Second-order conditioning: Different outcomes in fear and eyelid conditioning

Second-order conditioning has been frequently observed with the fear response but not with the eyelid response. The present experiments manipulated the temporal relationship between the second-order and first-order stimulus on second-order conditioning trials. Our results indicated that a trace second-order procedure is not effective with either response system. Second-order fear conditioning was most prominent when the second-order CS terminated at the onset of the first-order CS. This arrangement, however, did not produce second-order eyelid CRs. In eyelid conditioning, the second-order CS appears to inhibit responding to the first-order CS which immediately follows it.     

Second-order sexual conditioning in male Japanese quail (Coturnix japonica)

Second-order conditioning of social approach to a female conspecific in male Japanese quail was investigated in four experiments. Subjects that received paired first- and second-order trials acquired second-order conditioning in both Experiments 1 and 2. In contrast, subjects that received paired first-order but unpaired second-order trials, and subjects that received unpaired first-order but paired second-order trials, did not acquire second-order conditioning. In Experiment 3, subjects for whom the first-order conditioned stimulus was presented in extinction showed second-order conditioning comparable to that shown by subjects in a control group that did not receive the extinction procedure. In Experiment 4, subjects approached a second-order stimulus less when sexually satiated than when sexually deprived. These findings suggest that second-order sexual conditioning in quail is mediated by an association of the second-order stimulus with a representation of the unconditioned stimulus.     

Second-order conditioning of the pigeon’s keypeck

Pigeons learned to peck a keylight (S2) when it was paired with a stimulus (S1) that already evoked keypecking. Control procedures showed that S2 acquired control over responding because it was paired with S1 and because S1 had a conditioning history, thereby supporting the claim that S2 was a second-order conditioned stimulus. Second-order conditioning occurred as rapidly when S1 was a keylight as when it was a tone. Test procedures showed that after second-order conditioning, responding to S2 was markedly debilitated by the extinction of responding to S1, indicating that the ability of S2 to evoke a response importantly depends upon the continued ability of S1 to do so. Our demonstration that directed motor action in the pigeon is susceptible to second-order conditioning suggests a new interpretation of conditioned reinforcement in instrumental learning. Our demonstration that the effectiveness of S2 depends upon the continued effectiveness of S1 indicates that S-S associations are formed in this version of the second-order conditioning experiment.     

Effect of number of trials,interstimulus interval,and dishabituation during CS habituation on subsequent conditioning in a CER paradigm

Three experiments were conducted to investigate the influence of variables during habituation of a CS on a subsequent conditioning to that CS in a conditioned emotional response paradigm. Experiment I varied the number of habituation trials received before conditioning, and it was found that additional habituation trials resulted in a further attenuation of conditioning. Experiment II varied the interstimulus interval of the habituating stimulus, and it was found that the longer intervals produced the greater attenuation in conditioning. Experiment III examined the effect of interpolating another stimulus between habituation and conditioning (dishabituation), and it was found that the dishabituating stimulus augmented subsequent conditioning. It was concluded that manipulations during habituation training that produce the greatest decrement in response to a stimulus, when assessed under equivalent conditions for all animals, have the greatest attenuating effect on subsequent conditioning to that stimulus.     

Second-order conditioning of the rabbit’s nictitating membrane response as a function of the CS2-CS1 and CS1-US intervals

Three experiments investigated conditioning of the rabbit’s nictitating membrane response (NMR) in a second-order conditioning procedure which intermixed first-order trials (CS1-US) and second-order trials (CS2-CS1) from the outset of training. Experiment 1 provided a controlled demonstration that substantial levels of second-order conditioning can be obtained with the NMR preparation. Experiment 2 showed that the level of CR acquisition to CS2 was an inverse function of the CS2-CS1 interval over the values of 400, 800, and 2,400 msec. Experiment 3 found that CR acquisition to CS2 and CS1 in second-order conditioning varied in a parallel fashion across CS1-US intervals. Similarly, Experiment 3A found that the level of CR acquisition to the two components of a serial compound (CSA-CSB-US) varied in a parallel fashion as a function of the CSB-US interval. The results of the CS2-CS1 and CS1-US interval manipulations were all predictable from the known CS-US interval effects in NMR conditioning with a single CS. The present results are discussed with regard to their implications for accounts of serial compound conditioning and second-order conditioning.     

Trial number and compound stimuli temporal relationship as joint determinants of second-order conditioning and conditioned inhibition

Two conditioned lick suppression experiments with rats used feature-negative training (A-footshock trials intermixed with nonreinforced XA presentations) to analyze the role of the number of XA compound presentations and the temporal relationship of the elements within the compound (simultaneous or serial) as determinants of the resulting behavioral control. Second-order conditioning (i.e., excitatory behavioral control by X) was observed to decline as the number of XA compound trials was increased. This decline was more rapid if X and A were presented simultaneously, as opposed to serially (i.e., X before A; Experiment 1). Conditioned inhibition to X, as assessed by a summation test (Experiment 1) and a retardation test (Experiment 2), increased with the number of XA trials and did so more quickly for simultaneous than for serial pairings of X and A. The results help to clarify previously discrepant findings regarding factors that promote excitation versus inhibition with this protocol.     

Summation of undetected excitation following extinction of the CER

The effects of compounding two conditioned stimuli (CSs), each of which had been extinguished to varying degrees in different groups of rats given identical acquisition training, was examined within the conditioned emotional response paradigm. Greater suppression to the compound than to the individual CSs was observed following 6, 12, 48, 72, and 96 stimulus extinction trials, although after only 12 extinction trials suppression to the individual stimuli was no longer observed. The amount of compound suppression decreased progressively as the number of extinction trials increased until, after 120 extinction trials on each stimulus, the compound no longer elicited observable suppression. Control group data indicated that the observed summation effect could not be attributed to disinhibition. The possible role of the summation of undetected excitation in studies examining configurai conditioning, avoidance conditioning and reinstatement is discussed briefly.     

Motivational vs. associative role of the US in classical conditioning of the rabbit’s nictitating membrane response

Two experiments tested the motivational role of the US in classical conditioning of the rabbit’s nictitating membrane (NM) response. In Experiment 1, subjects were trained to an intermediate performance level and then given a series of (1) CS-US trials, (2) “backwards” US-CS trials, (3) CS-alone trials, (4) US-alone trials, or (5) no-stimulus presentations. Interpolated presentations of the US, either alone or in a backwards contingency, tended to produce an impairment of subsequent acquisition. In Experiment 2, subjects were trained with strong or weak US intensity on paired or interpolated trials. US intensity on interpolated trials had only a very small effect, whereas the effect of US intensity on paired trials was quite large. Shifts in paired-trial US intensity produced corresponding shifts in performance, but shifts in the intensity of the interpolated US produced no apparent effect. We conclude that the arousal of motivation is not sufficient to maintain performance in classical NM conditioning.     

Pentobarbital-induced place aversion learning

Pentobarbital is self-administered by rats but has also been reported to produce a conditioned place aversion. Since the self-administration and place preference paradigms both are considered to assess drug reward, we further examined the hedonic properties of pentobarbital, using place conditioning. In Experiment 1, a dose of 15 mg/kg (intraperitoneal) of pentobarbital produced a conditioned place aversion after 4 conditioning trials of various durations (5, 15, 30, or 60 min). Since rats are typically drug experienced in the self-administration paradigm, in Experiments 2 and 3, we examined the effect of drug history on pentobarbital-induced place conditioning. Although preexposure to pentobarbital attenuated the place aversion, it never resulted in a place preference. As has been reported with alcohol, pentobarbital is hedonically aversive in rats, when novel.     

Dominance,the bidirectional hypothesis,and Pavlovian backward conditioning in the US-US paradigm

Based upon considerations raised by Soviet research, the role of relative stimulus intensity, or dominance, in the unconditioned stimulus-unconditioned stimulus (US-US) paradigm was investigated under circumstances presumed favorable to the backward conditioned response (CR). Using the classically conditioned forelimb response of the cat, a brief shock (USD delivered to one forepaw preceded a shock (US2) to the opposite forepaw in paired conditioning fashion; subjects in the control group received explicitly unpaired presentations of the stimuli. Conditioning in both the forward and backward directions was evaluated by the appearance of contralateral CRs on test trials to each of the USs. In Experiment 1, a ratio of the intensities between US1 and US2 of 100:80 was used to create a relative dominance in favor of the backward CR. In addition, to evaluate the suggestion that the appearance of the backward CR is retarded in the Pavlovian paradigm, overtraining was provided to a forward conditioning criterion of 200%. In Experiment 2, the cats were exposed to successive reductions in the intensity of US2 to verify manipulations of dominance reportedly involved in the reactivation of a latent backward CR. Although forward conditioning was readily established to USl, there was no evidence of back-ward conditioning to US2 under any of the conditions.     

Some determinants of second-order conditioning

In a Pavlovian conditioning situation, an initially neutral stimulus may be made excitatory by nonreinforced presentations in compound with an established conditioned excitor [i.e., second-order conditioning (SOC)]. The established excitor may be either a punctate cue or the training context. In four conditioned suppression experiments using rats, we investigated whether SOC phenomena parallel other cue interaction effects. In Experiment 1, we found that the response potential of a target stimulus was directly related to the intertrial interval when SOC was mediated by a punctate cue, and inversely related to the intertrial interval when SOC was mediated by the training context. Experiment 2 demonstrated that punctate- and context-mediated SOC are oppositely affected by posttraining context extinction, and Experiments 3 and 4 demonstrated that context- and punctate-mediated SOC are differentially affected by conditioned stimulus (Experiment 3) and unconditioned stimulus (Experiment 4) preexposure treatments. These findings parallel phenomena in conditioned inhibition and cue competition situations.     

Temporal integration and instrumental conditioned reinforcement

Stimuli associated with primary reinforcement for instrumental behavior are widely believed to acquire the capacity to function as conditioned reinforcers via Pavlovian conditioning. Some Pavlovian conditioning studies suggest that animals learn the important temporal relations between stimuli and integrate such temporal information over separate experiences to form a temporal map. The present experiment examined whether Pavlovian conditioning can establish a positive instrumental conditioned reinforcer through such temporal integration. Two groups of rats received either delay or trace appetitive conditioning in which a neutral stimulus predicted response-independent food deliveries (CS1→US). Both groups then experienced one session of backward second-order conditioning of the training CS1 and a novel CS2 (CS1–CS2 pairing). Finally, the ability of CS2 to function as a conditioned reinforcer for a new instrumental response (leverpressing) was assessed. Consistent with the previous demonstrations of temporal integration in fear conditioning, a CS2 previously trained in a trace-conditioning protocol served as a better instrumental conditioned reinforcer after backward second-order conditioning than did a CS2 previously trained in a delay protocol. These results suggest that an instrumental conditioned reinforcer can be established via temporal integration and raise challenges for existing quantitative accounts of instrumental conditioned reinforcement.     

Second-order conditioning detects unexpressed morphine-induced salt aversion

Morphine failed to condition a salt taste aversion at a dose (15 mg/kg) sufficient to produce a robust aversion to a saccharin taste. Indeed, three different concentrations of salt (1%, 1.5%, and 2%) paired with the same morphine dose yielded no direct evidence for conditioned aversion. Yet, when a novel saccharin taste was paired in compound with the previously conditioned salt conditioned stimulus, we found evidence for a conditioning to the saccharin cue alone in three separate experiments. Control groups eliminated alternative accounts such as neophobia and differential exposure to morphine. Combined, these findings indicate that morphine conditioned a salt aversion. Although this aversion was not directly expressed, a second-order conditioning procedure was able to provide a more sensitive index of conditioning.     

Classically conditioned heart rate in rats following preconditioning exposure to the CS

The influence of preexposure to the CS on classically conditioned heart rate was examined in three groups of rats receiving either 0, 10, or 50 CS-alone trials prior to the beginning of acquisition training. The conditioned response was a deceleration in heart rate for all groups. Compared to the 50 group, the 0 and 10 groups both showed a lower overall level of conditioning performance and a slower rate of development of the conditioned response. It was suggested that the presence of the nonhabituated orienting response may have interfered with the conditioning process.     

Differential conditioned suppression in the Konorski-Lawicka paradigm

In conditioned suppression discriminations, the Konorski-Lawicka paradigm involves A+ trials, where conditioned stimulus A is followed by shock, and sh → A? trials, where A is preceded by shock Rats easily mastered this A+/sh → A? discrimination, as indicated by suppression of food-reinforced barpressing on A+ trials and acceleration of barpressing on sh → A? trials. A history of A+ conditioning resulted in nearly perfect discrimination performance on the very first day of A+/sh → A ? training, but a history of sh→ A? conditioning retarded development of the discrimination. The basis for the development of the discrimination was discussed in terms of an inferred stimulus (sh′) arising from the aftereffects of shock.     

Bidirectional heart rate responses in rats associated with excitatory and inhibitory stimuli

Following classical conditioning to a shock-reinforced tone CS (T₁+), heart rate (HR) of three groups of rats was examined in response to a new reinforced tone (T₂+) and a nonreinforced light (L?) CS given during conditioned inhibition (T₂+ vs. T₁L?), discrimination conditioning (T₂+ vs. L?), or explicitly unpaired (T₂/L? vs. US alone) procedures. Decelerative HR reactions occurred to the reinforced T₂+ and T₂+ CSs. To the respective nonreinforced CSs, the conditioned inhibition group displayed diminished but sizable HR deceleration, the discrimination group showed near-zero responding, and the explicitly upaired group showed HR acceleration. Subsequent reversal conditioning to L was retarded in the conditioned inhibition and explicitly unpaired groups relative to the discrimination group. Group differences on combined-cue [T₂/L?) trials were not found. Both the HR responses during inhibitory training and the reversal-conditioning impairments suggest that inhibition may have been established to L? in the conditioned inhibition and explicitly unpaired groups.     

Conditioning of tentacle lowering in the snail (Helix aspersa): Acquisition, latent inhibition, overshadowing, second-order conditioning, and sensory preconditioning

In a series of related experiments, we studied associative phenomena in snails (Helix aspersa), using the conditioning procedure of tentacle lowering. Experiments 1A and 1B demonstrated a basic conditioning effect in which the pairing of an odor (apple) as the conditioned stimulus (CS) with the opportunity to feed on carrot as the unconditioned stimulus (US) made snails exhibit increased levels of tentacle lowering in the presence of the CS. Experiments 2 and 3 showed that the magnitude of the conditioning was reduced when snails were exposed to the CS prior to the conditioning trial (a latent inhibition effect). Experiment 4 examined the effects produced by pairing a compound CS (apple—pear) with food presentations and demonstrated the existence of an overshadowing effect between the two odors. Experiment 5 revealed that pairing one CS with another previously conditioned stimulus increased tentacle lowering to the new CS (a second-order conditioning effect). Finally, Experiment 6 showed that pairing two odors prior to conditioning of one of them promoted an increase in tentacle lowering in response to the other (a sensory preconditioning effect). The results are discussed in terms of an associative analysis of conditioning and its implications for the study of cognition in invertebrates.     

Signaling and affective functions in Pavlovian conditioning

The present study employed a Pavlovian-instrumental-transfer paradigm to investigate the role of conditioned fear in appetitive discrimination learning. Each of three Pavlovian training procedures was used to establish a conditioned fear excitor (CS+), a “neutral” CS (CSo), and a conditioned fear inhibitor (CS?). Then, the CSs were administered to rats in the three groups contingent upon the rewarded response in a difficult visual discrimination. In addition, half of each group received shock punishment for each incorrect response. Relative to CSo, CS+ facilitated performance in contrast to the usual interfering effect of conditioned suppressors; conversely, CS? retarded performance even when its reinforcing action (fear inhibition) was potentiated by punishment for the incorrect response. These results, together with other findings showing a reversed outcome when the CSs are administered for the incorrect response, indicate that Pavlovian conditioning comprises both general signaling and affective functions, the former reflecting a basic “expectancy” or nominal type of cognitive processing in the rat.     

Circadian-temporal context and latent inhibition of conditioned taste aversion: Effect of restriction in the intake of the conditioned taste stimulus

Latent inhibition of conditioned taste aversion (CTA) is sensitive to changes in the temporal context. A change in the time of day of conditioning with respect to the time of day of the preexposure can disrupt the latent inhibition. This contextual change in the time of day may reveal a temporal specificity of latent inhibition. The optimum procedure to induce this temporal specificity is not well established. For example, it has been shown that a long period of habituation to temporal contexts is one factor that can determine the effect. However, the experimental conditions on the conditioning day that facilitate this phenomenon are unknown. The aim of this study is to elucidate whether a restriction in the intake of the conditioned taste stimulus affects the temporal specificity of latent inhibition. Two main groups of Wistar rats were tested in a latent inhibition of CTA paradigm, in which the temporal specificity of this phenomenon was analyzed by a change in the time of day of conditioning. The intake of the taste stimulus was restricted in the conditioning day in one of the groups, but this restriction was not applied in the other group. The results indicated temporal specificity of latent inhibition only in the group without restriction, but not in the group with limitation in the intake of the taste stimulus during conditioning. These findings can help to elucidate the characteristics of the procedure to induce temporal specificity of latent inhibition.     

Discriminated and nondiscriminated avoidance conditioning of the rearing response in rats

Experiment 1 employed a shock box in which light beams ran at 10, 15, 20, or 25 cm above the floor level of the box. Four groups of nine rats each were trained to avoid shock by cutting the light beams or letting them pass by, which the animal accomplished by upward or downward change of its posture. Training employed a discriminated avoidance paradigm, 60 trials per day for 5 days, with a 5-sec CS-US interval. Acquisition of the rearing avoidance response was observed only in the 15-cm condition. Using the same apparatus as in Experiment 1 and with a beam height of 15 cm, the rearing avoidance response was successfully conditioned in five rats using a nondiscriminated avoidance conditioning paradigm. There was good evidence of temporal discrimination in these animals.