Determinants of tonic and phasic reactions in transswitching

Forty-eight college students were assigned randomly to four groups in a 2 X 2 factorial arrangement of phasic conditional stimuli (same vs. different) and tonic conditional stimuli (same vs. 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.(ABSTRACT TRUNCATED AT 250 WORDS)     

Contextual conditioning. A comparison of eastern and western views

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.     

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.     

Effects of an extinguished CS on competition with another CS

Three experiments were conducted using a conditioned taste aversion procedure with rats to examine the effect of nonreinforced presentations of a conditioned stimulus (CS) on its ability to compete with a target stimulus for manifest conditioned responding. Two CSs (A and B) were presented in a serial compound and then paired with the unconditioned stimulus. CS A was first paired with the US and then presented without the US (i.e., extinction) prior to reinforced presentation of the AB compound. Experiment 1 showed that A was poor at competing with B for conditioned responding when given conditioning and extinction prior to reinforcement of AB relative to a group that received both A and B for the first time during compound conditioning. That is, an extinguished A stimulus allowed greater manifest acquisition to B. Experiment 2 found that extinction treatment produced a poor CR to the pretrained and extinguished CS itself following compound conditioning. Experiment 3 found that interposing a retention interval after extinction of A and prior to compound conditioning enhanced A's ability to compete with B. The results of these experiments are discussed with regard to different theories of extinction and associative competition.     

Kainate receptor activation enhances both tonic and phasic GABA-ergic inhibition in CA1 hippocampal interneurons

Kainate receptor agonists are powerful convulsants and excitotoxins. It has been a lot of controversy around functions of these receptors in the brain. It is shown in this article that kainate enhances evoked GABAergic IPSC (phasic currents) in CA1 interneurons in concentration-dependent manner. The phenomenon is likely to be due to kainate-mediated lowering of the threshold for action potential generation in interneuron axons and increased number of terminals responding to the same stimulus strength. Kainate application also induced an enhancement in tonic GABAergic conductance. This phenomenon can be attributed to massive extracellular GABA accumulation caused by interneuron firing in the presence of kainate. Extracellular GABA also shunts synaptic currents by activating tonic conductance as well as desensitizing synaptic GABAA receptors. Thus, the enhancement of the evoked IPSCs by 1 microM kainate was complicated by early and transient decrease. The kainate receptor-mediated enhancement of GABAergic tonic and phasic signalling to interneurons can contribute to the depression of GABAergic transmission to pyramidal neurons. The consequence of this phenomenon may play a major role in the epileptogenic action of this agent.     

Contextual fear conditioning differs for infant, adolescent, and adult rats

Contextual fear conditioning was tested in infant, adolescent, and adult rats in terms of Pavlovian-conditioned suppression. When a discrete auditory-conditioned stimulus (CS) was paired with footshock (unconditioned stimulus, US) within the largely olfactory context, infants and adolescents conditioned to the context with substantial effectiveness, but adult rats did not. When unpaired presentations of the CS and US occurred within the context, contextual fear conditioning was strong for adults, weak for infants, but about as strong for adolescents as when pairings of CS and US occurred in the context. Nonreinforced presentations of either the CS or context markedly reduced contextual fear conditioning in infants, but, in adolescents, CS extinction had no effect on contextual fear conditioning, although context extinction significantly reduced it. Neither CS extinction nor context extinction affected responding to the CS–context compound in infants, suggesting striking discrimination between the compound and its components. Female adolescents showed the same lack of effect of component extinction on response to the compound as infants, but CS extinction reduced responding to the compound in adolescent males, a sex difference seen also in adults. Theoretical implications are discussed for the development of perceptual-cognitive processing and hippocampus role.     

Separation of Forearm Hemodynamics Into Skin and Muscle Components by Means of I-Epinephrine Iontophoresis

By a combination of iontophoresis of I-epinephrine into the skin of one arm and simultaneous venous occlusion plethysmography in both treated (muscle only) and untreated forearms (muscle plus skin), we examined in 16 normal volunteers forearm blood flow, capillary filtration coefficient and venous capacity at cuff pressure of 40 mm of mercury (VC₄₀) at rest, during tonic finger exercise and after interrupted repetitive finger exercise. Blood pressure did not change during the testing procedure. Forearm muscle conductance was about 60% to 70% of total conductance and was positively correlated with total conductance during rest and exercise. With standard exercises muscle conductance rises to 1½ to 2½ times resting level, and skin conductance rises to 2½ to 4½ times resting level. The capillary filtration coefficient is almost entirely in the muscle. It doubles in value with tonic exercise but decreases to half its resting value after interrupted repetitive exercise despite greatly increased conductance. Therefore, repetitive exercise-induced dissociation between conductance and filtration surface occurs in striated muscle. The mechanism is yet unknown. VC₄₀ in muscle is about 84% of total forearm VC₄₀. During tonic exercise muscle VC₄₀ was reduced, and during interrupted repetitive exercise the values for muscle and skin returned to resting values. A high correlation between muscle only and muscle-plus-skin for forearm blood flow and the identify between arms for measuring capillary filtration coefficient makes iontophoresis unnecessary for determining these values in forearm striated muscle under these experimental conditions.     

Multimodal Assessment of Long-Term Memory Recall and Reinstatement in a Combined Cue and Context Fear Conditioning and Extinction Paradigm in Humans

Learning to predict danger via associative learning processes is critical for adaptive behaviour. After successful extinction, persisting fear memories often emerge as returning fear. Investigation of return of fear phenomena, e.g. reinstatement, have only recently began and to date, many critical questions with respect to reinstatement in human populations remain unresolved. Few studies have separated experimental phases in time even though increasing evidence shows that allowing for passage of time (and consolidation) between experimental phases has a major impact on the results. In addition, studies have relied on a single psychophysiological dimension only (SCRs/SCL or FPS) which hampers comparability between different studies that showed both differential or generalized return of fear following a reinstatement manipulation. In 93 participants, we used a multimodal approach (fear-potentiated startle, skin conductance responses, fear ratings to asses fear conditioning (day 1), extinction (day 2) as well as delayed memory recall and reinstatement (day 8) in a paradigm that probed contextual and cued fear intra-individually. Our findings show persistence of conditioning and extinction memory over time and demonstrate that reinstated fear responses were qualitatively different between dependent variables (subjective fear ratings, FPS, SCRs) as well as between cued and contextual CSs. While only the arousal-related measurement (SCRs) showed increasing reactions following reinstatement to the cued CSs, no evidence of reinstatement was observed for the subjective ratings and fear-related measurement (FPS). In contrast, for contextual CSs, reinstatement was evident as differential and generalized reinstatement in fear ratings as well as generally elevated physiological fear (FPS) and arousal (SCRs) related measurements to all contextual CSs (generalized non-differential reinstatement). Returning fear after reinstatement likely depends on a variety of variables (experimental design, dependent measurements) and more systematic investigations with respect to critical determinants of reinstatement in humans are required.     

Extinction and Renewal of Conditioned Sexual Responses

Introduction

Extinction involves an inhibitory form of new learning that is highly dependent on the context for expression. This is supported by phenomena such as renewal and spontaneous recovery, which may help explain the persistence of appetitive behavior, and related problems such as addictions. Research on these phenomena in the sexual domain is lacking, where it may help to explain the persistence of learned sexual responses.

Method

Men (n = 40) and women (n = 62) participated in a differential conditioning paradigm, with genital vibrotactile stimulation as US and neutral pictures as conditional stimuli (CSs). Dependent variables were genital and subjective sexual arousal, affect, US expectancy, and approach and avoid tendencies towards the CSs. Extinction and renewal of conditioned sexual responses were studied by context manipulation (AAA vs. ABA condition).

Results

No renewal effect of genital conditioned responding could be detected, but an obvious recovery of US expectancy following a context change after extinction (ABA) was demonstrated. Additionally, women demonstrated recovery of subjective affect and subjective sexual arousal. Participants in the ABA demonstrated more approach biases towards stimuli.

Conclusions

The findings support the context dependency of extinction and renewal of conditioned sexual responses in humans. This knowledge may have implications for the treatment of disturbances in sexual appetitive responses such as hypo- and hypersexuality.     

Failure to condition evaluative and electrodermal responses to neutral stimuli by means of prepared cues

To test Gray's proposal (1979, 1982) that prepared cues (snakes, spiders) are unconditioned, innate fear stimuli, a classical conditioning procedure was carried out in which a neutral, conditioned stimulus (CS; slides of flowers, mushrooms or apples) was repeatedly followed by a prepared, unconditioned stimulus (US; slides of snakes or spiders). Dependent variables were subjective evaluation of, and skin conductance responses to the CS slides, while respiration was used as a control variable. In one group of subjects (n = 12), skin conductance was recorded from the left hand, whereas in a second group (n = 11) it was recorded from the right hand. No evidence of skin conductance response - or of evaluative conditioning to the neutral CS as a result of its being paired with the prepared US was found. Thus, the results lend no support to Gray's proposal. No differences in skin conductance responses as a function of left versus right hand recording were found.     

Red - Take a Closer Look

Color research has shown that red is associated with avoidance of threat (e.g., failure) or approach of reward (e.g., mating) depending on the context in which it is perceived. In the present study we explored one central cognitive process that might be involved in the context dependency of red associations. According to our theory, red is supposed to highlight the relevance (importance) of a goal-related stimulus and correspondingly intensifies the perceivers’ attentional reaction to it. Angry and happy human compared to non-human facial expressions were used as goal-relevant stimuli. The data indicate that the color red leads to enhanced attentional engagement to angry and happy human facial expressions (compared to neutral ones) - the use of non-human facial expressions does not bias attention. The results are discussed with regard to the idea that red induced attentional biases might explain the red-context effects on motivation.     

Contrasting the overexpectation and extinction effects

After many target stimulus (X)-unconditioned stimulus (US) pairings, further conditioning of X in the presence of another well-established signal for the US (A) disrupts X's behavioral control. Some researchers have argued that the mechanism underlying this so-called overexpectation effect is similar to that underlying extinction (a reduction in X's behavioral control due to X-alone presentations). Three conditioned suppression experiments with rats as subjects compared overexpectation and extinction. Experiment 1 replicated the basic overexpectation effect by showing that A disrupts responding to X more than does a previously neutral stimulus. Experiment 2 found that posttraining context exposure disrupts extinction but not overexpectation. Experiment 3 suggested that overexpectation and extinction are differentially sensitive to the effects of overtraining (compound reinforced or nonreinforced, respectively), such that extinction is enhanced by increases in the amount of nonreinforced trials and overexpectation is unaffected. These results are inconsistent with the view that overexpectation and extinction are driven by a common mechanism.     

Responses of caudate neurons to clicks in chronic experiments on cats

Spontaneous and evoked unit activity in response to repeated application of clicks at a frequency of 0.3–2.0 Hz in the caudate nucleus was studied by an extracellular recording technique in chronic experiments on cats. Four types of spontaneous unit activity in the caudate nucleus were distinguished. Altogether 44% of neurons tested responded by changes in spontaneous activity to clicks. Five types of responses of caudate neurons to clicks were discovered: phasic excitation, phasic inhibition, tonic activation, tonic inhibition, and mixed tonic responses; the commonest type was tonic activation. During prolonged stimulation by clicks extinction of the phasic responses was not observed. Complete or partial extinction of tonic responses in the course of frequent repetition of stimulation was observed in 33% of responding neurons. The question of possible convergence of specific and nonspecific influences on caudate neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 28–35, January–February, 1980.     

Unconditioned Stimulus Revaluation to Promote Conditioned Fear Extinction in the Memory Reconsolidation Window

The retrieval-extinction paradigm, which disrupts the reconsolidation of fear memories in humans, is a non-invasive technique that can be used to prevent the return of fear in humans. In the present study, unconditioned stimulus revaluation was applied in the retrieval-extinction paradigm to investigate its promotion of conditioned fear extinction in the memory reconsolidation window after participants acquired conditioned fear. This experiment comprised three stages (acquisition, unconditioned stimulus revaluation, retrieval-extinction) and three methods for indexing fear (unconditioned stimulus expectancy, skin conductance response, conditioned stimulus pleasure rating). After the acquisition phase, we decreased the intensity of the unconditioned stimulus in one group (devaluation) and maintained constant for the other group (control). The results indicated that both groups exhibited similar levels of unconditioned stimulus expectancy, but the devaluation group had significantly smaller skin conductance responses and exhibited a growth in conditioned stimulus + pleasure. Thus, our findings indicate unconditioned stimulus revaluation effectively promoted the extinction of conditioned fear within the memory reconsolidation window.     

Resting Heart Rate Variability Predicts Safety Learning and Fear Extinction in an Interoceptive Fear Conditioning Paradigm

This study aimed to investigate whether interindividual differences in autonomic inhibitory control predict safety learning and fear extinction in an interoceptive fear conditioning paradigm. Data from a previously reported study (N = 40) were extended (N = 17) and re-analyzed to test whether healthy participants'' resting heart rate variability (HRV) - a proxy of cardiac vagal tone - predicts learning performance. The conditioned stimulus (CS) was a slight sensation of breathlessness induced by a flow resistor, the unconditioned stimulus (US) was an aversive short-lasting suffocation experience induced by a complete occlusion of the breathing circuitry. During acquisition, the paired group received 6 paired CS-US presentations; the control group received 6 explicitly unpaired CS-US presentations. In the extinction phase, both groups were exposed to 6 CS-only presentations. Measures included startle blink EMG, skin conductance responses (SCR) and US-expectancy ratings. Resting HRV significantly predicted the startle blink EMG learning curves both during acquisition and extinction. In the unpaired group, higher levels of HRV at rest predicted safety learning to the CS during acquisition. In the paired group, higher levels of HRV were associated with better extinction. Our findings suggest that the strength or integrity of prefrontal inhibitory mechanisms involved in safety- and extinction learning can be indexed by HRV at rest.     

Renewal and reinstatement of the conditioned but not the unconditioned response following habituation of the unconditioned stimulus

Research on the inhibition of learned fear currently relies almost exclusively on one specific procedure, namely extinction of the conditioned stimulus (CS). Importantly, however, learned fear responses can be reduced by a number of other procedures, including habituation of the unconditioned stimulus (US). We recently demonstrated that reductions in learned fear following US habituation, like CS extinction, were subject to both renewal and reinstatement (Storsve et al., 2010). The present study further investigates the associative and non-associative processes shared between habituation and extinction. Given that habituation is typically context-independent (Mackintosh, 1987), in the present study we directly compared renewal and reinstatement of both a conditioned response (CR; freezing) and an unconditioned response (UR; startle) following habituation. It was found that the reduction in conditioned freezing resulting from habituation was context specific (i.e., a change in context led to a renewal of the conditioned fear response; Experiment 1) and was attenuated when a pre-test shock was given (i.e., reinstatement of conditioned fear was observed; Experiment 2). In contrast, habituation of an unconditioned response elicited by the US (i.e., a startle response) was unaffected by either a change in test context or administration of a pre-test shock. This dissociation in the effects of habituation on learned and unlearned responses is discussed in relation to theories of fear extinction.     

Fear Conditioning in an Abdominal Pain Model: Neural Responses during Associative Learning and Extinction in Healthy Subjects

Fear conditioning is relevant for elucidating the pathophysiology of anxiety, but may also be useful in the context of chronic pain syndromes which often overlap with anxiety. Thus far, no fear conditioning studies have employed aversive visceral stimuli from the lower gastrointestinal tract. Therefore, we implemented a fear conditioning paradigm to analyze the conditioned response to rectal pain stimuli using fMRI during associative learning, extinction and reinstatement.In N = 21 healthy humans, visual conditioned stimuli (CS⁺) were paired with painful rectal distensions as unconditioned stimuli (US), while different visual stimuli (CS⁻) were presented without US. During extinction, all CSs were presented without US, whereas during reinstatement, a single, unpaired US was presented. In region-of-interest analyses, conditioned anticipatory neural activation was assessed along with perceived CS-US contingency and CS unpleasantness.Fear conditioning resulted in significant contingency awareness and valence change, i.e., learned unpleasantness of a previously neutral stimulus. This was paralleled by anticipatory activation of the anterior cingulate cortex, the somatosensory cortex and precuneus (all during early acquisition) and the amygdala (late acquisition) in response to the CS⁺. During extinction, anticipatory activation of the dorsolateral prefrontal cortex to the CS⁻ was observed. In the reinstatement phase, a tendency for parahippocampal activation was found.Fear conditioning with rectal pain stimuli is feasible and leads to learned unpleasantness of previously neutral stimuli. Within the brain, conditioned anticipatory activations are seen in core areas of the central fear network including the amygdala and the anterior cingulate cortex. During extinction, conditioned responses quickly disappear, and learning of new predictive cue properties is paralleled by prefrontal activation. A tendency for parahippocampal activation during reinstatement could indicate a reactivation of the old memory trace. Together, these findings contribute to our understanding of aversive visceral learning and memory processes relevant to the pathophysiology of chronic abdominal pain.     

The ratio between the phasic and tonic components of the frontal midline Θ rhythm in the attention test

The frontal midline Θ rhythm in the GO/NOGO paradigm was studied in a group of apparently healthy children at ages of 7–13 years. Calculated event-related synchronization in response to stimulus presentation in tests was used as an index of the phasic component, and the relative change in the EEG power in the Θ band (compared to the activity in the state of quiet wakefulness) in response to test performance was used as an index of the tonic component. Subjects were divided into two groups according to the characteristics of the baseline Θ activity. A statistically significant correlation between the phasic and tonic components of the Θ rhythm was found in the group of children characterized by the absence of the frontal midline Θ rhythm in the baseline EEG. No such correlation was found in the group of children characterized by a pronounced baseline Θ rhythm. The results testify to the functional heterogeneity of the phasic and tonic components of the human midline Θ rhythm.     

Short-Latency Afferent Inhibition Modulation during Finger Movement

When somatosensory input via electrical stimulation of a peripheral nerve precedes a transcranial magnetic stimulation (TMS) pulse over the primary motor cortex (M1) the corticospinal output is substantially reduced, a phenomenon known as short-latency afferent inhibition (SAI). The present study investigated SAI during rest and during pre-movement, phasic and tonic components of movement. Participants were required to perform an index finger flexion reaction time task in response to an auditory cue. In a series of experiments, SAI was evoked from the mixed, median nerve at the wrist or the cutaneous, digital nerve stimulation of the index finger. To assess the spinal versus cortical origin of movement-related modulation of SAI, F-wave amplitudes were measured during rest and the three movement components. Results indicated that SAI was reduced during all movement components compared to rest, an effect that occurred for both nerves stimulated. Pre-movement SAI reduction was primarily attributed to reduced cortical inhibition, while increased spinal excitability additionally contributed to reduced SAI during tonic and phasic components of movement. SAI was differentially modulated across movement components with mixed but not cutaneous nerve stimulation. These findings reveal that SAI is reduced during movement and this reduction begins as early as the preparation to move. Further, these data suggest that the degree of SAI reduction during movement may be specific to the volume and/or composition of afferent input carried by each nerve.     

Effects of vasopressin on the habituation of the orienting reaction in men

Effects of arginine-vasopressin (AVP) on the habituation of the orienting reaction and response to stimulus mismatch were investigated in a between-group design with 40 healthy male volunteers using skin conductance and heart rate responses as dependent measures. Twenty-one 1000 Hz tones of 90 dB(A) intensity and 2 s duration were presented with alternating intervals of 20 and 140 s. Stimulus mismatch responses were analyzed to the tones after the long intervals and to a change of the interval duration. The expected prevention of habituation as an indicator of a general stimulus-related increase of phasic arousal under AVP could not be confirmed. There were no differences between the AVP and the placebo group in the skin conductance and heart rate responses. The interval change did not provoke a dishabituation reaction, but responses to the tones after the long intervals were reliably enhanced. However, AVP did not increase the reaction to stimulus mismatch. It is concluded that autonomic indicators of the habituation of the OR remain unaffected by AVP.