Second-order conditioning of the rabbit's nictitating membrane response. Interstimulus interval and frequency of CS-CS pairings.

Second-order conditioning of the rabbit's nictitating membrane response (NMR) was investigated when second-order trials (CS1-CS2) were intermixed with first-order trials (CS2-US) from the outset of training. Experiment 1 showed that CR acquisition to CS1 was inversely related to the CS1-CS2 interval but nevertheless extended to an interval of 8,400 ms. Experiment 2 revealed that CR acquisition of CS1 was an inverted-U function of the number of CS1-CS2 trials relative to a fixed number of CS2-US trials. Experiment 3 directly contrasted second-order conditioning with a reinforced serial compound procedure (CS1-CS2-US) and a mixed procedure in which second-order trials were intermixed with the reinforced serial compound. Second-order conditioning was about half the strength of either the reinforced serial compound or the mixed procedure, which were similar. The present results are discussed with respect to the relative strength of excitatory and inhibitory processes in second-order conditioning.     

Temporal discrimination using different feature--target intervals in classical conditioning of the rabbit's nictitating membrane response

In a typical conditional discrimination, a target stimulus (X) is reinforced during one feature cue (A-->X+), but not during another feature cue (B-->X-). The present experiments used only a single "feature" cue (a 66-sec tone). On half of the trials, the target stimulus (a 400-msec light) was paired with the reinforcer when the feature-target interval was one duration (e.g., 5 sec). On the remaining trials, the interval was different (e.g., 45 sec), and the target stimulus was presented without the reinforcer. All the animals acquired this temporal discrimination, and subsequent testing with other feature-target intervals yielded generalization-like gradients. These results provide solid evidence that each portion of a feature cue is encoded in a distinctive fashion. Had temporal encoding not occurred, the feature cue would have been just as ambiguous a predictor of the reinforcer as was the target stimulus, and discrimination would not have been possible. The integration of real-time temporal encoding mechanisms into models of conditional discrimination is 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.     

Extinction revisited: similarities between extinction and reductions in US intensity in classical conditioning of the rabbit's nictitating membrane response

The mechanisms of extinction were examined by reducing the intensity of the unconditioned stimulus (US) after acquisition training to determine whether such reductions lie on a continuum with CS-alone extinction. The experiments revealed that reductions in US intensity yielded extinction-like effects. Specifically, there were proportional reductions in the daily mean level of responding across sessions. There were also persistent within-session declines and between-session increases of responding analogous to spontaneous recovery. Surprisingly, even when US intensity was held constant, within-session declines and between-session increases were apparent. The results are discussed with respect to possible contributions from unlearning, new learning, generalization decrement, and nonassociative loss, especially CS-specific attentional changes and CR-specific reactive inhibition.     

Role of the hippocampus in blocking and conditioned inhibition of the rabbit's nictitating membrane response.

Bilateral aspiration of the dorsal hippocampus produced a disrupttion of blocking of the rabbit's nictitating membrane response in Kamin's two-stage paradigm (Experiment 1) but had no effect on the formation of a Pavlovian conditioned inhibitor (Experiment 2). The results of Experiment 1 indicated that normal animals and those with cortical lesions given conditioning to a light-plus-tone conditioned stimulus (CS) gave conditioned responses (CRs) to both the light and the tone during nonreinforced presentations of each (test phase). If, however, compound conditioning was preceded by tone acquisition, only the tone elicited a CR during testing; that is, blocking was observed. In rabbits with hippocampal lesions, however, CRs were given to both the light and the tone during testing whether or not compound conditioning was preceded by tone acquisition. The data from Experiment 2 showed that rabbits with hippocampal lesions could discriminate as well as normal rabbits and those with cortical lesions between a light (CS+) and light plus tone (CS-). In addition, when the inhibitory tone was subsequently paired with the unconditioned stimulus in retardation testing, animals in all three lesion conditions acquired the CR at the same rate. Thus, it appears that hippocampal lesions do not disrupt conditioned inhibition. The results of these experiments were taken as support for the view that the hippocampus is responsible for "tuning out" stimuli that have no adaptive value to the organism.     

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.     

Conditioning the unconditioned response: modification of the rabbit's (Oryctolagus cuniculus) unconditioned nictitating membrane response

Conditioning-specific reflex modification (CRM) occurs when classical conditioning modifies responding to an unconditioned stimulus (US) in the absence of a conditioned stimulus (CS). Three experiments monitored rabbit nictitating (Oryctolagus cuniculus) membrane unconditioned responses to 5 intensities and 4 durations of periorbital electrical stimulation before and after CS or US manipulation. CRM occurred after 12 days of CS-US pairings but not following unpaired CS/US presentations or restraint. CRM survived CS-alone and CS/US-unpaired extinction of the conditioned response (CR) but not presentations of the US alone, although CRs remained intact. Thus, CRs could be weakened without eliminating CRM and CRM could be weakened without eliminating CRs. Data indicate CRM is a reliable, associative effect that is more than a generalized CR and may not be explained by habituation, stimulus generalization, contextual conditioning, or bidirectional conditioning.     

Repeated acquisitions and extinctions in classical conditioning of the rabbit nictitating membrane response

The rabbit nictitating membrane (NM) response underwent successive stages of acquisition and extinction training in both delay (Experiment 1) and trace (Experiment 2) classical conditioning. In both cases, successive acquisitions became progressively faster, although the largest, most reliable acceleration occurred between the first and second acquisition. Successive extinctions were similar in rate. The results challenge contextual control theories of extinction but are consistent with attentional and layered-network models. The results are discussed with respect to their implications for the interaction between cerebellar and forebrain pathways for eyeblink conditioning.     

Conditioning of the rabbit's nictitating membrane response to a CSA-CSB-US serial compound: manipulations of CSB's associative character.

Response acquisition to a trace conditioned stimulus (CSA) can be facilitated by insertion of a second stimulus (CSB) at the end of the trace interval just before the unconditioned stimulus (US). This effect may arise from serial mediation of trace conditioning, second-order conditioning, or both. Whereas serial mediation relies only on the presence of CSB, associative transfer relies on CSB's associative strength. In the present experiments, the presence of CSB was fixed, whereas CSB's associative strength was manipulated by (a) extinction of CSB, (b) latent inhibition of CSB, and (c) prior CSB-US pairings. In the first 2 cases, the level of responding to CSA was reduced in a fashion parallel to that of CSB. However, in the third case, partial blocking of conditioned response (CR) acquisition to CSA was observed. The results are discussed with reference to the role of associative transfer to both facilitating and blocking CR acquisition to CSA.     

Heart rate changes during conditioning-specific reflex modification of the rabbit's (Oryctolagus cuniculus) nictitating membrane response

Conditioning-specific reflex modification (CRM) of the rabbit's nictitating membrane response (NMR) involves changes in responding to an unconditioned stimulus (US) when the US is tested in the absence of the conditioned stimulus. Previous experiments have shown that CRM is a function of the type and intensity of the aversive US used during classical conditioning. As a result, it has been suggested that CRM may be mediated, at least in part, by the aversiveness of the US. Here, we show that by using a moderately intense electrical pulse to the skin as a US, CRM of the rabbit NMR is accompanied by an increase in heart rate. The largest changes in heart rate occur at US intensities that produce the strongest levels of CRM. The heart rate data show that there may be an increased emotional/arousal component to the US that is correlated with CRM and support the use of CRM as a potential model for posttraumatic stress disorder.     

Classical conditioning of the nictitating membrane response of the lemon shark (Negaprion brevirostris)

An experimental group of lemon sharks received 100 daily presentations of light flash as the conditioned stimulus (CS) and electric shock as the unconditioned stimulus (US) in a classical conditioning situation. The conditioned responses (CRs) and unconditioned responses (URs) under observation consisted of extensions of the nictitating membrane. Separate control groups received either (a) no CS or US, (b) CS-alone, or (c) completely random presentations of CS and US. Few CRs occurred in the experimental group at the outset of conditioning, but the percentage of CRs during the second half of the first acquisition session exceeded 95%. Conditioning stabilized above 95% CRs during Acquisition Sessions 3 through 7. These responses could not be attributed to pseudoconditioning, sensitization, or other nonassociative factors. When the experimental group was subsequently given six CS-alone sessions, the course of extinction was gradual. Most results seemed similar to those previously obtained during classical conditioning of the nictitating membrane in rabbits.     

The CS-US interval function in rabbit nictitating membrane response conditioning: Single vs multiple trials per conditioning session

The effect of varying trials per day conditions on the CS-US interval or interstimulus interval (ISI) function in rabbit nictitating membrane response conditioning was studied in two experiments. Experiment 1 showed that a 1250-msec ISI was more effective than a 250-msec ISI when there was 1 trial/day. Experiment 2 showed that as the number of trials per day decreased from 20 to 1, the superiority of the 250-msec ISI group over the 1250-msec ISI group declined, with a reversal at 1 trial/day. Results are interpreted in terms of the role of a hypothesized CS-elicited short-duration orienting response in CR performance.     

Effect of preacquisition UCS exposure on classical conditioning of the rabbit's nictitating membrane response

Conditioning of the rabbit nictitating membrane response was found to be inversely related to the number (0, 50, 200, 350) and intensity, (0, 1, 3, 5 mA) of preacquisition UCS exposures and directly related to the interval between preexposure and acquisition (0 or 24 hr). The results were discussed in terms of sensory adaptation, habituation, blocking, and emotional reactivity.     

Classical conditioning of the rabbit's nictitating membrane response under fixed and mixed CS-US intervals

The rabbit's nictitating membrane response was classically conditioned to a tone CS, followed by an electric shock US, at two randomly alternating CS-US interstimulus intervals (ISIs). Different groups were exposed to different proportions of 200- and 700-msec ISIs. The study revealed: (a) CR percentage was a direct function of the proportion of 200-msec ISIs; (b) trials to the first test-trial CR were an inverse function of the proportion of 200-msec ISIs; (c) CR onset latencies decreased over training; (d) CR peak latencies coincided with the temporal loci of the US; and (e) groups exposed to both ISIs acquired double-peaked CRs. Moreover, the amplitudes of nictitating membrane responses in the present experiment paralleled certain quantitative aspects of the free operant key peck response rates of pigeons trained under two mixed inter-reinforcement intervals. The results were discussed with respect to the micromolar response shaping and macromolecular CS-trace accounts of classical conditioning.