Electrodermal discrimination with visual feedback in two incompatible stressful situations. A yoked control comparison

One group (N = 14) of human volunteers received three sessions of discriminated avoidance and punishment with the skin resistance response (SRR) as the instrumental behavior. Each session consisted of three 7 minute periods of Sidman avoidance (response-stimulus [R-S] and stimulus-stimulus [S-S] = 40 sec) of a 0.5 second, 15 Hz square wave shock mixed with three periods of punishment with the same shock. The avoidance and punishment periods were differentially signaled by red and green lights, and a circle appeared superimposed on the discriminative stimuli during a criterion SRR. A second group (N = 14) was yoked to the first by recording shock events on magnetic tape and shocking the yoked subject in the same temporal sequence. In both groups the visual feedback stimulus was consistently related to the subject's ongoing electrodermal behavior. As in previous research, the contingent shock subjects made significantly more SRRs during avoidance than during punishment in all three sessions. For yoked subjects the discrimination did not appear until the third session. The findings imply that the discrimination is at least partly independent of the avoidance and punishment contingencies, and they raise questions about the role of the feedback stimulus.     

Tolerance to the anti-avoidance properties of cholecystokinin-octapeptide

Previous investigations have suggested a neuroleptic-like action of cholecystokinin-octapeptide (CCK8) on conditioned-avoidance behavior. This study was initiated to test tolerance to this effect. Rats were trained to avoid electric shock in a shuttle box under a free-operant (Sidman) avoidance paradigm. Each shuttle response postponed a 0.2 sec, 1 mA shock for 20 sec. If the rat failed to respond, shock was delivered every 5 sec until a response occurred. After avoidance training, half of the rats received two daily injections of CCK8 (0.320 mg/kg, IP) and half received saline for 7 days. Rats were then tested on the Sidman avoidance 1 min after receiving CCK8 (0.640 mg/kg, IP) or saline. CCK8 depressed avoidance responding if rats received saline for 7 days prior to the test. Rats pretreated with CCK8 for 7 days were not significantly affected by CCK8 during the avoidance test. Thus, repeated injections of CCK8 result in tolerance to its anti-avoidance properties.     

Response-cost punishment via token loss with pigeons

Two experiments were conducted to investigate punishment via response-contingent removal of conditioned token reinforcers (response cost) with pigeons. In Experiment 1, key pecking was maintained on a two-component multiple second-order schedule of token delivery, with light emitting diodes (LEDs) serving as token reinforcers. In both components, responding produced tokens according to a random-interval 20-s schedule and exchange periods according to a variable-ratio schedule. During exchange periods, each token was exchangeable for 2.5-s access to grain. In one component, responses were conjointly punished according to fixed-ratio schedules of token removal. Response rates in this punishment component decreased to low levels while response rates in the alternate (no-punishment) component were unaffected. Responding was eliminated when it produced neither tokens nor exchange periods (Extinction), but was maintained at moderate levels when it produced tokens in the signaled absence of food reinforcement, suggesting that tokens served as effective conditioned reinforcers. In Experiment 2, the effect of the response-cost punishment contingency was separated from changes in the density of food reinforcement. This was accomplished by yoking either the number of food deliveries per component (Yoked Food) or the temporal placement of all stimulus events (tokens, exchanges, food deliveries) (Yoked Complete), from the punishment to the no-punishment component. Response rates decreased in both components, but decreased more rapidly and were generally maintained at lower levels in the punishment component than in the yoked component. In showing that the response-cost contingency had a suppressive effect on responding in addition to that produced by reductions in reinforcement density, the present results suggest that response-cost punishment shares important features with other forms of punishment.     

Characterization of radiation-induced performance decrement using a two-lever shock-avoidance task

Rats were trained to perform a task involving responses on two levers. Responding on an avoidance lever delayed the onset of electrical footshock for 20 sec and responding on a warning lever turned on a light for 60 sec. When the light was on, the task on the avoidance lever was changed from unsignaled shock avoidance to signaled shock avoidance by preceding the shocks with 5-sec warning tones. The animals preferred the signaled avoidance condition. After 100 Gy of 60Co irradiation, the animals were less able to avoid shock, an effect from which the animals recovered somewhat over 90 min. The response rate on the avoidance lever remained at or above control rates, while the response rate on the warning lever showed an initial increase, followed by a decrease below baseline. The increase in responding on the avoidance lever occurred in bursts just after presentation of the shocks. The data suggest that under these experimental conditions a subject will not respond appropriately to avoid shock or acquire cues that can facilitate the avoidance of shock. The effects, however, do not reflect an inability to perform the required movements but instead appear to reflect some characteristic of the task associated with a particular lever.     

Dopamine and octopamine influence avoidance learning of honey bees in a place preference assay

Biogenic amines are widely characterized in pathways evaluating reward and punishment, resulting in appropriate aversive or appetitive responses of vertebrates and invertebrates. We utilized the honey bee model and a newly developed spatial avoidance conditioning assay to probe effects of biogenic amines octopamine (OA) and dopamine (DA) on avoidance learning. In this new protocol non-harnessed bees associate a spatial color cue with mild electric shock punishment. After a number of experiences with color and shock the bees no longer enter the compartment associated with punishment. Intrinsic aspects of avoidance conditioning are associated with natural behavior of bees such as punishment (lack of food, explosive pollination mechanisms, danger of predation, heat, etc.) and their association to floral traits or other spatial cues during foraging. The results show that DA reduces the punishment received whereas octopamine OA increases the punishment received. These effects are dose-dependent and specific to the acquisition phase of training. The effects during acquisition are specific as shown in experiments using the antagonists Pimozide and Mianserin for DA and OA receptors, respectively. This study demonstrates the integrative role of biogenic amines in aversive learning in the honey bee as modeled in a novel non-appetitive avoidance learning assay.     

Specificity of finger pulse volume feedback during relaxation

The effect of biofeedback during brief periods of relaxation was examined. Two groups (10 subjects in each group) were asked to relax as completely as possible during a series of six 3-minute relaxation periods in each of two 1-hr sessions. One group received biofeedback based on finger pulse volume (FPV) during the relaxation trials, while the other group received no biofeedback. Measures of heart rate, respiration rate, skin conductance level, and FPV were recorded during the sessions, and subjective ratings concerning relaxation were obtained after each session. The results showed that FPV scores for the groups differed during the relaxation trials of the second session, but other measures failed to distinguish between the groups. The group that received FPV feedback revealed a significantly higher level of FPV (relative to baseline) than the group that received no feedback.     

Pre-test administration of beta-endorphin, or of electroconvulsive shock reverses the memory disruptive effect of posttraining electroconvulsive shock

Memory disruption by posttraining electroconvulsive shock was studied in adult Wistar rats using three different tasks: step-down inhibitory avoidance, two-way active avoidance, and habituation of rearing to an open field. The animals were given training and test sessions 24 hours apart in each of these tasks. Immediate posttraining transcorneal, 15 mA, 60 Hz, 2 sec electroconvulsive shock disrupted memory of the three tasks. The effect was completely reversed by the IP administration of beta-endorphin (2.0 micrograms/kg), 6 min prior to testing, or of another electroconvulsive shock, 30 min prior to testing. These findings indicate that the posttraining electroconvulsive shock did not affect memory storage. In view of the fact that electroconvulsive shock has been previously shown to cause a pronounced decrease of brain beta-endorphin immunoreactivity, attributable to a release of the peptide, the present findings can be interpreted as showing that memory disruption by posttraining electroconvulsive shock results from the induction of state dependency on beta-endorphin.     

Instrumental conditioning of skin potential responses in curarized cats: non specific effects of a punishment procedure

This study evaluates the effects of a punishment paradigm upon electrodermal activity (EDA) in curarized cats. In ten experimental subjects, any skin potential response (SPR) exceeding threshold value was punished by an electric shock. Ten control subjects received an electric shock or an electric stimulation of the mesencephalic reticular formation unrelated to the EDA. The results show that: a) No significant modifications of EDA occurred in subjects characterized by low initial activity; b) The frequency of SPR significantly decreased in high activity subjects used both for the experiments and the controls. Therefore, this diminution, in experimental subjects, cannot be due to a learning process. Neither can it be related to level of curarization or to rate of artificial ventilation. It is suggested that this effect results from a non-specific cortical inhibitory mechanism acting on the reticular activating system. The results obtained in three decorticated cats submitted to a punishment procedure are in accordance with such an hypothesis. Consequently the existence of operant autonomic learning remains unproven.     

Voluntary heart rate deceleration: A critical evaluation

This experiment was designed as a test of the view that the human heart rate (HR) deceleration response can be brought under voluntary control, when some form of exteroceptive feedback is available. Sixteen female volunteers were randomly assigned to two groups. The first group received instructions to decrease their HR plus a continuous negative (failure) binary feedback, while the second group received only the instructions. Each subject was given four sessions of HR deceleration training. Two identical tests were presented, one before and the other after the series of training sessions. These tests were divided into two parts. In the first part, subjects attempted to decrease their HR while undergoing an ischemic arm pain stress. In the second part, subjects performed a 40-trial HR discrimination task. The results indicate that all subjects decrease HR during both rest and voluntary control periods within each training session, but there are no significant group differences, no improvement in HR deceleration control over the four training sessions, and no difference in performance between rest and voluntary control periods. Similarly HR, blood pressure (BP), and the HR×BP product levels during the ischemic stress condition and the HR discrimination performance do not show group differences. It is suggested that the HR deceleration response may not meet the criteria generally applied to the definition of a voluntary response.     

Cognitive and physiologic responses to EMG biofeedback and three types of pseudofeedback during a muscular relaxation task

Four groups of normal human subjects were tested for their ability to reduce frontal muscle tension levels during presentation of veridical auditory biofeedback or auditory pseudofeedback. A double-blind methodology was used. Three groups of subjects assigned to the pseudofeedback conditions received a feedback signal that was not contingent on EMG activity but that followed one of three different patterns. One group received a truly random signal, the second received a signal that gradually increased in frequency (apparent failure), and the third received a signal that gradually decreased in frequency (apparent success). Dependent measures included both physiologic (frontal and neck EMG) and subjective reactions to the relaxation task. The different patterns of pseudofeedback did produce reliably different subjective responses, suggesting that the manipulations succeeded in producing unequal nonspecific effects that were unrelated to the feedback contingency specifically. However, these differential subjective effects were not strongly reflected in the physiologic responses since the differences in EMG levels among the four groups did not differ significantly at any stage of training. An analysis of the integrity of the double-blind procedure showed that although experimenters were effectively kept blind to group assignment, subjects' responding suggested a response bias as well as the possibility that the double-blind was breached. The utility of the double-blind methodology in biofeedback experiments is discussed and suggestions for future research are offered.     

Learning to Obtain Reward,but Not Avoid Punishment,Is Affected by Presence of PTSD Symptoms in Male Veterans: Empirical Data and Computational Model

Post-traumatic stress disorder (PTSD) symptoms include behavioral avoidance which is acquired and tends to increase with time. This avoidance may represent a general learning bias; indeed, individuals with PTSD are often faster than controls on acquiring conditioned responses based on physiologically-aversive feedback. However, it is not clear whether this learning bias extends to cognitive feedback, or to learning from both reward and punishment. Here, male veterans with self-reported current, severe PTSD symptoms (PTSS group) or with few or no PTSD symptoms (control group) completed a probabilistic classification task that included both reward-based and punishment-based trials, where feedback could take the form of reward, punishment, or an ambiguous “no-feedback” outcome that could signal either successful avoidance of punishment or failure to obtain reward. The PTSS group outperformed the control group in total points obtained; the PTSS group specifically performed better than the control group on reward-based trials, with no difference on punishment-based trials. To better understand possible mechanisms underlying observed performance, we used a reinforcement learning model of the task, and applied maximum likelihood estimation techniques to derive estimated parameters describing individual participants’ behavior. Estimations of the reinforcement value of the no-feedback outcome were significantly greater in the control group than the PTSS group, suggesting that the control group was more likely to value this outcome as positively reinforcing (i.e., signaling successful avoidance of punishment). This is consistent with the control group’s generally poorer performance on reward trials, where reward feedback was to be obtained in preference to the no-feedback outcome. Differences in the interpretation of ambiguous feedback may contribute to the facilitated reinforcement learning often observed in PTSD patients, and may in turn provide new insight into how pathological behaviors are acquired and maintained in PTSD.     

The effects of naloxone,diprenorphine, and diazepam on responding suppressed by pre-shock and pre-food stimuli

The lever-pressing of rats was reinforced with food according to a variable-interval 1-min schedule. In one group, occasional illumination of cue lights for 30-sec periods was followed by a brief electric shock; responding was suppressed during these periods. Naloxone (0.01–10 mg/kg) did not change the degree to which responding was suppressed during the pre-shock stimulus. Diprenorphine (0.1–10 mg/kg) slightly attenuated suppression, and diazepam (1.0–3.0 mg/kg) increased responding during the stimulus to normal levels. These results confirm that opiate antagonists do not always enhance the effects of shock on behavior. In a second group, occasional illumination of the cue lights for 20-sec periods was followed by delivery of free food pellets. Responding was also suppressed during the pre-food stimulus. Neither naloxone nor diprenorphine had any effect on response rate during this stimulus. In contrast to the results of earlier studies using benzodiazepines, diazepam (1.0–3.0 mg/kg) produced a marked attenuation of response suppression during the pre-food stimulus.     

Segmentation and the pairing hypothesis

The effect of stimulus contiguity and response contingency on responding in chain schedules was examined in two experiments. In Experiment 1, four pigeons were trained on two simple three-link chain schedules that alternated within sessions. Initial links were correlated with a variable-interval 30s schedule, and middle and terminal links were correlated with interdependent variable-interval 30s variable-interval 30s schedules. The combined duration of the interdependent schedules summed to 60s. The two chains differed with respect to signaling of the schedule components: a two-stimulus chain had one stimulus paired with the initial link and one stimulus paired with both the middle and the terminal link, while a three-stimulus chain had a different stimulus paired with the each of the three links. The results showed that the two-stimulus chain maintained lower initial-link responding than the three-stimulus chain. In Experiment 2, four pigeons were exposed to three separate conditions, the two- and three-stimulus chains of Experiment 1 and a three-stimulus chain that had a 3s delay to terminal-link entry from the middle-link response that produced it. The two-stimulus chain maintained lower initial-link responding than the three-stimulus chain, as in Experiment 1, and a similar initial-link responding was maintained by the two-stimulus chain and the three-stimulus chain with the delay contingency. The results demonstrate that a stimulus noncontiguous with food can maintain responding that is sometimes greater than a stimulus contiguous with food, depending on the response contingency for terminal-link entry. The results are contrary to the pairing hypothesis of conditioned reinforcement.     

The Role of Informative and Ambiguous Feedback in Avoidance Behavior: Empirical and Computational Findings

Avoidance behavior is a critical component of many psychiatric disorders, and as such, it is important to understand how avoidance behavior arises, and whether it can be modified. In this study, we used empirical and computational methods to assess the role of informational feedback and ambiguous outcome in avoidance behavior. We adapted a computer-based probabilistic classification learning task, which includes positive, negative and no-feedback outcomes; the latter outcome is ambiguous as it might signal either a successful outcome (missed punishment) or a failure (missed reward). Prior work with this task suggested that most healthy subjects viewed the no-feedback outcome as strongly positive. Interestingly, in a later version of the classification task, when healthy subjects were allowed to opt out of (i.e. avoid) responding, some subjects (“avoiders”) reliably avoided trials where there was a risk of punishment, but other subjects (“non-avoiders”) never made any avoidance responses at all. One possible interpretation is that the “non-avoiders” valued the no-feedback outcome so positively on punishment-based trials that they had little incentive to avoid. Another possible interpretation is that the outcome of an avoided trial is unspecified and that lack of information is aversive, decreasing subjects’ tendency to avoid. To examine these ideas, we here tested healthy young adults on versions of the task where avoidance responses either did or did not generate informational feedback about the optimal response. Results showed that provision of informational feedback decreased avoidance responses and also decreased categorization performance, without significantly affecting the percentage of subjects classified as “avoiders.” To better understand these results, we used a modified Q-learning model to fit individual subject data. Simulation results suggest that subjects in the feedback condition adjusted their behavior faster following better-than-expected outcomes, compared to subjects in the no-feedback condition. Additionally, in both task conditions, “avoiders” adjusted their behavior faster following worse-than-expected outcomes, and treated the ambiguous no-feedback outcome as less rewarding, compared to non-avoiders. Together, results shed light on the important role of ambiguous and informative feedback in avoidance behavior.     

Temporal threshold and response bias in the discrimination of empty intervals by cats

Cat's differential duration threshold was investigated by the method of limits in a schedule of discrimination of empty durations. The standard stimulus was 4 sec long throughout the experiment. The comparison stimulus was reduced from 10 to 5 sec by 1 sec steps in successive blocks of 5 sessions. Standard and comparison stimuli, delimited by 50 msec auditory signals, were equiprobably distributed, in a random sequential order of presentation in each trial. After a 2 sec delay, an auditory signal indicated that reinforcement was available upon a response on one of two levers. Weber fractions around .25 were obtained. Strong response bias developed in most cats. Some consequences of the inhibition of responding induced by the procedure were considered.     

A further study of choice and percentage reinforcement

In Experiment I four pigeons were trained in a concurrent chains procedure with fixed-ratio schedules (FR1) in the initial components and fixed-time schedules in the terminal components. Pecking one of the keys when both keys were white initiated a fixed time schedule on that key. A peck to the left key produced three stripes on the key. At the termination of the fixed-time component food always occurred. Pecking the other key produced either a circle or a triangle. If a circle appeared, reinforcement occurred. If a triangle appeared a brief timeout was given. Initially the stripes appeared on the left key and the circle and triangle on the left. This was reversed during the course of the experiment. In addition, sessions were conducted in which both circle and triangle sometimes preceded reinforcement and sometimes timeout. For most birds under most conditions there was a preference for the key that produced the circle and triangle. When these were uncorrelated with reinforcement and time out three of the birds preferred the key producing 100% reinforcement.

In Experiment II three factors were varied and VI 20 sec schedules were used in the initial links instead of FR1. The results showed that pigeons preferred the 50% condition more 1) the greater the duration of the terminal links, 2) the smaller the value on the initial link VI schedules and 3) the less the probability of food in the terminal link with stripes on the key.     

Common Neural Mechanisms Underlying Reversal Learning by Reward and Punishment

Impairments in flexible goal-directed decisions, often examined by reversal learning, are associated with behavioral abnormalities characterized by impulsiveness and disinhibition. Although the lateral orbital frontal cortex (OFC) has been consistently implicated in reversal learning, it is still unclear whether this region is involved in negative feedback processing, behavioral control, or both, and whether reward and punishment might have different effects on lateral OFC involvement. Using a relatively large sample (N = 47), and a categorical learning task with either monetary reward or moderate electric shock as feedback, we found overlapping activations in the right lateral OFC (and adjacent insula) for reward and punishment reversal learning when comparing correct reversal trials with correct acquisition trials, whereas we found overlapping activations in the right dorsolateral prefrontal cortex (DLPFC) when negative feedback signaled contingency change. The right lateral OFC and DLPFC also showed greater sensitivity to punishment than did their left homologues, indicating an asymmetry in how punishment is processed. We propose that the right lateral OFC and anterior insula are important for transforming affective feedback to behavioral adjustment, whereas the right DLPFC is involved in higher level attention control. These results provide insight into the neural mechanisms of reversal learning and behavioral flexibility, which can be leveraged to understand risky behaviors among vulnerable populations.     

Modification of a stimulus-reinforcer interaction by blocking

With tone-light compound-stimulus training, light will gain control of responding under an appetitive schedule while tone will gain considerable control with avoidance training. The present experiment sought to determine whether this stimulus-reinforcer interaction could be modified by a “blocking” procedure. Initially, one group of rats was trained to respond for food in tone while tone-off was food free, and a second group was trained to avoid shock in light while light-out was shock free. Following attainment of a 10:1 discrimination ratio, food and avoidance schedules were signaled, for the respective groups, by a tone-light compound stimulus. An element test demonstrated that the blocking procedure overcame the stimulus-reinforcer interaction previously reported with rats in both the appetitive and avoidance groups. Contrary to previous stimulus-reinforcer interaction research, here, the tone, rather than the light, controlled responding after food training while the light, rather than the tone, controlled responding after avoidance training. These results obey predictions from associative theory. The current experiment is also the first demonstration of blocking within an instrumental avoidance procedure.     

Stimulus specificity in the acquisition and extinction of conditioned taste aversion

An experiment evaluated whether the acquisition and extinction of conditioned taste aversion in the rat is stimulus-specific by testing the degree of response transfer between sweet and salty tastes. Animals in the paired-same and paired-different groups received a presentation of a gustatory CS and a cyclophosphamide injection US. Nonconditioned control groups received unpaired CS /US presentations or the CS followed by a vehicle injection. Taste avoidance was evaluated in three nonreinforced test sessions. In the paired-same, unpaired and vehicle groups, all test sessions were conducted with the same flavor as originally used in training, whereas the paired-different group was tested with a novel flavor on the first and second sessions and with the originally trained flavor in last session. Stimulus specific acquisition was apparent in the first test session, when the animals in the group paired-same exhibited lower fluid intake than the other three groups. Evidence of specificity of extinction was apparent in the last test session, when animals in the group paired-different exhibited lower fluid intake than the other three groups. These results provide further evidence of stimulus specificity in acquisition and extinction of conditioned taste aversion, supporting the associative interpretation of these phenomena.     

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)