Aspects of the reinforcer learned in second-order Pavlovian conditioning

Four experiments used an autoshaping procedure in pigeons to explore learning about the reinforcer in a second-order conditioning paradigm. Experiment 1 conditioned two visual second-order stimuli (S2), using as reinforcers two visual first-order stimuli (S1), each of which had previously been paired with food. Animals for which the S2 stimuli were each consistently paired with one particular S1 developed second-order responding more rapidly than did animals for which the identity of S1 varied from trial to trial. Moreover, following consistent pairings, extinction of an S1 had a depressive effect upon second-order responding which was peculiar to the S2 with which it had been paired. Both results suggest that in this preparation the organism identifies a particular S1 as the reinforcer for each S2. The remaining experiments examined the details of that identification. A compound S1, itself composed of two separable elements, was used to reinforce an S2. Subsequent extinction of either element of S1 led to a depression in the responding to S2, which indicates that both elements were involved in the second-order conditioning. Moreover, the use of several complex discriminations, which produced different behavior to S1 and to its elements, suggested that the organism had associated the S2 with the compound S1 rather than with its separate elements. However, even complete extinction of the response to S1 left some residual behavior to S2, which indicates that a portion of the second-order conditioning is independent of the current state of the reinforcer. These results demonstrate that in some situations the organism associates a conditioned stimulus with a rich representation of the reinforcer.     

Differential effects of two ways of devaluing the unconditioned stimulus after Pavlovian appetitive conditioning

Three experiments with rat subjects investigated the effects of two methods of devaluing a food unconditioned stimulus (US) after pairings of an auditory conditioned stimulus (CS) with that US. Experiment 1 found no effect of postconditioning pairings of the food US with lithium chloride (LiCl) on general activity to a tone CS, even though those pairings substantially reduced food consumption. Experiments 2 and 3 compared the effects on conditioned responding of postconditioning pairings of food with LiCl and with high-speed rotation. In these experiments the general activity measure was supplemented by a detailed visual analysis of the rats' behavior. Experiment 2 found that food-rotation pairings had larger effects than food-LiCl pairings on general activity responding and on two detailed behavioral measures but that food-LiCl pairings had larger effects on food consumption and on one behavioral measure. Experiment 3 replicated the findings of Experiment 2 and found that the ability of the CS to serve as a reinforcer for second-order conditioning after US devaluation was reduced more by food-LiCl pairings.     

Posttraining glucocorticoid receptor agonist enhances memory in appetitive and aversive Pavlovian discrete-cue conditioning paradigms

Glucocorticoid modulation of emotional memory has repeatedly been shown in aversive learning paradigms, but has received little attention in appetitive tasks. It has also been suggested that it may be selective for contextual cues. In order to investigate if glucocorticoids can modulate memory in discrete-cue conditioning of both appetitive and aversive tasks, two experiments were carried out. Male Lister-Hooded rats received pairings of an auditory cue and either food-reward (experiment 1) or footshock (experiment 2), followed immediately by posttraining injections of the glucocorticoid receptor agonist dexamethasone (0.3, 0.6, and 1.2 mg/kg) or vehicle. Dexamethasone (1.2 mg/kg) led to significantly enhanced learning. These results give support to the notion that glucocorticoids play a role in the modulation of both appetitive and aversive emotional memories and show that their role in learning goes beyond the construction of context representations. The modulation of appetitive and aversive discrete-cue learning may be subserved by a common mechanism.     

Analysis of us-preexposure effects in appetitive conditioning

In two experiments, rats received preexposure to one type of food followed by autoshaping in which presentation of one lever was associated with the preexposed food, and presentation of another lever with a novel type of food pellet. In both it was found that acquisition of the leverpress response occurred more readily on the lever associated with the novel food. This example of the US (unconditioned stimulus) preexposure effect is not to be explained in terms of the development of competing responses during preexposure. Explanations in terms of blocking by contextual cues and of habituation to the US are considered.     

Post-training amphetamine administration enhances memory consolidation in appetitive Pavlovian conditioning: Implications for drug addiction

It has been suggested that some of the addictive potential of psychostimulant drugs of abuse such as amphetamine may result from their ability to enhance memory for drug-related experiences through actions on memory consolidation. This experiment examined whether amphetamine can specifically enhance consolidation of memory for a Pavlovian association between a neutral conditioned stimulus (CS-a light) and a rewarding unconditioned stimulus (US-food), as Pavlovian conditioning of this sort plays a major role in drug addiction. Male Long-Evans rats were given six training sessions consisting of 8 CS presentations followed by delivery of the food into a recessed food cup. After the 1st, 3rd, and 5th session, rats received subcutaneous injections of amphetamine (1.0 or 2.0 mg/kg) or saline vehicle immediately following training. Conditioned responding was assessed using the percentage of time rats spent in the food cup during the CS relative to a pre-CS baseline period. Both amphetamine-treated groups showed significantly more selective conditioned responding than saline controls. In a control experiment, there were no differences among groups given saline, 1.0 or 2.0 mg/kg amphetamine 2 h post-training, suggesting that immediate post-training amphetamine enhanced performance specifically through actions on memory consolidation rather than through non-mnemonic processes. This procedure modeled Pavlovian learning involved in drug addiction, in which the emotional valence of a drug reward is transferred to neutral drug-predictive stimuli such as drug paraphernalia. These data suggest that amphetamine may contribute to its addictive potential through actions specifically on memory consolidation.     

A comparison of latent inhibition and learned irrelevance pre-exposure effects in rabbit and human eyeblink conditioning

The learning of an association between a CS and a US can be retarded by unreinforced presentations of the CS alone (termed latent inhibition or LI) or by un-correlated presentations of the CS and US (termed learned irrelevance or LIRR). In rabbit eyeblink conditioning, there have been some recent failures to replicate LI. LIRR has been hypothesized as producing a stronger retardation effect than LI based on both empirical studies and computational models. In the work presented here, we examined the relative strength of LI and LIRR in eyeblink conditioning in rabbits and humans. In both species, a number of pre-exposure trials sufficient to produce LIRR failed to produce LI (Experiments 1 & 3). Doubling the number of CS pre-exposures did produce LI in rabbits (Experiment 2), but not in humans (Experiment 4). LI was demonstrated in humans only after manipulations including an increased inter-trial interval or ITI (Experiment 5). Overall, it appears that LIRR is a more easily producible pre-exposure retardation effect than LI for eyeblink conditioning in both rabbits and humans. Several theoretical mechanisms for LI including the conditioned attention theory, stimulus compression, novelty, and the switching theory are discussed as possible explanations for the differences between LIRR and LI. Overall, future work involving testing the neural substrates of pre-exposure effects may benefit from the use of LIRR rather than LI.     

Recent learned helplessness/irrelevance research in Japan: Conceptual framework and some experiments on learned irrelevance

In Section 1, a conceptual framework within which to study learned helplessness (LH), learned irrelevance (LI) and the related phenomena is introduced. In Section 2, three rat experiments on LI conducted in our laboratory using a conditioned suppression of licking preparation are introduced. In Experiment 1, the phenomena of LI and general LI were confirmed. In Experiment 2, random presentations of CSt/US were found to interfere with subsequent initial excitatory conditioning to CSl under random CSl/US presentations. In Experiment 3, pre-paired presentations of CSn or CSt with US before random CSt/US presentations was found to have an attenuating effect on the development of general LI.     

Theoretical mechanisms underlying the trial-spacing effect in Pavlovian fear conditioning.

Contemporary explanations of the trial-spacing effect (TSE) were evaluated. Experiment 1 revealed, in rats given tone-footshock trials with 15-, 60-, or 900-s intertrial intervals (ITIs), a direct relationship between freezing to the tone and ITI (the TSE) but an inverted U-shaped relationship between freezing to the training context and ITI. In Experiment 2, footshock preexposure eliminated the TSE that otherwise occurs across 15- to 60-s ITIs but had no effect on the TSE that occurs across 60- to 900-s ITIs. In Experiments 3 and 4, neither (a) increasing posttraining exposure to the training context in rats trained with 60-s ITIs nor (b) reducing between-trial exposure to this context in rats trained with 900-s ITIs influenced freezing to the tone. These findings suggest that the TSE obtained in this research is due to more than 1 mechanism: 1 responsible for the TSE that occurs with ITIs less than approximately 60 s and another responsible for the TSE that occurs with ITIs greater than this. Although the perceptual-defensive-recuperative model may correctly describe the former mechanism, none of the theories tested seems to correctly describe the latter.     

The effect of contingency upon the appetitive conditioning of free-operant behavior

The problem of maintaining independence between response rates and reinforcement probabilities when determining the effect of varying the response-reinforcement contingency upon free-operant behavior was solved by programming local reinforcement probabilities for response and no response on a second-by-second basis. Fifty-seven rats were trained to lever-press on schedules of water reinforcement involving different values of contingency. All rats were first trained on a high positive contingency and then shifted to less positive, zero, or negative contingencies. Under these conditions, rate of lever-pressing declined appropriately when the contingency between response and reinforcement decreased or was made negative. The decline in rate produced by a zero contingency cannot be attributed to extinction, since the probability of reinforcement given the occurrence of a response was the same as for the positive contingency from which the shift to zero was made. That is, there was no change in the opportunity for response-reinforcement contiguity. It was concluded that the technique of programming local reinforcement probabilities offers promise for more critical examinations of the effects of contingency upon free-operant behavior.     

Pavlovian compound conditioning in the rabbit

Two experiments are reported on eyelid conditioning in the rabbit involving compounds of isolable CSs. In Experiment 1 it was demonstrated that subjects could be trained to respond discriminatively on the basis of specific configurations when no other reliable component cues were available. In Experiment II, a novel test procedure failed to provide further evidence that subjects utilized specific configurational cues when reliable isolable components were available. The findings were discussed in terms of the assumption that any stimulus compound involves both isolable and configurational components with the former being more “salient” than the latter.     

Unblocking in Pavlovian fear conditioning

Six experiments used rats to study blocking and unblocking of fear learning. An excitatory stimulus (A) blocked fear learning to a neutral stimulus (B). Unblocking of B occurred if the AB compound signaled an increase in unconditioned stimulus (US) intensity or number. Assessments of associative change during blocking showed that more was learned about B than A. Such assessments during unblocking revealed that more was learned about B than A following an increase in US intensity but not US number. These US manipulations had no differential effects on single-cue learning. The results show that variations in US intensity or number produce unblocking of fear learning, but for each there is a different profile of associative change and a potentially different mechanism.     

Contextual Pavlovian conditioning in the crab Chasmagnathus

In contextual conditioning, a complex pattern of information is processed to associate the characteristics of a particular place with incentive or aversive reinforcements. This type of learning has been widely studied in mammals, but studies of other taxa are scarce. The context-signal memory (CSM) paradigm of the crab Chasmagnathus has been extensively used as a model of learning and memory. Although initially interpreted as habituation, some characteristics of contextual conditioning have been described. However, no anticipatory response has been detected for animals exposed to the training context. Thus, CSM could be interpreted either as an associative habituation or as contextual conditioning that occurs without a context-evoked anticipatory response. Here, we describe a training protocol developed for contextual Pavlovian conditioning (CPC). For each training trial, the context (conditioned stimulus, CS) was discretely presented and finished together with the unconditioned stimulus (US). In agreement with the CSM paradigm, a robust freezing response was acquired during the 15 training trials, and clear retention was found when tested with the US presentation after short (2 and 4 h) and long (1–4 days) delays. This CPC memory showed forward but not simultaneous presentation conditioning and was context specific and protein synthesis dependent. Additionally, a weak CPC memory was enhanced during consolidation. One day after training, CPC was extinguished by repeated CS presentation, while one presentation induced a memory labilisation–reconsolidation process. Finally, we found an anticipatory conditioned response (CR) during the CS presentation for both short-term (4 h) and long-term memory (24 h). These findings support the conditioning nature of the new paradigm.     

Contemporary study of Pavlovian conditioning

Pavlov's first report on conditioning emphasized its role in allowing the animal to adjust to its environment. Contemporary theories have seen this adjustment in terms of developing accurate knowledge of the environment. Three aspects of that thinking are explored: how the animal acquires initial knowledge, how it changes its knowledge when conditions of the world change, and how it makes use of multiple knowledge representations.     

Pavlovian conditioning: the CS-UR relation

A new Pavlovian procedure used fluid-elicited throat-movement responses of the pigeon (N=66) to study the effects on conditioning of the temporal relation of the conditioned stimulus (CS) to the unconditioned stimulus-unconditioned response (US-UR). Because the throat-movement response has a substantial latency and duration, the relation of the CS to the US and UR could be independently evaluated. Four experiments indicated that, operationally, the relation of the CS to the UR--not to the US--is critical for conditioning in this preparation. The conventional emphasis on CS-US relations is based on procedures that confound the occurrence of the US with the UR and that foster generalization decrement between training and testing. The authors indicate how several conditioning phenomena may be reinterpreted in terms of CS-UR relations.     

Backward conditioning in Pavlovian type studies

Authors of almost all learning texts have indicated that unconditional/ stimulus-conditional stimulus (UCS-CS) trials (backward conditioning) utilized in classical conditioning experiments will not result in establishing a conditional response (CR) (excitation). Recently, it has been proposed that backward conditioning can take place although whether UCS-CS trials result in excitation or inhibition has been left unspecified. It is proposed that the diversity of findings can be attributed to inadequate methodology and the use of a classical-instrumental transfer paradigm. When only traditional Pavlovian conditioning studies are examined, the experimental findings suggest that UCS-CS trials will result in inhibition.     

One-trial appetitive conditioning in the sexual behavior system

One-trial Pavlovian conditioning has been demonstrated with aversive reinforcers such as electric shock and lithium chloride. The present experiments using male Japanese quail were conducted to determine if an appetitive unconditioned stimulus (US), the opportunity to copulate with a female quail, can also generate one-trial learning. In Experiment 1, subjects were exposed to a contextual conditioned stimulus (CS) for 35 sec and then given copulatory opportunity. In Experiment 2, male birds were given 0, 2, or 4 min of exposure to the contextual CS before presentation of a female. In both cases, control groups received comparable exposure to both the contextual CS and the sexual US, but in an unpaired fashion. Subjects were tested for sexual learning by reintroducing them to the experimental context in the presence of a test object that incorporated the taxidermically prepared head of a female quail. In both Experiments 1 and 2, subjects for whom the contextual CS and sexual US had been paired displayed significantly more approach to the test object than did unpaired subjects.     

Asymmetrical effects of Pavlovian excitatory and inhibitory aversive transfer on Pavlovian appetitive responding and acquisition

Predictions of a theory of Pavlovian motivational transfer, which incorporates principles of both the theory of reciprocal inhibition and the Rescorla-Wagner model, were tested in several Pavlovian aversive to Pavlovian appetitive transfer tasks. As predicted, the presence of a signal for an aversive event, conditioned stimulus (AV CS+), reliably suppressed performance of appetitive conditioned responses (CRs) whether imposed during acquisition or on independently established responding. Acquisition of appetitive responding to a novel CS reinforced in compound with an AV CS+, however, was enhanced (“superconditioning”). This observation suggests that the effects of a discrepancy between expectation and actual outcome on a conditioning trial are influenced by the affective value of both the expectation and the reinforcer. These transfer effects were not symmetrical for an inhibitory aversive stimulus (AV CS?). An AV CS? did not enhance appetitive responding compared to a random control condition, nor did the AV CS? reduce (i.e., block) appetitive conditioning to a novel CS when appetitive reinforcement occurred in the presence of the AV CS?. Comparison of the two shock-exposed conditions with a naive control condition suggests that previous results that were apparently consistent with inhibitory aversive enhancement and blocking of appetitive conditioning may have been due to aversive context conditioning.     

Second-order conditioning of Pavlovian conditioned inhibition

Two experiments are reported which investigated the conditioning of inhibition to a neutral stimulus as a result of its repeated pairing with a previouslyconditioned inhibitor. Both experiments employed a conditioned suppression technique with rat subjects. Experiment 1 detected the second-order inhibition through the retarded acquisition of concurrently administered excitatory conditioning. Experiment 2 found similar retardation in the acquisition of excitatory fear conditioning following previous second-order conditioning of inhibition to the stimulus. Implications are discussed for theories of the nature of inhibition and for second-order conditioning as an assessment technique.