Shock signals and the development of stress-induced analgesia

We report five experiments in which we investigated the effects of "feedback signals" on the pattern of hypoalgesia produced by inescapable shocks. A 5-s lights-out stimulus coincident with shock termination had no effect on the naltrexone-insensitive (nonopioid) hypoalgesia, which occurred after 10 inescapable shocks, but completely blocked the naltrexone-sensitive (opioid) hypoalgesia, which followed 100 inescapable shocks. The stimulus prevented the development of the opioid hypoalgesia rather than merely masking its measurement. This effect did not depend on the use of lights-out as the stimulus but did depend on the temporal relation between the stimulus and shock. Stimuli immediately preceding or simultaneous with shock had no effect. Surprisingly, stimuli randomly related to shock also blocked the opioid hypoalgesia. Simultaneous measurement of both hypoalgesia and fear conditioned to contextual cues revealed that the level of fear did not predict the blockade of hypoalgesia. Different backward groups received different temporal gaps between shock termination and the signal. An interval between 2.5 s and 7.5 s eliminated the effect of the signal on fear, but 12.5-17.5 s were required to eliminate the effect of the signal on hypoalgesia. The opioid hypoalgesia blocking power of the random stimulus was entirely attributable to those stimuli occurring within 15 s of the termination of the preceding shock. The implications of these results for the explanation of stimulus feedback effects and for stress-induced analgesia are discussed.     

Feedback during exposure to inescapable shocks and subsequent shock-escape performance

In two experiments, we assessed the ability of a feedback stimulus during helplessness training to reduce the performance deficits common to inescapable shock. In each experiment, four groups of rats were exposed to either escapable shock (E), inescapable shock with a feedback stimulus following shock termination (Y-FS), inescapable shock with no feedback stimulus (Y-NFS), or no shock (N). The feedback stimulus eliminated the interference effects of inescapable shock when tested with an FR-3 lever press escape task (Experiment 1) or on an FR-1 task with a 3-s delay between the response and shock termination (Experiment 2). These results suggest that stress-induced biochemical changes may mediate the interference effects seen in inescapably shocked rats.     

Warning signal termination and a feedback signal may not serve the same function

Three experiments were conducted to test the possibility that a feedback signal (FS) and warning signal termination (WST), while equally reinforcing in the avoidance learning situation, reinforce through different underlying mechanisms. The first experiment showed that the reinforcing properties of an FS are reduced more than those of WST when these stimulus changes are made unreliable by the presence of shock following a CR on specified trials throughout acquisition. Experiment 2 confirmed this effect on avoidance performance when only a few punishment trials were administered following asymptotic avoidance acquisition. Experiment 3 demonstrated this effect during avoidance extinction with and without the presentation of punishment trials between acquisition and extinction performance. The results provided no support for the expectancy explanation of avoidance learning and were interpreted as consistent with the assumption that WST reinforces by permitting fear to dissipate and that the FS reinforces through fear inhibition.     

Effects of signaling inescapable shock on subsequent escape learning: implications for theories of coping and "learned helplessness"

The present experiments reveal that shuttle-escape performance deficits are eliminated when exteroceptive cues are paired with inescapable shock. Experiment 1 indicated that, as in instrumental control, a signal following inescapable shock eliminated later escape performance deficits. Subsequent experiments revealed that both forward and backward pairings between signals and inescapable shock attenuated performance deficits. However, the data also suggest that the impact of these temporal relations may be modulated by qualitative aspects of the cues because the effects of these relations depended upon whether an increase or decrease in illumination (Experiment 2) or a compound auditory cue (Experiment 4) was used. Preliminary evidence suggests that the ability of illumination cues to block escape learning deficits may be related to their to reduce contextual fear (Experiment 3). The implications of these data for conceptions of instrumental control and the role of fear in the etiology of effects of inescapable shock exposure are discussed.     

Reversibility of single-incentive selective associations.

Rats were trained to press a lever in the presence of a tone-light compound stimulus and not to press in its absence. In each of two experiments, schedules were designed to make the compound a conditioned punisher for one group and a conditioned reinforcer for the other. In Experiment 1, one group's responding produced food in the presence of the compound but not in its absence. The other group's responding terminated the compound stimulus, and food was presented only in its absence. When tone and light were later presented separately, light controlled more responding than did tone in the former group, but tone gained substantial control in the latter. The same effects were also observed within subjects when the training schedules were switched over groups. In Experiment 2, two groups avoided shock in the presence of the compound stimulus. In the absence of the compound, one group was not shocked, and the other received both response-independent and response-produced shock. When tone and light were presented separately, the former group's responding was mainly controlled by tone, but the latter group's responding was almost exclusively controlled by light. These effects were also observed within subjects when the training schedules were switched over groups. Thus, these single-incentive selective association effects (appetitive in Experiment 1 and aversive in Experiment 2) were completely reversible. The schedules in which the compound should have been a conditioned reinforcer consistently produced visual control, and auditory control increased when the compound should have become a conditioned punisher. Currently accepted accounts of selective associations based on affinities between shock and auditory stimuli and between food and visual stimuli (i.e., stimulus-reinforcer interactions) do not adequately address these results. The contingencies of reinforcement most recently associated with the compound and with its absence, rather than the nature of the reinforcer, determined whether auditory or visual stimulus control developed.     

Mechanisms of sentence context effects in reading: Automatic activation and conscious attention

In. Experiment I subjects named target words that were preceded by a congruous sentence context, an incongruous sentence context, or no sentence context, under stimulus conditions that were either normal or degraded by contrast reduction. Under normal stimulus conditions, a contextual facilitation effect, but no contextual inhibition effect, was observed. When the target word was degraded, both contextual facilitation and inhibition were observed. Experiment 2 replicated the increase in contextual inhibition under degraded conditions and also demonstrated that inhibition increased as the interval between contextual processing and target-word onset was lengthened. The results were interpreted within the framework of the Posner and Snyder two-process theory of expectancy. Thus, when target-word recognition is rapid, only the fast-acting automatic activation component of context effects has time to operate. When target-word processing is delayed, the conscious-attention mechanism, which is responsible for inhibition effects, becomes operative. The relevance of these results to developmental investigations of the interaction of word recognition and contextual processing is discussed.     

Contextual Control of Latent Inhibition by the Reinforcer

Three experiments examined the contextual control of latent inhibition (LI) by the unconditioned stimulus (US) using a within-subjects conditioned suppression procedure with rats. The effect of reducing the context change produced by the introduction of the shock US was investigated by presenting this US during preexposure to the conditioned stimulus (CS). Although limited CS preexposure in the absence of the US had no impact on subsequent conditioning, preexposure in the presence of the shock retarded both excitatory and inhibitory conditioning. We conclude that the introduction of the US during the conditioning phase of a normal LI experiment can produce a contextual change that reduces the observed magnitude of LI.     

An immediate-shock freezing deficit with discrete cues: a possible role for unconditioned stimulus processing mechanisms.

Five experiments with C57BL/6 mice (Mus musculus) investigated whether failures in shock processing might contribute to deficits in freezing that occur after an animal receives a shock immediately on exposure to a conditioning context. Experiment 1 found that more contextual freezing resulted from delayed shocks than from immediate shocks across 4 shock intensities. Experiment 2 extended the immediate-shock freezing deficit to discrete stimuli. Experiment 3 found that preexposure to the to-be-conditioned cue did not facilitate immediate cued conditioning. Experiment 4 found that context preexposure enhanced context-evoked fear after an immediate shock. Experiment 5 found that context preexposure also enhanced immediate cued conditioning. These findings are problematic for current theories of the immediate-shock freezing deficit that focus exclusively on processing of the conditioned stimulus, and they suggest that failures in shock processing may contribute to the deficit.     

No sex difference in contextual control over the expression of latent inhibition and extinction in Pavlovian fear conditioning in rats

Three experiments with Wistar rats searched for a sex difference in contextual control over the expression of latent inhibition and extinction. Experiment 1 used a latent inhibition procedure; Experiments 2 and 3 employed an extinction preparation. All experiments used a shock as the unconditioned stimulus, a tone as the conditioned stimulus, and suppression of food magazine visits as the measure of conditioned responding to the tone. Each experiment revealed a reliable context effect on conditioned responding to the tone; after conditioning in a separate context, conditioned responding in the former latent inhibition or extinction context was attenuated relative to conditioned responding in a control context. There was no sex difference in the magnitude of this effect. These results are discussed in the framework of sex differences in the hippocampus and of the putative role of this structure in various instances of contextual learning.     

What are human express saccades?

When a fixation point is removed 200 msec prior to target onset (the gap condition), human subjects are said to produce eye movements that have a short latency (80–120 msec), that form the early peak of a bimodal latency distribution, and that have been labeled “human express saccades” (see, e.g., Fischer, 1987; Fischer & Breitmeyer, 1987; Fischer & Ramsperger, 1984, 1986). In three experiments, we sought to obtain this express saccade diagnostic pattern in the gap condition, We orthogonally combined target location predictability with the presence versus absence of catch trials (Experiment 1). When target location was fixed and catch trials were not used, we found mostly anticipations. In the remaining conditions, where responses were under stimulus control, bimodality was not frequently observed, and, whether it was or not, latencies were not in the express saccade range. Using random target locations, we then varied stimulus luminance and the mode of stimulus presentation (LEDs vs. oscilloscope) in the gap and overlap (fixation is not removed) conditions (Experiment2). Bimodality was rarely observed, the gap effect (overlap minus gap reaction time) was additive with luminance, and only the brightest targets elicited saccades in the express range. When fixed locations and no catch trials were combined with latency feedback (Experiment 3), we observed many responses in the express saccade range and some evidence for bimodality, but the sudden introduction of catch trials revealed that many early responses were not under stimulus control. Humanscan make stimulus-controlled saccades that are initiated very rapidly (80–120 msec), but unless catch trials or choice reaction time is used, it is not possible to distinguish such saccades from anticipatory responses that are prepared in advance and timed to occur shortly after target onset. Because the express saccade diagnostic pattern is not a characteristic feature of human saccadic performance, we urge investigators to focus their attention on the robustgap effect     

Stimulus specificity in avoidance learning

The present experiments address the issue of stimulus specificity in fear and avoidance learning. First, it was established that light stimuli are effective warning signals (WS) in shuttle avoidance. Then light stimuli were shown to produce conditioned suppression in the conditioned emotional response situation comparable to that produced by noise conditioned stimuli. Finally, the effectiveness of noise onset and noise offset as feedback signals was tested. This was assessed under conditions of immediate and delayed termination of a light WS. Delayed termination of a light WS interfered with avoidance learning and the introduction of noise offset as a feedback signal enhanced it. The only demonstration of stimulus specificity was the failure of noise onset to function as a feedback signal.     

US habituation,like CS extinction,produces a decrement in conditioned fear responding that is NMDA dependent and subject to renewal and reinstatement

Just as fear can be learned, it can also be inhibited. The most common way of reducing learned fear is through extinction, where the conditioned stimulus (CS) previously paired with an aversive unconditioned stimulus (US) is repeatedly presented on its own. Another, much less commonly studied, way to inhibit learned fear is by habituating, or devaluing, the US. In this procedure, fear responding to a CS is reduced by repeatedly presenting the US in the absence of the CS following the conditioning phase. The purpose of the present study was to directly compare the effects of US habituation and CS extinction on a learned fear response (freezing). Experiment 1 demonstrated that US habituation given either after (Experiment 1A) or before (Experiment 1B) fear conditioning reduced freezing to the CS at test. We then showed that the reduction in freezing resulting from either US habituation or CS extinction was context-specific (i.e., a change in context led to a renewal of the learned fear response; Experiment 2) and, furthermore, was attenuated when a pre-test shock was given (i.e., reinstatement of fear was observed in both cases; Experiment 3). Finally, Experiment 4 demonstrated that an injection of the NMDA antagonist MK-801 prior to US habituation impaired long-term retention of the learning that takes place during this procedure. Together, these results suggest that the decrement in conditioned fear responses produced by US habituation and CS extinction could rely on overlapping processes.     

Investigations of the reinstatement of extinguished fear

In two experiments, fear was conditioned to the situational cues in one compartment of a hurdle-jumping apparatus and was then extinguished. Subsequently, either one shock (Experiment 1) or three or nine shocks (Experiment 2) were given in a situation distinctively different from that in which conditioning and extinction had taken place. Although some associative strength between the situational cues and fear was shown to have remained after extinction, in neither experiment did the postextinction-shock treatment increase the fear elicited by these cues: Escape-from-fear performance was no better in the shocked groups than in control groups given no additional shock. Thus, the nonassociative hypothesis which postulates that inflating the value of the representation of the UCS with shock-alone presentations can reinstate the extinguished fear of a stimulus was not supported. Rather, the results showed that, after extinction, an increase in fear of a simulus depended on further conditioning to that stimulus. The data also indicated that the nonvisual components of the situational cues predominated over the visual component.     

Conflict adaptation depends on task structure

The dependence of the Simon effect on the correspondence of the previous trial can be explained by the conflict-monitoring theory, which holds that a control system adjusts automatic activation from irrelevant stimulus information (conflict adaptation) on the basis of the congruency of the previous trial. The authors report on 4 experiments showing that conflict adaptation is mediated by task structure as well as previous congruency. Experiment 1 showed that for 2 completely segregated sets of stimulus-response pairs there was no conflict adaptation across sets. However, if the stimulus sets overlapped for 2 separate response sets, conflict adaptation could be observed across the response sets. Experiment 2 showed that this effect was not due to the use of stimulus-response sets lateralized to 1 hemisphere each. Experiment 3 showed that if response sets are common for 2 separate stimulus sets, then conflict adaptation can again be observed across sets. Finally, Experiment 4 showed local control effects in the absence of confounding feature-overlap effects. These results indicate that deployment of control as evidenced by conflict adaptation reflects task structure.     

Naloxone and Pavlovian fear conditioning

Previous research has shown that pretreating rats with the opiate antagonist naloxone increases the freezing that follows painful electric shock. Three experiments, using freezing behavior as a dependent variable, were carried out to determine whether the drug might cause this effect by enhancing fear conditioning. Two of the studies employed a differential context fear-conditioning paradigm. Naloxone did not affect freezing behavior during the preshock adaptation period. In Experiment 1, naloxone was found to increase resistance to extinction in the S+ context. In Experiment 2, naloxone was found to increase freezing in the S+ context. This effect was dependent upon administering naloxone during training but not dependent on administering it during testing. The third study employed a generalization paradigm. It was found that naloxone's effect on postshock freezing was dependent on the place of testing; as the contextual cues of the test chamber were changed from those of the conditioning chamber, the effect of naloxone on freezing was reduced. The results of these experiments lend strong support to the hypothesis that naloxone increases freezing by enhancing the conditioning of fear to contextual stimuli associated with shock.     

Pavlovian processes and response competition as determinants of avoidance response-prevention effects

Two experiments investigated the facilitation of avoidance extinction by exposure to lengthy (5-sec) shock during avoidance response prevention. In Experiment 1, animals exposed to light only or to light-shock pairings during response prevention showed equal facilitation of extinction relative to shock-only animals or to animals receiving no response prevention. Preshock rearing, directly antagonistic to the avoidance response, developed for shocked animals during response prevention and persisted during extinction for light-shock animals. Immediately before extinction, half of each group was permitted a single escape from a light-shock compound by means of the response previously required for avoidance. The only effect was upon the extinction performance of light-shock animals. Rearing was eliminated and extinction responding increased to a level far above that for any of the other animals. Experiment 2 demonstrated that the shock-only treatment affected the extinction performance and rearing of nonescape and escape animals in a manner entirely equivalent to the effects of the light-shock treatment of Experiment 1, provided stimulus conditions (light absent) were the same for all experimental phases. Thus, lengthy shock during avoidance response prevention simultaneously leads to the acquisition of competing behavior and enhances control by a warning signal or contextual stimuli over the avoidance response. Implications for the CS-only response-prevention treatment and the transfer of aversive control are discussed.     

Conditioned-aversive aspects of electric shock

Two experiments were reported in which it was observed that a weak shock became more aversive as a result of appropriate pairing of that stimulus with a stronger shock. The enhanced aversiveness of the weak shock was reflected in Experiment 1 by an apparent increase in the reinforcement provided by response-contingent termination of that shock. In Experiment 2, the weak shock was observed to be more suppressive of a consummatory-response baseline following appropriate pairings of the weak and strong shocks. The results of these studies were discussed in terms of their implications for certain aversive-conditioning situations in which shocks may serve, in part, as conditioned stimuli.     

Effects of contextual predictability and transitional probability on eye movements during reading

In 2 eye-movement experiments, the authors tested whether transitional probability (the statistical likelihood that a word precedes or follows another word) affects reading times and whether this occurs independently from contextual predictability effects. Experiment 1 showed early effects of predictability, replicating S. A. McDonald and R. C. Shillcock's (2003a) finding that words with a high transitional probability (defeat following accept) are read faster than words with a low transitional probability (losses following accept). However, further analyses suggested that the transitional probability effect was likely due to differences in predictability rather than transitional probability. Experiment 2, using a better controlled set of items, again showed an effect of predictability, but no effect of transitional probability. The authors conclude that effects of transitional probability are part of regular predictability effects. Their data also show that predictability effects are detectable very early in the eye-movement record and between contexts that are weakly constraining.     

The effects of context changes on the reinstatement of extinguished conditioned behavior in a conditioned suppression task with humans

Reinstatement refers to the return of previously extinguished conditioned responses to test trials of a conditional stimulus (CS) when presentations of the unconditional stimulus (US) alone are given following extinction. Four experiments were conducted to determine whether reinstatement could be found in a conditioned suppression task with humans and whether contextual changes can abolish it. Experiment 1 demonstrated the reinstatement of conditioned suppression when acquisition, extinction, US alone, and test trials were all given in the same context. Experiments 2 and 3 suggested that the reinstatement effect was still present when the US alone presentations were given in a different context to the subsequent test trials. Experiment 4 replicated this effect using additional controls over the amount of exposure to the various contexts. The results suggest that reinstatement can be robust across changing contexts. Aspects of the conditioned suppression task that promote the transfer of learning across contexts or the establishment of configural context-CS stimuli may underlie the apparent lack of contextual control over reinstatement.