The dynamics of cardiac defense: From attention to action

The attentional and motivational significance of cardiac defense is examined in two studies. In Study 1, cardiac defense was evoked by an intense acoustic stimulus in the context of either a visual search or a memory search task using letters as stimuli. Results showed a potentiation of the long latency acceleration of cardiac defense in the visual search task. In Study 2, participants performed the same visual search task using pleasant, neutral, and unpleasant pictures as stimuli. Results showed a further potentiation of the long latency acceleration of cardiac defense when the visual search task was performed with unpleasant, compared to pleasant or neutral pictures. These results indicate that cardiac defense has both attentional and motivational contributions, where the attentional significance is related to increased sensory processing, whereas the motivational significance is associated with preparation for active defense.     

Cardiovascular and Neurophysiologic Correlates of Sensory Intake and Rejection. II. Interview Behavior

As part of a three task study of the influence of attentional style on cardiovascular response, 19 normal volunteers were given a 15-min interview during which systolic and diastolic blood pressure, digital pulse volume, heart rate, and forearm blood flow were recorded. At the same time two observers independently assessed five elements of the subjects' interview behavior; arousal, eye contact with the interviewer, self-revelation of interview center, attentiveness to the interviewer, and overall transactional engagement in the interview task. When subjects were divided into groups of interview at tenders and nonattenders on the basis of interviewer ratings, attenders had a mean decrease in forearm blood flow and nonattenders a mean increase. These group differences extended across a word identification (sensory intake) and mental arithmetic (sensory rejection) task as well. When subjects were divided into groups of forearm blood flow increasers and decreasers, increasers displayed less attentiveness to the interviewer, less self-revelation, greater arousal, and less transactional engagement than did decreasers (N=9). Attentiveness to the interviewer and transactional engagement were the two most sensitive behavioral discriminators in comparing the increaser and decreaser groups.     

Stimulus-focused attention speeds up auditory processing

Stimulus-focused attention enhances the processing of auditory stimuli, which is indicated by enhanced neural activity. In situations where fast responses are required, attention may not only serve as a means to gain more information about the relevant stimulus, but it may provide a processing speed gain as well. In two experiments we investigated whether attentional focusing decreased the latency of the auditory N1 event related potential. In Experiment 1 slowly emerging, soft (20dB sensation level) sounds were presented in two conditions, in which participants performed a sound-detection task or watched a silent movie and ignored the sounds. N1 latency was shorter in the sound-detection task in comparison to the ignore condition. In Experiment 2 we investigated whether the attentional N1 latency-decrease was caused by a frequency-specific attentional preparation or not. To this end, tone sequences were presented with a single tone frequency or with four different frequencies. N1 latency was shorter in the sound-detection task in comparison to the ignore condition regardless the number of frequencies. These results suggest that stimulus-focused attention increases stimulus processing speed by generally increasing sensory gain.     

Single bouts of exercise selectively sustain attentional processes

This study examined how single bouts of exercise may differentially modulate neuroelectric correlates of attentional orienting and processing. Using a within‐participants design, ERPs and task performance were assessed in response to a perceptually challenging three‐stimulus oddball task prior to and following a bout of exercise or seated rest during two separate, counterbalanced sessions. Findings revealed that, following a single bout of exercise, attentional processing was sustained relative to pretest whereas prolonged sitting resulted in attentional decrements. Focal attention resulting from attentional orienting, in contrast, does not appear to be sensitive to the influences of single bouts of physical activity. These findings suggest that acute exercise‐induced changes in cognition do not originate from an overall modulation of attention but instead are specific to aspects of attentional processing.     

Lateralized frontal activity elicited by attention-directing visual and auditory cues

In event-related potential studies of voluntary spatial attention, lateralized activity observed over anterior scalp sites prior to an impending target has been interpreted as the activity of a supramodal attentional control mechanism in the frontal lobes. However, variability in the scalp topography and presence of this activity across studies suggests that multiple neural generators contribute to the lateralized activity recorded at the scalp. Using distributed source modeling we found two distinct frontal lobe sources following attention-directing cues, one dependent on the sensory modality of the eliciting stimulus and one dependent on the response requirements of the task. Differential activity of these sources depending on task parameters suggests that neither source reflects activity necessary for controlling attention.     

Cognitive and emotional modulation of the cardiac defense response in humans

The cognitive and emotional modulation of the cardiac defense response was investigated in this study. One hundred forty-four participants were exposed to three presentations of an intense auditory stimulus while performing one of four attentional tasks: a control task, an external perceptual tracking task, and two internal tasks presented at either easy or difficult memory loads. State anxiety was also manipulated by requiring each group to perform either with or without the threat of shock. Heart rate and vasomotor activity were recorded. Results indicated that only the externally directed tracking task led to potentiation of the cardiac response. No predicted effects for attentional demands were obtained and the anxiety manipulation did not appear to have an effect. Differences between measures were also observed, particularly with respect to response habituation. Unlike cardiac activity, vasomotor responses displayed resistance to habituation. The results are discussed in relation to contemporary accounts of defensive responding.     

The Relationship Between Skin Conductance Orienting the Allocation of Processing Resources

Information processing models of autonomic orienting suggest that the elicitation of an orienting response is associated with either the call for, or the actual allocation of, limited attentional processing resources. However, Dawson, Filion, and Schell (1989) reported a directional dissociation between elicitation of the skin conductance orienting response and resource allocation, as indexed by reaction time slowing on a secondary task. Although larger skin conductance responses were elicited by a task-relevant stimulus than by a task-irrelevant stimulus, reaction time showed the opposite pattern (i.e., greater slowing to secondary task probes presented shortly following the onset of the task-irrelevant stimulus). In the present report, we describe three experiments which examine the generality of this dissociation effect and test specific hypotheses regarding its nature. Results of the first two experiments revealed that the dissociation effect is observed reliably when the task-relevant and task-irrelevant stimuli consist of left ear and right ear tones or high and low pitched binaural tones, across a range of secondary task probe presentation times. However, the third experiment demonstrated that when task-relevant and task-irrelevant stimuli are presented to different sensory modalities (auditory and visual), orienting and resource allocation are both greater during the task-relevant than the task-irrelevant stimuli, thus eliminating the dissociation effect. These results support the hypothesis that the dissociation effect is due to a switch of attention initiated because of the physical similarity of the task-relevant and task-irrelevant stimuli, and suggest that there is a fundamentally positive relationship between skin conductance orienting and resource allocation under selective attention conditions.     

Temporal dynamics of lateralized ERP components elicited during endogenous attentional shifts to relevant tactile events

To investigate the temporal dynamics of lateralized event-related brain potential (ERP) components elicited during covert shifts of spatial attention, ERPs were recorded in a task where central visual symbolic cues instructed participants to direct attention to their left or right hand in order to detect infrequent tactile targets presented to that hand, and to ignore tactile stimuli presented to the other hand, as well as all randomly intermingled peripheral visual stimuli. In different blocks, the stimulus onset asynchrony (SOA) between cue and target was 300 ms, 700 ms, or 1,100 ms. Anterior and posterior ERP modulations sensitive to the direction of an attentional shift were time-locked to the attentional cue, rather than to the anticipated arrival of a task-relevant stimulus. These components thus appear to reflect central attentional control rather than the anticipatory preparation of sensory areas. In addition, attentional modulations of ERPs to task-irrelevant visual stimuli were found, providing further evidence for crossmodal links in spatial attention between touch and vision.     

Synchrony: a neural correlate of somatosensory attention

We investigated whether synchrony between neuronal spike trains is affected by the animal's attentional state. Cross-correlation functions between pairs of spike trains in the second somatosensory cortex (SII) of three macaque monkeys trained to switch attention between a visual task and a tactile task were computed. We previously showed that the majority of recorded neuron pairs (66%) in SII cortex fire synchronously while the animals performed either task and that in a subset of neuron pairs (17%), the degree of synchrony was affected by the animal's attentional state. Of the neuron pairs that showed changes in synchrony with attention, about 80% showed increased synchrony when the animal attended to the tactile stimulus. Here, we show that peak correlation typically occurred at a delay <25 ms; most commonly the delay was close to zero. Half-widths of the correlation peaks were distributed between a few milliseconds and hundreds of milliseconds, with the majority lying <100 ms and the mode of the distribution around 20-30 ms. Maximal change in synchrony occurred mainly during the periods when the stimulus was present, and synchrony usually increased when attention was on the tactile stimulus. If periods of elevated firing rates around the motor response times were removed from the analysis, the percentage of pairs that changed the degree of synchrony with attention more than doubled (from 35 to 72%). The observed effects did not depend on details of the statistical criteria or of the time window used in the analysis.     

Early interaction between perceptual load and involuntary attention: An event-related potential study

Whether selective attention affects C1, the first (earliest) visual cortical component of the event-related potential (ERP), remains controversial. We used a cued, involuntary attention task requiring discrimination of targets under low and high levels of perceptual load to examine early attentional modulation in visual cortex. Potential confounds due to physical stimulus differences between load conditions and cue–target sensory interaction were minimized. An interaction between perceptual load and involuntary attention was observed for the P1m component (peak latency between 100 and 140 ms). Furthermore, the parieto-central C1 component (peak latency 80 ms) was modulated by attention, but only under the high-load condition. Thus, whereas attention typically modulates the later P1 component, attentional modulation of C1 is possible under optimal conditions. Specifically, a high perceptual load is necessary for eliciting this earliest attentional effect on cortical processing.     

Focused attention modulates visual responses in the primate prefrontal cortex

Several current models propose an important role of the prefrontal cortex (PFC) in attention. To test the effects of attention in PFC, we recorded from PFC neurons in monkeys performing a task in which they had to attend to one hemifield and wait for a single stimulus that matched a previously presented cue. Neurons exhibited a slight decrease in their initial response and an enhanced activity late in the response to a stimulus at the cued location. The data demonstrate attentional effects on the activity of PFC neurons but they also show that single visual stimuli are initially represented in the activity of PFC neurons even when they are behaviorally irrelevant.     

Attentional modulation of short- and long-lead-interval modification of the acoustic startle eyeblink response: comparing auditory and visual prestimuli.

Studies in our laboratory have shown that modification of startle by lead stimuli with short- and long-lead-intervals is modulated by stimulus significance. The significant stimulus in a tone duration judgement task generates enhanced short-lead-interval startle inhibition as well as pronounced long-lead-interval startle facilitation. The present study was designed to compare tones with simple visual stimuli as lead stimuli in a counterbalanced within-subjects design (Experiment I) or between-subjects design (Experiment II). The results show that auditory compared to visual lead stimuli generate more short-lead-interval inhibition but comparable amounts of long-lead-interval startle facilitation, which was significantly enhanced on to-be-attended trials independent of sensory modality. The attentional manipulation did not yield short-lead-interval effects in Experiment I, but previously reported attention effects were replicated in Experiment II. The results suggest early modality effects on startle modification, reflected by the differing levels of inhibition. Late effects of both modality and attention, however, seem to reflect a sensory modality independent process in startle modification.     

Physiological response patterns during "intake" and "rejection" tasks

Two 2-min tasks, one requiring outer-directed attention (intake task), the other a mental arithmetic task (rejection task) were presented to 20 subjects. According to Lacey the distinguishing feature of the response to intake tasks are cardiac deceleration and blood pressure decreases. None of these responses was obtained. Some variables, however, were only affected by one of the tasks; the intake task led to a decreased hand (skin) blood flow and an increased hand vascular resistance, whereas a heart rate acceleration and increased forearm (muscle) blood flow and decreased forearm vascular resistance were observed during the rejection task. Blood pressure, respiration rate and skin conductance were similarly affected by the two tasks. Inconsistencies among previous studies of the response to the two types of tasks were discussed and found to be partially explained by methodological differences.     

The effect of stimulus modality and task difficulty on attentional modulation of blink startle

The effects of the sensory modality of the lead stimulus and of task difficulty on attentional modulation of the electrical and acoustic blink reflex were examined. Participants performed a discrimination and counting task with either two acoustic, two visual, or two tactile lead stimuli. In Experiment 1, facilitation of the electrically elicited blink was greater during task-relevant than during task-irrelevant lead stimuli. Increasing task difficulty enhanced magnitude facilitation for acoustic lead stimuli. In Experiment 2, acoustic blink facilitation was greater during task-relevant lead stimuli, but was unaffected by task difficulty. Experiment 3 showed that a further increase in task difficulty did not affect acoustic blink facilitation during visual lead stimuli. The observation that blink reflexes are facilitated by attention in the present task domain is consistent across a range of stimulus modality and task difficulty conditions.     

The orienting of attention during eye and hand movements: ERP evidence for similar frame of reference but different spatially specific modulations of tactile processing

To investigate which frames of reference guide shifts of attention triggered during eye and hand movement preparation and the specificity of their effects on somatosensory processing, we recorded event-related brain potentials (ERPs) in a Go/Nogo task where a cue indicated to prepare an eye movement toward - or a hand movement with - the left or right hand. Before the imperative stimulus, a tactile probe was presented to one hand. Spatially selective modulations of tactile processing were more sustained for hand than eye movements, indicating stronger attentional modulations for the modality of the effector's sensory organ. Importantly, attentional modulations of somatosensory processing as well as lateralized ERP components in the preparation interval were virtually identical under uncrossed and crossed hands conditions, suggesting that shifts of attention triggered during hand and eye movement preparation are guided by a common external reference frame.     

Neural correlates of attention and distractibility in the lateral intraparietal area

We examined the activity of neurons in the lateral intraparietal area (LIP) during a task in which we measured attention in the monkey, using an advantage in contrast sensitivity as our definition of attention. The animals planned a memory-guided saccade but made or canceled it depending on the orientation of a briefly flashed probe stimulus. We measured the monkeys' contrast sensitivity by varying the contrast of the probe. Both subjects had better thresholds at the goal of the saccade than elsewhere. If a task-irrelevant distractor flashed elsewhere in the visual field, the attentional advantage transiently shifted to that site. The population response in LIP correlated with the allocation of attention; the attentional advantage lay at the location in the visual field whose representation in LIP had the greatest activity when the probe appeared. During a brief period in which there were two equally active regions in LIP, there was no attentional advantage at either location. This time, the crossing point, differed in the two animals, proving a strong correlation between the activity and behavior. The crossing point of each neuron depended on the relationship of three parameters: the visual response to the distractor, the saccade-related delay activity, and the rate of decay of the transient response to the distractor. Thus the time at which attention lingers on a distractor is set by the mechanism underlying these three biophysical properties. Finally, we showed that for a brief time LIP neurons showed a stronger response to signal canceling the planned saccade than to the confirmation signal.     

Effect of spatial attention on the responses of area MT neurons

This study examines the influence of spatial attention on the responses of neurons in the middle temporal visual area (MT or V5) of extrastriate cortex. Two monkeys were trained to perform a direction-discrimination task. On each trial, two apertures of random-dot stimuli appeared simultaneously at two spatially separated locations; the monkeys were required to discriminate the direction of stimulus motion at one location while ignoring the stimulus at the other location. After extensive training, we recorded the responses of MT neurons in two configurations: 1) Both apertures placed "within" the neuron's receptive field (RF) and 2) one aperture covering the RF while the other was presented at a "remote" location. For each unit we compared the responses to identical stimulus displays when the monkey was instructed to attend to one or the other aperture. The responses of MT neurons were 8.7% stronger, on average, when the monkey attended to the spatial location that contained motion in the "preferred" direction. Attentional effects were equal, on average, in the within RF and remote configurations. The attentional modulations began approximately 300 ms after stimulus onset, gradually increased throughout the trial, and peaked near stimulus offset. An analysis of the neuronal responses on error trials suggests that the monkeys failed to attend to the appropriate spatial location on these trials. The relatively weak attentional effects that we observed contrast strikingly with recent results of Treue and Maunsell, who demonstrated very strong attentional modulations (median effect >80%) in MT in a task that shares many features with ours. Our results suggest that spatial attention alone is not sufficient to induce strong attentional effects in MT even when two competing motion stimuli appear within the RF of the recorded neuron. The difference between our results and those of Treue and Maunsell suggests that the magnitude of the attentional effects in MT may depend critically on how attention is directed to a particular stimulus and on the precise demands of the task.     

Quantitative modeling of visual attention processes in patients with Parkinson's disease: effects of stimulus integrality on selective attention and dimensional integration.

Parkinson's disease (PD) patients and normal controls (NCs) were administered a series of visual attention tasks. The dimensional integration task required integration of information from 2 stimulus dimensions. The selective attention task required selective attention to 1 stimulus dimension while ignoring the other stimulus dimension. Both integral- and separable-dimension stimuli were examined. A series of quantitative models of attentional processing was applied to each participant's data. The results suggest that (a) PD patients were not impaired in integrating information from 2 stimulus dimensions, (b) PD patients were impaired in selective attention, (c) selective attention deficits in PD patients were not due to perceptual interference, and (d) PD patients were affected by manipulations of stimulus integrality and separability in much the same way as were NCs.     

Dimensional weighting and task switching following frontal lobe damage: Fractionating the task switching deficit

Deficits in task switching can be found after frontal lobe damage. Here we demonstrate an impairment in task switching specifically linked to when perceptual weights have to be moved between different dimensions of the same stimulus. A patient (DS) with left frontal lobe damage showed normal performance when he responded to the meaning (a word task) or location (a location task) of a word presented to the left or right of fixation when there was no switching between the tasks. However, when the two tasks were switched every 16 trials in a block, DS showed severe difficulty in performing both tasks (Experiment 1). There were then abnormally large switch costs and effects of stimulus–response congruency. The difficulty was not simply due to switching tasks per se: There were no costs of switching when one of the tasks was modified to have different stimulus displays from the other (Experiment 2). The deficit was also not greater when the switch had to be made from a well-practised task to an unpractised task with more arbitrary stimulus–response mappings, indicating no particular problem in disengaging from a learned task or in configuring new stimulus–response links (Experiment 4). We suggest instead that DS was impaired at shifting attentional weights across different dimensions of the same stimulus, a process required with practised and unpractised tasks alike. The results link this process of shifting attention across stimulus dimensions to the left frontal lobe.     

Cardiovascular and Neurophysiologic Correlates of Sensory Intake and Rejection. I. Effect of Cognitive Tasks

In this study of the relationship between sensory processing and cardiovascular function, five cardiovascular parameters were monitored during baseline periods and during tasks requiring either sensory intake or sensory rejection behavior on the part of 19 subjects. Sensory intake behavior was associated with a pattern of response similar lo that seen with activation of peripheral sympathetic nerves-vasoconstriction in both the digit (skin) and forearm (skeletal muscle). In contrast, sensory rejection behavior was associated with vasodilatation in the forearm and vasoconstriction in the digit. Individual differences in an EEG measure of characteristic ways of processing sensory information were predictably associated with differences in resting cardiovascular function. The association of sensory intake with a skeletal muscle vasoconstriction may help to extend our understanding of the physiology of sensory processing, since heretofore only heart rate and somatic motor activity have reliably differentiated sensory intake from sensory rejection behavior.