Rabbit Nictitating Research Articles

Real-time processing of serial stimuli in classical conditioning of the rabbit's nictitating membrane response.

Real-time models contend that a conditioned stimulus (CS) gives rise to a cascade of hypothetical stimuli that govern conditioned responses (CRs) on a moment-by-moment basis. Experiments with the rabbit nictitating membrane response successfully extended these models to external stimuli. CSs were trained in sequence with an unconditioned stimulus (CSA-CSB-US). When the CSA-CSB interval was shortened, the CR was compressed; when the CSA-CSB interval was lengthened, the CR was broadened. Peaks appeared at 2 places, namely, 1 following CSA by a period equal to its CS-US interval and another following CSB by its CS-US interval. Outside the sequence, the individual CSs evoked CRs located between their respective CS-US intervals. When, however, the 2 CSs were trained separately, the CRs were appropriate to their respective CS-US intervals when tested alone or in sequence. The results are discussed in terms of the J.E. Desmond and J.W. Moore (1988) and S. Grossberg and N.A. Schmajuk (1989) models.

Real-time processing of serial stimuli in classical conditioning of the rabbit's nictitating membrane response.

Real-time processing of serial stimuli in classical conditioning of the rabbit's nictitating membrane response.

6-Hydroxydopamine induced impairment of Pavlovian conditioning in the rabbit.

This study employed bilateral, intraventricular injections of 6-hydroxydopamine (6-HDA) to examine the effects of monoamine depletion on Pavlovian conditioning of the rabbit's nictitating membrane response. 6-HDA produced dose-dependent and highly correlated decreases in the rate of acquisition of conditioned responses and in the telencephalic content of 5-HT, DA, and NE. At the highest dose of 6-HDA (1340 micrograms), 5-HT, DA, and NE were reduced by 42, 48, and 89%, respectively, and the number of trials required to achieve criterion acquisition was increased by 123%. Control experiments established that the highest dose of 6-HDA: 1) had no effect on the unconditioned nictitating membrane reflex; 2) had no effect on the threshold of the conditioned stimulus for eliciting conditioned responses; and 3) produced only a small, less than 5%, decrease in nonassociative responding. It was concluded that decreases in 5-HT, DA, and NE can impair associative learning without altering sensory or motor function.

Effects of MDA upon differential serial compound conditioning and reflex modification of the rabbit's nictitating membrane response

The present investigations sought to determine the effects of 3,4-methylenedioxyamphetamine (MDA) on: 1) differential conditioning of the rabbit's nictitating membrane response to the serial compounds A-X-US (tone-light-reinforced compound) and B-X (white noise-light-unreinforced compound) by examining differential responding to A and B and their conditional control over responding to X within the compounds (Experiment 1); and 2) the ability of the compound stimuli and their components to modify the amplitude of the unconditioned nictitating membrane response (Experiment 2). Experiment 1 revealed that MDA decremented differential responding to the serial compounds and their A and B components, while enhancing conditioned responding to the X component. In addition, Experiment 2 indicated that MDA attenuated reflex modification to the compounds and their A and B components, but facilitated reflex modification to X alone. The results of these experiments indicated that MDA operated to alter the intensity, distinctiveness, and persistence (short-term memory) of stimulus representations.

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

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

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.

Second-order conditioning of the rabbit’s nictitating membrane response

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.

Trace conditioning of the rabbit's nictitating membrane response as a function of CS-US interstimulus interval and trials per session

The present experiment examined modification of URs and acquisition of CRs in the rabbit's nictitating membrane preparation as a function of the CS-US interstimulus interval (ISIs = 400, 800, 1600, and 3200 ms) and trials per session (1, 5, and 25). In both paired CS-US groups and corresponding unpaired groups ( n = 12), UR amplitude was greater under 1 and 5 trials per session than under 25 trials per session. In early CR acquisition, ISI interacted with trials per session such that CRs emerged rapidly for all ISIs under 1 and 5 trials per session but emerged more slowly as an inverse function under 25 trials per session. Later in training, CR frequency tended to be inversely related to ISI under 1, 5, and 25 trials, but training under 1 and 5 trials per session differentially raised the asymptotic CR frequency at the longer ISIs, especially the 3200-ms value. The results are discussed in terms of their implications for accounts of trial spacing effects and the time course of CS representations in trace conditioning.

A unique enhancement of associative learning produced by methylenedioxyamphetamine

The effects of methylenedioxyamphetamine (MDA) on classical conditioning of the rabbit's nictitating membrane response were assessed in four experiments. Experiment 1 established a dose-effect curve for MDA doses of 0, 1, 3, and 10µmol/kg. Both the 3 and 10µmol/kg doses significantly enhanced the rate of learning. Experiment 2 established that the 10µmol/kg dose of MDA had no effect on non-associative determinants of responding to the conditioned stimulus (CS). However, this dose of MDA apparently sensitized the response to the unconditioned stimulus (US). Experiments 3 and 4 assessed the effects of the 10µmol/kg dose of MDA on sensory processing of the CS and US, respectively. MDA had no effect on the intensity thresholds for eliciting responses to either of these stimuli. Thus, the enhanced rate of acquisition observed in experiment 1 cannot be attributed to an increase in nonassociative responding to the CS or to enhanced sensory processing of the CS or US. MDA apparently enhanced the rate of acquisition in experiment 1 by facilitating the association between the CS and US. These findings are unique in that no other hallucinogen we have examined has enhanced acquisition without also affecting either non-associative responding or sensory processing of the CS and US.

Effects of cocaine on conditioning of the rabbit nictitating membrane response

Two experiments were conducted to determine cocaine's (0, 1, 3, and 6 mg/kg) effects on associative, nonassociative, and motor processes in classical conditioning of the rabbit's nictitating membrane response (NMR). In experiment 1, acquisition training consisted of tone- and light-conditioned stimuli (CSs) each paired on separate trials with a shock unconditioned stimulus (UCS). Cocaine injected prior to each session significantly impaired acquisition of conditioned responses (CRs). In Experiment 2, rabbits received cocaine injections prior to each training session involving explicitly unpaired CS-alone presentations. Cocaine had no significant effects upon: base rate of NMRs; frequency of NMRs during presentations of the CSs; and frequency, amplitude, and latency of the UCRs. Consequently, cocaine's impairment of CR acquisition could not be attributed to its effects upon the nonassociative processes of base rate, sensitization, and pseudoconditioning, nor upon the sensory processing of the UCS and/or motor functioning of the UCR. Rather, cocaine's effects upon CR acquisition were mediated by the drug's effect upon associative processes. It appears likely that the drug affected the ability of the CS to enter into the associative conditioning process.

Learning-specific differences in Purkinje-cell dendrites of lobule HVI (Lobulus simplex): intracellular recording in a rabbit cerebellar slice

The cerebellum has been directly implicated in the associative learning exemplified by classical conditioning of the rabbit nictitating membrane response. Lesions of cerebellar lobule HVI (Lobulus simplex) can disrupt or even abolish classical conditioning. Electrical stimulation of inputs that converge at cerebellar lobule HVI and the deep cerebellar nuclei has been shown to support classical conditioning of the responses elicited by such stimulation. Here we report that intracellular recording in a slice of rabbit cerebellar lobule HVI revealed a conditioning-specific increase in the excitability of Purkinje-cell dendrites without significant changes in dendritic membrane potential or input resistance. The finding of a conditioning-specific change in lobule HVI lends support to the theory that associative learning involves local dendritic modification and provides further evidence for the involvement of the cerebellar cortex in classical conditioning of the rabbit's nictitating membrane response.

Effects of MDA on classical conditioning of the rabbit nictitating membrane response

In Experiment 1, classical conditioning of the rabbit's nictitating membrane response (NMR) was accomplished by pairing tone and light conditioned stimuli (CSs) with a shock unconditioned stimulus (UCS). MDA impaired the acquisition of conditioned responses (CR) to a tone-CS, while significantly enhancing CR acquisition to a light-CS. Experiment 2, employing explicitly unpaired CS, UCS training, revealed no reliable effects of MDA upon nonassociative processes. Subsequent efforts determined if MDA's CR acquisition effects resulted from alterations in sensory processing of the CS, UCS, and/or UCR motor functioning. Specifically, it was determined that MDA: (a) increased the tone-CS intensity threshold for eliciting CRs (Experiment 3); (b) attenuated the tone-induced reflex modification of the unconditioned NMR (Experiment 4); and (c) enhanced UCR frequency at varying UCS intensities (Experiment 5). It was concluded that MDA's effect upon CR acquisition reflected the drug's effects upon CS and UCS/UCR processing and thereby altered the ability of these components of conditioning to enter into associative learning.

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

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

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.

A neural network approach to hippocampal function in classical conditioning.

Hippocampal participation in classical conditioning in terms of Grossberg's (1975) attentional theory is described. According to the present rendition of this theory, pairing of a conditioned stimulus (CS) with an unconditioned stimulus (US) causes both an association of the sensory representation of the CS with the US (conditioned reinforcement learning) and an association of the sensory representation of the CS with the drive representation of the US (incentive motivation learning). Sensory representations compete among themselves for a limited-capacity short-term memory (STM) that is reflected in a long-term memory storage. The STM regulation hypothesis, which proposes that the hippocampus controls incentive motivation, self-excitation, and competition among sensory representations thereby regulating the contents of a limited capacity STM, is introduced. Under the STM regulation hypothesis, nodes and connections in Grossberg's neural network are mapped onto regional hippocampal-cerebellar circuits. The resulting neural model provides (a) a framework for understanding the dynamics of information processing and storage in the hippocampus and cerebellum during classical conditioning of the rabbit's nictitating membrane, (b) principles for understanding the effect of different hippocampal manipulations on classical conditioning, and (c) numerous novel and testable predictions.

Electrical stimulation of brainstem nuclei: Elicitation, modification, and conditioning of the rabbit nictitating membrane response.

Electrical stimulation of the reticular formation, pars oralis of the spinal trigeminal, abducens, and accessory abducens nuclei was used to assess the role of these sites in the elicitation, reflex modification, and classical conditioning of the rabbit's nictitating membrane response (NMR). Although electrical brain stimulation of the targeted sites revealed comparable levels of unconditioned responses, the spinal trigeminal nucleus was the only site at which reflex modification and conditioned response acquisition occurred reliably. These findings suggest that a locus of conditioned stimulus and unconditioned stimulus interaction, mediating either or both reflex modification and NMR conditioning, is on the sensory side of the reflex arc, at the pars oralis of the spinal trigeminal nucleus.

Temporal patterns of the rabbit's nictitating membrane response to compound and component stimuli under mixed CS-US intervals.

The acquisition of the rabbit's nictitating membrane response to a tone and light compound and to its components was examined when compound presentations were reinforced at one conditioned stimulus-unconditioned stimulus (CS-US) interval and individual component presentations were reinforced at another CS-US interval. Examination of the time course of the CRs revealed that (a) despite the mixture of CS-US intervals, conditioned response (CR) timing remained accurate, that is, CRs reached their peaks at the alternative points of US delivery; (b) the momentary magnitude of the CR to the compound was predominantly an additive function of the CR magnitude to the individual components; but (c) there was modest evidence of differentiation between the compound as a unit and the individual components. The results are discussed in terms of their implications for the study of the neural substrates of temporal and sensory integration as they modulate CR acquisition.

Temporal patterns of the rabbit's nictitating membrane response to compound and component stimuli under mixed CS–US intervals.

Temporal patterns of the rabbit's nictitating membrane response to compound and component stimuli under mixed CS–US intervals.

Asymmetric uptake of 2-deoxy- d-[ 14C]glucose in the dorsal cochlear nucleus during Pavlovian conditioning in the rabbit

Uptake of 2-deoxy- d-[ 14C]glucose was measured during Pavlovian conditioning of the rabbit's nictitating membrane response by both qualitative autoradiography and by quantitative measurement of radioactivity in samples of brain tissue. Conditioning was accomplished by pairing a tone stimulus delivered to both ears with an air-puff stimulus delivered to the right eye. Infusion of 2-deoxy- d-[ 14C]glucose during the first day of conditioning when there was no evidence of acquisition or during the 7th day of conditioning when animals demonstrated 68% conditioned responses resulted in a significantly greater uptake of radioactivity by the caudal portions of the left as compared with the right dorsal cochlear nucleus. Similar changes were not observed in other auditory and non-auditory nuclei. Rabbits that had acquired conditioned responses across 6 days of training and were then exposed only to the tone-conditioned stimulus on the 7th day of testing exhibited 69% conditioned responses but no asymmetry in the uptake of 2-deoxy- d-[ 14C]glucose. Control animals receiving unpaired presentations of tone and air puff or no stimulation did not acquire conditioned responses and did not demonstrate asymmetric uptake of radioactivity in the dorsal cochlear nucleus. These results indicate that the asymmetric uptake of radioactivity by the dorsal cochlear nucleus did not result from the effects of stimulation per se or the prior occurrence of learning but was due to the explicit pairing of the tone stimulus with the asymmetric delivery of the air puff. It would appear that the caudal dorsal cochlear nucleus not only serves as a signal transducer for auditory stimuli but also receives input from other sensory systems thus allowing it to both recognize when an auditory stimulus is followed by a biologically significant event and to transmit such information to other brain regions that are, in turn, responsible for learning.

Stimulation of the spinal trigeminal nucleus supports classical conditioning of the rabbit's nictitating membrane response.

Acquisition, extinction, and differential conditioning of the rabbit's nictitating membrane response to a tone conditioned stimulus were supported by electrical stimulation of the spinal trigeminal nucleus. Stimulation of the accessory abducens nucleus, the abducens nucleus, and the reticular formation at the level of the spinal trigeminal nucleus supported lower, transient levels of conditioning. The results are discussed in terms of stimulation of sensory inputs to the brainstem and cerebellum.