Cardiac pacing and the law of initial value in rhesus monkeys

A cardiac pacemaker was used to produce a wide range of pre-CS heart rates in Pavlovian conditioning, and correlations were computed between pre-CS and CS heart rates. Two rhesus monkeys were presented trials in which 30 sec of light (CS) was followed by a brief electric shock (UCS). Pacing rates during the inter-trial intervals were varied from 150 to 350 bpm. Pacing was discontinued during CS. Unpaced control sessions showed that the maximum heart rate in CS on each trial was proportional to the pre-CS heart rate. Following pacing, however, heart rate in CS did not depend on the pacing rate; rather, the rates were not different from those of the unpaced control sessions. Pacing at high rates shifted the maximum heart rate toward the end of CS.     

Punishment of shock-induced aggression

Squirrel monkeys were restrained in a chair equipped with a tail-shock apparatus and a pneumatic bite hose located in front of the subject's face. An aggressive response was recorded when the monkey bit the hose. Initial sessions in which no shocks were delivered produced some biting. When biting during these sessions stabilized at a near-zero level, regularly scheduled shocks were delivered to the monkey's tail, causing a consistently higher rate of biting. After several sessions under these conditions, a punishment phase was introduced in which the previous shock conditions were maintained, and every bite was followed immediately by another, more intense shock. Biting under these conditions was suppressed to a near-zero level. When the punishment contingency was removed, biting increased. With one subject, two additional bite-contingent stimuli were examined: (1) a milder shock that, when made contingent upon hose biting, also suppressed that response, and (2) a contingent tone that had no obvious suppressing or facilitating effect. Individual differences among subjects were extreme, but the effect of bite-contingent shock was consistent. Observations of the subjects during the punishment sessions indicated the existence of certain side effects that resulted from the use of punishment to suppress shock-induced aggression.     

Classical conditioning and extinction of human systolic blood pressure

Fifty-five male and female undergraduates underwent 3 sessions of conditioning and one of extinction. Shock (UCS) occurred intermittently during light-on (CS+) and never in light-off (CS-). Results showed that (1) pressure in CS+ was significantly higher than CS-throughout all sessions; (2) within session adaptation was significantly retarded while incoming pressure levels did not adapt across sessions; (3) conditioned effects persisted 6 months after the last shock session; (4) verbal information as to non-appearance of shock significantly dropped pressure levels and reduced the discriminative light effect; (5) the post-shock unconditioned pressure response was a peak rise at 52 seconds, then a decrease which stabilized to the pre-shock level at 2 minutes. Pavlovian control of blood pressure was demonstrated and the present methodology offers an explicit procedure for further laboratory exploration.     

Constraint, Alcoholism, and Electrodermal Response in Aversive Classical Conditioning and Mismatch Novelty Paradigms

This study investigated whether low levels of the personality trait of constraint and early-onset alcoholism would be associated with deficits in aversive conditioning and smaller responses to novelty in a stimulus mismatch protocol. Personality traits (constraint and socialization) and skin conductance responses (SCRs) during conditioning and novelty paradigms were assessed in alcoholics (n=41) and non-alcoholics (n = 32). The conditioning protocol involved measuring SCRs after conditioned stimuli (CS+: tones) paired with shock, CS- tones unpaired with shock, and CS+ probes unpaired with shock. The mismatch protocol involved measuring SCRs to auditory stimuli consisting of a series of 5 pure tones of the same pitch followed a shorter white noise stimulus (the novel stimulus). Contrary to the hypothesis, alcoholics did not differ from non-alcoholics in SCRs to CS+ probes or on the mismatch measure (SCR novel tone-SCR to 5th tone). Higher levels of constraint and self-reports of fear during conditioning were associated with smaller responses to both the CS+ probes and the CS- tones as well as the mismatch measure within non-alcoholics, but not within alcoholics. In alcoholics, low constraint was associated with greater habituation to CS+ probes, and poor differential conditioning on measures of change across trials in SCR to CS+ probes and CS- stimuli. The results suggest that different processes influence levels of constraint in non-alcoholics and alcoholics. The data indicate that low constraint in non-alcoholics is associated with allocating fewer processing resources to potentially significant stimuli, rather than being associated with a specific deficit in aversive conditioning per se.     

SOME RELATIONS BETWEEN CLASSICALLY CONDITIONED AGGRESSION AND CONDITIONED SUPPRESSION IN SQUIRREL MONKEYS

During three experiments with squirrel monkeys, stimulus and shock pairings were given in the presence of a bite tube. Experiments 1 and 2 used a conditioned-suppression procedure in which bar pressing was reinforced with food. A transparent shield prevented biting of the bar. When the stimulus was paired with shock, bar pressing decreased (conditioned suppression) and tube biting increased during the stimulus (classically conditioned aggression). When the bite tube was removed on alternate sessions in Experiment 2, there was more suppression when the tube was present, thus suggesting that biting competed with bar pressing. However, this simple competing-response interpretation was complicated by the findings of Experiment 3 where, with naive monkeys, bar pressing was never reinforced with food, yet bar pressing was induced during the stimulus and was highest when the bite tube was absent. The fact that stimulus-induced bar pressing developed indicates that bar pressing in conditioned-suppression procedures, suppressed or not, may be maintained by two types of control—the food reinforcer and induced CS control. The higher rate of induced bar pressing during the stimulus with the bite tube absent confounds a simple competing response interpretation of conditioned suppression. It suggests that shock-induced responses during conditioned suppression could be both contributing to and competing with responding maintained by food, with the net effect depending on specific but ill-defined features of the situation.     

Pavlovian conditioning of shock-elicited aggression: a discrimination procedure

Two auditory stimuli, separated by a fixed intertrial interval, were alternately presented to two rats in a closed environment. The positive conditioned stimulus (CS+) terminated with the offset of a 2-mA, 0.75-sec shock. The negative conditioned stimulus (CS-) terminated without shock. The incidence of the "stereotyped fighting posture" was recorded during the CS+, the CS-, the intertrial interval, and shock. The results showed an increase in the percentage of conditioned responses during the CS+, and a decrease during both the CS- and the intertrial interval, when the duration of the conditioned stimuli and the intertrial interval was 16 sec. Appropriate changes in the incidence of aggression during the two stimuli were obtained following the reversal of the stimulus functions. During the acquisition and reversal phases there was a between-session decrement and a within-session improvement in the incidence of aggression during the CS+, defined as warm-up. The presentation of free shocks before the conditioning sessions was effective in reducing the warm-up only when the interval between shocks was 64 sec. These data were interpreted as demonstrating classical conditioning of shock-elicited aggression, with little chance of non-associative factors contributing to the measurement of the conditioned response.     

Constraint, alcoholism, and electrodermal response in aversive classical conditioning and mismatch novelty paradigms

This study investigated whether low levels of the personality trait of constraint and early-onset alcoholism would be associated with deficits in aversive conditioning and smaller responses to novelty in a stimulus mismatch protocol. Personality traits (constraint and socialization) and skin conductance responses (SCRs) during conditioning and novelty paradigms were assessed in alcoholics (n=41) and non-alcoholics (n=32). The conditioning protocol involved measuring SCRs after conditioned stimuli (CS+: tones) paired with shock, CS− tones unpaired with shock, and CS+ probes unpaired with shock. The mismatch protocol involved measuring SCRs to auditory stimuli consisting of a series of 5 pure tones of the same pitch followed a shorter white noise stimulus (the novel stimulus). Contrary to the hypothesis, alcoholics did not differ from non-alcoholics in SCRs to CS+ probes or on the mismatch measure (SCR novel tone—SCR to 5^th tone). Higher levels of constraint and self-reports of fear during conditioning were associated with smaller responses to both the CS+ probes and the CS− tones as well as the mismatch measure within non-alcoholics, but not within alcoholics. In alcoholics, low constraint was associated with greater habituation to CS+ probes, and poor differential conditioning on measures of change across trials in SCR to CS+ probes and CS− stimuli. The results suggest that different processes influence levels of constraint in non-alcoholics and alcoholics. The data indicate that low constraint in non-alcoholics is associated with allocating fewer processing resources to potentially significant stimuli, rather than being associated with a specific deficit in aversive conditioning per se.     

Relations between behavioral arousal and plasma cortisol levels in monkeys performing repeated free-operant avoidance sessions.

The commonly believed hypothesis that increases in adrenocorticoid levels sensitively reflect behavioral arousal was tested by subjecting monkeys to repeated sessions of free-operant avoidance; these sessions produced varying degrees of behavioral arousal over time, which were quantified by a behavioral scoring technique. Cortisol was reliably elevated only in the most aroused subjects early in the first avoidance session. Although subjects were still aroused later in the session, cortisol had returned to basal levels. During subsequent avoidance sessions, cortisol returned to basal levels and did not increase significantly even when arousal was further manipulated by the superimposition of unavoidable shock during the final avoidance session. The addition of unavoidable shock was associated with a significant correlation between arousal, as reflected by increases in response rate, and magnitude of change, usually decreases, in mean cortisol levels. The frequent occurrence of dissociations between cortisol levels and behavioral arousal, as reflected by behavioral score, operant rate, and shock frequency, indicates that cortisol levels are of little use as a neuroendocrine index of arousal.     

Stock optimizing: maximizing reinforcers per session on a variable-interval schedule.

In Experiment 1, 2 monkeys earned their daily food ration by pressing a key that delivered food according to a variable-interval 3-min schedule. In Phases 1 and 4, sessions ended after 3 hr. In Phases 2 and 3, sessions ended after a fixed number of responses that reduced food intake and body weights from levels during Phases 1 and 4. Monkeys responded at higher rates and emitted more responses per food delivery when the food earned in a session was reduced. In Experiment 2, monkeys earned their daily food ration by depositing tokens into the response panel. Deposits delivered food according to a variable-interval 3-min schedule. When the token supply was unlimited (Phases 1, 3, and 5), sessions ended after 3 hr. In Phases 2 and 4, sessions ended after 150 tokens were deposited, resulting in a decrease in food intake and body weight. Both monkeys responded at lower rates and emitted fewer responses per food delivery when the food earned in a session was reduced. Experiment 1's results are consistent with a strength account, according to which the phases that reduced body weights increased food's value and therefore increased subjects' response rates. The results of Experiment 2 are consistent with an optimizing strategy, because lowering response rates when food is restricted defends body weight on variable-interval schedules. These contrasting results may be attributed to the discriminability of the contingency between response number and the end of a session being greater in Experiment 2 than in Experiment 1. In consequence, subjects lowered their response rates in order to increase the number of reinforcers per session (stock optimizing).     

Allocation of cognitive processing capacity during human autonomic classical conditioning

In each of two experiments, allocation of cognitive processing capacity was measured in college-student subjects during autonomic discrimination classical conditioning. A 7.0-sec delay paradigm was used to establish classically conditioned responses to a reinforced visual conditioned stimulus (CS+). Electrodermal responses were the primary measures of autonomic classical conditioning. Allocation of processing capacity was measured by monitoring performance on a secondary reaction-time (RT) task. The auditory secondary-task RT signal was presented before, and 300, 500, 3500, 6500, and 7500 msec following CS onset. The RT signal was also presented following properly and improperly cued shock unconditioned stimuli (UCSs). Significant discrimination classical conditioning was obtained in both experiments. Comparison with control subjects who did not receive the RT signals indicated that the presence of the RT signals did not interfere with the development of classical conditioning. Four principal findings were obtained with the secondary-task RT measure. First, RTs to signals presented during CS+ were consistently slower than RTs to signals presented during CS-. This finding indicates that greater capacity allocation occurred during CS+ than CS- and is consistent with recent cognitive interpretations of classical conditioning. Second, the largest capacity allocation (i.e., slowing of RTs) occurred 300 msec following CS+ onset. This finding is consistent with the notion that subjects are actively processing the signal properties of the CS+ at 300 msec following CS+ onset. Third, presentation of the UCS when improperly cued (following CS-) significantly increased capacity allocation, whereas presentation of the same UCS when properly cued (following CS+) did not affect capacity allocation. These findings indicate that subjects were actively prepared for the UCS following CS+ but not following CS- and that a surprising UCS elicits greater capacity allocation than does an expected UCS. Fourth, large electrodermal responders to the CSs exhibited patterns of capacity allocation during the CSs, particularly during the CS+, different from those of small electrodermal responders. In particular, they exhibited significantly longer RTs at 300 msec after CS+ onset than did the small responders, followed by a shortening of RT at 500 msec relative to the small responders. This finding suggests that large electrodermal responders devote greater processing capacity to significant environmental stimuli than do small responders and that their processing may begin and be completed more rapidly. All in all, the data indicate the complexity of the cognitive processes that occur during human classical conditioning and the usefulness of the secondary-task technique in integrating conditioning theories and psychophysiology with cognitive psychology.     

Hippocampal associative cellular responses: dissociation with behavioral responses revealed by a transfer-of-control technique

Multiunit activity was recorded in the CA3 field of the dorsal hippocampus in freely moving rats during classical conditioning and subsequent presentation of the CS on operant baselines for food reward as well as shock avoidance. Rats were first trained in a nonsignaled bar-pressing-dependent shock omission task and in a food-motivated lever-pressing task (60-s VI). Five sessions with presentations of a previously habituated tone as a CS paired with footshock as a US were then given. Testing was carried out by presenting the CS alone while behavioral responses were maintained by reinforcement in both instrumental tasks on alternate sessions. As expected, the CS induced a marked suppression of lever pressing for food reward and a marked enhancement of bar-pressing for shock avoidance. The analysis of the frequency of multiunit discharges to the CS revealed that the hippocampal cellular responses established during classical conditioning were maintained while two different behavioral responses were exhibited to the CS. The results showed that the associative response of hippocampal neurons may be dissociated from the Pavlovian conditioned responses the CS elicits. They support the hypothesis that hippocampal cellular responses represent a neural index of the acquired CS-US associative representation.     

Neuronal nitric oxide synthase gene inactivation reduces the expression of vasopressin in the hypothalamic paraventricular nucleus and of catecholamine biosynthetic enzymes in the adrenal gland of the mouse

The impact of a lifelong absence of the neuronal nitric oxide synthase (nNOS) in the neuroendocrine stress response was investigated in nNOS knockout (KO) and wild type (WT) mice under basal conditions and in response to forced swimming. In the hypothalamic paraventricular nucleus oxytocin and corticotropin-releasing-hormone mRNA levels did not differ between these genotypes under resting conditions, whereas vasopressin mRNA levels were significantly lower in nNOS KO than in WT animals. Also, in the adrenal glands basal levels of tyrosine hydroxylase protein, the rate-limiting enzyme for catecholamine biosynthesis, and of phenylethanolamine N-methyltransferase, which converts norepinephrine to epinephrine, were significantly reduced in nNOS KO mice. Plasma adrenocorticotropin, corticosterone, norepinephrine and epinephrine levels were similar in the KO and WT genotypes under resting conditions. In response to forced swimming, a similar increase in plasma adrenocorticotropin and corticosterone was observed in KO and WT animals. Stressor exposure triggered also an increased epinephrine release in WT animals, but did not significantly alter plasma epinephrine levels in KO mice. These data suggest that the chronic absence of nNOS reduces the capacity of epinephrine synthesising enzymes in the adrenal gland to respond to acute stressor exposure with an adequate epinephrine release.     

Fear develops to the conditioned stimulus and to the context during classical eyeblink conditioning in rats

In classical eyeblink conditioning, non-specific emotional responses to the aversive shock unconditioned stimulus (US), which are presumed to coincide with the development of fear, occur early in conditioning and precede the emergence of eyeblink responses. This twoprocess learning model was examined by concurrently measuring fear and eyeblink conditioning in the freely moving rat. Freezing served as an index of fear in animals and was measured during the inter-trial intervals in the training context and during a tone conditioned stimulus (CS) presented in a novel context. Animals that received CS-US pairings exhibited elevated levels of fear to the context and CS early in training that decreased over sessions, while eyeblink conditioned responses (CRs) developed gradually during acquisition and decreased during extinction. Random CS-US presentations produced a similar pattern of fear responses to the context and CS as paired presentations despite low eyeblink CR percentages, indicating that fear responding was decreased independent of high levels of learned eyeblink responding The results of paired training were consistent with two-process models of conditioning that postulate that early emotional responding facilitates subsequent motor learning, but measures from random control animals demonstrate that partial CS-US contingencies produce decrements in fear despite low levels of eyeblink CRs. These findings suggest, a relationship between CS-US contingency and fear levels during eyeblink conditioning, and may serve to clarify further the role that fear conditioning plays in this simple paradigm.     

Extensive extinction in multiple contexts eliminates the renewal of conditioned fear in rats

Two studies examined whether nonreinforcement of a stimulus in multiple contexts, instead of a single context, would decrease renewal of conditioned fear in rats (as assessed by conditioned suppression of lever pressing). In Experiment 1, renewal was measured after 36 nonreinforced CS trials delivered during six extinction sessions in a single context or two extinction sessions in each of three different contexts. The number of extinction contexts did not have an effect on renewal. In Experiment 2, groups received either 36 or 144 nonreinforced CS trials during six or twenty-four extinction sessions in a single context or three different contexts. Again, renewal was not influenced by the number of extinction contexts when only 36 trials were given. However, when 144 trials were used, renewal was completely eliminated when extinction was divided between 3 contexts, but was not weakened when the sessions took place in a single context. The results suggest that the use of multiple treatment settings in exposure-based therapies is only likely to reduce relapse if a sufficient number of sessions are provided in each of the treatment settings.     

Fear Develops to the Conditioned Stimulus and to the Context during Classical Eyeblink Conditioning in Rats

In classical eyeblink conditioning, non-specific emotional responses to the aversive shock unconditioned stimulus (US), which are presumed to coincide with the development of fear, occur early in conditioning and precede the emergence of eyeblink responses. This two-process learning model was examined by concurrently measuring fear and eyeblink conditioning in the freely moving rat. Freezing served as an index of fear in animals and was measured during the inter-trial intervals in the training context and during a tone conditioned stimulus (CS) presented in a novel context. Animals that received CS-US pairings exhibited elevated levels of fear to the context and CS early in training that decreased over sessions, while eyeblink conditioned responses (CRs) developed gradually during acquisition and decreased during extinction. Random CS-US presentations produced a similar pattern of fear responses to the context and CS as paired presentations despite low eyeblink CR percentages, indicating that fear responding was decreased independent of high levels of learned eyeblink responding. The results of paired training were consistent with two-process models of conditioning that postulate that early emotional responding facilitates subsequent motor learning, but measures from random control animals demonstrate that partial CS-US contingencies produce decrements in fear despite low levels of eyeblink CRs. These findings suggest a relationship between CS-US contingency and fear levels during eyeblink conditioning, and may serve to clarify further the role that fear conditioning plays in this simple paradigm.     

Effects of hemispheric asymmetry on electrodermal conditioning in a dichotic listening paradigm

It was predicted that when an auditive verbal conditional stimulus (CS+), previously associated with an aversive unconditional stimulus (UCS), is presented to theright ear (initial left hemisphere input), in a dichotic listening task, with the CS-simultaneously presented in the left ear, the result would be greater resistance to extinction than when the same CS+ is presented to the left ear (initial right hemisphere input). In the same vein, it was predicted that, when a tonal CS+ is presented to theleft ear, greater resistance to extinction would occur as compared with when it is presented to the right ear. During a dichotic test phase, each group was split into two subgroups, and the CS+ and CS—repeatedly were presented simultaneously to each ear. Half of the subjects in each subgroup had the CS+ presented to the left ear. The other half had the CS+ presented to the right ear. Skin conductance responses were recorded from both hands. Results showed significantly larger responding in all groups to the CS+/UCS compound during the acquisition phase. However, during the dichotic extinction phase, the verbal CS+ right-ear subgroup showed significantly larger resistance to extinction as compared with the verbal CS+ left-ear subgroup. No differences were found between the two tonal subgroups.     

An interresponse-time analysis of responding maintained by schedules of response-produced electric shock

The present study investigated ratio contingencies to evaluate factors that may determine the maintenance of responding when electric shock is the consequent event. Initially, squirrel monkeys (Saimiri sciureus) were exposed to a continuous-avoidance schedule to initiate bar pressing. Subsequently, a multiple random-interval variable-ratio yoked schedule of response-produced shock was used to maintain and to compare interval and ratio performance. A microcomputer recorded and stored the number of responses and interresponse times occurring between successive shock presentations during a given random-interval component, and these numbers determined the ratio requirements during the subsequent ratio component. Responding was maintained for more than 80 sessions in two of three monkeys under the multiple schedule with the ratio yoked to the interval component. Responding during the ratio component persisted in only one monkey, however, when the components were no longer yoked. An analysis of the interresponse times immediately preceding shock under the multiple yoked schedule revealed that the terminal interresponse times were longer under the interval schedule than under the ratio contingency. The interresponse-time analysis indicated that differential interresponse-time relationships may be major determinants of the maintenance of behavior controlled by schedules of electric-shock presentation.     

Varying temporal location of a conditioned stimulus in heart rate conditioning ofMacaca mulatta

The several functions that a stimulus can assume were investigated in a Pavlovian conditioning procedure. The subjects were six rhesus monkeys; the response under observation was heart rate. The conditioning began with a temporal separation of zero between a signal and a regularly repeating electric shock; the signal was then moved to a series of earlier locations in the inter-shock interval. After six sessions at each location, two sessions followed in which only the shock was delivered periodically. The findings included: (1) A two-phased conditioned cardiac rate response seen at the first location became more multiphasic and irregular during longer intervals between signal and shock; (2) the location where the conditioned response peaked became increasingly variable as the signal was moved back, but this variability maintained a constant proportion to the signal-shock interval; and, (3) heart rate during a presignal period, and during a comparable period in shock only sessions, was generally deceleratory early in training and acceleratory thereafter. Sessions with the signal showed heart rate in the presignal period to have become acceleratory earlier in training than sessions with shock only. The data pertain to stimulus control over heart rate as a function of: (A) the temporal proximity of a signal to an aversive stimulus; and, (B) the presence or absence of the signal. The use of appropriate response units in cardiac conditioning is also discussed.     

Effects of long-term shock and associated stimuli on aggressive and manual responses

Squirrel monkeys were exposed to response-independent, fixed-frequency shock that produced biting attack upon a pneumatic hose. Attacks decreased within and across sessions at low intensities and high frequencies of shock, but increased within and across sessions at higher intensities and lower shock frequencies. Stimuli paired with shock, when presented alone, came to produce biting, and stimuli correlated with shock parameters that produced increases in responding within sessions produced similar increases when presented alone. Further experiments showed that continuing exposure to shock also produced lever pressing or chain pulling, with longer shock exposure again producing higher response rates. Whereas biting generally decreased throughout the intershock interval, manual responding generally increased as shock time approached, but immediately before shock was often suppressed. Following shock, biting attack predominated over manual behavior. The results suggest a possible explanation for the extreme resistance of avoidance behavior to extinction, and may also partially explain the persistence of responding during schedules of response-produced shock. Relationships of the present findings to naturalistic observations of relations between fleeing, freezing, and fighting performances are discussed.     

The role of gross locomotor restriction in the acquisition of motor and cardiac conditional discrimination in the dog

The effects of gross locomotor restriction and lack of restriction, respectively, on the acquisition of a motor and cardiac discrimination task was investigated. Litter-mate dogs were used in a counterbalanced design involving two treatment conditions (free and restrained) and two discrimination learning problems. Each learning problem included eight daily training sessions with 10 CS+ tones and ten CS—tones presented randomly. The US was a shock to the left foreleg. Degree of foot flexion and percentage change in heart rate in response to CS tones were recorded. Dogs subjected initially to the free condition manifested a superior and more rapid motor and cardiac discrimination than Ss initially restrained. The free condition was not as facilitatory when presented to Ss which first had been exposed to the restraint condition. The results are discussed in terms of activation and negative transfer.