Goal-tracking behavior in the pond snail, Lymnaea stagnalis

The occurrence of goal-tracking, an unconditioned stimulus (US)-directed autoshaping behavior, was studied in open-field tests with control and classically conditioned pond snails, Lymnaea stagnalis. In an appetitive classical conditioning paradigm with a tactile stimulus as conditioned stimulus (CS) and a localized food stimulus as US a conditioned feeding response built up in the experimental but not in the control animals. In the post-training open-field tests the experimental group alone showed an enhanced attraction toward the source of water current in the environment which previously signalled the arrival of the US but did not act as CS in the classical conditioning procedure. We suggest that this stimulus-directed goal-tracking behavior in Lymnaea is the result of a classical-operant interaction, described so far only in vertebrate animals, and that neurophysiological analysis of this behavior is possible in this snail.     

Contextually Based Conditional Discrimination of the Rabbit Eyeblink Response

Rabbits received conditional discrimination training using contextual stimuli to set the occasion for stimulus pairings during eyelid conditioning. Specifically, animals were exposed to either the presence or the absence of an oscillating chamber light throughout the intertrial interval (50 ± 10 s). For half the animals, this light signaled paired presentations of a discrete tone conditioned stimulus (CS) and air puff unconditioned stimulus (US) while darkness signaled presentations of only the tone CS. The remaining animals experienced the opposite contextual relationship to the conditioning stimuli. These trial types occurred pseudo-randomly across a session, with all transitions between contextual settings (i.e., light or dark) taking place immediately at the CS–US offset. Under these conditions, animals successfully utilized the contextual stimuli as conditional cues for differential responding to the shared CS. Moreover, both light and dark were equally effective as discriminative stimuli. A subset of animals received further training in which the contextual contingency was removed by restricting all conditioning to the CS-alone context. Without the contingency in place, subsequent CS presentations (paired and CS-alone) evoked equivalent conditioned responding across three sessions of training. Following the reinstatement of the contextual contingencies, discriminatory responding was immediately observed and returned to previous levels within three sessions. Finally, animals appeared to use the static representation of the conditional cue, rather than the phasic transition between cues, for discriminatory responding. These findings are discussed in terms of current neurobiological models of eyelid conditioning.     

The potentiation effect during serial conditioning

The development of suppression in rats to a target conditioned stimulus (CS) was compared in trace and serial conditioning procedures. The interval between the end of the target CS and the shock unconditioned stimulus (US) was filled by a second CS in the serial, but not the trace, procedure. In five experiments the serial procedure produced superior conditioning. This potentiation effect, however, depended critically upon the level of conditioning to the stimulus interpolated between the target CS and the US. When conditioning to the interpolated CS was either reduced by giving independent nonreinforced trials with this CS alone or enhanced by independent reinforced trials, the potentiation effect was abolished. In addition, the insertion of a trace interval between the target and interpolated CSs reduced the effect. However, the magnitude of conditioning to the target CS was unaffected by post-conditioning changes in the conditioned strength of the interpolated CS. These findings are discussed in terms of the contribution of both the association between the CSs themselves, which is inherent in the serial procedure, and that between the target CS and the US.     

A new tool for assessing context conditioning induced by US-unpredictability in humans: The Martians task restyled

Unpredictability of an unconditioned stimulus (US) typically produces context conditioning in animals and humans. We modified the Martians task - a computer game measuring learning of Pavlovian associations through conditioned suppression - for assessing context conditioning in humans. One between-subjects and one within-subjects study are reported. Both experiments comprised four conditions: a predictable (Paired) condition in which the conditioned stimulus (CS) signaled the US, a neutral condition (No-US), one unpredictable condition in which the CS did not signal the US (Unpaired) and another one in which only unsignaled, temporally unpredictable USs were presented (US-only). Results showed more contextual conditioned suppression in the unpredictable conditions compared to the predictable condition. In contrast, more cued conditioned suppression occurred in the Paired condition than in the Unpaired condition. Consistent with animal research, context conditioning was increased by unpredictability. These data support the Martians task as a promising tool to extend research on human context conditioning.     

Conditioned taste aversion is enhanced when the unconditioned stimulus is presented in a different context

Using a conditioned taste aversion procedure with rats as the subjects, two experiments examined the effect of presenting a conditioned stimulus (CS saccharin solution) in one context followed by an unconditioned stimulus (US LiCl) in a different context. Experiment 1 showed that animals which received the above-mentioned procedure (Group D) showed a more marked conditioned aversion to the CS than animals which were given both the CS and the US in the same context (Group S). Experiment 2 found that in both Group D and Group S, aversion to the CS increased when the subjects were exposed to the conditioned context after the conditioning. These findings supported the argument that the strength of the CS-US association acquired during conditioning is compared with that of the context-US to determine the magnitude of aversion revealed to the CS.     

Trace and delay eyeblink conditioning: contrasting phenomena of declarative and nondeclarative memory

We tested the proposal that trace and delay eyeblink conditioning are fundamentally different kinds of learning. Strings of one, two, three, or four trials with the conditioned stimulus (CS) alone and strings of one, two, three, or four trials with paired presentations of both the CS and the unconditioned stimulus (US) occurred in such a way that the probability of a US was independent of string length. Before each trial, participants predicted the likelihood of the US on the next trial. During both delay ( n =20) and trace ( n = 18) conditioning, participants exhibited high expectation of the US following strings of CS-alone trials and low expectation of the US following strings of CS-US trials a phenomenon known as the gambler's fallacy. During delay conditioning, conditioned responses (CRs) were not influenced by expectancy but by the associative strength of the CS and US. Thus, CR probability was high following a string of CS-US trials and low following a string of CS-alone trials. The results for trace conditioning were opposite. CR probability was high when expectancy of the US was high and low when expectancy of the US was low. The results show that trace and delay eyeblink conditioning are fundamentally different phenomena. We consider how the findings can be understood in terms of the declarative and nondeclarative memory systems that support eyeblink classical conditioning.     

The effects of lesions in the central nucleus of the amygdala on appetitive classical conditioning in rats

Abstract: Lesions in the central nucleus of the amygdala (cAMY) have been known to interfere with the acquisition of fear classical conditioning when footshock is used as an unconditioned stimulus (US). The present study examined whether or not a similar interference would occur with an appetitive US. Five rats with lesions in the cAMY (the cAMY group), and eight unoperated control rats were trained in an appetitive classical conditioning paradigm, which did not include elements of operant learning, using a visual conditioned stimulus (CS) (5 W of light for 10 s duration) paired with a food pellet US (45 mg, cheese flavor). The behavioral index of appetitive conditioning was an increase in rearing approach behavior to the CS after CS and US pairings. During CS and US pairings, the movement of the rat was limited so that this approach behavior could not occur. As a result, all control rats showed an increase in rearing, but the cAMY group did not. These results suggest that the cAMY is critical for appetitive as well as fear classical conditioning.     

Generalisation of fear and avoidance along a semantic continuum

Directly conditioned fear and avoidance readily generalises to dissimilar but conceptually related stimuli. Here, for the first time, we examined the conceptual/semantic generalisation of both fear and avoidance using real words (synonyms). Participants were first exposed to a differential fear conditioning procedure in which one word (e.g., “broth”; CS+) was followed with brief electric shock [unconditioned stimulus (US)] and another was not (e.g., “assist”; CS–). Next, an instrumental conditioning phase taught avoidance in the presence the CS+ but not the CS–. During generalisation testing, synonyms of the CS+ (e.g., “soup”; GCS+) and CS– (e.g., “help”; GCS–) were presented in the absence of shock. Conditioned fear and avoidance, measured via skin conductance responses, behavioural avoidance and US expectancy ratings, generalised to the semantically related, but not to the semantically unrelated, synonyms. Findings have implications for how natural language categories and concepts mediate the expansion of fear and avoidance repertoires in clinical contexts.     

Simultaneous behavioral and neural (cochlear nucleus) measurement during signal detection in the rabbit

Classical conditioning of the rabbit nictitating membrane (NM) response was adapted as a model system for simultaneous measurement of behavioral and neural responses in auditory signal detection. Animals were trained to a 36-dB spectral-level white-noise CS paired with a corneal airpuff US. The CS intensity was then reduced to threshold, using a modified staircase procedure, and responses on detect and nondetect trials were compared for the same stimulus intensities. At threshold, the behavioral NM response was dichotomous, being present on detect trials and completely absent on nondetect trials. In marked contrast, evoked neural unit activity to the CS recorded from the anteroventral cochlear nucleus was clearly present and identical on both detect and nondetect trials, suggesting that the detection “decision” is made more centrally than the cochlear nucleus.     

Occasional reinforced trials during extinction can slow the rate of rapid reacquisition

Two appetitive conditioning experiments with rats examined reacquisition after conditioned responding was eliminated by either extinction or by a partial reinforcement procedure in which reinforced trials were occasionally presented among many nonreinforced trials. In Experiment 1, reacquisition to a conditional stimulus (CS) that had been conditioned and extinguished was more rapid than acquisition in a group that had received no prior conditioning. However, the addition of occasional reinforced trials to extinction slowed this rapid reacquisition effect. Experiment 2 replicated the result and showed that a procedure in which the CS and the unconditional stimulus (US) were unpaired in extinction interfered even further with reacquisition. The results suggest that rapid reacquisition is ordinarily produced when reinforced trials provide a contextual cue that can renew responding by signaling other acquisition trials (Ricker & Bouton, 1996). The effects of partial reinforcement in extinction are surprising from several theoretical perspectives and have useful clinical implications.     

Signal-centered action patterns of dogs in appetitive classical conditioning

The movements of five unrestrained dogs were monitored during discriminative Pavlovian conditioning. An auditory-visual stimulus from one source (CS+) was followed by meat morsels (US); the same stimulus from another source (CS?) was not. Sources were equidistant from the site of US delivery. Before each trial, animals were required to position themselves at a starting location equidistant from both sources and removed from the US site. The stable behavior pattern in most subjects included approach to and contact or near contact with CS+, followed by approach to the US site. Dogs showed individually distinctive action patterns to CS+, in some cases suggestive of soliciting, in another of sight-pointing. Similar actions were not evoked by CS?. In postacquisition tests, patterns were generally unaffected by increased deprivation or by relocating the starting position next to the US site. The auditory component of CS+ was more effective than the visual. With a response-contingent procedure animals were successfully trained to approach the US site directly, although many trials, and in one case a special procedure, were required. It was proposed that the experimental signaling arrangement mimics a natural signaling sequence and induces appetitive behavior to the CS that corresponds to the behavior induced by certain natural signals of the US.     

Compound conditioning: Elimination of the blocking effect

Prior conditioning employing one element of a compound stimulus as the CS blocked the acquisition of a Conditioned Emotional Response (CER) to the second element of a simultaneous compound stimulus that was subsequently used as the CS in further conditioning trials. Presentation of a brief “surprise” stimulus 3 or 5 sec after the occurrence of the US during the compound trials eliminated the blocking effect.     

Galantamine facilitates acquisition of a trace-conditioned eyeblink response in healthy, young rabbits

Previous work has demonstrated that drugs increasing brain concentrations of acetylcholine can enhance cognition in aging and brain-damaged organisms. The present study assessed whether galantamine (GAL), an allosteric modulator of nicotinic cholinergic receptors and weak acetylcholinesterase inhibitor, could improve acquisition and retention of an eyeblink (EB) classical conditioning task in healthy, young animals. We trained 24 rabbits (n = 8/group) in a 1000-msec trace Pavlovian EB conditioning paradigm in which a tone conditioned stimulus (CS) was presented for 500 msec, followed by a 500-msec trace period in which no stimuli were presented. A 100-msec corneal airpuff was the unconditioned stimulus (US). Acquisition sessions, consisting of 100 trials each, occurred daily for 10 consecutive days, followed by 3 d of extinction training. Animals were treated with one of three doses of GAL (0.0-3.0 mg/kg) prior to each session. Animals that received 3.0 mg/kg GAL showed significantly more EB conditioned responses (CRs) in fewer training trials than animals receiving either 1.5 mg/kg GAL or vehicle injections. GAL had no effect on CR performance during extinction. Pseudoconditioning control experiments, consisting of 200 explicitly unpaired tone-puff presentations indicated that GAL did not increase reactivity to the CS or US. These findings indicate that GAL may improve acquisition of moderately difficult associative learning tasks in healthy young organisms.     

Classical Delay Eyeblink Conditioning in in 4- and 5-Month-Old Human Infants

Abstract-Simple delay classical eyeblink conditioning, using a tone conditioned stimulus (CS) and airpuff unconditioned stimulus (US), was studied in cross-sectional samples of 4- and 5-month-old healthy, full-term infants. Infants received two identical training sessions, 1 week apart. At both ages, infants experiencing paired tones and air-puffs demonstrated successful conditioning over two sessions, relative to control subjects who had unpaired training. Conditioning was not evident, however, during the first session. Two additional groups of 5-month-olds received varied experiences during Session 1, either unpaired presentations of the CS and US or no stimulus exposure, fol-lowed by paired conditioning during Session 2. Results from these groups suggest that the higher level of conditioning observed following two sessions of paired conditioning was not the result of familiarity with the testing environment or the stimuli involved but, rather, the result of retention of associative learning not expressed during the first conditioning session.     

Trace fear conditioning is reduced in the aging rat

Auditory trace fear conditioning is a hippocampus-dependent learning task that requires animals to associate an auditory conditioned stimulus (CS) and a fear-producing shock-unconditioned stimulus (US) that are separated by an empty 20-s trace interval. Previous studies have shown that aging impairs learning performance on hippocampus-dependent tasks. This study measured heart rate (HR) and freezing fear responses to determine if aging impairs hippocampus-dependent auditory trace fear conditioning in freely moving rats. Aging and Young rats received one long-trace fear conditioning session (10 trials). Each trial consisted of a tone-CS (5 s) and a shock-US separated by an empty 20-s trace interval. The next day rats received CS-alone retention trials. Young rats showed significantly larger HR and freezing responses on the initial CS-alone retention trials compared to the Aging rats. A control group of aging rats received fear conditioning trials with a short 1-s trace interval separating the CS and US. The Aging Short-Trace Group showed HR and freezing responses on the initial CS alone retention trials that were similar to the Young Long-Trace Group, but greater than the Aging Long-Trace Group. A second aging control group received unpaired CSs and USs, and showed no HR or freezing responses on CS-alone retention trials. These data show that HR and freezing are effective measures for detecting aging-related deficits in trace fear conditioning.     

Autonomic nervous control of heart rate orienting and alpha responses in dog

The object of this experiment is to study the autonomic nervous control of alpha responses elicited in classical conditioning. Twenty-two mongrel dogs were trained in classical discriminative conditioning. Typical two-phase tachycardic responses were observed in positive (CS+) trials while only the earliest, phase 1, response was observed in negative (CS-) trials. The phase 1 response, which was identical in CS+ and CS-trials, was compared in dogs before and after selective SA-nodal parasympathectomy (N=7) and beta-adrenergic blockade (N=11). The phase 1 tachycardic response was eliminated by selective SA-nodal parasympathectomy but not by beta-adrenergic blockade. We conclude that the phase 1 response observed in both CS+ and CS-trials with similar time sequence and magnitude is an alpha response. The heart rate orienting response results from a withdrawal of parasympathetic activity with little or no change in sympathetic tone.     

Similarity of cardiac CR forms in the rhesus monkey during several experimental procedures

Cardiac rates of rhesus monkeys were observed in a variety of different conditioning procedures, each of which involved a visual stimulus (CS) followed by an electric shock (US). With a 30-sec CS, cardiac rate accelerated rapidly after CS onset, reached a maximum in the middle of CS, and decelerated thereafter, with a terminal CS rate often at the level of, or below, pre-CS levels. A similar biphasic cardiac rate response in CS was also observed under subsequent exposure to intermittent pairings of CS and US, avoidance of US, response-produced termination of US, and when CS-US pairings were superimposed upon an avoidance baseline, even when CS duration was varied from 12 to 60 seconds. The unusual regularity of cardiac rate responses in several different procedures may result from one or more of these factors: (a) characteristics of the rhesus monkey, (b) initial exposure to Pavlovian conditioning, or (c) the uniformity of measurement of cardiac rate employed in this study.     

Timed excitatory conditioning under zero and negative contingencies

Rats (rattus norvegicus) anticipated the arrival of a food pellet unconditioned stimulus (US) even when the conditioned stimulus (CS) signaled no overall change or a substantial decrease in the overall rate of US occurrence. Pellet USs were scheduled probabilistically in the intertrial interval at either an equivalent rate (Experiment 1) or a four times higher rate (Experiments 2 and 3) than in the CS, which included one fixed-time target US. Conditioning has been said to involve learning "whether" (contingency) the CS signals a change in the US, and if so, "when" (contiguity) the US is scheduled to arrive. Our results suggest that "when" trumps "whether," challenging the received view that a positive CS-US contingency is necessary for successful conditioning.