Conditioned inhibition training of the competing cue after compound conditioning does not reduce cue competition

In each of two experiments, we studied Pavlovian fear conditioning (as assessed by barpress conditioned suppression) in 32 albino rats. Following a two-stage cue-competition procedure (A+ then AX+), we subjected the competing cue (A) to conditioned inhibition training (B+, BA?) before testing the target cue (X). Conditioned inhibition training was designed to weaken the putative A-unconditioned stimulus (US) association, perhaps changing it to an A-no-US association. Performance-deficit theories of cue competition, such as comparator theory and retrieval-interference theory, predict that such procedures should weaken cue competition, causing Conditioned Stimulus X (CS X) to evoke strong responding. The same prediction can be deduced from recent acquisition-focused models (Dickinson &; Burke, 1996; Van Hamme &; Wasserman, 1994). In opposition to this prediction, however, we found in both experiments that conditioned inhibition training had no detectable effect on cue competition even though it successfully abolished conditioned responding to CS A. In Experiment 2, moreover, we found evidence against the hypothesis that the weak response to CS X was due to generalization decrement rather than to cue competition. Results favor early learning-deficit theories of cue competition over performance-deficit theories and over the recent acquisition-focused models.     

Protection from extinction provided by a conditioned inhibitor

Three conditioned suppression experiments with rats as subjects investigated the influence of higher order associations in determining the response potential of a target stimulus. In these experiments, a Pavlovian conditioned inhibitor was compounded with the target cue during extinction treatment. In Experiment 1, strong suppression was observed to the target cue that was given extinction treatment in the presence of a conditioned inhibitor, relative to a target that was extinguished with an associatively neutral cue or was extinguished alone, suggestive of enhanced protection from extinction provided by a conditioned inhibitor. This effect was replicated in a sensory preconditioning preparation in Experiment 2; in Experiment 3, in a sensory preconditioning preparation, this protection effect was retroactively attenuated when the conditioned excitor used to train the conditioned inhibitor was extinguished following extinction of the target. This provides evidence that, at least in a sensory preconditioning preparation, stimuli that are only indirectly associated with the target cue can contribute to the response potential of that target.     

Biological significance attenuates overshadowing, relative validity, and degraded contingency effects

Miller and Matute (1996) showed that blocking is attenuated when the blocked conditioned stimulus (CS) is “biologically significant” (i.e., when the CS has the potential to elicit vigorous responding of any kind). To the extent that blocking is representative of cue competition, this finding suggests that biological significance protects CSs against cue competition effects in general. In the present experiments, we tested this possibility by examining the influence of biological significance of CSs on other examples of cue competition, namely, overshadowing, the relative stimulus validity effect, and the degraded contingency effect in rats. In Experiment 1, we found that intense auditory stimuli induced transient unconditioned lick suppression, thereby indicating that intense sounds were of high inherent biological significance. In Experiment 2A, we found that cues with high inherent biological significance were protected from overshadowing. In Experiment 2B, this finding was extended to cues with high acquired biological significance, which was obtained through prior pairings with a reinforcer of the valence opposite to that used in the overshadowing treatment. In Experiments 3 and 4, we found that cues with high inherent biological significance attenuated the relative validity effect and the degraded contingency effect, respectively. These results lend support to the view that biological significance (inherent and acquired) protects stimuli from cue competition effects, a finding that is problematic for many contemporary theories of learning.     

A comparison of learned aversions to gustatory and exteroceptive cues in rats

The experiment provides a direct comparison of the ability of subjects (rats) to associate gustatory and exteroceptive stimuli with illness. Previous experiments which have made similar comparisons between gustatory and exteroceptive cues have suffered from certain methodological problems involving stimulus control and compounding. The present experiment utilized a between-subjects design wherein half of the subjects had an auditory cue associated with poisoning and half had a taste cue. In both cases, the other cue was present, but was not predictive of poisoning, The auditory cue, like the taste cue, occurred only during drinking. This comparison was made in both an immediate and a delayed poisoning situation. The experiment found that while subjects were able to quickly associate a taste cue with illness, they were unable to form a similar association between poisoning and the exteroceptive stimulus. Results also showed that subjects will fail to acquire a taste aversion to a novel and salient gustatory cue when that cue is followed by illness only 50% of the time. This latter effect was more pronounced in the delayed poisoning situation.     

Further evidence that conditioned inhibition is not the mechanism of an extinction cue’s effect: A reinforced cue prevents spontaneous recovery

Previous research has demonstrated that discrete cues presented during extinction of a conditioned response (extinction cues) subsequently reduce spontaneous recovery and the renewal effect. In order to assess whether an extinction cue’s effect is due to conditioned inhibition, the effect of pairing an extinction cue with food on the cue’s capacity to modulate spontaneous recovery was investigated in an appetitive conditioning experiment with rats. Conditioned inhibitors paired with the unconditioned stimulus lose their potential to inhibit responding. The food-paired extinction cue did not lose its potential to reduce spontaneous recovery. In fact, it reduced spontaneous recovery more than did an extinction cue that was not paired with food. The results indicate that extinction cues attenuate postextinction recovery of conditioned responding through a mechanism other than conditioned inhibition. The cue’s action appears to be similar to that of serial negative occasion setters. Theories of spontaneous recovery and the relationship of extinction cues to other modulatory stimuli are discussed.     

CS modality effects in one-trial backward and forward excitatory conditioning as assessed by conditioned suppression of licking in rats

Rats received a single 4-sec 1-mA grid-shock US either preceded or followed by a 4-sec tone or light CS. Conditioning was later assessed by comparing the amount of lick suppression evoked by the forward- or backward-paired CS versus an explicitly unpaired CS. The backward-paired CS produced more suppression than the unpaired CS only when both were tone; the light evoked strong suppression whether paired or not. In the forward procedure, tone produced more suppression when paired and less when unpaired than did light; conditioning thus appeared stronger with the tone. In one experiment, observations showed that rats froze during the forward-paired tone but not during the light. Increasing CS duration from 4 to 12 sec had no effect for the forward-paired light but increased freezing to the forward-paired tone. Another experiment showed similar unconditioned suppression to tone and light but faster habituation to tone. Problems that these results create for interpreting evidence for excitatory backward conditioning in the conditioned suppression procedure are discussed.     

Taste potentiates color-sickness associations in pigeons and quail

The present experiments assessed cue utilization in pigeons and quail on similar tests of poison-based aversion learning. In Experiments 1 and 2, three groups of pigeons were given colored water, flavored water, or colored flavored water prior to induction of sickness; these experiments differed only as to the specific colors and flavors used as stimuli. In both experiments, the birds trained with flavored water exhibited reliable taste aversions when tested with uncolored flavored water. Similar degrees of aversion were observed whether the flavored water had been colored or uncolored during training, suggesting that the color cue had little or no effect on the conditioning of the flavor cue. In contrast, the flavor cue had a pronounced effect on the conditioning of the color cue. When tested with unflavored colored water, the birds trained with colored flavored water exhibited significantly stronger color aversions than those trained with unflavored colored water. That is, the flavor cue enhanced or potentiated the conditioning of the color cue. In a third experiment, quail were trained in the same way as the pigeons with virtually the same result. The pattern of cue utilization observed in the present experiments with pigeons and quail differs markedly from that proposed by Wilcoxon, Dragoin, and Kral (1971) for quail. However, a reexamination of the results obtained by Wilcoxon et al. suggested that they are susceptible to an alternative interpretation consistent with the present results.     

Associative foundation of causal learning in rats

Are humans unique in their ability to interpret exogenous events as causes? We addressed this question by observing the behavior of rats for indications of causal learning. Within an operant motor–sensory preconditioning paradigm, associative surgical techniques revealed that rats attempted to control an outcome (i.e., a potential effect) by manipulating a potential exogenous cause (i.e., an intervention). Rats were able to generate an innocuous auditory stimulus. This stimulus was then paired with an aversive stimulus. The animals subsequently avoided potential generation of the predictive cue, but not if the aversive stimulus was subsequently devalued or the predictive cue was extinguished (Exp. 1). In Experiment 2, we demonstrated that the aversive stimulus we used was in fact aversive, that it was subject to devaluation, that the cue–aversive stimulus pairings did make the cue a conditioned stimulus, and that the cue was subject to extinction. In Experiments 3 and 4, we established that the decrease in leverpressing observed in Experiment 1 was goal-directed instrumental behavior rather than purely a product of Pavlovian conditioning. To the extent that interventions suggest causal reasoning, it appears that causal reasoning can be based on associations between contiguous exogenous events. Thus, contiguity appears capable of establishing causal relationships between exogenous events. Our results challenge the widely held view that causal learning is uniquely human, and suggest that causal learning is explicable in an associative framework.     

In a daily time–place learning task,time is only used as a discriminative stimulus if each daily session is associated with a distinct spatial location

It is difficult for rats to acquire daily time–place (TP) learning tasks. One theory suggests that rats do not use time of day as a stimulus signaling a specific response. In the present study, we tested rats’ ability to use time of day as a discriminative stimulus. A fixed-interval procedure was used in which one lever provided reinforcement on a FI-5-s schedule in morning sessions, and the same lever provided reinforcement on a FI-30-s schedule in afternoon sessions. Because only one place was used in this paradigm, the rats could only use time of day to acquire the task. Mean responses during the first 5 s of the first trial in each session indicated that the rats did not discriminate between the two sessions. In Phase II, a different lever location was used for each of the two daily sessions, which meant that both spatial and temporal information could be used to acquire the task. The rats readily acquired the task in this phase, and probe trials indicated that the rats were using a combination of spatial and temporal information to discriminate between the two different trial types. When the spatial cue was removed in Phase III, rats no longer discriminated the two sessions, suggesting that time can only be used as a discriminative stimulus when each daily session is associated with a distinct spatial location.     

Interaction of retention interval with CS-preexposure and extinction treatments: Symmetry with respect to primacy

Imposition of a retention interval between cue-outcome pairings and testing can alleviate the retardation of conditioned responding induced by pretraining exposure to the cue (i.e., the CS-preexposure effect). However, recent studies have reported an enhanced effect of CS-preexposure treatment with longer retention intervals (De la Casa & Lubow, 2000, 2002; Lubow & De la Casa, 2002). In a series of conditioned barpress suppression studies with rats, we examined the effects of imposing a retention interval just prior to testing following either CS-preexposure (cue alone before cue-outcome pairings) or extinction (cue alone after cue-outcome pairings) treatments. Experiment 1 replicated in a different preparation recent reports of CS-preexposure treatment effects increasing with longer retention intervals. Experiment 2 showed that spontaneous recovery of stimulus control of behavior after extinction can be obtained with the same parameters as those used to observe the augmented effect of CS-preexposure treatment. In Experiment 3, both the augmented effect of CS-preexposure treatment and spontaneous recovery from extinction were found when we used, in place of a retention interval, an associative priming manipulation.     

Coherence of retrieval cues,rather than additivity,determines prior cuing effectiveness in the rat

In previous studies, we have shown that adding a supplementary cue in a prior-cuing treatment may dramatically reduce its initial effectiveness. This series of experiments was aimed at testing the hypothesis that such a lack of additivity among retrieval cues could be due to the combined reactivation of memory attributes related to incompatible behavioral responses. For that purpose, rats were partially trained to reach the lighted arm (light condition; usual training condition) or the darkened arm (dark condition) of a Y-maze in order to avoid an electric foot-shock. The first experiment demonstrates that animals learned the two versions of the task similarly and that an exposure to the light similarly acts as an effective retrieval cue after a 24-h interval. In the second experiment, the effectiveness of exposure to the grid alone and to the light alone as well as to the combined light + grid cue was investigated after a 21-day training-to-test interval in the light condition, where light was related to safety and grid was associated with the shock, and in the dark condition, where both light and grid were related to the shock. The results indicate that whereas the compound light + grid cue was an ineffective retrieval cue in the light condition, it acted as an effective retrieval cue in the dark condition, supporting the hypothesis that coherence between retrieval cues is a necessary condition for effective cuing.     

Partial reinforcement and latent inhibition effects on stimulus–outcome associations in flavor preference conditioning

Two experiments with thirsty rats explored the harmful effects of non-reinforced exposures to a flavor cue in the control by sensory-specific flavor–sucrose associations in a conditioned flavor preference paradigm. Experiment 1 demonstrated that rats learned to prefer a flavor cue that was consistently paired with sucrose over one that was paired with sucrose the same number of times but was also presented without sucrose on other occasions. However, rats for which sucrose was devalued following the conditioning phase preferred the partially reinforced flavor cue over the consistently reinforced flavor, suggesting that non-reinforcement weakened the ability of that flavor cue to evoke a specific representation of sucrose during the preference test. Experiment 2 demonstrated comparable effects of non-reinforcement in a latent inhibition procedure, although relatively more non-reinforced pre exposures to the flavor, in conjunction with fewer flavor–sucrose pairings, were required to see the effect. Together, the results suggest, as is often found with more traditional learning paradigms, that non-reinforcement of a flavor cue has deleterious effects on preference learning and/or performance.     

Performance factors in associative learning: Assessment of the sometimes competing retrieval model

Previous simulations revealed that the sometimes competing retrieval model (SOCR; Stout & Miller, Psychological Review, 114, 759-783, 2007), which assumes local error reduction, can explain many cue interaction phenomena that elude traditional associative theories based on total error reduction. Here, we applied SOCR to a new set of Pavlovian phenomena. Simulations used a single set of fixed parameters to simulate each basic effect (e.g., blocking) and, for specific experiments using different procedures, used fitted parameters discovered through hill climbing. In simulation 1, SOCR was successfully applied to basic acquisition, including the overtraining effect, which is context dependent. In simulation 2, we applied SOCR to basic extinction and renewal. SOCR anticipated these effects with both fixed parameters and best-fitting parameters, although the renewal effects were weaker than those observed in some experiments. In simulation 3a, feature-negative training was simulated, including the often observed transition from second-order conditioning to conditioned inhibition. In simulation 3b, SOCR predicted the observation that conditioned inhibition after feature-negative and differential conditioning depends on intertrial interval. In simulation 3c, SOCR successfully predicted failure of conditioned inhibition to extinguish with presentations of the inhibitor alone under most circumstances. In simulation 4, cue competition, including blocking (4a), recovery from relative validity (4b), and unblocking (4c), was simulated. In simulation 5, SOCR correctly predicted that inhibitors gain more behavioral control than do excitors when they are trained in compound. Simulation 6 demonstrated that SOCR explains the slower acquisition observed following CS-weak shock pairings.     

The influence of cue type and configuration upon radial-maze performance in the rat

The influence of cue type and cue configuration on radial-maze performance in rats was examined in two experiments. In the first experiment, it was found that rats provided with both salient intramaze and extramaze cues acquired the task faster than rats given only one set of cues. No difference in acquisition was found between a group trained with intramaze cues alone and a group trained with extramaze cues alone. In a cue-preference test, it was found that groups that had been trained with extramaze cues, intramaze cues, or both sets of cues relied on extra-maze cues to avoid visited arms when given both types of cues concurrently. When all groups were transferred to intramaze-cue-alone trials, only the group that had been originally trained with extramaze cues alone showed any disruption. Also, during the second half of the intramaze-cue-alone trials, the arrangement of these cues was randomly changed on each trial. This disruption in cue configuration did not deleteriously affect performance in any of the three groups; all remained above chance performance, although the performance of the group originally trained with extramaze cues alone was inferior to that of the other two groups. In Experiment 2, groups of rats were trained on daily alternating trials under intramaze-cue-alone and extramaze-cue-alone conditions. For one group, the configuration of intramaze cues was altered randomly on each trial; the other group had intramaze cues always presented in the same configuration over trials. It was found that acquisition was more rapid on intramaze trials in the group given static configurations. Also, acquisition of the extramaze task was faster than the intramaze task in the group given variable intramaze cue configurations. No difference was found between the intramaze and extramaze conditions in the group given static intramaze cue configurations. These data suggest that a static cue configuration may influence radial maze performance, but is not a necessary condition for such performance.     

Reminder-induced attenuation of the effect of relative stimulus validity

The roles of deficient acquisition and deficient expression of learned information in the effect of relative stimulus validity were examined using rats in a conditioned lick suppression paradigm. Recovery from the effect without further pairings of the conditioned stimulus (CS) and the unconditioned stimulus (US) would favor an interpretation of the relative validity effect based on a latent CS-US association as distinct from a failure to acquire the CS-US association. As a potential recovery manipulation, “reminder” treatments, consisting of the US alone (Experiment 1) or the CS alone (Experiment 2), were administered following relative validity training. In both cases, subjects for which the CS target was of low relative predictive validity exhibited enhanced responding relative to appropriate controls. Additionally, Experiment 2 showed that the amelioration of the relative validity deficit was stimulus specific. Thus, the results of these experiments support previous suggestions that the performance deficit resulting from low relative stimulus validity is due, at least in part, to a failure to express acquired information (Cole, Barnet, & Miller, 1995a).     

The Choice of Response Probability in Bookmark Standard Setting: An Experimental Study

Evidence of the internal consistency of standard-setting judgments is a critical part of the validity argument for tests used to make classification decisions. The bookmark standard-setting procedure is a popular approach to establishing performance standards, but there is relatively little research that reflects on the internal consistency of the resulting judgments. This article presents the results of an experiment in which content experts were randomly assigned to one of two response probability conditions: .67 and .80. If the standard-setting judgments collected with the bookmark procedure are internally consistent, both conditions should produce highly similar cut scores. The results showed substantially different cut scores for the two conditions; this calls into question whether content experts can produce the type of internally consistent judgments that are required using the bookmark procedure.     

Overshadowing not potentiation of illness-based contextual conditioning by a novel taste

In four experiments using rat subjects, we investigated the effects of presenting a novel flavor cue at the time of pairing an environmental context with illness. In each experiment, the subjects were allowed to spend time in a distinctive cage before receiving an injection of LiCl. For some, plain water was made available on these conditioning trials; for others, a novel taste (HCl) was presented. We measured the strength of the context aversion by assessing the ability of the contextual cues to block the acquisition of an aversion to sucrose in a further phase of training. We found that initial training with HCl present made the context less effective as a blocking cue and concluded that the HCl had overshadowed learning about the context. We suggest that this blocking procedure provides a more accurate assessment of contextual aversion than does the consumption test that has more usually been used and that taste-context overshadowing may be a more robust phenomenon than has thus far been thought.     

“counting” by pigeons: Discrimination of the number of biologically relevant sequential events

Numerical competence has been studied in animals under a variety of conditions, but only a few experiments have reported animals’ ability to detect absolute number. Capaldi and Miller (1988) tested rats’ ability to detect absolute number by using biologically important events—the number of reinforced runs followed by a nonreinforced run—and found that the rats ran significantly slower on the nonreinforced run. In the present experiments, we used a similar procedure. Pigeons were given a sequence of trials in which responding on the first three trials ended in reinforcement but responding on the fourth trial did not (RRRN). When the response requirement on each trial was a single peck (Experiment 1), we found no significant increase in latency to peck on the fourth trial. When the response requirement was increased to 10 pecks (Experiment 2), however, the time to complete the peck requirement was significantly longer on the nonreinforced trial than on the reinforced trials. Tests for control by time, number of responses, and amount of food consumed indicated that the pigeons were using primarily the number of reinforcements obtained in each sequence as a cue for nonreinforcement. This procedure represents a sensitive and efficient method for studying numerical competence in animals.     

Extended exposure to environmental cues,but not to sucrose,reduces sucrose cue reactivity in rats

In the present study, we examined the effects of extinction of sucrose-predictive contextual cues and/or sucrose satiation on the expression of sucrose cue reactivity in a rat model of relapse. Context extinction was imposed by housing rats in their home cage or in the operant conditioning chamber for 17 h prior to testing. For sucrose satiation, rats were allowed unlimited access to water or sucrose for 17 h prior to testing. Cue reactivity was assessed after either one (Day 1) or 30 (Day 30) days of forced abstinence from sucrose self-administration. An abstinence-dependent increase in sucrose cue reactivity was observed in all conditions (“incubation of craving”). Context extinction dramatically reduced lever responding on both Day 1 and Day 30. Sucrose satiation had no significant effect on cue reactivity in any condition. These results demonstrate that the context in which self-administration occurs maintains a powerful influence over cue reactivity, even after extended forced abstinence. In contrast, the primary reinforcer has little control over cue reactivity. These findings highlight the important role of conditioned contextual cues in driving relapse behavior.     

Nonassociative habituation,US preexposure,and backward inhibitory conditioning with signaled and unsignaled USs

In a conditioned suppression experiment, rats received a single, massed session of conditioning in which one backward conditioned inhibitory stimulus (CS-) followed shocks that were signaled by a visual cue, and a second backward CS-followed shocks that were unsignaled. Conditioning was preceded by a preexposure phase in which some groups of rats were preexposed to unsignaled shock, while others were not preexposed and remained in the experimental apparatus in the absence of shock. The groups were further distinguished by whether US preexposure and conditioning occurred in the same or different contexts, and by whether conditioning began immediately or after a 24-h rest period in the home cage. Although the conditioning itself was effective in establishing the visual cue as a conditioned excitor in the nonpreexposed groups, it was not effective in establishing the two backward cues as reliable inhibitors with either signaled or unsignaled USs. After 210 US preexposures, however, the same conditioning sessions did yield conditioned inhibition to both CS-s. A 24-h rest period in the home cage reduced the magnitude of, but did not completely abolish, the facilitative effect of US preexposure on inhibitory conditioning. Other tests demonstrated that US preexposure had retarded excitatory conditioning to the visual cue. This interference with excitatory conditioning was unchanged in magnitude after the 24-h rest period. The facilitative effect of US preexposure on backward inhibitory conditioning, and the interference effect on excitatory conditioning, were both eliminated by a change in context between US preexposure and conditioning. These observations encourage predominantly associative accounts of the effects, but allow for a small nonassociative habituation component.