Posttraining shifts in the overshadowing stimulus–unconditioned stimulus interval alleviates the overshadowing deficit.

Two conditioned lick suppression experiments explored the effects on overshadowing of a posttraining change in the temporal relationship between the overshadowing conditioned stimulus (CS) and the unconditioned stimulus (US). Rats received either trace (Experiment 1) or delay (Experiment 2) overshadowing training. Then pairings of the overshadowing CS and US were given with either a trace or delay temporal relationship. Overshadowing was alleviated by shifting the overshadowing CS–US temporal relationship so that it no longer matched the overshadowed CS–US temporal relationship. These outcomes are explicable in terms of an integration of the comparator hypothesis, which states that cue competition effects (e.g., overshadowing) will be maximal when the information potentially conveyed by competing CSs is equivalent, and the temporal coding hypothesis, which states that CS–US intervals are part of the information encoded during conditioning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of posttraining damage to the pedunculopontine tegmental nucleus on conditioned stimulus transfer in two-way active avoidance in rats.

The effects of posttraining excitotoxic lesions of the pedunculopontine tegmental nucleus (PPTg) on two-way active avoidance after changing the conditioned stimulus (CS) used during prelesion training were examined. Prelesion training was carried out with either a tone or a light as the CS, and this CS was changed during postlesion training. Replacing the tone with a light reduced the performance of control and lesioned rats, but the degree of reduction was higher in the latter. Replacing the light with a tone had slight detrimental effects in lesioned rats but not in controls. Thus, posttraining PPTg lesions slowed down the reacquisition of shuttle-box avoidance under conditions of CS transfer, an effect that may be attributable to disruption of attention and/or gating of sensory stimuli. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Single neurons in the medial prefrontal cortex of the rat exhibit tonic and phasic coding during trace fear conditioning.

Trace fear conditioning is a learning task that requires the association of an auditory conditioned stimulus (CS) and a shock unconditioned stimulus (US) that are separated by a 20-s trace interval. Single neuron activity was recorded from the prelimbic and infralimbic areas of the medial prefrontal cortex in rats during trace fear conditioning or nonassociative unpaired training. Prelimbic neurons showed learning-related increases in activity to the CS and US, whereas infralimbic neurons showed learning-related decreases in activity to these stimuli. A subset of prelimbic neurons exhibited sustained increases in activity during the trace interval. These sustained prelimbic responses may provide a bridging code that allows for overlapping representations of CS and US information within the trace fear conditioning circuit. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Timing during inhibitory conditioning.

Analyses of second-by-second conditioned responding into a food receptacle by hungry rats (Rattus norvegicus) found that inhibition varies across the duration of a conditioned stimulus (CS) in a manner consistent with initial training. Variations in the arrival time of the unconditioned stimulus (US) supported temporally specific suppression of responding (Experiment 1). Summation and retardation tests (Experiments 2 and 3, respectively) revealed that points of greatest inhibition coincided with US omission at the time normally specified by the excitor. Our data provide a clear demonstration of fine-grained changes in the time course of inhibitory conditioning for the first time. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

A Role for CS-US Contingency in Pavlovian Conditioning.

Two experiments evaluated the role of conditioned stimulus-unconditioned stimulus (CS-US) contingency in appetitive Pavlovian conditioning in rats. In both experiments, some groups received a positively contingent CS signaling an increased likelihood of the US relative to the absence of the CS. These groups were compared with control treatments in which the likelihood of the US was the same in the presence and absence of the CS. A trial marker served as a trial context. Experiment 1 found contingency sensitivity. There was a reciprocal relationship between responding to the CS and the trial marker. Experiment 2 showed that this result was not stimulus or response specific. These results are consistent with associative explanations and the idea that rats are sensitive to CS-US contingency. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporally specific extinction of conditioned responses in the rabbit (Oryctolagus cuniculus) nictitating membrane preparation.

Extinguishing a conditioned response (CR) has entailed separating the conditioned stimulus (CS) from the unconditioned stimulus (US). This research reveals that elimination of the rabbit (Oryctolagus cuniculus) nictitating membrane response occurred during continuous CS-US pairings. Initial training contained a mixture of 2 CS-US interstimulus intervals (ISIs): 200 ms and 1,200 ms. The CRs showed double peaks, one for each ISI. When 1 ISI was removed, its CR peak showed the hallmarks of extinction: a decline across sessions, spontaneous recovery between sessions, and rapid reacquisition when the absent ISI was reintroduced. These results support real-time models of conditioning that segment the CS into microstimuli while challenging theories that rely on contextual control, US representations, CS processing, and response inhibition. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal Specificity of Extinction in Autoshaping.

Three experiments investigated the effects of varying the conditioned stimulus (CS) duration between training and extinction. Ring doves (Streptopelia risoria) were autoshaped on a fixed CS-unconditioned stimulus (US) interval and extinguished with CS presentations that were longer, shorter, or the same as the training duration. During a subsequent test session, the training CS duration was reintroduced. Results suggest that the cessation of responding during an extinction session is controlled by generalization of excitation between the training and extinction CSs and by the number of nonreinforced CS presentations. Transfer of extinction to the training CS is controlled by the similarity between the extinction and training CSs. Extinction learning is temporally specific. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Time and stimulus specificity in extinction of the conditioned nictitating membrane response in the rabbit (Oryctolagus cuniculus).

The present experiments demonstrated that, in the rabbit nictitating membrane preparation, a conditioned response (CR) can be selectively eliminated in one portion of a conditioned stimulus (CS) while it is still paired with the unconditioned stimulus (US). Rabbits were initially trained with two stimuli (tone, light). Each was paired with the US by using a mixture of two CS-US interstimulus intervals (ISIs): 200 ms and 1,200 ms in Experiment 1; 150 ms and 500 ms in Experiment 2. The CRs showed double peaks, one for each ISI. Subsequently, one CS (A) was trained with only the longer ISI, whereas the other CS (B) continued to be trained with both ISIs. Consequently, the CR peak based on the shorter ISI disappeared for CSA but not for CSB. The later CR peaks during both CSA and CSB were maintained. These results support time-based models of conditioning. Implications for proposed mechanisms of extinction are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hippocampectomy disrupts auditory trace fear conditioning and contextual fear conditioning in the rat

The hippocampus is believed to be an important structure for learning tasks that require temporal processing of information. The trace classical conditioning paradigm requires temporal processing because the conditioned stimulus (CS) and the unconditioned stimulus (US) are temporally separated by an empty trace interval. The present study sought to determine whether the hippocampus was necessary for rats to perform a classical trace fear conditioning task in which each of 10 trials consisted of an auditory tone CS (1 5-s duration) followed by an empty 30-s trace interval and then a fear-producing floor-shock US (0.5-s duration). Several weeks prior to training, animals were anesthetized and given aspiration lesions of the neocortex (NEO; n = 6), hippocampus and overlying neocortex (HIPP; n = 7), or no lesions at all (control; n = 6). Approximately 24 h after trace conditioning, NEO and control animals showed a significant decrease in movement to a CS-alone presentation that was indicative of a conditioned fear response. Animals in the HIPP group did not show conditioned fear responses to the CS alone, nor did a pseudoconditioning group (n = 7) that was trained with unpaired CSs and USs. Furthermore, all groups except the HIPP group showed conditioned fear responses to the original context in which they received shock USs. One week later, HIPP, NEO, and control animals received delay fear-conditioning trials with no trace interval separating the CS and US. Six of seven HIPP animals could perform the delay version, but none could perform the trace version. This result suggests that the trace fear task is a reliable and useful model for examining the neural mechanisms of hippocampally dependent learning.     

Odor-guided fear conditioning in rats: 1. Acquisition, retention, and latent inhibition.

Three experiments examined the acquisition, retention, and latent inhibition of odor-guided fear conditioning in rats. The results of Experiment 1 indicate that forward conditioned stimulus (CS)–unconditioned stimulus (US) pairings resulted in robust freezing responses to subsequent presentation of the CS alone. In Experiment 2, rats in one group (PRE) received unreinforced preexposures to the odorant CS, and those in a second group (NON) were not preexposed to the odorant. All rats then received forward CS–US pairings. PRE rats exhibited a marked attenuation of freezing to subsequent exposure to the CS relative to NON rats. All rats were then retested at one of the following posttraining delays: 17, 24, or 31 days. Freezing behavior of the NON rats declined significantly across these delays, whereas rats in the PRE group froze no more at any delay than they had 24 hr after training. Experiment 3 examined the contextual specificity of latent inhibition. Only those rats that were preexposed and were trained in the same context exhibited latent inhibition. These results indicate that odor-guided fear conditioning is a robust and useful paradigm suitable for future studies of the neural bases of associative learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Using context to resolve temporal ambiguity.

Three conditioned lick suppression experiments with rats examined the role of the context in the selection and integration of independently acquired interval relationships. In Experiment 1, rats were exposed to separate conditioned stimuli 1 and 2 (CS1–CS2) pairings with 2 different interval relationships, each in its own distinctive context, X or Y. The resultant integration was determined by the training context (X or Y) in which unconditioned stimulus (US)–CS2 backward pairings occurred, as assessed in a third neutral context (Z). In Experiment 2, rats experienced CS1–CS2 pairings with 2 different interval relationships as in Experiment 1, and then received US–CS2 pairings in both contexts X and Y. The testing context (i.e., X or Y) determined the resultant integration. In Experiment 3, rats were exposed to CS1–CS2 pairings in 2 different interval relationships each in different phases (i.e., Phases 1 and 2), and then in Phase 3 received US–CS2 pairings. The temporal context of testing (i.e., short or long retention interval) determined the resultant integration. Thus, both physical and temporal context can be used to disambiguate conflicting temporal information. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Motor cortex lesions do not affect learning or performance of the eyeblink response in rabbits.

The possible modulatory role of motor cortex in classical conditioning of the eyeblink response was examined by ablating anterior neocortex in rabbits and training them with an auditory conditioned stimulus (CS) and an airpuff unconditioned stimulus (US) in either a delay (Experiment 1) or a trace (Experiment 2) conditioning paradigm. Topographic measures such as amplitude and onset latency were assessed during conditioning sessions for conditioned responses (CRs) and on separate test days for unconditioned responses (URs) by using a range of US intensities. No lesion effects were observed for learning or performance measures in acquisition or retention of either delay or trace conditioning. During trace conditioning, lesioned rabbits did, however, exhibit a trend toward impairment and demonstrated significantly longer CR latencies. Damage to motor and frontal cortex does not significantly affect eyeblink response performance or learning in either a delay or a trace conditioning paradigm. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Topography of spatially directed conditioned responding: Effects of context and trial duration.

Domesticated quail (Coturnix japonica) received a discrete conditioned stimulus (CS) at one end of the experimental chamber paired with the opportunity to copulate with a female quail (the unconditioned stimulus) in a goal box located 112 cm away. Approach to the CS (sign tracking) and approach to the goal area (goal tracking) were measured. The duration of exposure to the experimental context (C) was varied in Experiment 1, and the duration of the conditioning trials (T) was varied in Experiment 2 for independent groups, creating C/T ratios of 1.0, 1.5, 4.5, 45, and 180. Contrary to previous reports of a direct relation between the C/T ratio and conditioned responding, in the present experiments, a shift in the topography and stimulus control of conditioned behavior occurred. Low C/T ratios (1.0–4.5) produced goal tracking controlled by contextual cues, whereas high C/T ratios (45 and 180) produced sign tracking controlled by the discrete CS. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Pavlovian Conditioning: The CS-UR Relation.

A new Pavlovian procedure used fluid-elicited throat-movement responses of the pigeon (N=66) to study the effects on conditioning of the temporal relation of the conditioned stimulus (CS) to the unconditioned stimulus-unconditioned response (US-UR). Because the throat-movement response has a substantial latency and duration, the relation of the CS to the US and UR could be independently evaluated. Four experiments indicated that, operationally, the relation of the CS to the UR--not to the US--is critical for conditioning in this preparation. The conventional emphasis on CS-US relations is based on procedures that confound the occurrence of the US with the UR and that foster generalization decrement between training and testing. The authors indicate how several conditioning phenomena may be reinterpreted in terms of CS-UR relations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Aware and (Dis)Liking: Item-Based Analyses Reveal That Valence Acquisition via Evaluative Conditioning Emerges Only When There Is Contingency Awareness.

Evaluative conditioning (EC) refers to changes in the liking of an affectively neutral stimulus (the conditioned stimulus, or CS) following the pairing of that stimulus with another stimulus of affective value (the unconditioned stimulus, or US). In 3 experiments, the authors assessed contingency awareness, that is, awareness of the CS-US associations, by relying on participants' responses to individual items rather than using a global method of assessment. They found that EC emerged on contingency aware CSs only. Of note, whether the CSs were evaluated explicitly (Experiments 1 and 2) or implicitly (Experiment 3) did not make a difference. This pattern supports the idea that awareness of the CS-US associations may be required for valence acquisition via EC. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Amygdala lesions block conditioned enhancement of the early component of the rat eyeblink reflex.

A tone-conditioned stimulus (CS) previously paired with a grid shock unconditioned stimulus (US) can greatly enhance the early electromyographic (EMG) component (RI) of the rat eyeblink reflex. The hypothesis that the central nucleus of the amygdala (ACe) is an essential part of the circuitry mediating conditioned RI enhancement was tested. After bilateral ACe lesions (L) or a sham operation (S), rats received paired presentations of the CS and US (P) or explicitly unpaired CS and US presentations (U), resulting in 4 groups: P/S, P/L, U/S, and U/L. ACe lesions completely prevented conditioned RI enhancement, which was only exhibited in Group P/S. In the latter group, the "preextinction" conditioned enhancement effect was roughly a 2-fold increase in the RI magnitude. Circuit-level mechanisms are discussed, and some advantages of the eyeblink EMG response in this general conditioning paradigm are considered. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Conditional Discrimination Learning in Patients With Bilateral Medial Temporal Lobe Amnesia.

The ability of bilateral medial temporal lobe amnesic patients (MT; n=8) and normal participants (NC; n=8) to acquire a conditional discrimination in trace and delay eyeblink conditioning paradigms was investigated. Experiment 1 assessed trace conditional discrimination learning by using a light conditional stimulus (S+/S-) and tone conditioned stimulus (CS) separated by a 1-s trace. NCs responded differentially on S+ trials (mean percent conditioned responses=66) versus S- trials (30). Whereas MTs were impaired in their acquisition of the conditional discrimination (S+=51, S-=43). In Experiment 2, the temporal separation was eliminated. NCs acquired the conditional discrimination (S+=70, S-=29). MTs were unable to respond differentially (S+=42, S-=37). The findings indicate that the hippocampal system is essential in acquiring a conditional discrimination, even in a delay paradigm. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Opioid Receptors Regulate the Extinction of Pavlovian Fear Conditioning.

Rats received a single pairing of an auditory conditioned stimulus (CS) with a footshock unconditioned stimulus (US). The fear (freezing) that had accrued to the CS was then extinguished. Injection of naloxone prior to this extinction significantly impaired the development of extinction. This impairment was mediated by opioid receptors in the brain and was not observed when naloxone was injected after extinction training. Finally, an injection of naloxone on test failed to reinstate extinguished responding that had already accrued to the CS. These experiments show that opioid receptors regulate the development, but not the expression, of fear extinction and are discussed with reference to the roles of opioid receptors in US processing, memory, and appetitive motivation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Classical conditioning and brain systems: the role of awareness

Classical conditioning of the eye-blink response, perhaps the best studied example of associative learning in vertebrates, is relatively automatic and reflexive, and with the standard procedure (simple delay conditioning), it is intact in animals with hippocampal lesions. In delay conditioning, a tone [the conditioned stimulus (CS)] is presented just before an air puff to the eye [the unconditioned stimulus (US)]. The US is then presented, and the two stimuli coterminate. In trace conditioning, a variant of the standard paradigm, a short interval (500 to 1000 ms) is interposed between the offset of the CS and the onset of the US. Animals with hippocampal lesions fail to acquire trace conditioning. Amnesic patients with damage to the hippocampal formation and normal volunteers were tested on two versions of delay conditioning and two versions of trace conditioning and then assessed for the extent to which they became aware of the temporal relationship between the CS and the US. Amnesic patients acquired delay conditioning at a normal rate but failed to acquire trace conditioning. For normal volunteers, awareness was unrelated to successful delay conditioning but was a prerequisite for successful trace conditioning. Trace conditioning is hippocampus dependent because, as in other tasks of declarative memory, conscious knowledge must be acquired across the training session. Trace conditioning may provide a means for studying awareness in nonhuman animals, in the context of current ideas about multiple memory systems and the function of the hippocampus.     

CS-US preexposure effects on trace eyeblink conditioning in young rats: Potential implications for functional brain development.

Recent studies of delay eyeblink conditioning (EBC) in young rats have demonstrated different effects of various conditioned and unconditioned stimulus (CS-US) preexposure conditions on learning at different ages. The present study extends this research to trace EBC. Subjects experienced 1 of 3 preexposure conditions (paired CS-US, unpaired CS-US, or no stimuli) at either 20 or 24 days of age. Four days later, they were conditioned using either trace (Experiment 1) or delay (Experiment 2) EBC parameters. Results were similar at both ages tested. Paired preexposure facilitated acquisition of delay but not trace relative to context preexposure. Unpaired preexposure impaired acquisition of both delay and trace. These behavioral findings provide a foundation for hypotheses about the functional maturation of cerebellar, hippocampal, and entorhinal learning circuits. (PsycINFO Database Record (c) 2010 APA, all rights reserved)