Appetitive and aversive olfactory learning induce similar generalization rates in the honey bee

Appetitive and aversive learning drive an animal toward or away from stimuli predicting reinforcement, respectively. The specificity of these memories may vary due to differences in cost–benefit relationships associated with appetitive and aversive contexts. As a consequence, generalization performances may differ after appetitive and aversive training. Here, we determined whether honey bees show different rates of olfactory generalization following appetitive olfactory conditioning of the proboscis extension response, or aversive olfactory conditioning of the sting extension response. In both cases, we performed differential conditioning, which improves discrimination learning between a reinforced odor (CS+) and a non-reinforced odor (CS?) and evaluated generalization to two novel odors whose similarity to the CS+ and the CS? was different. We show, given the same level of discriminatory performance, that rates of generalization are similar between the two conditioning protocols and discuss the possible causes for this phenomenon.     

Fatty acid amide hydrolase (FAAH) knockout mice exhibit enhanced acquisition of an aversive, but not of an appetitive, Barnes maze task

It is well established that genetic deletion or pharmacological inhibition of the CB₁ receptor disrupts extinction learning in aversive conditioning tasks, but not in appetitive tasks. Consistent with these findings is that genetic deletion or pharmacological inhibition of fatty acid amide hydrolase (FAAH), the primary catabolic enzyme of the endogenous cannabinoid anandamide (AEA), accelerates acquisition as well as extinction in aversive conditioning tasks. However, it is unknown whether FAAH blockade will affect acquisition in an appetitive conditioning task. Therefore, in the present study, we assessed FAAH (−/−) and (+/+) mice in appetitive and aversive Barnes maze conditioning procedures. Here we report that FAAH (−/−) mice displayed accelerated acquisition rates in an aversively-motivated, but not in the appetitively-motivated, Barnes maze task. The CB₁ receptor antagonist, rimonabant attenuated enhanced acquisition in the aversive procedure, consistent with the idea that elevated AEA levels mediate this apparent nootropic effect. These findings support the hypothesis that stimulation of the endocannabinoid system enhances learned behavior in aversive, but not appetitive, conditioning paradigms.     

Asymmetrical effects of Pavlovian excitatory and inhibitory aversive transfer on Pavlovian appetitive responding and acquisition

Predictions of a theory of Pavlovian motivational transfer, which incorporates principles of both the theory of reciprocal inhibition and the Rescorla-Wagner model, were tested in several Pavlovian aversive to Pavlovian appetitive transfer tasks. As predicted, the presence of a signal for an aversive event, conditioned stimulus (AV CS+), reliably suppressed performance of appetitive conditioned responses (CRs) whether imposed during acquisition or on independently established responding. Acquisition of appetitive responding to a novel CS reinforced in compound with an AV CS+, however, was enhanced (“superconditioning”). This observation suggests that the effects of a discrepancy between expectation and actual outcome on a conditioning trial are influenced by the affective value of both the expectation and the reinforcer. These transfer effects were not symmetrical for an inhibitory aversive stimulus (AV CS?). An AV CS? did not enhance appetitive responding compared to a random control condition, nor did the AV CS? reduce (i.e., block) appetitive conditioning to a novel CS when appetitive reinforcement occurred in the presence of the AV CS?. Comparison of the two shock-exposed conditions with a naive control condition suggests that previous results that were apparently consistent with inhibitory aversive enhancement and blocking of appetitive conditioning may have been due to aversive context conditioning.     

The effect of Pavlovian stimuli on the acquisition of a new response

Three experiments explored the effects of response dependent stimuli which have had different types of contingent relationships with positive reinforcement. Results showed consistent differences in the effects of response dependent appetitive Pavlovian CS + s, CS ? s, and truly random control (TRC) stimuli on the acquisition of a new response. These differences were shown in both a free operant and a discrete-trial learning situation. Appetitive Pavlovian CS?s consistently produced lower levels of responding than did TRC stimuli, even in an experimental situation free of possible contrast effects. The results of these experiments extended a contingency analysis of acquired associations to appetitive conditioning and provided a clear demonstration of a conditioned reinforcement effect against a variety of control procedures.     

Differential involvement of the central amygdala in appetitive versus aversive learning

Understanding the function of the distinct amygdaloid nuclei in learning comprises a major challenge. In the two studies described herein, we used c-Fos immunolabeling to compare the engagement of various nuclei of the amygdala in appetitive and aversive instrumental training procedures. In the first experiment, rats that had already acquired a bar-pressing response to a partial food reinforcement were further trained to learn that an acoustic stimulus signaled either continuous food reinforcement (appetitive training) or a footshock (aversive training). The first training session of the presentation of the acoustic stimulus resulted in significant increases of c-Fos immunolabeling throughout the amygdala; however, the pattern of activation of the nuclei of the amygdala differed according to the valence of motivation. The medial part of the central amygdala (CE) responded, surprisingly, to the appetitive conditioning selectively. The second experiment was designed to extend the aversive versus appetitive conditioning to mice, trained either for place preference or place avoidance in an automated learning system (INTELLICAGE). Again, much more intense c-Fos expression was observed in the medial part of the CE after the appetitive training as compared to the aversive training. These data, obtained in two species and by means of novel experimental approaches balancing appetitive versus aversive conditioning, support the hypothesis that the central nucleus of the amygdala is particularly involved in appetitively motivated learning processes.     

Partial reinforcement effects in instrumental escape conditioning

In Experiment I acquisition and extinction of instrumental escape conditioning with rats (N = 64) were studied as a function of reinforcement magnitude under conditions of partial and continuous reinforcement. In Experiment II the effects of partial and continuous reinforcement were studied in rats (N = 96) during acquisition followed by small, medium, and large reductions in reinforcement magnitude. A water-tank escape apparatus was used with temperature as the relevant variable. It was found that (1) with large reinforcement magnitude a continuously reinforced group was superior in acquisition to one that was partially reinforced; there were no differences with small reinforcement; (2) disruptive effects of a nonreinforced trial (a) appear early in learning, (b) are quite strong after each nonreinforced trial, and (c) persist through several succeeding reinforced trials; (3) major competing behaviors persist throughout acquisition for small reinforcement magnitude regardless of schedule, decline with large reinforcement (more so with continuous than with partial), and return to a high level in extinction for all conditions; (4) the partial reinforcement extinction effect occurs after large reinforcement but not after small, and it appears only with large reductions in reinforcement magnitude which approach extinction conditions. Only the first part of the last finding appears to be consistent with the appetitive conditioning literature.     

Effects of an infusion of morphine into the accumbens nucleus upon acquisition of hypoalgesic and fear responses in rats exposed to a heat stressor

Four experiments examined the effects of an infusion of morphine into the accumbens nucleus upon the aversive conditioning that can occur in rats exposed to the heated floor of a hot-plate apparatus. An infusion of morphine into the accumbens nucleus but not into the caudoputamen or into the prefrontal cortex impaired the acquisition of a conditioned hypoalgesic (Experiment 1) and fear (Experiment 4) response. This impairment was dose-dependent (Experiment 2) and mediated by opioid receptors in the accumbens nucleus, because it was removed by a systemic (Experiment 3a) or by an accumbal (Experiment 3b) infusion of naloxone. The results were attributed to an antagonism between the reinforcement process for aversive conditioning and the appetitive properties of an accumbal infusion of morphine.     

Transfer of conditioned appetitive stimuli to conditioned aversive excitatory and inhibitory stimuli

The transfer of Pavlovian appetitive stimuli to Pavlovian aversive stimuli was examined in three experiments. In Experiment 1, rats received appetitive (Ap) conditioning designed to establish a flashing-light stimulus as either a CS+, CSo, or CS? for food, or to maintain it as a novel stimulus for US-alone subjects. Then, the stimulus was employed as a signal for weak shock in conditioned-emotional-response (CER) training. Both acquisition and extinction results showed that the ApCS+ facilitated and the ApCS? retarded aversive excitatory conditioning relative to the ApCSo and US-alone controls. Experiment 2 replicated the findings of Experiment 1 with both a moderate and a severe shock in CER training. In Experiment 3, different groups received the same appetitive conditioning as before, but to a flashing-light stimulus which was then employed as a signal for no shock in CER training. The ApCS? facilitated and the ApCS+ retarded aversive inhibitory conditioning relative to ApCSo and US-alone controls. Collectively, these findings establish that, in Pavlovian conditioning, transfer of an appetitive CS to an aversive excitor or inhibitor is facilitated by maintaining the initial conditioning contingency.     

Acquisition and discrimination of appetitively and aversively conditioned heart rate responses in rhesus monkeys

This report describes the acquisition of a conditional heart rate response to both classical aversive and appetitive conditioning in eight rhesus monkeys(Macaca mulatta). The behavioral paradigm consisted of two discrete one minute tones followed by the corresponding unconditional stimuli,i.e., electric shock or Purina monkey chow. A third tone followed by neither food nor shock served as a discriminative stimulus (DS). The conditional heart rate responses developed in two phases. The first phase was characterized by similar responses to both conditional stimuli and the DS. Control heart rate rose during this period. During the second phase, control heart rate decreased (five animals), the DS response disappeared, and different conditional heart rate patterns to food and shock emerged. The existence of distinct conditional response patterns indicates probable differences in the autonomic nervous regulation of the heart during aversive and appetitive conditioning.     

The effects of language conditioning on various aspects of anxiety

Staats' three-function learning theory provides the basis for investigating the effects of emotionally-relevant self-verbalizations (SV) on the physiological, subjective-affective and behavioural aspects of anxiety. Using aversive electric UCSs and slides of snakes (CS), anxiety was classically conditioned in 88 volunteer Ss. In 20 subsequent language-conditioning trials (without aversive electric UCSs), the same snakes slides were paired with UCS verbalizations having either positive or negative connotative meanings. Half of the Ss were exposed to a living snake prior to language conditioning. The results show a complete extinction of the conditioned anxiety response in groups with positive SV whereas negative SV impeded extinction; the latter effect could only be found in groups without exposure to snakes prior to language conditioning. In general, the affective evaluation of snakes improved in groups with positive SV and deteriorated in groups with negative SV. However, these effects were more pronounced in groups without exposure to snakes. Although the results indicate that Ss in all groups with positive SV exhibited more approach behaviours than Ss with negative SV, this trend was not statistically significant. The possible relevance of the results for a language-conditioning approach to anxiety reduction is briefly discussed.     

Defining reinforcess — A problem in communication for consultation in behavior modification

This paper reviews the respondent (Hull-Spence) and operant (Skinnerian) conditioning definitions of reinforcers and reinforcement and demonstrates the need to keep the systems separate when consulting about behavior modification. The two systems are shown to lead to different modification procedures.One important distinction between the systems is whether a reinforcer is simply associated with a response (respondent) or whether it must follow the response (operant). A second important distinction is the definition of negative reinforcement. In respondent conditioning, negative reinforcement entails presenting an aversive stimulus in association with the response and results in a decrease in response rate. In operant conditioning, negative reinforcement entails the removal of an aversive stimulus following a correct response, which results in an increase in response rate.     

Nitric oxide is involved in appetitive but not aversive olfactory learning in the land mollusk Limax valentianus

The land slug Limax performs both aversive and appetitive olfactory learning, and we investigated neurotransmitters involved in each type of learning. Slugs were conditioned by presenting a vegetable juice (appetitive conditioning) or a mixture of vegetable juice and quinidine (aversive conditioning), and the latency to reach the juice became shorter (appetitive conditioning) or longer (aversive conditioning) after conditioning. L-NAME injected either before conditioning or testing blocked the reduction in latency in appetitive conditioning but had no significant effects in aversive conditioning. 5,7-dihydroxytryptamine had no significant effects in appetitive conditioning. These results suggest different mechanisms for appetitive and aversive learning.     

Effects of schedule and magnitude of reinforcement under conditions of thirst motivation

In contrast to a recent finding (Macdonald, G. E., & De Toledo, L. Learning and Motivation, 1974, 5, 288–298.) the results of three experiments investigating various partial reinforcement (PRF) manipulations under conditions of thirst motivation demonstrated strong similarity to analogous manipulations involving food reward. Specifically, for animals receiving water reinforcement, PRF was shown to generate greater resistance to extinction than continuous reinforcement (Expt 1 & 3), the schedule of reinforcement was shown to interact with level of acquisition (Expt 1 and 2), and the magnitude of the partial reinforcement extinction effect was shown to be a function of reward magnitude (Expt 3). These results provide strong evidence that mechanisms which operate in partial reinforcement situations are highly similar, regardless of the type of appetitive reinforcement.     

Aversive and appetitive electrodermal classical conditioning using pictures as stimuli

The principal goal of this study was to verify whether it was possible to obtain both aversive and appetitive electrodermal classical conditioning, using pictures as conditioned stimuli (CS), and unconditioned stimuli (US). Additionally, we tried to verify whether, as a consequence of such conditioning, diminution of the unconditioned response (UR) was observed. With this aim, IAPS (?International Affective Picture System?) pictures were selected as stimuli. A picture showing a burnt face was used as the aversive US (USav), and a picture showing a scene with erotic content was used as the appetitive US (USap). As the aversive CS (CSav), and appetitive CS (CSap), two images with intermediate values of valence and arousal showing male faces were selected. In the experimental group, 10 CSav/USav and 10 CSap/USap trials were presented. In the control group 10 CSav, CSap, USav, and USap trials were presented in pseudorandom order. Skin conductance response (SCR) elicited by both the CSs and the USs was scored. Results showed aversive conditioning, but neither appetitive conditioning nor UR diminution. Problems to obtain conditioning using pictures as stimuli and possible options to overcome them in future research are discussed.     

Impaired appetitively as well as aversively motivated behaviors and learning in PDE10A-deficient mice suggest a role for striatal signaling in evaluative salience attribution

Phosphodiesterase 10A (PDE10A) hydrolyzes both cAMP and cGMP, and is a key element in the regulation of medium spiny neuron (MSN) activity in the striatum. In the present report, we investigated the effects of targeted disruption of PDE10A on spatial learning and memory as well as aversive and appetitive conditioning in C57BL/6 J mice. Because of its putative role in motivational processes and reward learning, we also determined the expression of the immediate early gene zif268 in striatum and anterior cingulate cortex. Animals showed decreased response rates in scheduled appetitive operant conditioning, as well as impaired aversive conditioning in a passive avoidance task. Morris water maze performance revealed not-motor related spatial learning and memory deficits. Anxiety and social explorative behavior was not affected in PDE10A-deficient mice. Expression of zif268 was increased in striatum and anterior cingulate cortex, which suggests alterations in the neural connections between striatum and anterior cingulate cortex in PDE10A-deficient mice. The changes in behavior and plasticity in these PDE10A-deficient mice were in accordance with the proposed role of striatal MSNs and corticostriatal connections in evaluative salience attribution.     

Acquisition of cup drinking using previously refused foods as positive and negative reinforcement

We used previously refused foods as positive and negative reinforcement in the acquisition of cup drinking. Cup drinking increased with positive and negative reinforcement, both alone and in combination (without escape extinction), indicating that treatment of food refusal can establish some foods as appetitive stimuli whereas others remain aversive.     

Cardiac arrhythmias in the monkey during classically conditioned fear and excitement

Rhesus monkeys(Macaca mulatto) were tested in classic aversive and appetitive conditioning paradigms following complete coronary artery occlusion (CO) to test the hypothesis that “emotional stress” induces ventricular arrhythmias. Findings were based upon conditioning trials conducted for one or more weeks after occlusion in 13 animals. When all data from each animal for the week following CO were considered, there was no demonstrable tendency for arrhythmias to increase during “fear” conditioned to unavoidable electric shock or during “excitement” in anticipation of food. However, selected trials from six monkeys did reveal instances when changes in the frequency of occurrence of arrhythmias were coupled with behavioral conditioning. While analysis of these trials did not reveal any simple relationship between emotional stress and the development of ventricular arrhythmias after myocardial infarction, certain of the behavioral situations may be more potentially arrhythmogenic than others. For these selected trials, with respect to control, the number of arrhythmias may have increased or decreased upon presentation of the conditional stimulus; the exact response appears to depend upon the immediate physiologic status of the animal as well as on the behavioral condition. “More stressful” situations, such as aversive conditioning, are not necessarily associated with greater numbers of arrhythmias than were “less stressful” situations, such as appetitive conditioning. Arrhythmias appear to occur more frequently when an animal’s heart rate is within a given range; this may reflect underlying cardiac sympathetic and parasympathetic nerve activity.     

Sensitization and aversive conditioning: Effects on the startle reflex and electrodermal responding

Animal data suggest that shock sensitization as well as aversive learning potentiates the acoustic startle reflex. The present experiment tested, whether this shock sensitization also occurs in human subjects and whether it precedes aversive conditioning. Sixty subjects viewed—prior to conditioning—a series of slides of different emotional contents including the to be conditioned stimuli (CSs). Afterwards, the experimenter attached the shock electrodes and initiated shock exposure. Then, subjects were randomly assigned to view a series of two slides, each for eight acquisition trials in which one slide was followed by a shock. Subsequently, extinction trials (12 for each slide) were administered. During preconditioning, acquisition, and extinction, startle probes occurred unpredictably during and between slide viewing. Preconditioning data replicated previous results by Lang and his associates, showing that the startle response magnitude is directly related to the affective valence induced by the slides. Shock exposure strongly facilitated the startle reflex magnitude. This shock sensitization was absent for the skin conductance response. Course of learning also varied for both response systems. The data suggest that startle reflex potentiation indexes the acquisition of an avoidance disposition, which is preceded by a general sensitization of the protective reflexes. Skin conductance learning follows arousal changes and is modulated by cognitive processes.     

Cardiovascular dynamics during classical appetitive and aversive conditioning in laboratory primates

This report describes changes in the rate of rise of left and right intraventricular pressures during aversive and appetitive conditioning procedures in chair-restrained rhesus monkeys. The conditioning paradigm consisted of a one-minute tone followed, in the one case, by an electric shock, and in the other, by the delivery of Purina monkey pellets. The conditional cardiovascular response was characterized by short latency, highly significant elevations in the derivatives of both ventricular pressures as well as a marked arterial pressor response and tachycardia. The magnitude of the conditional response to the classical aversive procedure was somewhat larger than that to appetitive conditioning. These alterations in the rate of development of intraventricular pressure can be attributed largely to augmentation in the sympathetic neural input to the heart and contribute to an analysis of selective aspects of the nervous regulation of the heart in intact, behaviorally conditioned animals.     

Response-outcome contingency: Behavioral and judgmental effects of appetitive and aversive outcomes with college students

In two experiments, positive, negative, and zero response-outcome contingencies were responded to and rated by college students under a free-operant procedure. In Experiment 1, outcomes were either neutral or were associated with point gain. In Experiment 2, subjects were administered different outcome treatments: neutral outcomes, outcomes associated with money gain, or outcomes associated with money loss. In both experiments, subjects' judgments of response-outcome contingency and their operant responses were each strong linear functions of ΔP, the difference between the probability of an outcome given a response and the probability of an outcome given no response. Appetitive and aversive outcomes produced opposite and symmetrical response patterns. In Experiment 1, no differences in ratings occurred with neutral or appetitive outcomes; however, in Experiment 2, more potent appetitve outcomes led to somewhat more extreme ratings than either neutral or aversive outcomes. Increasing outcome probability produced only a slight bias in ratings of noncontingent problems in Experiment 1 and no bias in Experiment 2. Contrary to predictions derived from an analysis of superstitious behavior, increasing outcome probability in noncontingent problems decreased operant responding when outcomes were appetitive and increased operant responding when outcomes were aversive. Trend analyses revealed that Δ P was superior to several other metrics in predicting subjects' estimates of contingency and the behavioral effects of contingency. Operant responding was in closer accord with matching predictions than with maximizing predictions.