Stimulus Significance Effects in Habituation of the Phasic and Tonic Orienting Reflex

Phasic and tonic measures of electrodermal activity were examined in a simple habituation paradigm, using innocuous visual stimuli. Separate groups were used to investigate the effects of stimulus significance, as manipulated by instructions. One group had no stimulus-related task (indifferent group), while the other was asked to silently count the stimuli to report to the experimenter later (significant group). Prestimulus skin conductance levels were considered as measures of the arousal level at each stimulus presentation, and the subsequent electrodermal responses were taken as the phasic orienting reflex (OR) elicited by each stimulus. Changes in prestimulus arousal were taken as measures of the tonic OR to the experimental series. Marked group differences were found in both phasic and tonic components of the OR. Some, but not all, of the significance effects in the phasic OR were attributable to differences in arousal. The results are discussed in the context of theoretical accounts of the OR.     

Dishabituation and spontaneous recovery of the electrodermal orienting response: Effects of extraversion,impulsivity, sociability,and caffeine

The relationship between extraversion and psychophysiological indices of arousal is not yet fully understood. The present study attempted to further clarify this relationship by using caffeine to systematically manipulate arousal. Subjects pretested on the Eysenck Personality Inventory were given caffeine or a placebo, then underwent habituation, dishabituation, and spontaneous recovery of the electrodermal OR. Separate analyses examined the main and interactive effects of extraversion, impulsivity, and sociability with caffeine. Several tonic and phasic measures showed group reversal effects, with introverts having higher tonic levels and larger phasic responses under placebo conditions and extraverts having larger phasic responses and higher tonic levels under caffeine. Although both tonic and phasic responses were differentially affected, the group reversal effects occurred at different points in time on tonic than they did on phasic measures relative to the theoretical buildup of inhibition. The findings for impulsivity were quite consistent with those for extraversion, and both sets of results were generally supportive of the Eysenck hypothesis.     

Exploring the mechanism of dishabituation

In this study we explored elicitation and habituation of the orienting reflex (OR) in the context of indifferent and significant stimuli, particularly aiming to clarify the mechanism driving dishabituation. An in-depth analysis of the mechanisms of electrodermal habituation and dishabituation was conducted, focusing on the role of state measures as determinants of the phasic response profile. Twenty-four young adult participants completed an auditory dishabituation task while electrodermal activity was recorded. Participants listened to a series of 10 innocuous tones of the same frequency (standards), followed by a deviant tone of a different frequency, and succeeded by 2-4 tones of the same frequency as the initial 10 stimuli. All stimuli had a random stimulus onset asynchrony of 5-7 s. Participants completed an indifferent condition in which there was no task in relation to the stimuli, and a significant condition where instruction was given to count the stimuli silently; order was counterbalanced between participants. As predicted, both skin conductance responses (SCRs) and skin conductance levels (SCLs) were larger for the significant than the indifferent condition. The initial phasic ORs were dependent on pre-stimulus arousal level, and there were significant decreases in both SCR and SCL over the first 10 standards in both conditions. Phasic response recovery was apparent to the deviant stimulus, and dishabituation to the following standard stimulus; both effects were enhanced in the significant condition. Sensitisation was apparent in SCL following the initial and deviant stimuli, but the extent of this was confounded with incomplete resolution of the preceding phasic OR in the significant condition. In the indifferent condition, dishabituation was independent of deviant-related sensitisation; this could not be tested in the significant condition. These findings suggest that dishabituation is not a process of sensitisation, but rather, a disruption of the habituation process.     

Electrodermal measures of attention and effort to stimulus onset and offset for intramodal and intermodal tasks.

Two experiments were designed to assess the relationship between task difficulty and arousal. Electrodermal measures of tonic and phasic arousal to four levels of task difficulty at stimulus onset and offset were studied in college students for intramodal and intermodal tasks. The students were presented for 18 trails with visual-auditory or visual-visual stimuli with either 2.0-, 0.2-, or 0.02-sec difference between stimuli onset or offset, and asked to judge which stimulus came on or went off first or merely to observe the stimuli. Both frequency and amplitude of skin conductance responses reliably differentiated the levels of task difficulty for both the intramodal and the intermodal task. None of the measures of tonic level of arousal was reliable. Electrodermal measures of phasic responses accurately reflected the task demands.     

Determinants of tonic and phasic reactions in transswitching

Forty-eight college students were assigned randomly to four groups in a 2 × 2 factorial arrangement of phasic conditional stimuli (samevs. different) and tonic conditional stimuli (samevs. different) to receive 2 days of classical conditioning with a transswitching procedure. Tonic stimuli were a 5-minute projected white triangle or circle; phasic stimuli were a 5-second red or green square superimposed over the tonic stimuli. There were six tonic stimulus segments each day, separated by 20-second periods of no stimulus, three containing six trials of the phasic stimulus paired with shock and three containing six trials of the phasic stimulus alone, in the counterbalanced order. Tonic responding at the onset of the tonic stimuli or during brief periods following its onset were recorded, along with phasic responses to the phasic stimuli. Responses included magnitude of skin conductance responses, frequency of unelicited skin conductance responses, and tonic heart rate. Both skin conductance measures of responding to the tonic stimuli differentiated significantly between positive and negative tonic segments during Day 2, but only in the group with two different tonic stimuli and one phasic stimulus (“standard” transswitching). This supported the hypothesis that tonic stimulus differentiation would be absent when two different phasic stimuli were present. The heart rate data did not support this hypothesis, showing tonic differentiation in both groups with two tonic stimuli. Phasic differentiation controlled by the different phasic stimuli was observed on Day 1; on Day 2, phasic differentiation was present only in the group with two tonic and one phasic stimuli and the group with one tonic and two phasic stimuli. The results were interpreted to mean that temporal contiguity between the tonic stimuli and shock or no shock is not sufficient to establish tonic response differentiation in transswitching.     

Insensitivity to stimulus validity in human pavlovian conditioning

Four experiments investigated the role of information value, or stimulus validity, in human electrodermal conditioning. Conditioned stimuli (CSs) were long, variable duration (10-50-sec) slides or sounds, the unconditioned stimulus (US) was shock, and the primary measures were change in tonic skin conductance level and self-report expectancy of shock. In Experiment 1 electrodermal responding to a target stimulus was marginally lower in a blocking group than in a superconditioning group. This difference failed to occur in Experiment 2, despite increased sensitivity of the electrodermal measure. Explicit instructions to pay attention to the relationship between each stimulus and shock improved learning, but did not lead to group differences (Experiment 3). In Experiment 4 expectancy ratings in a blocking group were lower than in an overshadowing group, but no electrodermal differences occurred. The results were interpreted in terms of non-additive learning processes such as occasion-setting, in addition to a general lack of transfer of learning about a stimulus from one context to another (element to compound or vice versa). It was suggested that sensitivity to stimulus validity might be observed if conditioning were embedded in a causally familiar task, as in contingency judgement research.     

Stimulus-dependent dopamine release in attention-deficit/hyperactivity disorder

Attention-deficit/hyperactivity disorder (ADHD) is related to an attenuated and dysfunctional dopamine system. Normally, a high extracellular dopamine level yields a tonic dopaminergic input that down-regulates stimuli-evoked phasic dopamine responses through autoreceptors. Abnormally low tonic extracellular dopamine in ADHD up-regulates the autoreceptors so that stimuli-evoked phasic dopamine is boosted. The authors propose that these boosted phasic responses yield hypersensitivity to environmental stimuli in ADHD. Stimuli evoking moderate brain arousal lead to well-functioning performance, whereas either too little or too much stimuli attenuate cognitive performance. Strong, salient stimuli may easily disrupt attention, whereas an environment with impoverished stimuli causes low arousal, which is typically compensated for by hyperactivity. Stochastic resonance is the phenomenon that makes a moderate noise facilitate stimulus discrimination and cognitive performance. Computational modeling shows that more noise is required for stochastic resonance to occur in dopamine-deprived neural systems in ADHD. This prediction is supported by empirical data.     

Contextual stimulus control of conditional vasomotor and electrodermal reactions to angry and friendly faces

An experiment with 42 human Ss used the transswitching procedure to examine tonic stimulus control of phasic and tonic conditioned vasomotor heart rate, and electrodermal reactions. The conditional stimulus (CSs) were photos of angry and friendly human faces, and the unconditional stimulus (US) was a human scream. In one tonic context (blue light), the CSs were paired with the US, in the other context (yellow light), the CSs were presented unpaired. Following acquisition, an extinction series was run with the US omitted during both tonic contexts. Phasic vasomotor and skin conductance reactions differed in the positive and negative tonic segments (stronger in positive). The skin conductance responses also differed during extinction, but the vasomotor responses did not. Tonic differences (following onset of the tonic stimuli) in unelicited skin conductance response frequency, finger pulse volume, and heart rate were also found, although these developed more slowly than the phasic differences. The finger pulse volume tonic difference was greater in extinction than the skin conductance response frequency. There was no effect of the angry-friendly facial expressions, either directly or in interaction with the transswitching effects. The results were interpreted to mean that the transswitching phenomenon is not limited to one another autonomic effector, but is more generalized across the ANS (sympathetic branch). The absence of influence of the facial expressions indicates the relative weakness of the “preparedness” hypothesis in comparison with more influential contextual factors.     

Arousal, working memory, and conscious awareness in contingency learning

There are wide individual differences in the ability to detect a stimulus contingency embedded in a complex paradigm. The present study used a cognitive masking paradigm to better understand individual differences related to contingency learning. Participants were assessed on measures of electrodermal arousal and on working memory capacity before engaging in the contingency learning task. Contingency awareness was assessed both by trial-by-trial verbal reports obtained during the task and by a short post-task recognition questionnaire. Participants who became aware had fewer non-specific skin conductance responses and tended to score higher on a digit span assessment. Skin conductance level was not significantly lower in the aware group than in the unaware group. These findings are consistent with studies showing that lower arousal and greater cognitive processing capacity facilitate conscious perception of a greater breadth of information within a scene or a task.     

Social stimuli increase physiological reactivity but not defensive responses

Emotional reactions are crucial in survival because they provide approach and withdrawal behaviors. However, an unsolved question is whether the social content of the affective stimuli has a specific effect on emotional responses. We studied whether the social content of affective pictures influenced the defensive response and response mobilization. For this purpose, we recorded startle blink reflex (a defensive response) and skin conductance responses (a measure of unspecific physiological reactivity or arousal) in 73 participants while they viewed a series of 81 pictures of varying affective valence and social content. Our results revealed that defense response, as indicated by increases in the magnitude of the startle blink reflex, was mainly dependent on threatening or unpleasant cues, but was unrelated to the social content of the pictures. The social content, however, had an influence on pleasant stimuli, provoking an increase in resource mobilization, as reflected by changes in electrodermal activity. Hence, the social content of the affective stimuli may increase the physiological arousal elicited by pleasant stimuli, and it appears to be unrelated to the defense reactivity provoked by unpleasant stimuli.     

Brain asymmetry and autonomic conditioning: Sensitization control

In the present experiment we report effects of cerebral asymmetry, or laterality, during classical conditioning to facial emotional stimuli. Twenty-five female subjects were presented with slides of a happy face in one visual half-field (VHF), and simultaneously a slide of an angry face in the other VHF, followed by shock as the unconditioned stimulus (UCS). To control for effects of sensitization, a new stimulus, never associated with the UCS, was introduced in the extinction phase. Dependent measures were phasic heart rate responses (HR) and skin conductance responses (SCR). The HR results showed a significant right hemisphere effect for the CS-UCS association, that was not attributable to UCS sensitization. No significant effects were found for the SCRs. The basic HR finding was a right hemisphere superiority for learning of a conditioned association.     

An investigation of bilateral asymmetries in electrodermal activity

Research in cerebral laterality supports the idea that functional differences between the left and right hemispheres exist with respect to cognitive style and perceptual ability. Related research, which has examined autonomic nervous system (ANS) correlates of cerebral laterality, suggests that a component of the ANS, electrodermal activity (EDA), is also lateralized. Some findings in the literature report the occurrence of bilateral asymmetries in phasic and tonic EDA as a function of a left or right hemisphere preference for information processing. This experiment used normal male subjects who were either left or right movers in a test of Conjugate Lateral Eye Movement (CLEM). This selection procedure served to maximize a subject’s preference for either a right or left hemisphere mode of information processing. Bilateral EDA was recorded continuously while subjects performed a visual recognition task using word (left hemisphere) and shape (right hemisphere) stimuli. The data do not support the contention that performance on a procedure chosen to selectively activate a given hemisphere elicits asymmetric tonic and/or phasic EDA. The results show no significant difference in the frequency of elicited skin conductance responses under either of the experimental conditions. Bilateral tonic EDA rose continuously over time and did not vary in either hand as a function of task. The present results fail to offer support for either of the hypotheses which argue for I) increased contralateralexcitation with selective hemispheric activation, or 2) increased contralateralinhibition of the EDR with hemispheric arousal.     

Tracking arousal state and mind wandering with pupillometry

In four experiments, the association between arousal state and different mind-wandering states was examined. Participants performed a sustained attention task while pupil responses were continuously recorded. Periodically during the task, participants were presented with thought probes to determine if they were on task or mind wandering. Across the four experiments, the results suggested that in situations that promoted on-task behaviors and focused external attention, mind wandering was associated with lowered arousal, as seen by smaller tonic pupil diameters and smaller phasic pupillary responses. However, in situations that promoted a more internal focus of attention, there were no differences between on-task states and mind wandering in tonic pupil diameter (although differences emerged for phasic pupillary responses), suggesting similar arousal levels. Furthermore, across the four experiments, mind blanking and mind wandering dissociated in terms of whether the situation promoted focused external attention or focused internal attention. These results are broadly consistent with the notion that mind wandering is a heterogeneous construct, with different forms of mind wandering being associated with different arousal states, and suggest that a combination of behavioral and pupillary measures can be used to track these various states.     

The Psychophysiology of Sensation Seeking

This article summarizes studies relating the trait of sensation seeking to electrodermal and heart-rate responses and cortical evoked potential arousal. Stimulus factors of novelty, intensity, and stimulus significance are important. High sensation seekers tend to give stronger physiological orienting responses than lows to novel stimuli of moderate intensity, particularly when such stimuli are of specific interest. Lows tend to show defensive responses as defined by heart-rate acceleration. The cortical reaction of the highs tends to be augmented by intense visual or auditory stimuli, while that of the lows tends to be reduced or unresponsive to variations in stimulus intensity. Differences between psychophysiological responses of high and low sensation seekers are interpreted as reflective of different evolved biological strategies for processing novel or intense stimulation.     

Electrodermal responses to movie stressors: Nicotine × extraversion interactions

Support was found for Eysenck's hypothesis that the effects of smoking sized doses of nicotine on cortical and electrodermal activity are functions of extraversion and the degree of externally-induced arousal. Responses to scenes that elicited moderate electrodermal and emotional responses followed the predicted extraversion ☆ nicotine interaction for electrodermal measures, but not for self-reported emotions, heart rate or electromyographic measures. The failure of the Ss to demonstrate an extraversion ☆ nicotine interaction when high-intensity stimuli were used (Gilbert and Hagen, 1980) is also interpreted as consistent with Eysenck's hypothesis.     

Tonic and phasic stimulus control of blood pressure responses using opposing unconditional stimuli

The transswitching paradigm was used in the present study to investigate the effects of tonic (long duration) environmental stimuli paired with opposing unconditional stimuli on human blood pressure. Sixty volunteers participated for two sessions, one week apart. Subjects were presented with a pseudorandom sequence of four trials of red lights (five minutes each) and four trials of blue lights (five minutes each) within each session. All subjects received the cold pressor test during the red lights. During the blue lights, half the subjects received warm water and the other half received a neutral temperature water. Subjects received the presentations of the water in one of three methods: as determined by the experimenter, preceded by a warning signal, or self-administered. The results indicated that conditional and unconditional responses during the red lights were progressive increases in blood pressure. During the blue lights associated with warm water, conditional and unconditional responses were often progressive decreases in blood pressure. The method of administration of the water affected the magnitude of the unconditional responses. Evidence for phasic stimulus control was not as unequivocal as tonic stimulus control. The present results indicated that it is possible to modify blood pressure in two directions (increases and decreases) when two high contrast, opposing unconditional stimuli are presented.     

Change in the event-related skin conductivity: an indicator of the immediate importance of elaborate information processing?]

In recent psychophysiological conceptualizations of the orienting response (OR) within the framework of information processing, the OR is increasingly considered a "call for processing resources", something which is especially inferred from variations in the event-related skin conductance response (SCR). The present study, therefore, was concerned with certain implications arising from this framework or perspective, particularly in regard to the question of whether stimuli eliciting skin conductance responses obligatorily receive/evoke processing priority or not. In order to examine whether these electrodermal responses denote a capturing of attention or merely a call for processing resources, short (1 s) pure sine tones of 65 dB with sudden onset (commonly used as orienting stimuli) were inserted in a reaction time paradigm with an additional memory load. This demand was primarily given because memory processes play a key role in theories of orienting and habituation. The task was run under two different conditions of complexity, factorially combined with a novelty variation of the added auditory stimuli. The results revealed a substantial deterioration of task performance subsequent to the occurrence of the tones, which, however, was dependent on task complexity and on novelty of the tones. The task impairment is particularly remarkable as subjects were asked to avoid distractions by paying attention to the task and as the tones were introduced as subsidiary and task-irrelevant. Together with the missing effects of task complexity on phasic and tonic electrodermal activity, results suggest that information-processing conceptualizations of the OR can only be a meaningful heuristic contribution to theoretical developments about human orienting and its habituation if the setting of processing priority, its conditions, as well as its implications are adequately taken into account. In addition, it seems to be promising to consider the strength of the SCR as an index of urgency of elaborate, attention-demanding processing and not as a peripheral physiological manifestation of the OR, or, respectively, of a call for unspecific processing resources. Such a view would also do justice to the aspect of prioritization. The sufficient conditions for an OR's occurrence could, in this context, be equated with, among others, some of those which activate a mechanism subserving selective attention and, as a possible result, which lead to further and more elaborate processing of potentially important information.     

Temporal kinetics of prefrontal modulation of the extrastriate cortex during visual attention

Single-unit, event-related potential (ERP), and neuroimaging studies have implicated the prefrontal cortex (PFC) in top-down control of attention and working memory. We conducted an experiment in patients with unilateral PFC damage (n=8) to assess the temporal kinetics of PFC-extrastriate interactions during visual attention. Subjects alternated attention between the left and the right hemifields in successive runs while they detected target stimuli embedded in streams of repetitive task-irrelevant stimuli (standards). The design enabled us to examine tonic (spatial selection) and phasic (feature selection) PFC-extrastriate interactions. PFC damage impaired performance in the visual field contralateral to lesions, as manifested by both larger reaction times and error rates. Assessment of the extrastriate P1 ERP revealed that the PFC exerts a tonic (spatial selection) excitatory input to the ipsilateral extrastriate cortex as early as 100 msec post stimulus delivery. The PFC exerts a second phasic (feature selection) excitatory extrastriate modulation from 180 to 300 msec, as evidenced by reductions in selection negativity after damage. Finally, reductions of the N2 ERP to target stimuli supports the notion that the PFC exerts a third phasic (target selection) signal necessary for successful template matching during postselection analysis of target features. The results provide electrophysiological evidence of three distinct tonic and phasic PFC inputs to the extrastriate cortex in the initial few hundred milliseconds of stimulus processing. Damage to this network appears to underlie the pervasive deficits in attention observed in patients with prefrontal lesions.     

Physiological patterns accompanying complex problem solving during warning and nonwarning conditions.

Physiological responses of 48 subjects were studied during both a warning and a nonwarning condition and during anagram problem solving. The warning condition was associated with heart rate deceleration and increased occurrence of galvanic skin responses (GSR), of skin conductance, and of tonic galvanic skin potentials (GSP). The warning condition also resulted in significantly faster solution rates. Problem solving was associated with increases in heart rate, occurrence of GSR, and skin conductance. The occurrence of phasic GSP habituated, but the occurrence of tonic GSP maintained a significant response frequency. Large increases in tonic GSP, with minimal phasic increases during problem solving, reflected the bidimensionality of electrodermal activity as a function of the task. These findings were interpreted to support Lacey's hypothesis of response fractionation and suggest the utility of physiological measures in the study of putative attentional functioning.     

Contextual conditioning

Eastern and Western interpretations of contextual control of phasic conditional responses (transswitching) are contrasted. The Eastern (Asratyan, 1965) approach emphasizes the role of the tonic conditional stimulus and the (hypothetical) tonic response it evokes. The Western (Lachnit, 1986) approach emphasizes the role of compound conditional stimuli. Although Lachnit showed that transswitching-like results can be obtained without a tonic stimulus, attempts to simulate transswitching experiments using a computer model of the Rescorla-Wagner theory (Kimmel and Lachnit, 1988) have shown that predictions from the theory approximate empirical results in human classical conditioning only when the tonic stimulus is given far greater weight than the phasic stimulus. In other words, only when the Rescorla-Wagner theory is made more like Asratyan’s theory, can the compound conditional stimulus approach account for real empirical transswitching data.