attention). The main results of the experiments were the following: First, exogenous attention markedly decreased the RTs when the required movement was directed toward the imperative stimulus location. This directional effect was much stronger for pointing than for ocular responses. Second, endogenously allocated attention did not influence differentially RTs of pointing responses directed toward or away the attended hemifield. In contrast, endogenous attention markedly favored the saccadic responses when made away from the cued hemifield. Third, regardless of cueing, the direction of movement affected both pointing and saccadic reaction times. Saccadic reaction times were faster when the required movement was directed upward, while manual reaction times were faster when the movement was directed downward. Fourth, lateralized spatial attention deviated the trajectory of the saccades contralateral to the attention location. This pattern of results supports the notion that spatial attention depends on the activation of the same sensorimotor circuits that program actions in space.

Task related modulation of the motor system during language processing

Recent neurophysiological and brain imaging studies have shown that the motor system is involved in language processing. However, it is an open question whether this involvement is a necessary requisite to understand language or rather a side effect of distinct cognitive processes underlying it. In order to clarify this issue we carried out three behavioral experiments, using a go-no go paradigm. Italian verbs expressing hand actions, foot actions or an abstract content served as stimuli. Participants used their right hands to respond. In Experiment 1, in which a semantics decision task with an early delivery of the go signal (during processing language material) was used, slower responses were found for hand action-related verbs than for foot action-related verbs. In Experiment 2, using the same task with either an early or a delayed delivery of the go signal (when language material had been already processed), no difference was found between responses to the two verb categories in the delayed delivery condition. In Experiment 3, in which a lexical decision task with an early delivery of the go signal was used, again no difference between the two verb categories was found. The present findings demonstrate that during language processing the modulation of the motor system crucially occurs while performing a semantics decision task, thus supporting the notion that this involvement is a necessary step to understand language rather than a side effect of upstream cognitive processes.