Chemosensory-based contextual conditioning inHermissenda crassicornis

In two experiments, the marine molluskHermissenda crassicornis was exposed to context discrimination training. In one context, defined by the presence of a diffuse chemosensory stimulus (shellfish extract A), brief, unsignaled, unconditioned stimuli (USs; high-speed rotation) were presented; in a second context, defined by the presence of shellfish extract B, no USs were presented. Animals were then tested (at both 1.5 and 24 h) by exposing them to small pieces of the shellfish meat used to define the two contexts. The latency to strike at the meat served as an index of the context-US association. In Experiment 1, the latency to strike at the cue associated with rotation was reduced relative to both preconditioning strike latencies and the associatively neutral cue. However, in a two-choice test where the animals could approach the conditioned or neutral stimulus, the animals regularly avoided the stimulus paired with rotation. Moreover, if, following conditioning, the animals were presented with an unsignaled rotation in the conditioned context or the neutral context, the animals exhibited more effective defensive clinging (an unconditioned reflex normally elicited by rotation) in the conditioned context, suggesting that it “prepared” the animal for the aversive US. In total, these results demonstrate thatHermissenda is capable of making associations to diffuse background (contextual) stimuli. Moreover, the results suggest that pairing the chemosensory cue with an aversive US elicits a strike response inHermissenda when the animal is placed in forced contact with the cue and an active avoidance response when the animal can choose between that cue and a neutral cue.     

Chemosensory conditioning in molluscs: II. A critical review

We critically review chemosensory conditioning studies with molluscs and find that, in many studies, the influence of nonassociative processes complicates, obscures, and renders ambiguous the unique contribution of associative learning. These nonassociative processes include sensory adaptation, habituation, sensitization, and changes in feeding motivation. They arise from both the food extracts that have often been used as conditioned stimuli and the aversive stimuli that have been used as unconditioned stimuli.     

Contextual associations in trace conditioning

Rats were given tone-footshock pairings with a 0-, 10-, or 30-sec trace interval between tone offset and shock onset. Half the rats within each trace interval were tested for their conditioned fear of the tone through a lick suppression procedure; the remaining rats were evaluated for their fear of the background or contextual cues through their avoidance of the compartment in which conditioning had occurred. Less conditioning was observed to the tone with increasing trace intervals. However, conditioned fear of the context increased with increases in the trace duration. The ability of the more predictive stimulus, the tone, to overshadow the contextual cues was determined by the tone’s temporal contiguity with the footshock. The need to incorporate temporal parameters within current theories of conditioning is discussed.     

Context effects on conditioning,extinction, and reinstatement in an appetitive conditioning preparation

Three experiments with rat subjects examined the effects of contextual stimuli on performance in appetitive conditioning. A 10-sec tone conditioned stimulus (CS) was paired with a food-pellet unconditioned stimulus (US); conditioning was indexed by the observation of headjerking, a response of the rat to auditory stimuli associated with food. In Experiment 1, a context switch following initial conditioning did not affect conditioned responding to the tone; however, when the response was extinguished in the different context, a return to the original conditioning context “renewed” extinguished responding. These results were replicated in Experiments 2 and 3 after equating exposure to the two contexts (Experiment 2) and massing the conditioning and extinction trials (Experiment 3). The results of Experiment 1 also demonstrated that separate exposure to the US following extinction reinstates extinguished responding to the tone; this effect was further shown to depend at least partly on presenting the US in the context in which testing is to occur (Experiments 2 and 3). Overall, the results are consistent with previous data from aversive conditioning procedures. In either appetitive or aversive conditioning, the context may be especially important in affecting performance after extinction.     

A history of morphine-induced taste aversion learning fails to affect morphine-induced place preference conditioning in rats

Drugs of abuse have both rewarding and aversive effects, as indexed by the fact that they support place preferences and taste aversions, respectively. In the present study, we explored whether having a history with the aversive effects of morphine (via taste aversion conditioning) impacted the subsequent rewarding effects of morphine, as measured in the place preference design. In Experiment 1, rats were exposed to a taste aversion procedure in which saccharin was followed by morphine. Place preference conditioning was then initiated in which animals were injected with morphine and placed on one side of a two-chambered apparatus. Animals with a taste aversion history acquired place preferences to the same degree as controls without such a history, suggesting that morphine’s affective properties condition multiple effects, dependent on the specific stimuli present during conditioning. To determine whether these results were a reflection of processes operating in traditional associative conditioning, in a modified blocking procedure, place preference conditioning was attempted in the presence of a taste previously associated with morphine (Exp. 2). Under these conditions, animals still acquired morphine-induced place preferences comparable to those of animals without a morphine or conditioning history. These results are consistent with the position that drugs of abuse have multiple stimulus effects (positive and negative) that are differentially associated with specific stimuli (environmental and taste) that drive different behavioral responses (approach and avoidance).     

Toxiphobia conditioning with exteroceptive cues

The types of conditioned properties acquired by novel (i.e., nonpreexposed) or familiar (i.e., preexposed) exteroceptive cues that were paired with toxicosis, in the absence of a flavor CS, were evaluated in four experiments. In Experiment 1, the conditioned properties of novel exteroceptive cues served to block the acquisition of an aversion to a flavor CS during flavor conditioning and to suppress the ingestion response during flavor testing; animals failed to suppress their ingestion of either the flavor CS or a neutral flavor when tested in the absence of the exteroceptive CS, but suppressed their ingestion of both the flavor CS and the neutral flavor when tested in the presence of the exteroceptive CS. In Experiment 2, preexposed exteroceptive cues that had been paired with toxicosis failed to provide evidence of such conditioned properties. Experiment 3 demonstrated that preexposed contextual cues that were reinforced in compound with novel exteroceptive cues failed to acquire the conditioned properties acquired by nonpreexposed contextual cues under the same conditions of reinforcement. Finally, in Experiment 4, the conditioned properties of the novel exteroceptive cues served to evoke a conditioned gastrointestinal response that gradually extinguished as a function of repeated nonreinforced exposures to the exteroceptive cues, and, in the absence of such extinction, the conditioned properties served to block the acquisition of a flavor aversion.     

Factors governing one-trial contextual conditioning

When a rat was placed in a chamber and shortly thereafter received a single footshock, it showed conditional freezing upon re-exposure to that chamber but not a different one (Experiment 1). Experiments 2–4 showed that the probability of this freezing decreased linearly with decreases in the delay between placement in the chamber and shock delivery. With very short delays (e.g., less than 27 sec), there was no freezing. Experiments 2 and 3 demonstrated that a 2-mm pre-exposure to the chamber, 24 h prior to shock delivery, reduced the minimum delay necessary to successfully condition freezing. Experiment 4 demonstrated that shorter delays were successful in conditioning freezing if a salient tone was a component of the contextual stimulus. The changes in freezing caused by delay interval and preexposure did not simply reflect the total time in the context, suggesting that there may be two requirements that place temporal restrictions on the conditioning of the freezing response. One is satisfied by sufficient exposure, whether or not that exposure is contiguous with shock. The second requirement is for a small amount of context exposure that is contiguous with shock.     

Solving Pavlov's puzzle: Attentional, associative, and flexible configural mechanisms in classical conditioning

This article introduces a new "real-time" model of classical conditioning that combines attentional, associative, and "flexible" configural mechanisms. In the model, attention to both conditioned (CS) and configural (CN) stimuli are modulated by the novelty detected in the environment. Novelty increases with the unpredicted presence or absence of any CS, unconditioned stimulus (US), or context. Attention regulates the magnitude of the associations CSs and CNs form with other CSs and the US. We incorporate a flexible configural mechanism in which attention to the CN stimuli increases only after the model has unsuccessfully attempted learn input-output combinations with CS-US associations. That is, CSs become associated with the US and other CSs on fewer trials than they do CNs. Because the CSs activate the CNs through unmodifiable connections, a CS can become directly and indirectly (through the CN) associated with the US or other CSs. In order to simulate timing processes, we simply assume that a CS is formed by a temporal spectrum of short-duration CSs that are activated by the nominal CS trace. The model accurately describes 94?% of the basic properties of classical conditioning, using fixed model parameters and simulation values in all simulations.     

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.     

US inflation with trace and simultaneous fear conditioning

Two experiments with rat subjects in a conditioned punishment paradigm are reported. These experiments attempted to determine if the events entering into association with the CS following conditioning with informative (forward) and noninformative (simultaneous) CSs were comparable. In Experiment 1, exposure to intense shock alone following trace (ISI = 10 sec) conditioning with moderate shock enhanced the suppressive effects of a 2-sec CS. A similar manipulation following explicitly unpaired CS-US presentations (ISI = 2 min) had no effect. These data were taken as evidence that the CS and US were associated during trace conditioning. Experiment 2 showed that exposure to intense shock following simultaneous conditioning also enhanced suppression to the CS. These results suggested that simultaneous and forward conditioning procedures yield similar forms of associative learning.     

Causal and predictive-value judgments,but not predictions,are based on cue-outcome contingency

In three experiments, we show that people respond differently when they make predictions as opposed to when they are asked to estimate the causal or the predictive value of cues: Their response to each of those three questions is based on different sets of information. More specifically, we show that prediction judgments depend on the probability of the outcome given the cue, whereas causal and predictivevalue judgments depend on thecue-outcome contingency. Although these results might seem problematic for most associative models in their present form, they can be explained by explicitly assuming the existence of postacquisition processes that modulate participants’ responses in a flexible way.     

Conditioning of tentacle lowering in the snail (Helix aspersa): Acquisition, latent inhibition, overshadowing, second-order conditioning, and sensory preconditioning

In a series of related experiments, we studied associative phenomena in snails (Helix aspersa), using the conditioning procedure of tentacle lowering. Experiments 1A and 1B demonstrated a basic conditioning effect in which the pairing of an odor (apple) as the conditioned stimulus (CS) with the opportunity to feed on carrot as the unconditioned stimulus (US) made snails exhibit increased levels of tentacle lowering in the presence of the CS. Experiments 2 and 3 showed that the magnitude of the conditioning was reduced when snails were exposed to the CS prior to the conditioning trial (a latent inhibition effect). Experiment 4 examined the effects produced by pairing a compound CS (apple—pear) with food presentations and demonstrated the existence of an overshadowing effect between the two odors. Experiment 5 revealed that pairing one CS with another previously conditioned stimulus increased tentacle lowering to the new CS (a second-order conditioning effect). Finally, Experiment 6 showed that pairing two odors prior to conditioning of one of them promoted an increase in tentacle lowering in response to the other (a sensory preconditioning effect). The results are discussed in terms of an associative analysis of conditioning and its implications for the study of cognition in invertebrates.     

Evaluating conditioning of related and unrelated stimuli using a compound test

Three experiments used a compound test procedure to evaluate whether superior conditioning results from the pairing of stimuli that are related to each other. In each case, a stimulus compound was tested after its component conditioned stimuli (CSs) had been conditioned by the same unconditioned stimuli (USs) arranged such that either related or unrelated CSs and USs were paired. Experiment 1 explored auditory and gustatory stimuli conditioned by LiCl or shock, using rats. Experiments 2 and 3 used second-order conditioning in pigeons to pair stimuli that were similar by virtue either of qualitative features or of shared physical location. In each case, the compound test provided clear evidence that pairing related stimuli produces superior associative learning.     

A transfer of control test for contextual associations

A transfer of control experiment measured the associative properties of contextual stimuli from three standard classical conditioning paradigms. After baseline training on a Sidman avoidance schedule, dogs received aversive conditioning using excitatory, inhibitory, or truly random conditioning procedures in the presence of a manipulable background stimulus. As predicted by current theory (Rescorla & Wagner, 1972; Wagner & Rescorla, 1972), the contextual stimulus was excitatory after serving as the background during conditioning of a CS? and was neutral when it had been part of the background for conditioning of a CS+. The background to the truly random procedure was also neutral. This last result contrasts with Rescorla and Wagner’s theory.     

Secondary extinction in Pavlovian fear conditioning

Pavlov (1927/1960) reported that following the conditioning of several stimuli, extinction of one conditioned stimulus (CS) attenuated responding to others that had not undergone direct extinction. However, this secondary extinction effect has not been widely replicated in the contemporary literature. In three conditioned suppression experiments with rats, we further explored the phenomenon. In Experiment 1, we asked whether secondary extinction is more likely to occur with target CSs that have themselves undergone some prior extinction. A robust secondary extinction effect was obtained with a nonextinguished target CS. Experiment 2 showed that extinction of one CS was sufficient to reduce renewal of a second CS when it was tested in a neutral (nonextinction) context. In Experiment 3, secondary extinction was observed in groups that initially received intermixed conditioning trials with the target and nontarget CSs, but not in groups that received conditioning of the two CSs in separate sessions. The results are consistent with the hypothesis that CSs must be associated with a common temporal context during conditioning for secondary extinction to occur.     

Normalization between stimulus elements in a model of Pavlovian conditioning: Showjumping on an elemental horse

Harris and Livesey. Learning & Behavior, 38, 1-26, (2010) described an elemental model of associative learning that implements a simple learning rule that produces results equivalent to those proposed by Rescorla and Wagner (1972), and additionally modifies in "real time" the strength of the associative connections between elements. The novel feature of this model is that stimulus elements interact by suppressively normalizing one another's activation. Because of the normalization process, element activity is a nonlinear function of sensory input strength, and the shape of the function changes depending on the number and saliences of all stimuli that are present. The model can solve a range of complex discriminations and account for related empirical findings that have been taken as evidence for configural learning processes. Here we evaluate the model's performance against the host of conditioning phenomena that are outlined in the companion article, and we present a freely available computer program for use by other researchers to simulate the model's behavior in a variety of conditioning paradigms.     

Symmetrical generalization decrements: Configural stimulus processing in human contingency learning

Models of associative learning differ in their predictions concerning the symmetry of generalization decrements. Whereas Pearce’s (1994) configural model predicts the same response decrement after adding elements to and after removing elements from a previously trained stimulus, elemental models, such as the replaced elements model and Harris’s (2006) model, anticipate more of a decrement for removing than for adding elements. In three contingency learning experiments, we manipulated the motion and the spatial arrangement of colored dots in order to induce configural or elemental processing by perceptual grouping. The results reliably showed symmetrical decrements for the added and removed groups. The manipulations of the stimuli had no effect on stimulus processing. This is in line with Pearce’s configural model, but it is at variance with the elemental models and previous studies.     

Biological constraints on instrumental and classical conditioning: Retrospect and prospect

The adequacy of traditional approaches to the study of animal learning to account fully for learning phenomena has been seriously questioned during the past decade. Critics of traditional analyses advocated a biological orientation to the interpretation of associative processes and introduced a variety of concepts intended to provide a new framework for the study of animal learning. This promise of a reorientation of the field has not been realized. The concepts of biological constraints, adaptive specializations, and situation specificity of learning have had a less profound influence on the general process approach to instrumental and classical conditioning than anticipated. The present paper makes explicit the conceptual bases of the original biological approaches to learning, identifies reasons why they failed to change fundamentally the study of instrumental and classical conditioning, and proposes an alternative approach to the use of ecological and evolutionary principles in studies of conditioning. We suggest a renewed comparative approach to the study of learning phenomena that avoids many of the difficulties inherent in earlier formulations by providing (1) a strategy for the discovery of adaptive specializations in learning, (2) an ecological framework for the discussion of these adaptive specializations, and (3) a renewed emphasis on the study of species differences in learning.     

Contextual conditioning during free-operant extinction: Unsignaled,signaled, and backward-signaled noncontingent food

Three experiments were designed to study the effects of contextual conditioning on the extinction of instrumental leverpressing that had been reinforced on a random-interval schedule. In Experiment 1, noncontingent food retarded extinction, but signaling food delivery, a treatment that should reduce contextual conditioning, reduced the interference. Experiment 2 replicated the results of Experiment 1 and demonstrated that if the food preceded rather than followed the signal, the retardation of extinction was not reduced but was enhanced. In Experiment 3, non-contingent leverpressing was used to directly verify that the three treatments—forward signaling, noncontingent food, and backward signaling—differentially influenced contextual conditioning. Forward signaling produced the least, and backward signaling produced the most, contextual conditioning. This monotonic relationship between contextual conditioning and interference with extinction was used as evidence to support the argument that context-food associations are important in controlling instrumental responding.     

Nonassociative habituation,US preexposure,and backward inhibitory conditioning with signaled and unsignaled USs

In a conditioned suppression experiment, rats received a single, massed session of conditioning in which one backward conditioned inhibitory stimulus (CS-) followed shocks that were signaled by a visual cue, and a second backward CS-followed shocks that were unsignaled. Conditioning was preceded by a preexposure phase in which some groups of rats were preexposed to unsignaled shock, while others were not preexposed and remained in the experimental apparatus in the absence of shock. The groups were further distinguished by whether US preexposure and conditioning occurred in the same or different contexts, and by whether conditioning began immediately or after a 24-h rest period in the home cage. Although the conditioning itself was effective in establishing the visual cue as a conditioned excitor in the nonpreexposed groups, it was not effective in establishing the two backward cues as reliable inhibitors with either signaled or unsignaled USs. After 210 US preexposures, however, the same conditioning sessions did yield conditioned inhibition to both CS-s. A 24-h rest period in the home cage reduced the magnitude of, but did not completely abolish, the facilitative effect of US preexposure on inhibitory conditioning. Other tests demonstrated that US preexposure had retarded excitatory conditioning to the visual cue. This interference with excitatory conditioning was unchanged in magnitude after the 24-h rest period. The facilitative effect of US preexposure on backward inhibitory conditioning, and the interference effect on excitatory conditioning, were both eliminated by a change in context between US preexposure and conditioning. These observations encourage predominantly associative accounts of the effects, but allow for a small nonassociative habituation component.