Perseveration of associative strength in rabbit nictitating membrane response conditioning

The present experiment used a transfer-of-training procedure in rabbit nictitating membrane response (NMR) conditioning to determine whether a retention interval and/or extinction could reduce associative strength. The experimental design required that groups receive 0, 3, 15, 45, 150, or 240 CS-US pairings in Stage 1. Next, the groups were given, in succession, a 10-day retention interval and 480 CS-alone trials. In Stage 2, all groups obtained 240 CS-US pairings for NMR conditioning. Another group was also employed and received only the 240 CS-US pairings in Stage 2. The results indicated that 15 to 240 CS-US pairings in Stage 1 substantially enhanced NM CR performance in Stage 2 despite the interpolation of the retention interval and CS-alone trials. When 3 CS-US pairings had been given in Stage 1, no augmentation in the NM CR performance rate occurred in Stage 2. However, the 3 CS-US pairings were effective in prohibiting the 480 CS-alone trials from retarding subsequent NM CR performance. Without any pairings in Stage 1, the 480 CS-alone trials produced strong latent inhibition of NMR conditioning in Stage 2. The data were used to support the theoretical view that associative strength resulting from CS-US pairings is relatively permanent. Moreover, the findings were relevant for an evaluation of Pearce and Hall’s (1980) recent statements concerning CS associability and the relationship between excitatory and inhibitory processes.     

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.     

Effects of an aversive CS+ and CS− under deprivation upon successive classical appetitive and aversive conditioning

Rabbits under high or moderate water deprivation received in Stage 1 either paired (CS+), unpaired (CS?), or no-tone/shock presentations, with the pairings being appropriate for nictitating membrane conditioning. In Stage 2, all groups were given paired tone and water deliveries for jaw-movement conditioning, while, in Stage 3, all group received the tone and shock paired together for membrane conditioning. In Stage 2, the previously established aversive CS+ suppressed jaw-movement conditioning under high deprivation, and membrane CR decrements were directly related to deprivation. Also in Stage 2, the aversive CS? raised jaw-movement conditioning under moderate deprivation. In Stage 3, membrane CR performance immediately returned in the aversive CS+ group. For the other groups, conditioning was faster under high, relative to moderate, deprivation; however, the initial membrane CR occurrence required more trials if unpaired presentations were used in Stage 1. These results suggest that CSs can acquire both opponent-process and associative effects expressed according to the prevailing training conditions.     

Classical-classical transfer: Effects of prior appetitive conditioning upon aversive conditioning in rabbits

An appetitive-ayersive transfer experiment with rabbits determined that prior paired and unpaired tone CS and water US presentations, given in jaw-movement (JM) conditioning, respectively facilitated and retarded the acquisition of the nictitating membrane (NM) CR when the tone was subsequently paired with a shock US. In addition, the unpaired tone and water deliveries reduced the level of JM conditioning that was undertaken following the completion of NM CR acquisition. Finally, the reacquisition of the NM CR was accompanied by a large savings effect in contrast to the failure of the JM CR to display reacquisition savings. When the present findings are compared to the results of previous work addressing the influence of prior NM conditioning procedures upon subsequent JM CR acquisition, an asymmetry in appetitive-aversive interactions is indicated. This asymmetry encourages a reinterpretation of the opponent-process explanation of appetitive-aversive interactions. Moreover, the observed effects of the unpaired CS suggest the operation of a salience factor.     

Evaluation of bidirectional interstimulus interval (ISI) shift in auditory delay eye-blink conditioning in healthy humans

Delay eye-blink conditioning is an associative learning task that can be utilized to probe the functional integrity of the cerebellum and related neural circuits. Typically, a single interstimulus interval (ISI) is utilized, and the amplitude of the conditioned response (CR) is the primary dependent variable. To study the timing of the CR, an ISI shift can be introduced (e.g., shifting the ISI from 350 to 850 ms). In each phase, a conditioned stimulus (e.g., a 400- or 900-ms tone) coterminates with a 50-ms corneal air puff unconditioned stimulus. The ability of a subject to adjust the CR to the changing ISI constitutes a critical timing shift. The feasibility of this procedure was examined in healthy human participants (N = 58) using a bidirectional ISI shift procedure while cortical event-related brain potentials were measured. CR acquisition was faster and the responses better timed when a short ISI was used. After the ISI shift, additional training was necessary to allow asymptotic responding at the new ISI. Interestingly, auditory event-related potentials to the CR were not associated with conditioning measures at either ISI.     

Incentive contrast produced by deprivation shifts

Deprivation shifts, using conditions analogous to those which produce strong incentive contrast when reward quantity is changed (extended preshift training, short interval from the last preshift trial to the first postshift trial, large preshift differences in deprivation) resulted in contrast of runway speed and choice behavior. In the first experiment, a downshift of hunger during rats’ runway training produced a slow lessening of speed below that of a group trained continually at low hunger. In the second experiment, rats were trained to traverse a runway to one food-containing goalbox when very hungry and to another, distinctively different food-containing goalbox when not very hungry. The rats were next given a series of choice trials between the two goalboxes. There was a brief preference for the high-hunger goalbox, followed by a preference for the low-hunger goalbox. The results of the second experiment suggest that deprivation affects the strength of conditioning of a cue-reinforcer expectancy, while the slow development of contrast in the first experiment indicates that deprivation also affects the development of either habit strength or a response-reinforcer expectancy.     

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.     

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.     

General transfer across sensory modalities survives reductions in the original conditioned reflex in the rabbit

Two experiments demonstrated that transfer of training between CSs from different sensory modalities survived substantial reductions in responding to the first CS. In both experiments, animals received three stages of training. Stage 1 entailed CS-US training with a CS from one modality (e.g., light), and Stage 3 entailed CS-US training with a CS from another modality (e.g., tone). The experiments differed in treatment during Stage 2. In Experiment 1, animals either remained in their home cages or received unreinforced exposures to the first CS, which extinguished the original CR. In Experiment 2, the animals received either continued CS-US training or exposure to the CS and US but at a long interval (2,800 msec), which eliminated the original CR. As the baseline for detection of transfer effects, each experimental group had a control group that received Stage 1 training with a 2,800-msec CS-US interval, which produced minimal CR acquisition. The results of both experiments revealed substantial positive transfer across CS modalities regardless of the treatment during Stage 2. The transfer did not appear immediately on test presentations of the second CS in Stage 3. Rather, the transfer appeared as an enhancement in the rate of CR acquisition after reinforced training with the second CS had commenced. The results are discussed with respect to stimulus generalization, neutralization of background stimuli, and learning processes superordinate to specific associations.     

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.     

Conditioning of the nictitating membrane response of the rabbit (Oryctolagus cuniculus) as a function of length and degree of variation of intertrial interval

The study involved three experiments. The first, a parametric investigation of nictitating membrane conditioning with eight constant intertrial intervals (ITIs) between 5 and 120 sec, orthogonal to interstimulus intervals (ISIs) of 250 and 750 msec plus three temporal conditioning control groups, revealed that performance improved rapidly with increasing ITI but stabilized at relatively low ITI values. At 750-msec ISI, a decrement in performance was found at 60-sec ITI. Experiment II, using constant ITIs of 45–75 sec in 5-sec steps, at 750-msec ISI confirmed the trend toward a performance decrement around 60 sec, although the trend was weak and highly variable. Experiment III evaluated the differences in performance between constant and variable ITI, using three ITI values and three conditions of variation at each value. Findings were discussed in terms of differences in conditioning resulting from both length and degree of variation of ITI and some subtle effects which may emerge only when constant ITIs are used.     

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.     

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.     

Concurrent development of excitatory and inhibitory associations during backward conditioning

After backward pairings with a paraorbital shock, a tone CS was an effective punisher of licking in rabbits, yet yielded retarded forward acquisition of the rabbit’s nictitating membrane response. The observation of both excitatory and inhibitory associative effects after a common training protocol challenges the assumption of a unidimensional associative continuum and supports approaches that assert a multidimensional structure to associations.     

The role of number and spacing of CS-US pairings in conditioned suppression

In four studies, with rats as Ss. acquisition and. where appropriate, extinction trials were presented against a baseline of ongoing licking. At shock intensities of 0.1. 0.5. 1.0, or 2.0 mA, acquisition performance was a function of number of CS-US pairings: spacing of trials (one or two per day) did not affect acquisition performance. Resistance to extinction could be predicted from terminal acquisition performance and reached a maximum after three CS-US pairings.     

Proactive interference of open field on consummatory successive negative contrast

Reactivity to a reward is affected by prior experience with the different reinforcer values of that reward, a phenomenon known as incentive relativity, which can be studied using the consummatory succesive negative contrast (cSNC) paradigm, in which the performance of animals that receive a 4 % sucrose solution after trials on which they were exposed to 32 % sucrose is compared with that of subjects that always receive the 4 % sucrose solution. The exploration of a novel open field can enhance or block the acquisition of associative and nonassociative memories. The effect of open field on cSNC has not yet been explored. The main result of the present study was that open-field exposure significantly modified the expression of cSNC. Exposure to an open field 1 h but not immediately before the downshift interfered with the expression of cSNC. These animals drank more of the downshifted reward than did controls that were not exposed to the apparatus, and this behavior persisted for up to three recovery trials. This phenomenon was observed even when the animals were given a more protracted preshift phase and when the discrepancy between the preshift and shift incentive values of sucrose were increased. An open field also interfered with incentive downshift when open-field exposure occurred 6 h before the downshift, and repeated exposure to the apparatus did not deteriorate this effect. The present study adds to a growing body of literature that indicates that open-field exploration can interfere with memory formation.     

Pain-elicited aggression in the squirrel monkey: An implicit avoidance contingency

Squirrel monkeys were given either forward pairings of a bite-tube CS and shock US or backward pairings of these stimuli. Backward pairings produced stronger control of biting by the bite tube alone than did forward pairings. In a second experiment, subjects received backward pairings of US and CS with either a fixed ITI or a random ITI. Conditioned biting was obtained only when trials were presented with a fixed ITI. The magnitude of unconditioned biting was also significantly greater with the fixed ITI. It was argued that these results demonstrate that conditioning in this situation depends upon the degree to which biting predicts a relatively long shock-free period. When trials occur randomly in time, biting predicts no definite shock-free period; hence, it is not learned.     

One-trial context fear conditioning as a function of the interstimulus interval

In two experiments, we examined the effects of a wide range of interstimulus intervals (2.5, 15, 45, 120, 135, and 405 sec) on one-trial context fear conditioning with rats. Here, the interstimulus interval (ISI) denotes the time between placement in a conditioning chamber and the onset of a single footshock. On the conditioning day, we observed that the rats’ behavior at the time of shock onset varied systematically across ISI values. On the subsequent test day, we used context-evoked freezing as a measure of context conditioning and found the well-known inverted U-shaped ISI function. We also found that conditioned freezing for the shortest ISI values was concentrated early in the test session, whereas freezing at longer ISIs was distributed more evenly throughout the test session. The freezing results found here are more consistent with the literature on conditioning with punctate cues than are previously described results from one-trial context fear-conditioning procedures.     

Retardation of rabbit nictitating membrane conditioning following US preexposures depends on the distribution and number of US presentations

Retarded conditioned response (CR) acquisition produced by unconditioned stimulus (US) preexposures has been attributed either to interference resulting from contextual conditioning or to habituation of the US. Both perspectives assume that the amount of retardation is directly related to the number of US preexposures. This assumption was examined in two experiments. In Experiment 1, separate groups of rabbits received 200 paraorbital shock US preexposures either in one session or spread equally over 10 daily sessions. Subsequently, all subjects received 150 CS-US pairings. Acquisition of nictitating membrane CRs was retarded relative to a naive control group only in the group that received the preexposures over 10 sessions. Thus, the number of US preexposure sessions, rather than the number of US preexposures, determined whether or not retarded acquisition was observed. In Experiment 2, four groups of rabbits received 1, 5, 20, or 40 shock US preexposures in each of 10 sessions. Over the subsequent 150 CS-US pairings, similar levels of retarded CR acquisition were observed in groups that received 20 and 40 US preexposures per session, a weak retardation effect was observed with 5 preexposures per session, and no retardation was observed with 1 preexposure per session. Thus, Experiment 2 suggested that retarded CR development was not greatly influenced by increasing the number of US preexposures above some minimum threshold number of exposures per session. Implications for current theories were discussed.     

Effects of a backward conditioning procedure following acquisition on extinction of conditioned suppression

A one-trial-a-day procedure was used to investigate the effects of US-CS pairings on extinction of conditioned suppression of licking by rats. Following acquisition trials, response suppression was immediately eliminated when US preceded CS, but it reappeared during subsequent CS-alone presentations. Ss that received backward pairings reached a significant level of extinction one trial before Ss that received conventional extinction trials.