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.     

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.     

Simultaneous and backward fear conditioning as a function of number of CS-UCS pairings.

Four experiments examined the possibility that the outcome of simultaneous and backward fear conditioning procedures might depend upon the number of CS-UCS pairings. A punishment procedure will rats as subjects and shock as the UCS was used; the amount of suppression produced by response-contingent CS presentations indexed the strength of acquired fear. Experiments 1, 3, and 4 examined the suppressive tendencies of simultaneous-and backward-trained CSs after 0, 10, 20, 40, 80, and 160 pairings. The pattern of data suggested that initial pairings have the effect of increasing suppressive tendencies of the CS, while subsequent pairings decrease them. In addition, evidence of fear inhibition was found after 160 pairings in the case of the backward paradigm. Experiment 2 examined several nonassociative accounts based upon differential shock exposure. Groups given 10 pairings or 80 pairings were compared to ggroups given 10 pairings plus 70 shock-alone presentations. The results indicated that number of pairings, rather than number of UCS occurrences, is the important factor in decreasing the initial suppression. The evidence for eventual inhibition in the backward paradigm suggests that this occurs through the acquisition of inhibitory tendencies which are antagonistic to the previously conditioned excitation.     

Sign-tracking behavior in aversive conditioning: Its acquisition via a Pavlovian mechanism and its suppression by operant contingencies

When rats are presented with Pavlovian backward pairings in which shock is the unconditioned stimulus (US) and an elevated platform the conditioned stimulus (CS), they show a strong tendency to approach the platform. Attempts were made at evaluating the nature of the associative mechanisms responsible for the acwuisition and maintenance of the approach behavior. In Experiment 1, following a baseline period when platform presentations were given in the absence of shocks, first (training I), approach to the platform was prevented and subjects were exposed to either backward, forward, or random presentations of shock and platform: this procedure attempted to minimize the effects of response learning on the acquisition of approach behavior. Then (training II), all subjects received backward shock-platform pairings and had access to the platform. Approach to the platform was stronger in animals previously exposed to backward pairings than in the animals of the other two groups. Also, animals preexposed to forward platform-shock pairings showed suppression of approach behavior on the first trial of training II relative to their performance on the last trial of baseline. These data suggest that a stimulus-reinforcer learning mechanism alone is sufficent for the acquisition of approach. Experiments 2a and 2b evaluated the effects of “punishing” operant contingencies on Pavlovian-produced approach behavior. The results showed that approach can be eliminated, but only after repeated exposure to an operant contingency which consisted of the immediate closure of the platform upon approach responding. These data suggest that approach behavior produced by Pavlovian contingencies can be modified by operant contingencies. Taken together, these data provide additional support for a sign-tracking notion in aversive conditioning.     

Backward conditioning as an extinction procedure

In two conditioned suppression experiments, rats received Pavlovian forward defense conditioning in which tonal conditioned stimuli (CSs) terminated with the onset of scrambled grid shock unconditioned stimuli (USs). After this experience, the rats then received a Pavlovian backward conditioning procedure in which the same USs now terminated with the onset of the same CSs. Although the two experiments differed greatly in terms of CS and US parameters, number of forward and backward pairings, and in terms of the general techniques used to establish and measure the Pavlovian conditioned response (CR), the results of both experiments agreed in showing that backward conditioning can indeed weaken a CR based on forward pairings. The results also show that, under some conditions, the backward procedure can be at least as effective in weakening an established CR as the traditional CS-alone extinction procedure; but, under other conditions, the backward procedure is less effective and leads to more spontaneous recovery than the CS-alone procedure.     

Acquisition of representation-mediated conditioned food aversions

Food aversions were established in rats by administering lithium chloride (LiCl) immediately after the presentation of an exteroceptive conditioned stimulus (CS) which previously had been paired with a food substance. In Experiment 1, rats which received first tone-food and then tone-LiCl pairings showed less food consumption in subsequent testing than rats which received only tone-food or tone-LiCl pairings. Experiments 2 and 3 demonstrated the stimulus specificity of aversions established in this manner. Rats which first received pairings of light and tone CSs with two different food substances and then received pairings of one of those CSs with LiCl showed greater aversion to the food previously associated with the LiCl-paired CS than to the other food substance. Experiment 3 also showed that specific aversions were not acquired if rats received CS-shock rather than CS-LiCl pairings. These results suggest that CS-evoked representations of events can substitute for those events themselves in the formation of new associations.     

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.     

Determinants of Response Recovery in Extinction Following Response Elimination

Elimination of autoshaped responding by random or negatively-contingent response-reductive procedures may leave a previously acquired association intact; however, two previous studies suggest that response elimination by backward conditioning may have more permanent effects. In Experiments 1 and 3, the autoshaped responding of pigeons was eliminated by a backward or negative contingency. Although speed of recovery during a subsequent extinction phase was greater following the backward contingency in Experiment 3, level of recovery did not differ as a function of response elimination procedure in either experiment. A possible basis for the discrepancy between the present and previous findings is the contingency arranged between the conditioned and unconditioned stimuli (CS and US, respectively) during forward conditioning. In Experiments 1 and 2, probability of US presentation following the CS (p(US|CS) = 0.4 or 1.0) was varied within (Experiment 1) or between (Experiment 2) groups of birds. In both cases, level of recovery was higher following training under (p(US|CS) = 0.4). The results are consistent with a memory retrieval theory in which unreinforced trials function as a retrieval cue for not responding.     

Backward conditioning: mediation by the context

The information acquired in backward conditioning (i.e., outcome-->cue) was assessed in 3 Pavlovian lick-suppression experiments with water-deprived rats as subjects. Experiment 1 confirmed previous research that few outcome-->cue pairings made the cue into a conditioned excitor and additionally showed that massive posttraining extinction of the training context attenuated a backward-trained cue's excitatory value. Experiment 2 found that many outcome-->cue pairings made the cue into a conditioned inhibitor and that the same context manipulation attenuated this inhibitory value. Experiment 3 confirmed the observations of Experiments 1 and 2 and demonstrated that these effects of context extinction were specific to backward-trained cues conditioned in the extinguished context. These results are interpreted in terms of cue-->context and context-->outcome associations.     

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.     

Within-compound associations between the context and the conditioned stimulus

Three experiments were conducted to test for the presence of associations between contextual cues and the nominal conditioned stimulus (CS) in fear conditioning. Rats were given tone-footshock pairings and were tested for their fear of the nominal CS, the tone, in a different context. Experiments 1 and 2 demonstrated that rats given nonreinforced exposure to the training context following conditioning were less fearful of the CS. Experiment 3 indicated that additional footshock presentations in the training context following conditioning produced greater fear of the CS. In both procedures postconditioning treatments designed to directly alter only the associative strength of the training context yielded parallel changes in the conditioned response to the CS. These data suggest that within-compound associations are formed between the context and the CS during classical conditioning.     

Associative effects of US preexposure: modulation of conditioned responding by an excitatory training context

In two experiments we examined factors that contribute to retarded emergence of conditioned responding to a conditioned stimulus (CS) trained in a context in which unsignaled unconditioned stimuli (USs) had previously been administered. In both experiments water-deprived rats were used in a conditioned lick suppression task to measure the conditioned response elicitation potential of the CS and the training context. From Experiment 1 we determined that nonreinforced exposure to the excitatory context after US preexposure and prior to CS-US pairings in that context eliminated the conditioned response deficit observed on a subsequent test of the CS. The recovery from the US preexposure deficit was nearly as great in animals that received nonreinforced exposure to the excitatory training context after the CS-US pairings but prior to the ultimate test of the CS. From Experiment 2 we determined that the recovery induced by contextual deflation after CS training was specific to deflation of the context in which the CS was trained as opposed to another excitatory context. In total, these experiments suggest that context-US associations partially mask the expression of a learned CS-US association. These results are discussed in terms of recent models of conditioned response generation.     

Pavlovian second-order conditioned analgesia

Three experiments with rat subjects assessed conditioned analgesia in a Pavlovian second-order conditioning procedure by using inhibition of responding to thermal stimulation as an index of pain sensitivity. In Experiment 1, rats receiving second-order conditioning showed longer response latencies during a test of pain sensitivity in the presence of the second-order conditioned stimulus (CS) than rats receiving appropriate control procedures. Experiment 2 found that extinction of the first-order CS had no effect on established second-order conditioned analgesia. Experiment 3 evaluated the effects of post second-order conditioning pairings of morphine and the shock unconditioned stimulus (US). Rats receiving paired morphine-shock presentations showed significantly shorter response latencies during a hot-plate test of pain sensitivity in the presence of the second-order CS than did groups of rats receiving various control procedures; second-order analgesia was attenuated. These data extend the associative account of conditioned analgesia to second-order conditioning situations and are discussed in terms of the mediation of both first- and second-order analgesia by an association between the CS and a representation or expectancy of the US, which may directly activate endogenous pain inhibition systems.     

Forward and backward second-order Pavlovian conditioning in honeybees

Second-order conditioning (SOC) is the association of a neutral stimulus with another stimulus that had previously been combined with an unconditioned stimulus (US). We used classical conditioning of the proboscis extension response (PER) in honeybees (Apis mellifera) with odors (CS) and sugar (US). Previous SOC experiments in bees were inconclusive, and, therefore, we attempted to demonstrate SOC in the following three experiments: (Experiment 1) After differential conditioning (pairing odor A with US and presenting odor B without US), the bees experienced two pairs of partially overlapping odors, either a new odor C followed by a previously reinforced odor A (C-A) or a new odor C followed by a previously nonreinforced odor B (C-B). (Experiment 2) After differential conditioning, bees were presented with C-A or A-C. (Experiment 3) Bees were first presented with C-A or A-C before differential conditioning and were tested with odor C. We observed: (Experiment 1) 40% of the bees showed PER to the C-A presentation, but only 20% showed PER to the C-B presentation. (Experiment 2) 40% of the bees showed PER to the C-A presentation, while only 20% showed PER to the reversed sequence A-C. Experiments 1 and 2 showed that a previously reinforced odor can be a secondary reinforcer for excitatory SOC only with forward-pairing. (Experiment 3) PER toward C was lower (15%) in bees presented with A-C than with C-A (25%). This showed that backward SOC is not as effective as forward SOC. These results help to delineate different conditions that are critical for the phenomenon of SOC.     

Comparing excitatory backward and forward conditioning

Three Pavlovian lick suppression studies with rats were conducted to compare the role of the conditioning context in excitatory backward and forward conditioning. The experiments explored the possibility that excitatory backward conditioning, but not forward conditioning, is mediated by the context. That is, in excitatory backward conditioning, the conditioning context may function as an excitatory mediator, which supports second-order conditioning of the target cue. This possibility contrasts with traditional accounts, which suggests that common processes underlie excitatory backward and forward conditioning. Experiment 1 found that conditioned responding following backward conditioning was attenuated as a result of posttraining extinction of the training context, but the same manipulation elevated responding after forward conditioning. Experiments 2 and 3 found that posttraining and pretraining associative inflation of the context (presenting unsignalled USs) increased conditioned responding to the target of a backward conditioning procedure but either had no effect or reduced responding to the target of a forward conditioning procedure. Thus, excitatory backward and forward conditioning appear to differ in their dependence on the status of the conditioning context.     

Transfer across unconditioned stimuli in serial feature discrimination training

The transfer of conditioned modulation across conditioned stimuli (CS) and unconditioned stimuli (US) was examined in 3 experiments that used Pavlovian appetitive training procedures with rats. In Experiment 1, after training in a positive patterning discrimination (X-->A+/X-/A-), X increased conditioned responding elicited by another trained-then-extinguished CS as long as that CS had been trained with the same US as was used in discrimination training. In Experiment 2, after training with a feature-negative discrimination (X-->A-/A+), X inhibited conditioned responding elicited by another trained-then-extinguished CS as long as that CS had been trained with the same US. Experiments 1 and 2 used a between-groups design, whereas Experiment 3 used a more powerful within-groups design. In Experiment 3, rats were trained in a feature-positive discrimination (X-->A+/A-). In transfer tests, X increased conditioned responding elicited by another CS trained then extinguished with the same US from training. This increase was greater than the X increased conditioned responding elicited by another CS trained then extinguished with a different US from training. The results supported the suggestion that features trained in serial discrimination tasks influence behavior indirectly by transiently raising or lowering the threshold for activation of the US representations by its target stimuli and by any other stimuli that may be associated with that US. Other interpretations of the findings were also considered.     

Implicit attitude formation through classical conditioning

We sought to demonstrate that attitudes can develop through implicit covariation detection in a new classical conditioning paradigm. In two experiments purportedly about surveillance and vigilance, participants viewed several hundred randomly presented words and images interspersed with critical pairings of valenced unconditioned stimuli (USs) with novel conditioned stimuli (CSs). Attitudes toward the novel objects were influenced by the paired USs: In a surprise evaluation task, the CS paired with positive items was evaluated more positively than the CS paired with negative items. This attitudinal conditioning effect was found using both an explicit measure (Experiments 1 and 2) and an implicit measure (Experiment 2). In a covariation estimation task involving the stimuli presented in the conditioning procedure, participants displayed no explicit memory for the pairings.     

Naloxone-induced hypoalgesia: Effects of heat, cold and novelty

Experiment 1 confirmed that pairings of the opioid antagonist naloxone and a heated floor served to induce and then to maintain a conditioned hypoalgesia in rats. Experiments 2a and b demonstrated that this was not specific to some property of the heated floor: pairings of the drug and a cold floor (a) resulted in conditioned hypoalgesia and (b) maintained the hypoalgesia provoked by pairings of the drug with a heated floor. Experiments 3a and 4 showed that pairings of naloxone with the non-heated floor of the apparatus provoked hypoalgesia and provided evidence that this was due to the drug's disruption of a familiarization process. Experiment 3b revealed that pairings of naloxone with the non-heated floor maintained the hypoalgesia induced by pairings of the drug with a heated one. The results were interpreted to mean that naloxone interacts selectively with a stressor so as to maintain or enhance its aversive properties.     

Overshadowing and stimulus duration

In 3 experiments, the authors investigated the effect of stimulus duration on overshadowing. Experiments 1 and 2 examined responding to a target conditioned stimulus (CS1) when it was conditioned in compound with a coterminating overshadowing stimulus (CS2) that was longer, shorter, or of the same duration (the long, short, and matched groups, respectively). Equal overshadowing of conditioning to CS1 was obtained in all 3 conditions relative to a control group conditioned to the light alone. There were, however, differences in responding to CS2 as a function of its absolute duration. Experiment 3 examined the contribution of the food-food interval/CS onset-food interval ratio to these findings. In Experiments 1 and 2, the ratio differed for the overshadowing CS but not for the target CS. In Experiment 3, this arrangement was reversed, but the pattern of results remained the same. The implications of these findings for trial-based and real-time models of conditioning are discussed.     

Malleability of conditioned associations: path dependence

Using conditioned suppression of barpressing to investigate the stability of a conditioned stimulus-unconditioned stimulus (CS-US) association, we gave water-deprived rats either a few pairings of the CS with a strong footshock US or many pairings with a weak footshock US so that barpress suppression in response to the CS was equated. Experiment 1 established training parameters that yielded this equivalence. Specifically, rapid acquisition to a preasymptotic level of responding with strong shock produced suppression comparable to the asymptotic level reached more slowly with weak shock. Experiment 2 showed that although equivalent performance was obtained from extensive conditioning with a weak shock or limited conditioning with strong shock, only extensive conditioning with weak shock resulted in retarded acquisition of an association between that same CS and a footshock level perceived as midway between the two initial training shock intensities as implied by asymptotic performance in Experiment 1. Experiment 3 demonstrated that the observed retardation in animals given many conditioning trials with weak shock was CS specific. Collectively, these findings indicate that the malleability of learned behavior is not simply a function of initial associative strength but is dependent on path during initial acquisition.