Reinforcement in an in vitro analog of appetitive classical conditioning of feeding behavior in Aplysia: blockade by a dopamine antagonist

In a recently developed in vitro analog of appetitive classical conditioning of feeding in Aplysia, the unconditioned stimulus (US) was electrical stimulation of the esophageal nerve (En). This nerve is rich in dopamine (DA)-containing processes, which suggests that DA mediates reinforcement during appetitive conditioning. To test this possibility, methylergonovine was used to antagonize DA receptors. Methylergonovine (1 nM) blocked the pairing-specific increase in fictive feeding that is usually induced by in vitro classical conditioning. The present results and previous observation that methylergonovine also blocks the effects of contingent reinforcement in an in vitro analog of appetitive operant conditioning suggest that DA mediates reinforcement for appetitive associative conditioning of feeding in Aplysia.     

Internal stimuli enhance feeding behavior in the mollusc Aplysia

The hypothesis that subsatiating levels of internal food stimuli can arouse and potentiate feeding behavior was examined in the mollusc Aplysia californica. Animals were fed a small quantity of seaweed and their latencies to show biting responses were determined after food arousal was permitted to partially decay. Control animals were stimulated with food, but were not permitted to ingest it, or were fed nonnutritive glass-fiber filter paper. Compared to controls, animals that were fed showed significantly shorter latencies to respond when tested up to 80 min after previous exposure to food. These results indicate that internal stimuli can function like external stimuli to enhance responsiveness to food and suggest the hypothesis that satiation may be viewed as an interactive process involving the interplay of excitatory and inhibitory influences arising from the alimentary system.     

Mechanisms underlying satiation of feeding behavior of the mollusc Aplysia

Animals filled almost to satiation by nonnutritive bulk do not satiate when they ingest a small amount of seaweed. This suggests that satiation is not triggered by chemostimulation of an anteriorly located "hot spot." Inflation of a balloon placed in the gut of the animal results in satiation as reflected in a number of different parameters of feeding behavior. The suppressive effect of a relatively brief inflation is rapidly and fully reversible, although repeated inflation and deflation appeared to produce slowly reversible or irreversible effects. The parameters of the changes in feeding during gut inflation are comparable to those of normal animals that are slowly fed individual pieces of food. The inflation volume needed to satiate the animal is a function of the rate of inflation--more rapid inflations requiring larger volumes. Cutting of the esophageal nerves results in a significant increase in the volume needed to satiate the animals, but nevertheless they eventually cease feeding and generally do not show a burst gut. The evidence indicates that the satiation that eventually occurs in nerve-sectioned animals, at least in part, is due to depression of feeding following very prolonged sensory stimulation. The data suggest that for a rapidly consumed meal, satiation results primarily due to distension-related gut signals conveyed by the esophageal nerves, whereas for very slowly consumed meals, the former factor interacts with a process associated with sensory stimulation, such as receptor adaptation. The current results indicate that balloon distension can serve as a reasonable stimulus in experiments in simplified preparations in which the nervous system can be studied.     

Rapid and persistent suppression of feeding behavior induced by sensitization training in Aplysia

In Aplysia, noxious stimuli induce sensitization of defensive responses. However, it remains largely unknown whether such stimuli also alter nondefensive behaviors. In this study, we examined the effects of noxious stimuli on feeding. Strong electric shocks, capable of inducing sensitization, also led to the suppression of feeding. The use of multiple training protocols revealed that the time course of the suppression of feeding was analogous to that of sensitization. In addition, the suppression of feeding was present only at the time points in which sensitization was expressed. These results suggest that, in Aplysia, noxious stimuli may produce concurrent changes in neural circuits controlling both defensive and nondefensive behaviors.     

One-trial appetitive conditioning in the sexual behavior system

One-trial Pavlovian conditioning has been demonstrated with aversive reinforcers such as electric shock and lithium chloride. The present experiments using male Japanese quail were conducted to determine if an appetitive unconditioned stimulus (US), the opportunity to copulate with a female quail, can also generate one-trial learning. In Experiment 1, subjects were exposed to a contextual conditioned stimulus (CS) for 35 sec and then given copulatory opportunity. In Experiment 2, male birds were given 0, 2, or 4 min of exposure to the contextual CS before presentation of a female. In both cases, control groups received comparable exposure to both the contextual CS and the sexual US, but in an unpaired fashion. Subjects were tested for sexual learning by reintroducing them to the experimental context in the presence of a test object that incorporated the taxidermically prepared head of a female quail. In both Experiments 1 and 2, subjects for whom the contextual CS and sexual US had been paired displayed significantly more approach to the test object than did unpaired subjects.     

Effects of conspecifics on feeding in Aplysia fasciata

The presence of other Aplysia in the environment increases the time spent feeding. This finding is consistent with the hypothesis that a common arousal mechanism modulates feeding and reproductive behaviors. The arousal level is presumably increased by pheromones released by additional animals.     

Common regulation of feeding and mating in Aplysia fasciata: pheromones released by mating and by egg cordons increase feeding behavior.

We examined whether pheromones released by reproductive behaviors (mating and egg-laying) affect feeding behavior. A preliminary experiment demonstrated that the quantity of food eaten can be used to measure the effects of pheromones on feeding. Using this measure, we then showed that Aplysia that were prevented from mating, but that were in the same aquarium as mating conspecifics, eat more food than do Aplysia in a medium lacking mating animals. Mating and feeding were not temporally correlated, indicating that pheromones released by mating probably do not initiate feeding, but rather modulate feeding after it has begun. Aplysia that were in the same aquarium as freshly deposited egg cordons also ate more than did animals in a medium lacking eggs.     

The control of feeding behavior by an imprinted stimulus

Imprinted ducklings were trained to peck a pole using brief presentations of the imprinted stimulus as the response-contingent (reinforcing) event. Subjects were then permitted to spend extended periods with continuous access to food and the imprinted stimulus (via a pole peck). For other (control) subjects the experimental situation was restricted to either responding for the stimulus, or feeding in the absence of the stimulus. For subjects in the control conditions, both activities occurred in cyclic fashion. When, however, there was continuous opportunity to respond for the stimulus and food was available, the tendency to respond was related to the tendency to feed. Other experiments showed that independent presentations of the stimulus could initiate feeding in imprinted ducklings with no prior pairing of the stimulus with food and with no prior pole-peck training. The most consistent control over feeding, however, was exhibited by ducklings that were imprinted and also accustomed to periodic removals of the stimulus. It is concluded that in ducklings, imprinting procedures are sufficient to endow an arbitrary stimulus with the capacity to release feeding behavior.     

The effect of contingency upon the appetitive conditioning of free-operant behavior

The problem of maintaining independence between response rates and reinforcement probabilities when determining the effect of varying the response-reinforcement contingency upon free-operant behavior was solved by programming local reinforcement probabilities for response and no response on a second-by-second basis. Fifty-seven rats were trained to lever-press on schedules of water reinforcement involving different values of contingency. All rats were first trained on a high positive contingency and then shifted to less positive, zero, or negative contingencies. Under these conditions, rate of lever-pressing declined appropriately when the contingency between response and reinforcement decreased or was made negative. The decline in rate produced by a zero contingency cannot be attributed to extinction, since the probability of reinforcement given the occurrence of a response was the same as for the positive contingency from which the shift to zero was made. That is, there was no change in the opportunity for response-reinforcement contiguity. It was concluded that the technique of programming local reinforcement probabilities offers promise for more critical examinations of the effects of contingency upon free-operant behavior.     

Facilitation of instrumental behavior by a Pavlovian appetitive conditioned stimulus

Three experiments examined appetitive Pavlovian-instrumental interactions by presenting separately trained conditioned stimuli (CSs) during reinforced instrumental responding in rabbits. Intra-oral reinforcement was used to minimize interference from peripheral responses such as magazine approach. In experiment 1, the rabbits were first trained to perform an instrumental head-raising response for sucrose reward. A conditioned jaw movement response was then established to a 2-sec CS by pairing it with sucrose; a control stimulus was unpaired with sucrose. Instrumental responding maintained by a variable-interval 40-sec schedule was enhanced during 10-sec presentations of the paired, but not the unpaired, CS. Responding on a variable-ratio 15 schedule was unaffected except on trials on which the pre-CS baseline response rate was low; in such cases the paired CS caused a long-lasting acceleration of responding. Noncontingent presentation of the sucrose reinforcer itself briefly suppressed responding but had no long-term effect. In Experiment 2, a CS that had been conditioned at a 10-sec duration produced the same pattern of effects as in the first study, indicating that facilitation resulted from CS presentation rather than from the frustrative effects of non-reinforcement of the CS. In Experiment 3 an inhibitory CS blocked facilitation by the excitatory CS but did not itself affect instrumental responding. These results support the view that Pavlovian processes play a positive role in instrumental performance and suggest that previous findings of suppression by a short-duration CS reflect peripheral interference. The dependence of facilitation on the baseline level of responding is discussed in terms of associative and motivational theories of Pavlovian mediation.     

Conditioning of appetitive and consummatory sexual behavior in male Japanese quail.

Two different types of stimulus objects, a live female quail artificially adorned with bright orange feathers and an inanimate toy dog, served as conditioned stimuli. For subjects in experimental groups, the conditioned stimuli were presented shortly before access to a sexually receptive normal female quail. For subjects in control groups, exposure to the conditioned stimuli was unpaired with copulatory opportunity. Subjects in the experimental but not in the control groups quickly came to approach the location of the conditioned stimulus objects. When an adorned female quail served as the conditioned stimulus, the conditioned approach behavior was controlled by a combination of the presence of the orange adornments and the visual cues of the head and neck of the female bird, and the approach behavior persisted as the adorned female moved to new locations. When the toy dog served as the conditioned stimulus, the conditioned approach behavior was limited to the spatial cues that surrounded the toy dog during conditioning trials. Although both types of stimulus objects evoked conditioned approach behavior, only the adorned female stimulus supported copulatory behavior. This last finding indicates that copulatory behavior can become redirected toward novel stimuli as a result of conditioning, but only under special circumstances. The results are consistent with the suggestion that appetitive components of reproductive behavior are more susceptible to conditioning than consummatory components. Possible reasons for this are discussed, together with implications of the results for the contribution of conditioning processes to sexual selection.     

Neocortex and feeding behavior in the rat.

Consummatory behavior and weight-regulation capacity were measured in 12 normal rats and in 43 rats that survived complete (C), sequential unilateral (U), anterolateral (A), or posterior (P) neocortical ablations. Groups C and A displayed aphagia and adipsia followed by a sequence of recovery stages gualitatively identical to, but shorter than, recovery typically seen following lateral hypothalamic lesions. After recovery, Group C displayed long-term effects of finickiness and pradial drinking. These effects as well as a measure of recovery of body-weight-regulation capacity were significantly intercorrelated with lesion size, and body-weight set point remained significantly lower than normal. Group U was relatively unaffected by the first unilateral ablation and showed, relative the second ablation but displayed the long-term effects. Group P, though significantly affected by the lesion, did not display the pattern or intensity of effects described for the other bilaterally ablated groups.     

Motivational control of sexual behavior in Aplysia fasciata: sequencing and modulation by sexual deprivation and by addition of partners

Variables similar to those affecting feeding behavior also modulate sexual behavior in Aplysia fasciata, indicating that mating is under motivational control. Motivated behaviors are often patterned into appetitive and consummatory components. Courtship, the appetitive phase of male sexual behavior, was temporally related to subsequent mating. Although many bouts of courtship failed to lead to mating, most incidents of mating were preceded by courtship. Motivation is also characterized by satiation after the goal of the behavior is achieved. We found an increase in likelihood to mate following a period of sexual isolation. Motivated behaviors are also modulated by environmental stimuli that induce arousal. Time spent mating was found to be a function of the number of potential mates accessible for mating. Number of A. fascinata participating in a mating group was also found to be a function of number of potential mates available.     

Drinking behavior in the chronic phase of alcoholism

40 patients in the chronic phase of alcohol addition were questioned in order to identify drinking stereotypes with regard to consumption in the course of the day and year. Alcohol intake reaches a maximum between 8 and 9 PM. No clear differences were found in this respect between days of abstinence for social reasons and days without such social pressures. In the course of the year, alcohol consumption reaches a maximum in autumn and a minimum in spring. The uniform, repetitive drinking rituals are suspected to have become habitual in the course of several years until they are finally unaffected by environment. This leads to a convergence of addiction forms, but the duration of the daily abstinence period still permits differentiation between gamma and delta addicts even when addiction has reached an advanced stage.