Appetitive differential conditioning of the rabbit’s jaw movement response

Two experiments examined appetitive differential conditioning of the rabbit’s jaw movement response (JMR) in a two-phase procedure. The first phase entailed reinforced training with one conditioned stimulus (CS+), and the second phase involved intermixed presentations of CS+ and an unreinforced stimulus (CS?). In Experiment 1, CS+ was a 600-Hz tone, and CS-was either a 660-,1,000-, or 2,100-Hz tone. In Experiment 2, the magnitude of the water unconditioned stimulus (US) paired with CS+ was either 1, 3, or 9 ml. The experiments revealed that 1) the level of responding to CS-rose for several days and then declined over the remainder of training; 2) the physical similarity between CS+ and CS?directly affected the level of responding to CS?but had no discernable effect on the level of responding to CS+ ; and 3) US magnitude positively affected the level of responding to CS + and, to a lesser extent, CS?. The results are discussed in terms of their implications for Spence’s gradient interaction theory and Pavlov’s external inhibition hypothesis.     

Facilitation of extinction by an increase or a decrease in trial duration

Five experiments examined the effects of altering the duration of a conditioned stimulus (CS) for extinction. For the first 3 experiments, rats received conditioning with a 10-s CS before different groups received extinction with a CS that was either the same duration or longer than that used for conditioning. For the remaining 2 experiments, conditioning was conducted with a 60-s CS before different groups received extinction with a CS of either the same duration or a shorter duration than that used for conditioning. In all experiments, extinction progressed more readily when the CS duration was different for the 2 stages than when it was constant. The results are discussed in terms of rate expectancy theory and associative learning theory.     

In the blink of an eye: Real-time stimulus factors in delay and trace conditioning of the rabbit's nictitating membrane response

Since Pavlov, theories of conditioning have assumed that CR evocation is governed by a series of internal stimuli generated by the CS. This hypothesis was tested in conditioning of the rabbit's nictitating membrane (NM) response by attempting to manipulate the internal sequence through truncating a delay CS and extending a trace CS on test trials. These perturbations of CS duration produced large deficits in CR likelihood and smaller alterations in the CR's time course. As predicted by many models of conditioning, the onset of the CS appeared to play a large but not exclusive role compared to CS duration and CS offset in both evocation and timing of the CR. The results are discussed with respect to their implications for real-time models of conditioning.     

Blocking with serial compound stimuli: The role of local context and second-order associations

Blocking was studied with rats in two serial conditioning experiments in which CS1 was followed by CS2 and then shock. Experiment 1 demonstrated that pretraining with CS1 was able to block conditioning to CS2 when the pretraining consisted of trace conditioning. But when serial conditioning was used for pretraining, with a third stimulus as the second element of the compound, then blocking was not detected during the subsequent phase. In Experiment 2 the effect of pretraining with CS2 on blocking with CS1 was examined. Blocking was effective, but only when steps were taken to minimize the growth of second-order associations resulting from the pairing of CS1 with CS2. These results are consistent with a principle stating that the ability of a pretrained stimulus to block the added stimulus in a compound depends on the relative contiguity of each stimulus to the US.     

Counterconditioning of an overshadowed cue attenuates overshadowing

In 3 Pavlovian conditioned lick-suppression experiments, rats received overshadowing treatment with a footshock unconditioned stimulus such that Conditioned Stimulus (CS) A overshadowed CS X. Subjects that subsequently received CS X paired with an established signal for saccharin (CS B) exhibited less overshadowing of the X-footshock association than subjects that did not receive the X-B pairings (Experiment 1). Experiment 2 replicated this effect and controlled for some additional alternative accounts of the phenomenon. In Experiment 3, this recovery from overshadowing produced by counterconditioning CS X was attenuated if CS B was massively extinguished prior to counterconditioning. These results are more compatible with models of cue competition that emphasize differences in the expression of associations than those that emphasize differences in associative acquisition.     

Mismatch between what is expected and what actually occurs triggers memory reconsolidation or extinction

In previous experiments on contextual memory, we proposed that the unreinforced re-exposure to the learning context (conditioned stimulus, CS) acts as a switch guiding the memory course toward reconsolidation or extinction, depending on reminder duration. This proposal implies that the system computes the total exposure time to the context, from CS onset to CS offset, and therefore, that the reminder presentation must be terminated for the switching mechanism to become operative. Here we investigated to what extent this requirement is necessary, and we explored the relation between diverse phases in the reconsolidation and extinction processes. We used the contextual memory model of the crab Chasmagnathus which involves an association between the learning context (CS) and a visual danger stimulus (unconditioned stimulus, US). Administration of cycloheximide was used to test the lability state of memory at different time points. The results show that two factors, no-reinforcement during the reminder (i.e., CS re-exposure) and CS offset are the necessary conditions for both processes to occur. Regardless of the reminder duration, memory retrieved by unreinforced CS re-exposure emerges intact and consolidated when tested before CS offset, suggesting that neither reconsolidation nor extinction is concomitant with CS re-exposure. Either process could only be triggered once the definitive mismatch between CS and US is confirmed by CS termination without the expected reinforcement.     

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.     

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.     

Cholinergic modulation of trace conditioning trained in serial compound: A developmental analysis

In four experiments the effects of serial compound conditioning on responding to a trace-conditioned CS were evaluated using a fear conditioning paradigm. The subjects were 18- and 25-day-old Sprague-Dawley rats, previously shown to exhibit little or no trace fear conditioning. Here, animals as young as 18 days of age were shown to be capable of trace conditioning between a visual CS1 and a shock US, provided the trace interval was filled with a non-target CS2 during serial conditioning trials (CS1-->CS2-->US). To explore cholinergic mechanisms involved in trace and serial conditioning, additional experiments assessed conditioned responding following pre-training administration of the muscarinic receptor antagonist scopolamine. Scopolamine produced a dose-dependent reduction in responding to the trace CS1, regardless of whether subjects were trained with standard trace (CS1-->trace interval-->US) or serial (CS1-->CS2-->US) trials. Responding to CS2 was unaffected by scopolamine. These data suggest that central cholinergic systems are functional in the young animals, but are not normally sufficiently activated by standard trace conditioning procedures. The results suggest that serial compound conditioning can promote trace conditioning in young rats, as it does in adults, perhaps by enhancing cholinergic activity during training. Implications for the late ontogenetic emergence of trace conditioning as it relates to maturation of neural pathways and their role in the potentiating effects of a gap filler are discussed.     

Associative transfer and stimulus selection in classical conditioning of the rabbit's nictitating membrane response to serial compound CSs

Four experiments were conducted to determine whether in conditioning to a serial compound, CS1-CS2-UCS, there are (a) associative mechanisms operating to extend conditioning beyond the bounds of a CS-UCS contiguity gradient and (b) stimulus selection processes acting to attenuate the potency of CS-UCS contiguity. In Experiments 1 and 2, the CS2-UCS interval was held at .35 sec while the CS1-UCS interval was varied across groups from .75 to 2.75 sec. CS1 test trials revealed substantial CR acquisition at all CS1-UCS intervals. Moreover, Experiment 2 indicated that when the contribution of cross-modal generalization from CS2 to CS1 was factored out, there still remained a substantial level of conditioning, which Experiment 3 indicated was attributable to an associative mechanism like higher-order or sensory conditioning. The observation of CR acquisition at CS1-UCS intervals of 4.75, 8.75, and 18.75 sec in Experiment 4 suggested that serial compound training yields conditioning to CSs located well beyond the single CS contiguity gradient for the rabbit's nictitating membrane response. Experiments 1 and 2 also indicated the presence of stimulus selection processes because, at the shorter CS1-UCS intervals (.75 and 1.25 sec), the levels of test-trial responding to CS2 fell below those observed to the less contiguous CS1.     

Olfactory fear conditioning paradigm in rats: Effects of midazolam,propranolol or scopolamine

In rodents, fear conditioned responses are more pronounced toward olfactory stimulus, since olfaction is a dominant sense in these subjects. The present study was outlined to investigate if the association between coffee odor (CS1) and electrical footshock (US) would be an effective model for the study of fear-induced behavior and whether compounds used in humans for emotional-related disorders such as midazolam, propranolol, or scopolamine, applied during the different stages of fear conditioning (acquisition, consolidation and expression), affect the defensive responses to both, the olfactory CS1, and the context (CS2) where the CS1 had been presented (second order conditioning). The results revealed that five pairings between coffee odor (CS1) and electrical footshock (US) were able to elicit consistent defensive responses and a second order conditioning to the context (CS2). Midazolam (0.375–0.5 mg/kg; i.p.) treatment was able to interfere with the CS1–US association and with the consolidation of the aversive information. The propranolol (5–10 mg/kg; i.p.) treatment interfered with the CS1–US association, with the retention of fear memory and with the CS1–CS2 association. Propranolol also attenuated the expression of conditioned fear responses when applied before the CS1 test session. Scopolamine (0.6–1.2 mg/kg; i.p.) treatment impaired the acquisition of CS1–US and CS1–CS2 associations, and also disrupted the expression of conditioned fear responses when injected prior to the CS1 test session. These findings have pointed out the usefulness for the olfactory fear conditioning paradigm to investigate drug effects on the acquisition, consolidation and expression of fear conditioned responses.     

The role of the CS and its relationship to the CR in avoidance conditioning

Two studies are reported which are concerned with the role of the CS and its relationship to the CR in the standard avoidance learning situation. Study I demonstrates that the decrement associated with a trace CS is greatly reduced in one-way avoidance as compared with two-way. The suggestion is made that the differences in CS effects is attributable to differences in task complexity. Study 2 varied CS duration and the CS-CR termination contingency. Rapid acquisition of the avoidance response was obtained only when the CS remained on for the entire CS-US interval unless terminated by a response. The results were discussed with respect to the theoretical aspects of maintenance and emergence of the avoidance response.     

Fear as a discriminative stimulus for an appetitive instrumental response

Two experiments tested the hypothesis that anticipation of shock could be established as a discriminative stimulus for an appetitive instrumental response. In multiphase experiments, bar pressing for food was brought under the discriminative control of intermittent and gradually increasing electric shock. In a second phase, tones were estalished as either a CS+, or CS? for shock. Subsequently, the CSs were introduced on to the operant baseline. Animals trained with shock as the S^D showed an increase in responding to the CS+ and a slight decrease to the CS?. Conversely, animals trained with shock as the S^Δ showed decreased responding to the CS+ and slight increase to the CS?. These findings are seen as supportive of the Discriminative Stimulus hypothesis of learned resistance to punishment.     

Learned safety and the CS-US delay gradient in taste-aversion learning

Decrements in taste-aversion learning produced by extensions of the interval between the CS flavor and poisoning (the US) may be attributed to decay of the CS trace during long CS-US intervals. Kalat and Rozin (1973) alternatively proposed that such decrements occur because during extended CS-US intervals subjects learn that the CS is safe. They sought to show that trace decay is not responsible for the CS-US delay gradient by demonstrating that learning is disrupted even if the CS is reintroduced during extended CS-US intervals. The present experiments show that such a second CS presentation during conditioning may (1) enhance subsequent intakes of the CS solution whether or not subjects are poisoned, and/or (2) facilitate aversion learning, the facilitation being greater the closer the second CS exposure is to poisoning. These results question the adequacy of previous evidence for the contribution of learned safety to the CS-US delay gradient and suggest that some other process is also involved.     

Is evaluative conditioning really resistant to extinction? Evidence for changes in evaluative judgements without changes in evaluative representations

Evaluative conditioning (EC) is defined as the change in the evaluation of a conditioned stimulus (CS) due to its pairing with a positive or negative unconditioned stimulus (US). Although several individual studies suggest that EC is unaffected by unreinforced presentations of the CS without the US, a recent meta-analysis indicates that EC effects are less pronounced for post-extinction measurements than post-acquisition measurements. The disparity in research findings suggests that extinction of EC may depend on yet unidentified conditions. In an attempt to uncover these conditions, three experiments (N = 784) investigated the influence of unreinforced post-acquisition CS presentations on EC effects resulting from simultaneous versus sequential pairings and pairings with single versus multiple USs. For all four types of CS–US pairings, EC effects on self-reported evaluations were reduced by unreinforced CS presentations, but only when the CSs had been rated after the initial presentation of CS–US pairings. EC effects on an evaluative priming measure remained unaffected by unreinforced CS presentations regardless of whether the CSs had been rated after acquisition. The results suggest that reduced EC effects resulting from unreinforced CS presentations are due to judgement-related processes during the verbal expression of CS evaluations rather than genuine changes in the underlying evaluative representations.     

Skin conductance response conditioning with CS intensities equal to and greater than UCS intensity

With .2-sec bursts of white noise as both conditioned stimulus (CS) and unconditioned stimulus (UCS), conditioning of first-interval skin conductance responses was obtained when the intensity of the CS equaled and exceeded that of the UCS. There was no evidence that second-interval response conditioning occurred. Nonspecific response frequencies were also affected by the variations in stimulus intensity, this raising some question about typical controls employed in SCR conditioning. There was some evidence that second interval responses were suppressed by the intense CS values. It was concluded that the existence of simple conditioning with a CS/UCS intensity ratio equal to or greater than unity was contrary to the Pavlovian proposition that a CS must be biologically less salient than the UCS in order for conditioning to occur. It was noted, however, that the suppression of second-interval responses might indicate that anticipatory CRs which are not confounded with orienting reflexes are prevented from exhibiting a conditioning effect when a high CS/UCS intensity ratio is employed.     

Acquisition and reversal of a discriminated autoshaped response in male and female rats: Effects of long or short and fixed or variable intertrial interval durations

Groups of male and female Wistar rats were exposed to autoshaping procedures in which the duration (50 vs 20 s) and the variablity (fixed vs variable) of the intertrial interval were manipulated. Pellet delivery occurred after one of the levers (CS+) had been presented for 10 s. Presentation of the other lever (CS−) was not associated with pellet delivery. CS+ contacts occurred more frequently and at higher rates when the intertrial interval was 50 s than when it was 20 s, while intertrial interval variability did not play an important role. Males contacted CS+ and CS− more frequently and at a higher rate than did females. Goal-directed activity (nose pokes in feeder tray) occurred more often when the intertrial interval was short than when it was long. Females showed higher goal-directed activity than did males, both during CS presentation and during the intertrial interval. When experimental contingencies were reversed, CS+ contacts decreased and CS− contacts increased for males and females, but the decrease in CS+ contacts occurred more slowly for males than for females. The results of these experiments add to a growing body of literature suggesting the existence of fundamental behavioral differences between the sexes. These differences are partly attributable to the influence of gonadal hormones during development and maturation and partly attributable to the behavioral requirements of the experimental procedures.     

An analysis of second-order autoshaping

Three mechanisms can explain second-order conditioning: (1) The second-order conditioned stimulus (CS2) could activate a representation of the first-order conditioned stimulus (CS1), thereby provoking the conditioned response (CR); The CS2 could enter into an excitatory association with either (2) the representation governing the CR, or (3) with a representation of the reinforcer evoked by the CS1. A series of experiments using second-order autoshaping with birds was performed to examine these possibilities. Following second-order autoshaping, birds’ responding to the CS2 was found to be unaffected by extinction of the CS1, a result interpreted as showing mechanism (1) to be unimportant and implicating either one or both of the other mechanisms. The CS2 was also shown to provide the birds with specific sensory information about the reinforcer, a pattern of results uniquely predicted by mechanism (3). Implications for the understanding of second-order conditioning and for the SOP model (Wagner, 1981) of associative learning are discussed.     

Classical discrimination conditioning of the rabbit's eyelid response using pontine stimulation as a conditioned stimulus

Classical discrimination conditioning of the nictitating membrane/eyelid response was performed on seven rabbits using stimulation of the pontine nuclei or middle cerebellar peduncle as the conditioned stimulus (CS) and an air puff as the unconditioned stimulus (US). The rabbits learned to discriminate between a CS paired with a US and delivered to one pontine nucleus (the CS+) and a CS presented alone and delivered to the contralateral pontine nucleus (the CS-). Subsequent reversal of the discrimination was also achieved when the CS+ and CS- stimulation sites were interchanged. The results are interpreted as support for the idea that essential plasticity for classical eyelid conditioning occurs efferent to the pontine nuclei, possibly in regions of the cerebellum.     

Transfer of the signaling properties of aversive CSs to an instrumental appetitive discrimination

Rats received Pavlovian conditioning in which, based on a shock US, white noise was established for different groups as an aversive CS+, CSo, or CS?. Then, half of each group received the CS contingent upon either the food-reinforced or nonreinforced response in an easy, choice discrimination. Correct responses, i.e., number of reinforcements, to criterion showed that a CS+ for the reinforced response facilitated learning, whereas a CS? retarded learning; conversely, a CS+ for the nonreinforced response retarded learning, whereas a CS? facilitated learning. The same contingency difference occurred with errors to criterion for CS? subjects but was obscured for CS+ subjects apparently by an avoidance-producing effect of the CS+. In support of the generality of the transfer effect, Z-score transforms of the correct-response data showed that the magnitude of transfer was comparable to that obtained in more difficult discriminations. Collectively, the findings indicate that an aversive CS can function as a transformed signal for the presence (CS+) or absence (CS?) of an appetitive reinforcer and that the signal's control of behavior is via a within-chain mediational process.