Independent effects of stimulus and cycle duration in conditioning: The role of timing processes

Rats received delay conditioning procedures with a white-noise conditioned stimulus (CS), a food unconditioned stimulus (US), and head entries into the food cup as the conditioned response. The stimulus duration (S) and the interval between food deliveries (C) were varied between groups:S=15, 30, 60, and 120 sec;C=90, 180, and 360 sec. The stimulus/cycle duration ratio was negatively related to the asymptotic level of conditioning but had no effect on the rate of acquisition. Conditioning and timing of responses emerged together in training. Timing occurred during the CS-US interval (ISI) and the US-US interval (ITI), as evidenced by increasing response rate gradients that were steeper for shorter intervals. The effects of the stimulus/cycle ratio on conditioning were attributed to independent timing of theS andC durations. Serial-, parallel-, and single-process accounts of conditioning and timing are compared.     

Comparison of auditory and visual conditioning stimuli in delay eyeblink conditioning in healthy young adults

Classical eyeblink conditioning (EBC) has been widely used to probe cerebellar function in humans and nonhuman mammals. Although the neural pathways governing behavior in this task are well understood and fairly discrete, it remains unclear in the human literature how conditioned stimuli (CSs) of different modalities (e.g., visual and auditory) influence the exhibition of conditioned responses (CRs). In the present study, therefore, CRs to a visual CS and an auditory CS were examined with the single-cue delay EBC procedure. An initial experiment (N=61) was conducted to identify visual and auditory stimuli that had equal perceived intensities. Using these perceptually equivalent stimuli, a second group of 25 subjects completed auditory and visual EBC procedures in two testing sessions 5–8 days apart. Whereas the acquisition of CRs was similar between the CS modality conditions, the timing of the CRs differed such that earlier CR onset and peak latencies were associated with the visual CS. In addition, CR timing improved across testing sessions, as indicated by the later CR peak latencies exhibited during the second testing session, as compared with the first.     

Temporal encoding in trace conditioning

Conditioned lick suppression in rats was used to explore the role of timing in trace conditioning. In Experiment 1, two groups of rats were exposed to pairings of a CS (CS1) with a US, under conditions in which the interstimulus interval (ISI) that separated CS1 offset and US onset was either 0 or 5 sec. Two additional groups were also exposed to the same CS1→US pairings with either a 0 or a 5-sec ISI, and then received “backward” second-order conditioning in which CS1 was immediately followed by a novel CS2 (i.e., CS1→CS2). A trace conditioning deficit was observed in that the CS1 conditioned with the 5-sec gap supported less excitatory responding than the CS1 conditioned with the 0-sec gap. However, CS2 elicited more conditioned responding in the group trained with the 5-sec CS1-US gap than in the group trained with the 0-sec CS1-US gap. Thus, the CS1-US interval had inverse effects on first- and second-order conditioned responding. Experiment 2 was conducted as a sensory preconditioning analogue to Experiment 1. In Experiment 2, rats received the CS1?CS2 pairings prior to the CS1→US pairings (in which CS1 was again conditioned with either a 0 or a 5-sec ISI). Experiment 2 showed a dissociation between first- and second-order conditioned responding similar to that observed in Experiment 1. These outcomes are not compatible with the view that differences in responding to CSs conditioned with different ISIs are mediated exclusively by differences in associative value. The results are discussed in the framework of the temporal coding hypothesis, according to which temporal relationships between events are encoded in elementary associations.     

The CS-US interval (ISI) function in rabbit nictitating membrane response conditioning with very long intertriai intervals

In Experiment 1, four values of conditioned stimulus-unconditioned stimulus interval, or interstimulus interval (ISI) (200, 500, 800, and 1,100 msec) were studied in one trial/day acquisition of the conditioned nictitating membrane response of the rabbit. The 1,100-msec ISI group was superior to the other groups, contrary to the usual findings of ISI studies in the conditioning literature. In Experiment 2, effective conditioning levels were attained with an ISI as long as 2,200 msec, when the intertriai interval was set at either 24 or 48 h. Results are interpreted in terms of the role of orienting responses in the conditioning process.     

Excitatory and inhibitory consequences of explicitly unpaired and truly random conditioning procedures on heart rate in rats

A comparison was made of heart-rate (HR) responses of restrained rats to CSs that were part of an explicitly unpaired or a truly random control procedure. Subsequent to these procedures, an assessment was made of the relative capacities of these CSs to affect an established HR CR in a combined-cue paradigm and to impede the development of a HR CR in a reversal-conditioning situation. The principal findings were (1) that the explicitly unpaired and truly random CSs generated HR responses of opposite direction, i.e., HR acceleration vs. HR deceleration, respectively, and (2) that conditioning of a decelerative HR CR to the CS that had earlier been employed in the explicitly unpaired procedure was retarded compared to what was obtained to the truly random CS. The two CSs did not have reliably different effects in the combined-cue test. It was suggested that the truly random CS may have produced both associative and nonassociative influences on HR. It was hypothesized further than the explicitly unpaired CS may have acquired the capacity to function as a conditioned inhibiting stimulus.     

Trace and long-delay fear conditioning in the developing rat

In two experiments with rats, we examined the developmental emergence of conditioned freezing following trace and short-delay conditioning and also included a long-delay comparison group. In the short-delay and trace groups, a 10-sec conditioned stimulus (CS) was paired with shock; for the trace rats, a 10-sec trace interval followed CS termination. The long-delay groups received a 20-sec CS paired with shock, to equate for the longer interstimulus interval (ISI) in the trace group. Trace conditioning emerged later in development than did short-delay conditioning (see Moye & Rudy, 1987). Importantly, long-delay conditioning emerged in parallel with trace conditioning, at a similar time, and with similar strength. These findings suggest a role for the longer ISI, as opposed to the unfilled gap per se, in the late emergence of trace conditioning. The role of the hippocampus in trace conditioning and the possibility that young rats encode the temporal relationship between CSs and unconditioned stimuli are also considered.     

Recovery of conditioned suppression after backward pairings

In four conditioned suppression experiments with rats (Rattus norvegicus), backward pairings of a shock unconditioned stimulus (US) and a tone conditioned stimulus (CS) eliminated an already established conditioned response (CR), but there was recovery of the CR if the shock was later withheld. In Experiment 1, there was recovery after backward pairings, regardless of whether the period after the US was normally shock free or not. In Experiment 2, the occurrence of recovery depended on the CS’s being presented closely after the US in response elimination. Levels of recovery were positively correlated with the resistance of the response to elimination during backward pairings (Experiments 3 and 4). Taken together, these data support the notion that recovery after backward pairings is a form of renewal (see, e.g., Bouton, 1991) and is not due toprotection from extinction.     

Frustrative nonrelief in instrumental escape conditioning

The generality of the frustration effect in an aversive stimulus conditioning procedure was examined by training and testing 40 rats in a double cold-waterway escape conditioning apparatus. Experimental and control procedures analogous to appetitive conditioning experiments (e.g., Amsel & Roussel, 1952; Wagner. 1959) indicated that frustrative nonrelief (i.e., reinforcement omission) in the first goal tank yielded significant increments in swimming speed in the second waterway, and that these increments in performance were dependent upon initial training with continuous relief (i.e., reinforcement) in the first goal tank. Extensions of the generality of the frustration effect are discussed.     

A Pavlovian analysis of food-attraction conditioning in the snailHelix aspersa

Following brief pairing of an odor with a feeding experience (food-attraction conditioning), snails will lower their tentacles when subsequently presented with that odor alone. Three experiments investigated the possible behavioral mechanism mediating food-attraction conditioning in the snailHelix aspersa. It is suggested that food-attraction conditioning is an example of Pavlovian conditioning. In this case, the odor (conditioned stimulus) is paired with oral stimulation (unconditioned stimulus), which elicits lowering of the tentacles (unconditioned response). Following conditioning, the odor comes to elicit lowering of the tentacles (conditioned response). Experiment 1 ruled out nonassociative effects, such as habituation and sensitization, using an unpaired control group. Experiment 2 provided further evidence against a role for habituation of neophobia, through the demonstration of extinction following conditioning. In Experiment 3, an omission procedure was used to rule out the possible role of instrumental contingencies.     

A surprising extension of the preconditioned stimulus beyond the cotermination of the US and the added element does not alleviate blocking to the added element

Rats were subjected to the Kamin two-stage blocking procedure. First, a stimulus, A, was conditioned and then reinforced in compound with a target stimulus, B. During compound training, an attempt was made to alleviate blocking of conditioning to B by presenting a “surprise” stimulus for 5 sec following the reinforced AB compound. The surprise stimulus consisted of the continued presentation of the previously reinforced element, A. During A training, A had never extended beyond the moment of reinforcement; thus, its extension beyond reinforcement during AB training was hypothesized to be “surprising” and, therefore, expected to alleviate blocking. Blocking in this condition was, however, just as strong as in a standard blocking condition. The results do not favor a surprise interpretation of unblocking but do seem to be consistent with other theoretical views (e.g., Rescorla & Wagner, 1972).     

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.     

Overshadowing and CS duration: counteraction and a reexamination of the role of within-compound associations in cue competition

In the present experiments, we examined the role of within-compound associations in the interaction of the overshadowing procedure with conditioned stimulus (CS) duration, using a conditioned suppression procedure with rats. In Experiment 1, we found that, with elemental reinforced training, conditioned suppression to the target stimulus decreased as CS duration increased (i.e., the CS duration effect), whereas, with compound reinforced training (i.e., the overshadowing procedure), conditioned suppression to the target stimulus increased as CS duration increased. In subsequent experiments, we replicated these findings with sensory preconditioning and demonstrated that extinction of the overshadowing stimulus results in retrospective revaluation with short CSs and in mediated extinction with long CSs. These results highlight the role of the duration of the stimulus in behavioral control. Moreover, these results illuminate one cause (the CS duration) of whether retrospective revaluation or mediated extinction will be observed.     

Outcome-specific conditioned inhibition in Pavlovian backward conditioning

In the present experiments, the outcome specificity of learning was explored in an appetitive Pavlovian backward conditioning procedure with rats. The rats initially were administered Pavlovian backward training with two qualitatively different unconditioned stimulus conditioned-stimulus (US-CS) pairs of stimuli (e.g., pellet --> noise or sucrose --> light), and then the effects of this training were assessed in Pavlovian-to-instrumental transfer (Experiment 1) and retardation-of-learning (Experiment 2) tests. In the transfer test, it was shown that during the last 10-sec interval, the CSs selectively reduced the rate of the instrumental responses with which they shared a US, relative to the instrumental responses with which they did not share a US. The opposite result was obtained when the USs (in the absence of the CSs) were presented noncontingently. In the retardation test, conditioned magazine approach, responding to the CSs was acquired more slowly when the stimulus-outcome combinations in the backward and the forward conditioning phases were the same, as compared with when they were reversed. These results are collectively in accord with the view that Pavlovian backward conditioning can result in the formation of outcome-specific inhibitory associations. Alternative views of backward conditioning are also examined.     

Effects of US parameters on classical conditioning of cat hindlimb flexion

Unconditioned stimulus (US) intensity and duration were manipulated to determine their effects on cat hindlimb flexion conditioning. Seven consecutive days of acquisition training of a hindlimb flexor response to a tone conditioned stimulus (CS) were followed by 2 days of extinction. Eight animals in each of 12 groups received a 1-, 2-, 3-, or 4-mA, 60-Hz shock delivered to the right hindleg for 25, 50, or 100 msec as the US. Analysis of conditioned-response (CR) frequency indicated that conditioned responding was a positive function of both the intensity and duration of shock, although these variables did not interact with one another. CR latency and amplitude were decreased and increased, respectively, by increases in US intensity. The pattern of results reported here may support a contiguity notion of conditioning, and are discussed in the context of other conditioning preparations.     

Rabbit nictitating membrane conditioning: Lower limit of the effective interstimulus interval

In three experiments, the nictitating membrane response of rabbits was conditioned for 10 daily sessions at interstimulus intervals (ISIs) ranging from 48 to 125 msec, followed by a shift to 250 msec for 5 days. At tested ISIs shorter than 67 msec, there was no evidence of conditioning, and postshift performance revealed neither facilitation nor interference as a result of the first 10 conditioning sessions. Postshift performance of groups conditioned at preshift ISIs of 67 msec or longer revealed a gradient of increasing savings with increasing ISI. One of the groups in Experiment 1, initially conditioned at 250 msec ISI and then shifted to 48 msec, exhibited extinction of the previously well-conditioned response. Analysis of CR-onset latencies substantiated the absence of associative effects at very short ISIs. It was concluded that there is a temporal limit below which classical conditioning of the nictitating membrane response of rabbits employing forward CS-US pairing does not occur.     

The role of signaled periods of nonreinforcement in responding on a random schedule in autoshaping

Little responding develops to a conditioned stimulus (CS) that is placed in a random relation to an unconditioned stimulus (US). However, if the USs not preceded by that CS are themselves signaled by another stimulus, then the CS does come to elicit responding. This result has been attributed (e.g., by Durlach, 1983) to the signal’s blocking of conditioning to background cues that otherwise would prevent conditioning of the CS. However, Goddard and Jenkins (1987) have suggested the alternative that signaling the USs promotes responding due to the adventitious creation of periods of signaled nonreinforcement. Two experiments were conducted to assess this alternative, involving an autoshaping preparation in pigeons. In Experiment 1, little responding to a keylight CS presented in a random relation to a food US occurred, despite the explicit presentation of a discrete noise signaling periods of no food in the intertrial interval (ITI). Experiment 2 was designed to replicate the procedure of Goddard and Jenkins, in which an auditory stimulus extended throughout the ITI of a random schedule, terminating only prior to extra USs and during the CS. Contrary to their findings, little responding developed to the target CS. However, responding did develop when the sound-free period occurred only prior to the extra USs. These results offer little support for the hypothesis that signaled periods of nonreinforcement promote responding on random schedules. However, they are consistent with the view that signaling of ITI USs acts by preventing conditioning of potentially competitive background cues.     

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 control during maintenance and extinction of conditioned keypecking in ring doves

Timing of conditioned keypecking was investigated using a delay autoshaping procedure. In two experiments, the CS-US interval (ISI) was varied and the temporal pattern of responding during unreinforced probe trials extending beyond the reinforced ISI was recorded. Both experiments showed temporal control of keypecking at ISIs of 4, 8, and 16 sec, regardless of whether the probe duration was held constant (Experiment 1) or covaried with the ISI (Experiment 2). Analyses showed that the timing of keypecking was scalar. Increased responding near the end of the probe was due possibly to independent timing of the probe duration. In a third experiment, a group of subjects trained at an ISI of 8 sec were extinguished by presentation of only probe trials. Although the maximal response rate declined progressively, timing of keypecking did not change.     

Memory priming and trial spacing effects in Pavlovian learning

Conditioning trials that are massed in time produce less conditioning than those that are spaced in time. Four experiments with rat subjects examined whether a recent conditioning trial interferes with conditioning on the next trial by temporarily “priming” information in short-term memory (e.g., Wagner, 1978, 1981). We used appetitive conditioning procedures in which priming trials preceded target trials by 60 sec. When the priming trials were nonreinforced presentations of a conditioned stimulus (CS), the CS had to be the same CS as the one on the target trial to interfere with conditioning. When priming trials were actual CS-unconditioned stimulus (US) pairings, the CS identity did not matter; the US was the event that interfered with conditioning on the next trial. Reinforced trials reduced performance in a way that did not depend on context blocking. The results suggest that CS and US priming effects do contribute to conditioning deficits observed with massed trial procedures. The results are consistent with Wagner’s (1981) “sometimes opponent process,” or SOP, model, although a result that is paradoxical for the model suggests that recent USs may have motivational as well as memory effects.     

One-trial simultaneous and backward fear conditioning as reflected in conditioned suppression of licking in rats

In three experiments, groups of albino rats received one strictly simultaneous pairing of a 4-sec auditory conditioned stimulus (CS) and a 4-sec 1-mA shock unconditioned stimulus (US). Other groups received a backward pairing, in which the US began before the CS, or a forward pairing, in which the CS began before the US. Control groups received only the US or received both the CS and the US but widely separated in time. Later, the CS was presented while the rats licked a drinking tube for water, and CS-elicited suppression of licking was taken as an index of the Pavlovian conditioned response (CR). It was found that groups receiving a single forward or a single simultaneous pairing suppressed more than groups that had received a backward pairing; and the backward groups, in turn, suppressed more than the control groups. It appears, then, that excitatory fear conditioning, as reflected in conditioned suppression of licking in rats, can be produced in a single trial by both backward and simultaneous conditioning procedures.