Formation of excitatory and inhibitory associations between absent events

Considerable evidence indicates that associations may be formed between two events even when one or both of them is absent at the time of learning. Previously, some researchers asserted that excitatory associations are formed when associatively activated representations for two events are paired, whereas others claimed that inhibitory associations are formed. In three experiments, the authors investigated the nature of tone-sucrose learning when associatively activated representations of those events were paired in the absence of either of the events themselves. Experiment 1 found substantial excitatory learning when the tone surrogate preceded the sucrose surrogate in training. Experiment 2 evaluated other accounts for the results of Experiment 1, and Experiment 3 found evidence for inhibitory tone-sucrose learning when the tone and sucrose surrogates were presented in simultaneous or backward order. The results indicated that the nature of representation-mediated learning is influenced by some of the same variables as more standard associative learning.     

Contrast effects in maintained generalization gradients

In Experiment I, pigeons were given equal reinforcement (variable-interval 1-min) for responding during randomized presentations of eight line-orientation stimuli. Then, only responding in the vertical orientation was reinforced. Stable generalization gradients soon formed and persistent behavioral and local (transient) contrast effects appeared. Local contrast effects were not a function of relative reinforcement frequency or of any other variable known to produce contrast. Instead, they were related to average response rates associated with each stimulus. Experiment II showed that local contrast effects represent increases and decreases in response rates relative to baseline responding, and that these effects are relative; a given stimulus might enhance responding during a subsequent presentation of one stimulus, but depress responding when followed by another. These data indicate that discrimination learning is not adequately described as the acquisition of excitatory properties by some stimuli and inhibitory properties by others. A more adequate account implies that stimuli exert both excitatory and inhibitory effects related to their value.     

Contralateral transfer of inhibitory and excitatory eyelid conditioning in the rabbit

In Experiment I, rabbits received training to establish a clicker as a conditioned inhibitor. In a subsequent test phase this stimulus was used as a signal for shock either to the eye reinforced during initial training or to the opposite eye. Learning to the clicker was slower in both conditions than in the appropriate control groups. The second experiment replicated the results of those subjects trained and tested with opposite eyes and ruled out the possibility that the slower learning was due to the effects of latent inhibition. Experiment III demonstrated that excitatory conditioning to a clicker to one eye facilitated future excitatory conditioning to that stimulus to the opposite eye. These results are consistent with the view that inhibitory and excitatory conditioning both involve the acquisition of a general, motivational conditioned response which is capable of mediating the transfer of conditioning across different response systems.     

Overshadowing of subsequent events and recovery thereafter

Four experiments using a conditioned lick suppression preparation with rats were conducted to examine whether overshadowing of subsequent events could be obtained in Pavlovian backward conditioning (i.e. unconditioned stimulus [US] before conditioned stimulus [CS]), and to determine whether such overshadowing could be reversed without further training with the overshadowed CS, as has been reported in overshadowing of antecedent events. In Experiment 1, a backward-conditioned CS overshadowed a second backward-conditioned CS. Two posttraining manipulations, extinction of the overshadowing CS (Experiment 2) and shifting of the temporal relationship of the overshadowing CS to the US (Experiment 3), increased responding to the overshadowed CS. These results constitute the first unambiguous demonstration of stimulus competition between subsequent events using first-order conditioning, and they show that, like overshadowing with forward conditioning, such overshadowing is due, at least in part if not completely, to a failure to express information that had been acquired.     

Predictive learning in nutrient-based flavor conditioning

This paper presents two experiments on nutrient-based flavor conditioning with rats as subjects and sucrose as the unconditioned stimulus (US). Experiment 1 was aimed at establishing an optimal control for conditioning, comparing simultaneous and serial presentations of a flavor and the US. The results showed that simultaneous, but not serial training, produced conditioning. Experiment 2 was designed to obtain evidence of summation as an index of both conditioned inhibition and predictive learning. Group Simultaneous received Pavlovian conditioned inhibition training during which flavor A was simultaneously paired with sucrose on excitatory trials (A+), and forming an unreinforced compound with flavor B on inhibitory trials (AB-). An independent excitor for the summation test was also trained by simultaneous pairings with sucrose (C+). In the control group (Blocked), the AB- trials were presented forming a block at the beginning of training to avoid a negative contingency-relationship with sucrose, and flavor A received serial rather than simultaneous pairing with sucrose (A --> +). On the summation test, only in group Simultaneous was consumption of the CB compound lower than that of flavor C alone, suggesting that, during training, flavor A activated an expectancy of the US occurrence.     

Concurrent learning of temporal and spatial sequences

In a serial reaction time task, stimulus events simultaneously defined spatial and temporal sequences. Responses were based on the spatial dimension. The temporal sequence was incidental to the task, defined by the response-to-stimulus intervals in Experiment 1 and stimulus onset asynchronies in Experiment 2. The two sequences were either of equal length and correlated or of unequal length. In both experiments, spatial learning occurred regardless of sequence length condition. In contrast, temporal learning occurred only in the correlated condition. These results suggest that timing is an integrated part of action representations and that incidental learning for a temporal pattern does not occur independently from the action. Interestingly, sequence learning was enhanced in the correlated condition, reflecting the integration of spatial-temporal information.     

Classical conditioning mechanisms can differentiate between seeing and doing in rats

We show that the attentional-associative SLG model of classical conditioning, based on the 1996 research of Schmajuk, Lam, and Gray, correctly describes experimental results regarded as evidence of causal learning in rats: (a) interventions attenuate responding following common-cause training but do not interfere on subsequent responding during observation, and (b) interventions do not affect responding after direct-cause training or (c) causal-chain training. According to the model, responding to the weakly attended test stimulus is strongly inhibited by the intervention in the common-cause case. Instead, in the direct-cause and causal-chain cases, the strongly attended test stimulus becomes inhibitory, thereby overshadowing the inhibitory effect of interventions. Most importantly, the model predicted that with relatively few test trials (a) the 2008 results of Experiment 3 by Leising, Wong, Waldmann, and Blaisdell should be similar to those of Dwyer, Starns, and Honey's 2009 Experiment 1, showing that interventions equally affect responding after common-cause and direct-cause training; and (b) the 2006 results of Experiment 2a by Blaisdell, Sawa, Leising, and Waldmann should be similar to those of Dwyer, Starns, and Honey's 2009 Experiment 2, showing that interventions equally affect responding after common-cause and causal-chain training. When those data were made available to us, we confirmed those predictions. In agreement with the SLG associative model, but not with causal model theory, this evidence supports the notion that the attenuation of responding by interventions only following common-cause training is the consequence of well-known learning processes-latent inhibition, sensory preconditioning, conditioned inhibition, protection from extinction, and overshadowing.     

On the timescale of stimulus-based action-effect learning

When performing an action, people pick up associations between their actions and the resulting consequences of that action, a phenomenon that has been termed response (R)-effect (E) learning. In the present study, we investigated incidental R-E learning in a forced-choice-that is, a stimulus (S)-based-acquisition mode. Specifically, the study examined at which timescale R-E learning evolves-that is, how many encounters are actually needed to form stable R-E associations. The learning of R-E associations was assessed in a subsequent test phase via effect-based response priming. Experiment 1 tested 4 different numbers of S-R-E repetitions for a 2-2-2 S-R-E mapping. Experiment 2 disentangled the contributions of S-E and R-E associations to the facilitating impact of effect-based response priming by means of a 4-2-4 S-R-E mapping. Experiment 3 investigated whether R-E associations can be picked up even when a given E cannot be unequivocally predicted based on the antecedent S in case of inconsistent S-R-E couplings. Together, the results of the present study clearly show that R-E learning occurs in a stimulus-based action mode and that it evolves very rapidly after only 12 S-R-E repetitions. Moreover, the present findings suggest that at least in this initial phase of learning, complete S-R-E consistency seems to be relevant for R-E learning.     

Conditioned inhibition is not the symmetrical opposite of conditioned excitation: A test of the Rescorla-Wagner model

Rescorla and Wagner's (1972) linear associative model of learning predicts that a neutral stimulus will become weakly excitatory if it is paired with a conditioned inhibitor and the compound is not reinforced. If the inhibitor is not allowed to extinguish during these pairings, the model predicts that the neutral stimulus will become strongly excitatory. Rescorla (1971) has presented some evidence to support the former prediction of a weak effect. Experiment I tested the latter prediction of a strong effect using rats in a conditioned emotional response paradigm, and found no evidence that the pairings made the neutral stimulus excitatory. Experiment II replicated Rescorla's results, but showed that even on a relearning test the experimental group learned no faster than a novel stimulus control group.     

Analysis of the learned irrelevance effect in appetitive Pavlovian conditioning

In Experiments 1 and 2 rats received uncorrelated presentations of a light conditioned stimulus (CS) and a food unconditioned stimulus (US) on each day of a preexposure phase. Control subjects received the same number of USs during the first half of preexposure and the same number of CSs during the second. Uncorrelated preexposure retarded inhibitory conditioning. Experiment 3 showed, however, that the different patterns of US preexposure experienced by the two groups could in itself influence the course of subsequent inhibitory conditioning. When this factor was equated by restricting the uncorrelated treatment to the first half of the pre-exposure phase (Experiment 2) or by extending the control treatment throughout the phase (Experiment 4) it was found that uncorrelated preexposure retarded excitatory conditioning, but facilitated inhibitory conditioning. This outcome challenges an interpretation in terms of the concept of learned irrelevance, which predicts that uncorrelated preexposure should retard both forms of conditioning.     

Conditioning across the duration of a backward conditioned stimulus

Five conditioned suppression experiments examined the extent to which an appetitively motivated lever-press response can be punished by different components of a backward conditioned stimulus (CS). Using a 0-s unconditioned stimulus (US)-CS interval, Experiments 1 and 2 showed that the initial 3 s of a normally 30-s backward CS served as a more effective punisher than the CS as a whole. Experiment 3 found no such effect if the US-CS interval were 3 s rather than 0 s. Experiments 4A and 4B found that if the US-CS interval were 0 s, the initial part of the backward CS acquired excitatory properties although the CS as a whole passed a summation test for conditioned inhibition. By contrast, the 3-s US-CS interval supported inhibitory conditioning across the whole duration of the backward CS. Taken together, these findings support a modified version of Wagner's sometimes opponent process model, which suggests that different components of a backward CS become either excitatory or inhibitory depending on the components' temporal proximity to the US.     

The comparator hypothesis of conditioned response generation: manifest conditioned excitation and inhibition as a function of relative excitatory strengths of CS and conditioning context at the time of testing

In the present research water-deprived rats were used in a conditioned lick suppression paradigm to test and further develop Rescorla's (1968) contingency theory, which posits that excitatory associations are formed when a conditioned stimulus (CS) signals an increase in unconditioned stimulus (US) likelihood and that inhibitory associations develop when the CS signals a decrease in US likelihood. In Experiment 1 we found that responding to a CS varied inversely with the associative status of the context in which the CS was trained and that this response was unaltered when testing occurred in a distinctively dissimilar context with a different conditioning history, provided associative summation with the test context was minimized. These results suggest that manifest excitatory and inhibitory conditioned responding is modulated by the associative value of the training context rather than that of the test context. In Experiment 2 it was demonstrated that postconditioning decreases in the associative value of the CS training context reduced the effective inhibitory value of the CS even when testing occurred outside of the training context. Moreover, this contextual deflation effect was specific to the CS training context as opposed to any other excitatory context. Collectively, these studies support the comparator hypothesis, which states that conditioned responding is determined by a comparison of the associative strengths of the CS and its training context that occurs at the time of testing rather than at the time of conditioning. This implies that all associations are excitatory and that responding indicative of conditioned inhibition reflects a CS-US association that is below (or near) the associative strength of its comparator stimulus. It is suggested that response rules which go beyond a monotonic relation between associative value and response strength can partially relieve learning theories of their explanatory burdens, thereby allowing for simpler models of acquisition.     

Resolution of outcome-induced response conflict by humans after extended training

Studies of incongruent discrimination learning, where the outcome event of one response acts as the discriminative stimulus for the opposite response, suggest that humans rely on habitual stimulus–response (S–R) associations when outcome–response (O–R) associations would cause response conflict. Here, two experiments were conducted to investigate the robustness of this habitual strategy. In Experiment 1, we found that extensive instrumental discrimination training supported learning about the incongruent R → O contingencies, as assessed by an outcome devaluation test. Differential representations of the stimulus and the (associatively retrieved) outcome may have allowed for goal-directed incongruent performance. Experiment 2 failed to provide evidence for this possibility; direct presentation as well as associative retrieval of the incongruent events (by Pavlovian stimuli) activated the response that was associated with each event in its role of stimulus as opposed to outcome. We did find that participants successfully acquired explicit knowledge of the incongruent contingencies, which raises the possibility that propositional encoding allowed them to overcome the response conflict caused by O–R associations. Alternative associative and propositional accounts of successful goal-directed incongruent performance with extensive training will be discussed.     

Blocking and unblocking in human causal learning

Three experiments sought to develop the suggestion that, under some circumstances, common associative learning mechanisms might underlie animal conditioning and human causal learning, by demonstrating, in humans, an effect analogous to the unblocking by reinforcer omission observed in animal conditioning. Experiment 1 found no such effect. Experiment 2, designed to prevent inhibitory influences that might have masked excitatory unblocking in Experiment 1, demonstrated unblocking, indicating common human-animal associative learning mechanisms in which the associability of a stimulus varies as a function of its predictive history. Experiment 3, using a similar design but with a procedure promoting application of rational inference processes, failed to detect the same unblocking effect, indicating that associative and cognitive mechanisms may influence human causal learning.     

On the timescale of stimulus-based action–effect learning

When performing an action, people pick up associations between their actions and the resulting consequences of that action, a phenomenon that has been termed response (R)–effect (E) learning. In the present study, we investigated incidental R–E learning in a forced-choice—that is, a stimulus (S)-based—acquisition mode. Specifically, the study examined at which timescale R–E learning evolves—that is, how many encounters are actually needed to form stable R–E associations. The learning of R–E associations was assessed in a subsequent test phase via effect-based response priming. Experiment 1 tested 4 different numbers of S–R–E repetitions for a 2–2–2 S–R–E mapping. Experiment 2 disentangled the contributions of S–E and R–E associations to the facilitating impact of effect-based response priming by means of a 4–2–4 S–R–E mapping. Experiment 3 investigated whether R–E associations can be picked up even when a given E cannot be unequivocally predicted based on the antecedent S in case of inconsistent S–R–E couplings. Together, the results of the present study clearly show that R–E learning occurs in a stimulus-based action mode and that it evolves very rapidly after only 12 S–R–E repetitions. Moreover, the present findings suggest that at least in this initial phase of learning, complete S–R–E consistency seems to be relevant for R–E learning.     

Initial excitatory conditioning with the truly random control procedure in rats: The effects of density of the conditioned stimulus

The present experiments were designed to test whether initial excitatory conditioning to a stimulus was obtained, using a within-subject design of the truly random control (TRC) procedure, and to examine the effects of the density of the conditioned stimulus on it. In Experiment 1, the effect of the three levels of CS density (50%, 25%, 20%) was assessed at six different stages of training, using the conditioned suppression of licking in rats. Although all of the groups given the TRC procedure showed evidence of initial excitatory conditioning, it was more marked in the group with the highest CS density than in the other groups. Experiment 2 confirmed the results of Experiment 1, using a between-subject design. The findings are in line with predictions from the Rescorla-Wagner model, and are discussed with reference to other contemporary theories of associative learning.     

Mechanisms underlying retarded emergence of conditioned responding following inhibitory training: evidence for the comparator hypothesis

The comparator hypothesis posits that conditioned responding is determined by a comparison at the time of testing between the associative strengths of the conditioned stimulus (CS) and stimuli proximal to the CS at the time of conditioning. The hypothesis treats all associations as being excitatory and treats conditioned inhibition as the behavioral consequence of a CS that is less excitatory than its comparator stimuli. Conditioned lick suppression by rats was used to differentiate four possible sources of retarded responding to an inhibitory CS. These include habituation to the unconditioned stimulus (US), latent inhibition to the CS, blocking of the CS-US association by the conditioning context, and enhanced excitatory associations to the comparator stimuli. Prior research has demonstrated the first three phenomena. Therefore, we employed parameters expected to highlight the fourth one--the comparator process. In Experiment 1, our negative contingency training was shown to produce a conditioned inhibitor that passed inhibitory summation and retardation tests. In Experiment 2 we found transfer of retardation from an inhibitory CS to a novel stimulus when the location where retardation-test training occurred was excitatory, which is indicative of contextual blocking and/or comparator effects. In Experiment 3, extinction of the conditioning context was found to attenuate retardation regardless of whether extinction occurred before or after the CS-US pairings of the retardation test. This indicates that much of the present retardation was due to the comparator process rather than to contextual blocking. Experiment 4 demonstrated that habituation to the US did not contribute to retardation in the present case. Collectively, these studies suggest that retardation following inhibitory training can be explained without recourse to any of the traditional mechanisms of conditioned inhibition.     

Representation-mediated extinction of conditioned flavor aversions

Conditioned flavor aversions were extinguished by presenting without consequence auditory stimuli that had been previously paired with the aversive flavor. In Experiment 1, rats that received tone-sucrose pairings, then sucrose-lithium chloride (LiCl) pairings, and finally repeated tone-alone presentations showed greater sucrose consumption in subsequent testing than rats that received similar sucrose-LiCl pairings and tone-alone presentations but no initial tone-sucrose pairings. Experiment 2 demonstrated the stimulus specificity of the mediated extinction observed in Experiment 1. In Experiment 3, rats that received first-order light-food and second-order tone-light pairings prior to sucrose-LiCl pairings did not show greater subsequent sucrose consumption when extinction of the second-order tone intervened. These results suggest that conditioned stimulus (CS)-evoked representations of events can substitute for those events themselves in the extinction of previously established associations.     

Stroop interference is affected in inhibition of return

In previous research, we have shown that the processing of targets that are presented to locations subject to inhibition of return (IOR) is affected by an inhibitory tagging mechanism. This mechanism acts by disconnecting activated representations of stimuli at inhibited locations from their associated responses. In two experiments, we assessed whether this inhibitory tagging mechanism of visual attention is also applied to task-irrelevant but prepotent dimensions of target stimuli, such as words in the Stroop task. To test this hypothesis, we examined the Stroop effect in an IOR procedure. The results showed that (1) IOR can be found in a color discrimination task, (2) the Stroop interference was reduced (Experiment 1) or eliminated (Experiment 2) when stimuli appeared at cued locations, as compared with cases in which they were presented at uncued locations, and (3) the effect of inhibitory tagging was limited to the shortest stimulus onset asynchrony value, replicating previous findings. These results agree with the idea that inhibitory tagging, occurring in IOR, affects the efficiency with which color words compete for responses in Stroop-like situations.     

Perirhinal cortex lesions impair feature-negative discrimination

The role of the perirhinal cortex in inhibitory eyeblink conditioning was examined. In Experiment 1, rats were given lesions of the perirhinal cortex or control surgery and subsequently trained with a feature-negative discrimination procedure followed by summation and retardation tests for conditioned inhibition. Perirhinal cortex lesions impaired, but did not prevent acquisition of feature-negative discrimination. Results from the summation test showed that rats with perirhinal cortex lesions could not generalize feature-negative discrimination to a new stimulus. There were no group differences during the retardation test. Experiment 2 showed that lesions of the perirhinal cortex did not impair simple excitatory conditioning. Experiment 3 showed that perirhinal cortex lesions had no effect on acquisition of a simple tone-light discrimination. The results suggest that the perirhinal cortex plays a role in eyeblink conditioning when using discrimination procedures involving overlapping stimuli.