Article of Kia Nobre
Article of Kia Nobre
Article of Kia Nobre
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TABLE OF CONTENTS
1. Abstract
2. Introduction
3. Temporal orienting
3.1. Behavioural studies
3.1.1. Experiment 1
3.1.2. Experiment 2
3.1.3. Experiment 3
3.1.4. Experiment 4
3.1.5. Experiment 5
3.1.6. Behavioural conclusions
3.2. Event-related potentials
3.3. Brain imaging
4. Comparison of spatial and temporal orienting
4.1. Brain imaging
4.2. Event-related potentials
5. Summary and conclusions
6. Acknowledgments
7. References
1. ABSTRACT
Temporal information is essential for effective studies manipulating foreperiod variability (see (2) for
perception and action in the dynamic environment in which review) have shown that reaction time decreases as a
we exist. However, our ability to use information about function of certainty about the time of occurrence of stimuli
time intervals flexibly to direct attention to an expected requiring detection (3,4,5) or choice decisions (6,7,8). Also,
point in time has until recently been unexplored. Here we psychophysical studies have found increased luminance
report a series of behavioural, neuroimaging and (9), orientation and stereoscopic thresholds (10) when there
electrophysiological experiments that investigate and was uncertainty about the time of stimulus presentation.
define the ability to orient attention in the temporal domain. Performance is improved when stimuli occur at constant
These studies reveal that we are able to orient attention and predictable, as opposed to variable, intervals after a
selectively to different time intervals, enhancing warning signal.
behavioural performance. These effects are mediated by a
left-hemisphere dominant frontal-parietal system, which Such benefits have been interpreted as suggesting
partially overlaps with the networks involved in spatial that warning signals can be used as a time cue to start some
orienting. The optimisation of behaviour by temporal preparatory/anticipatory adjustments (8) and thereby
orienting appears to be achieved via motor-related enhance behavioural performance. Posner and Boies (11)
mechanisms, in contrast to the typical perceptual considered such preparatory processes to be part of the
enhancements produced by spatial attention. From a more alertness component of attention, with the foreperiod
general perspective, these findings illustrate the flexibility between a warning signal and stimulus being likened to a
of attentional functions in the human brain. small-scale vigilance situation in which alertness must be
developed rapidly and maintained briefly in order to
2. INTRODUCTION maximise behavioural performance. Indeed, Wilkinson and
Haines (12) reported similarities in the brain processes
In our dynamic environment, the time of occurrence taking place during the foreperiod and those involved in the
of a stimulus is a crucial determinant of our behaviour performance of prolonged vigilance tasks.
toward it. In 1914 Woodrow (1) demonstrated that the time
interval between a warning signal and a stimulus requiring The question of the time course of preparation
detection (foreperiod) is one of the main factors influencing during the foreperiod, i.e. how long it takes such
the length of reaction time to the stimulus. Since then, preparatory processes to be built up, has also been
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3. TEMPORAL ORIENTING
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3.1.2. Experiment 2
Additional behavioural experiments have
extended the investigation of selective temporal orienting
by testing the roles contributed by specific SOAs,
perceptual discriminations, and response requirements.
Experiment 2 investigated more closely the time course of
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our ability to direct attention to temporal intervals, by alertness literature (e.g. (6)). Overall, the results showed
decreasing the range of time intervals at which the target that behavioural advantages from cueing information
stimulus could appear. It is possible that the short time remain even when there is uncertainty as to the type of
interval in Experiment 1 (600ms) was the inherently response to be given. Manipulating response demands does
optimal foreperiod for behavioural facilitation in this type not completely abolish the effects of temporal orienting.
of task. The result might therefore be a spurious This suggests that the processes underlying the orienting of
occurrence, rather than reflect cognitive control of orienting attention in the temporal domain are not solely linked to
attention to time intervals. We therefore tested subjects preparation of a specific motor response. However, there
(n=9) using the same task but a different time frame (300 may be some aspect of motor preparation or response
and 700ms). The stimuli and procedures were equivalent, timing involved in the effects seen, as the advantages of
except for the range of time intervals. The 9 possible SOAs being cued to expect the target after the short or long
were: 100, 250, 300, 350, 500, 650, 700, 750 and 900ms. interval are not as strong as in the detection experiments.
Subjects were cued to expect either a short (300ms) or long Direct experimental comparisons between detection and
(700ms) delay between cue onset and appearance of the discrimination procedures in the same subject group should
target. yield additional interesting information.
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were predictive or neutral regarding the time interval of the variables can be excluded from participating in temporal
target. The stimuli and procedures were based on orienting. Additional behavioural experimentation may
Experiment 1, but with the following modifications. The help shed light upon the levels at which temporal orienting
informative cue (150ms duration) predicted (80% validity) can exert its effect upon target processing. For example,
target appearance after 600ms. On invalid trials targets measurements of different types of perceptual thresholds
appeared at one of four alternative SOAs: 300, 450, 750 or may indicate whether and which perceptual levels may be
900ms (5% probability each). When the cue was neutral the affected. However, teasing apart what may be multiple
target appeared at one of 21 SOAs: 100, 150, 200, 250, effects of temporal orienting with behavioural
300, 350, 400, 450, 500, 550, 600, 650, 700, 750, 800, 850, manipulations alone can prove to be very difficult, since
900, 950, 1000, 1050, 1100ms (5% probability each, except response variables are influenced by rate-limiting steps and
for extreme SOAs where the probability was 2.5%). The bottlenecks of cognitive processes, which may interfere
task consisted of simple detection of the target stimulus with the readout of some levels of stimulus processing.
(50ms duration). Event-related potentials were therefore used, to provide a
measure of on-line information processing during visual
The results of this experiment (Figure 2e) temporal orienting experiments.
indicate that the behavioural advantages conferred by
attending to a particular point in time are best 3.2. Event-related potentials
conceptualised in terms of behavioural benefits of valid Event-related potentials (ERPs) have provided an
cueing, rather than costs of invalid cueing. The effect can important means of investigating attentional processing in
also be thought to correlate with the absolute probability of the human brain, giving a more complete picture of
target occurrence at a given interval, since the absolute stimulus processing than behavioural measures alone. Their
probability differed only between valid (80%) and neutral high temporal resolution provides information about the
(5%) trials, but not between invalid (5%) and neutral (5%) on-line modulation of brain activity by attention, and about
trials at comparable intervals. The behavioural advantages the level of stimulus processing at which attention acts.
of valid cueing were evident as mainly decreased reaction They also provide a measure of stimulus processing
times to validly cued targets at 600ms, as opposed to without any requirement for the subject to attend or
targets preceded by neutral cues. There was no increase in respond to that stimulus, thus allowing comparison of the
reaction times to invalid as opposed to neutral trials. In fact, processing of both attended and unattended stimuli during
the invalid trials with targets at 750ms actually showed conditions of focused selective attention.
decreased reaction times compared to neutral trials. This
could be due to a number of factors. Firstly, as mentioned Miniussi et al (19) used ERPs to investigate the
earlier, the conditional probability of the target appearing selective orienting of attention to time intervals. The
increases with longer SOAs (17). This is unlikely to be the mechanisms involved in orienting attention in the temporal
sole explanation, since no reaction time advantage was domain were reflected by ERPs elicited by the cues. The
found for the targets following predictive cues at 900ms. mechanisms for modulation of stimulus processing by
These targets have the highest conditional probability in temporal attention were reflected in ERPs elicited by
informative trials, which is higher than the conditional predicted and unpredicted target stimuli. The task used was
probability of targets at equivalent intervals during neutral similar to Experiment 4 (see Figure 1b). Cues were narrow
cueing. Alternatively, the temporal information in the or wide crosses (100ms duration) that predicted the
predictive trials may have spilled over into the adjacent appearance of the target after a short (600ms) or long
interval, due to limits of time estimation. Facilitation of (1400ms) SOA. The target consisted of the brightening of
reaction times to targets in nearby intervals is a consistent the circle surrounding the cue for 100ms. The cues had
pattern in these experiments (see Figure 2). 80% validity, with 10% invalid and 10% catch trials. The
EEG was recorded continuously from 54 electrode sites
3.1.6. Behavioural conclusions positioned according to the 10-20 International System
Together, the behavioural experiments have (20).
shown that predictive (probabilistic) information about the
time interval of target appearance can be used to optimise A behavioural benefit of temporal attention was
behavioural performance. Reaction times to detect or found (Figure 3a), similar to those seen in the purely
discriminate a target are facilitated by valid temporal behavioural experiments. Subjects were significantly faster
cueing, at multiple time frames. The effect survives to detect valid than invalid targets after the short interval.
manipulation of specific SOAs, response requirements and Analysis of ERPs elicited by the cueing stimuli (during the
perceptual judgements. There are many additional cue-target interval) revealed dynamic brain activity linked
parameters that remain to be tested behaviourally, such as to orienting attention in time. Differential processing of
the limits of the time frames that afford temporal orienting, cues predicting short and long intervals modulated the
the minimum difference required between possible CNV (contingent negative variation) component (see (21)).
intervals, the effect of stimulus modalities, etc. Another The CNV is a slow negative voltage change occurring
aspect for investigation is the neural mechanisms for between two stimuli, with the first stimulus being a
selective temporal orienting. The studies so far have shown warning signal and the second stimulus requiring a
that the effect does not depend upon specific motor response. It has been linked to expectancies and motor
programming, and can survive difficult perceptual preparation (see (22,23,24,25)). The CNV was significantly
thresholds. However, neither response nor perceptual accentuated when subjects expected targets at the short
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processing of foveal stimuli may already be optimised by Further fractionation of the processes that can
the visual system, so there may be no need to enhance be involved in orienting attention in time came from
resources further, thus explaining the absence of early VEP comparing brain activation during the two types of
modulation. Temporal orienting may therefore affect invalid trials. Trials in which a target is expected at the
stimulus processing by different, or additional, mechanisms short interval but does not appear may permit subjects to
to the early perceptual modulation seen in studies of spatial re-orient attention voluntarily to the later time point,
attention. thus emphasising top-down, endogenous control
mechanisms. Contrastingly, targets that appear sooner
Although yielding information about the than expected may automatically grab attention, thus
modulatory mechanisms of temporal orienting, ERPs are emphasising bottom-up, exogenous shifts of temporal
not particularly useful in identifying the neural system attention. Indeed, brain activation differed in the two
controlling the orienting of attention toward specific time types of trial, supporting the existence of endogenous
intervals, due to their poor spatial resolution. Brain imaging and exogenous control of temporal orienting.
techniques such as PET (positron emission tomography) Endogenous temporal orienting involved activations in
and fMRI (functional magnetic resonance imaging) afford a right frontal cortex (including dorsolateral and
much more detailed analysis of the neuroanatomical ventrolateral areas), and left superior parietal lobule.
substrates of temporal orienting, and are now considered. These prefrontal activations are coherent with the notion
of a higher order, “top down” attentional mechanism
3.3. Brain Imaging being involved in endogenous shifts. Exogenous shifts
An experiment using event-related fMRI revealed were associated with increased activation in visual
the brain areas that participate in temporal orienting, and extrastriate cortex, consistent with the notion of a
their sensitivity to trial validity and SOA (33). The task “bottom up” attentional mechanism being involved in
used was a simplified version of Experiment 1 (see Figure exogenous shifts of temporal attention, being subserved
1a). Inner or outer circular cues (100ms duration) predicted by sensory association cortex.
(80% validity) either a short (600ms) or long (1400ms)
cue-target SOA. The target (50ms duration) was an upright In conclusion, the brain system for temporal
cross and required a speeded detection response by pressing orienting appears to involve a left-hemisphere dominant
a button with the right index finger. The results supported frontal-parietal network, which also interacts with areas
the modulation of brain activity linked to motor preparation related to motor attention and motor readiness. The
and attention during temporal orienting. brain-imaging results thus support the suggestion from
the ERP experiments that modulation of motor-related
The comparison between invalid and valid trials variables plays an important role in temporal orienting.
isolated brain activity linked to attentional variables. Activity is also co-ordinated with other areas subserving
Sensory and motor variables are well controlled. During specific aspects of temporal orienting, such as prefrontal
invalid trials, temporal expectations are breached and control and visual areas. Studies of temporal orienting
attention is shifted from one time point to another. Invalid have not yet been conducted on neurological patients,
trials preferentially activated inferior parietal, inferior but we would predict that patients with lesions to left
premotor and prefrontal areas in the left hemisphere parietal-frontal areas would show significant deficits in
predominantly, as well as orbitofrontal cortex bilaterally orienting attention to time intervals. Patients with
(see (34)). Activations in similar left-hemisphere parietal lesions in areas related to motor preparation, such as
and premotor areas have been found during studies of basal ganglia and SMA are also likely to show
motor preparation (35,36). Furthermore, patients with left impairments. It will be interesting to test different
parietal lesions have deficits in a motor orienting task, in groups of patients on these tasks.
which the cue predicts the type of motor response to the
stimulus, rather than its spatial location (35). The role of 4. COMPARISON OF SPATIAL AND TEMPORAL
the left parietal cortex in motor control and attention is also ORIENTING
emphasised by the incidence of motor apraxic deficits after
lesions to this brain area (37). This suggests that orienting To test whether there exists a single attentional
attention in time is closely linked with motor preparation system, or rather different subsystems depending on the
and motor attention. information available to guide selection, we directly
compared temporal orienting with visual spatial orienting.
The comparison between valid trials with long Visual spatial orienting is the most well studied type of
and short SOAs emphasised brain activations linked to selective attention and therefore provides a useful
timing, anticipation, and motor readiness. Long-interval framework for comparison. The neural system for visual
trials enhanced activation in the medial premotor cortex in spatial orienting has been investigated by both brain
the region of the anterior supplementary motor area (SMA), imaging (e.g. (39,40,41)) and neuropsychological studies
left putamen and the thalamus bilaterally. These brain (see (42) for review). Convergent findings from these
regions have a well-known role in the timing and readiness experiments have suggested a large-scale frontal-parietal
of motor responses (e.g. (25,38)). Their modulation during network of brain regions that support spatial attention. ERP
temporal orienting suggests the ability to affect variables studies have suggested that spatial attention acts by
related to motor readiness to optimise behavioural modulating early visual processing in extrastriate areas (see
responding, as suggested by the ERP experiment (19). (29,30,43)).
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orienting condition, the appearance of the narrow or wide cross 5. SUMMARY AND CONCLUSIONS
cued subjects to expect the target after the first interval (short,
600ms from cue onset) or second interval (long, 1200ms from In conclusion, we have shown that we can use
cue onset). No information was given about probable target predictive information about time intervals to direct our
location (left, right). The spatial orienting condition used attention to a predicted point in time, and enhance
exactly the same stimuli and timing, except the narrow or wide behavioural performance. Temporal orienting appears to be
cross cued the subjects to expect the target at either the left or under flexible control, with behavioural advantages being
right location, with no information being given about probable seen for stimuli occurring over multiple time frames.
target interval. In both conditions the cues had 75% validity. Temporal orienting involves processes linked to motor
readiness and expectancies, and generally modulates
Behaviourally, valid cueing produced significant response-related stages of stimulus processing. The
reaction-time benefits in both the spatial and temporal neuroanatomical substrates of temporal orienting include a
orienting conditions (Figure 5b). Consistent with previous frontal-parietal network of areas with a left hemisphere
temporal attention experiments (e.g. (19)), benefits only bias, as well as co-ordinated activity in areas related to
occurred when targets appeared at the short interval in the motor attention and readiness. One important domain for
temporal orienting condition. further exploration is whether overt and speeded motor
responses are required to observe the effects of temporal
ERPs were analysed to identical, non-target orienting. Tasks in which no on-line responses are required
bilateral stimulus arrays occurring after the first interval. to temporally expected stimuli are needed to address this
These highly controlled conditions ensured that there were concern. In addition, it will be interesting to test whether
no attributes in the stimulus array that could automatically the patterns of brain activation depend specifically on the
“grab” attention selectively, making it possible to isolate type of motor response required (e.g., comparing button
purely endogenous attentional mechanisms (see (48,49)). presses, eye movements and verbal responses).
The ERP analysis revealed that the optimisation of
behaviour by spatial and temporal orienting is achieved via The comparison of temporal and spatial orienting
different attentional mechanisms (Figure 5c). reveals that there is not a single mechanism of action or a
ubiquitous attentional system in the brain. Rather, the
Spatial attention modulated the amplitude of information available for selection, and the demands of the
visual-evoked components P1 and N1. This is consistent task dictate which functionally specialised brain areas will
with previous studies of spatial attention (e.g. (29)), and support the optimisation of behaviour by attentional
confirms the ability of spatial attention to modulate early orienting. Such behavioural enhancement by predictive
stages of stimulus processing. information may be achieved by different modulatory
effects on stimulus processing, from perceptual to motor-
Temporal orienting involved a different pattern of related effects, depending on the nature of the task. This
modulation to spatial orienting, with the early perceptual illustrates the flexibility of attentional functions in the
P1 component being unaffected. The N1 component was human brain. Further fractionation of the neural systems
modulated by temporal attention, but with a different and and brain mechanisms that support the orienting of
non-lateralised scalp distribution. The N1 modulation by attention will shed light into the gamut of potential ways in
temporal attention demonstrated here, something not found which behavioural performance can be enhanced.
by Miniussi et al (19), suggests that temporal attention can
modulate visual processing, though in a different, and more 6. ACKNOWLEDGEMENTS
diffuse, manner than spatial orienting. The research was supported by a project grant
from the Wellcome Trust to ACN. Publication costs were
The later effects of temporal attention were partially covered by the International Multisensory
similar to those seen by Miniussi et al (19), that is, Research Forum.
modulation of the N2 and P300 components. N2
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