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.     

Consequences of response-contingent change in unconditioned stimulus intensity upon the rabbit (Oryctolagus cuniculus) nictitating membrane response.

In a between-groups design, an instrumental contingency was superimposed on classical conditioning of the rabbit nictitating membrane response. Subjects were exposed to a tonal conditioned stimulus (CS) and, if no conditioned response (CR) occurred, a 5.0-ma. paraorbital electric shock unconditioned stimulus (US). USs of 5.0, 3.3, 1.7, and 0.0 ma., respectively, were contingent on occurrence of a CR in the CS-US interval for subjects in the 4 groups of the experiment. The group exposed to contingent US omission differed from the other 3 groups in percentage CR, onset latency, amplitude, and 2 indices of CR form, and those 3 groups generally did not differ significantly among themselves on these dependent variables. Results were interpreted to be contrary to "law of effect" formulations of classical conditioning.     

Unconditional-stimulus locus and interstimulus-interval shift in rabbit(Oryctolagus cuniculus) nictitating membrane conditioning

The nictitating membrane response of rabbits was conditioned at a 200 msec interstimulus interval (ISI) with either circumorbital (C) or paraorbital (P) shock as the unconditional-stimulus locus. After 3 acquisition days half of each group was shifted to a 700 msec interstimulus interval. Results indicated: (1) more rapid acquisition for Group C, (2) postshift response decrements for both groups, (3) more rapid and stable, as well as complete return to preshift performance levels for Group C. Results were discussed in terms of the response-shaping hypothesis and the contiguity-substitution hypothesis in explaining both conditional response emergence and subsequent modifications of CR topography.     

Timing in trace conditioning of the nictitating membrane response of the rabbit (Oryctolagus cuniculus): scalar, nonscalar, and adaptive features

Using interstimulus intervals (ISIs) of 125, 250, and 500 msec in trace conditioning of the rabbit nictitating membrane response, the offset times and durations of conditioned responses (CRs) were collected along with onset and peak latencies. All measures were proportional to the ISI, but only onset and peak latencies conformed to the criterion for scalar timing. Regarding the CR's possible protective overlap of the unconditioned stimulus (US), CR duration increased with ISI, while the peak's alignment with the US declined. Implications for models of timing and CR adaptiveness are discussed.     

Latent inhibition and stimulus generalization of the classically conditioned nictitating membrane response in rabbits (Oryctolagus cuniculus) following dorsal hippocampal ablation.

Rabbits received 0 to 450 exposures of a tone conditioned stimulus (CS) prior to classical defensive conditioning of the nicitating membrane response based on an infraorbital eye shock unconditioned stimulus. Tone preexposure resulted in retarded conditioning in normal rabbits. This latent inhibition effect was not present in animals with bilateral dorsal hippocampectomy produced by aspiration. Control animals with bilateral neocortical and callosal aspiration lesions demonstrated a latent inhibition effect similar to that shown by normal nonoperated animals. The failure of CS preexposure to retard conditioning in hippocampal rabbits was not due to differences in threshold of the conditioned response to the CS or to differences in response mechanisms as determined by tests of habituation and dishabituation of the unconditioned response. A subsequent experiment employed combined-cue summation tests to confirm the fact that preexposure did not endow the tone with conditioned as well as latent inhibitiory properties. Finally, tests of stimulus generalization along the auditory frequency dimension indicated flatter relative gradients for hippocampals than for nonoperated controls, with cortical controls in between. These findings were discussed in terms of Douglas' model of hippocampal function.     

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.     

The effects of interpolated US alone (USa) presentations on classical nictitating membrane conditioning in rabbit(Oryctolagus cuniculus)

The nictitating membrane (NM) response of 36 rabbits was classically conditioned using a 1,000-Hz tone (CS) paired with circumorbital shock (US) at a 250-msec CS-US interval. Two experimental groups received additional presentations of the US given alone (USa), either 30 seconds (Group E30) or 60 seconds (Group E60) following each CS-US pairing, and their performance was compared to that of control Ss (Group C60) which never experienced USa. Presentations of USa resulted in significant and approximately equal performance decrements in both experimental groups relative to the C60 controls. However, no significant differences among the groups appeared during an extinction phase. The effects of USa presentations were discussed in terms of Papsdorf’s consolidation interpretation and Rescorla’s (1967) contingency theory of conditioning, and appeared to favor the latter view. Additional analyses led to the conclusions that (1) the increased amplitude of the NM UR on paired CS-US trials relative to USa trials can be attributed, at least in part, to the simple laws of the reflex, independent of the conditioning processper se, and (2) the partial reinforcement extinction effects previously reported by Leonard and Theios (1968) cannot be attributed solely to the presence of USa in their continuous reinforcement control Ss.     

Neuronal unit activity in the abducens nucleus during classical conditioning of the nictitating membrane response in the rabbit (Oryctolagus cuniculus).

Neuronal unit activity was recorded from the abducens (6th nerve) nucleus, the "final common path," during classical conditioning of the nictitating membrane (NM) response in the rabbit, with the use of a tone conditioned stimulus, an air puff unconditioned stimulus (UCS), 250-msec interstimulus interval, and 60-sec intertrial interval. Animals were given 2 days of conditioning training (104 trials in eight blocks per day) and 1 day of extinction. Control animals were given comparable periods of stimulus presentations, explicitly unpaired. Activity of small clusters of units--"multiple unit" recording--was compared with the amplitude-time course of the NM response. Between-blocks comparisons of neural and behavioral responses indicated an essentially perfect correlation during acquisition of the conditioned response (Day 1, r = .99; Day 2, r = .98) and a slightly lower correlation during extinction (r = .93) for the conditioning animals. Within-blocks comparisons indicated a close correspondence between the histograms of unit activity and the amplitude-time course of the NM response for the conditioning animals in all phase of training and for the control animals in the UCS trial blocks.     

Rapid reaquisition in conditioning of the rabbit's nictitating membrane response.

Reacquisition after extinction often appears faster than original acquisition. However, data from conditioned suppression studies indicate that this effect may arise from spontaneous recovery and reinstatement of unextinguished contextual stimuli related to the unconditioned stimulus (US). In the present experiments using the rabbit nictitating membrane preparation, spontaneous recovery was eradicated before reaquisition training. US contextual stimuli were controlled by retaining the US during extinction through explicit unpairings of the conditioned stimulus (CS) and US. Attempts were also made to drive the associative strength of the CS into the inhibitory region by differential conditioning and conditioned inhibition procedures. In all cases, reacquisition was very rapid in comparison with a rest control. The results are discussed with respect to their implications for CS and US processing models of conditioning.     

Differential conditioning of the rabbit's nictitating membrane response to serial compound stimuli

Three experiments were conducted to determine the time course and contents of CS representations through an examination of differential conditioning of the rabbit's nictitating membrane response to two serial compounds. One compound (A-X+) was always paired with the unconditioned stimulus, and the other (B-X-) was always presented alone. All three experiments entailed manipulation of the interstimulus interval between the initial distinctive element of each compound (A and B) and the second, shared element (X). The joint results revealed that (a) conditioned response acquisition to the initial elements depended on the presence of X in the A-X+ compound; (b) differentiation between A and B appeared across interstimulus intervals up to 4,600 ms; and (c) conditional control over responding following A and B appeared at interstimulus intervals of at least 4,600 ms and perhaps up to 12,600 ms. The results are discussed with respect to mechanisms of occasion setting, generalization, and configuration.     

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.     

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.     

Conditioning preparation for the nictitating membrane of the pigeon

A procedure is described for restraining the pigeon while recording movement of the nictitating membrane. The preparation provides a means of studying classical conditioning that combines the rich sensory capacities of the pigeon with the control achieved in nictitating membrane conditioning of the rabbit.