Hurdle-jump responding in the rat as a function of conspecific odor of reward and nonreward

A hurdle-jump escape response was employed to assess the laboratory rat’s aversion or attraction to different types of conspecific odor. Odorant donor subjects received 112 runway acquisition trials on a continuous reward schedule followed by 32 extinction trials, 112 acquisition trials on a 50% schedule of reward and nonreward followed by 32 extinction trials, or 144 “neutral” trials with no reward in the alley. Different groups of test subjects escaped from odor excreted by odorant subjects on (a) nonrewarded acquisition and extinction trials, (b) rewarded trials during continuous reinforcement, (c) rewarded trials during partial reinforcement, or (d) neutral trials; others escaped from a clean box. The principal findings were: (1) significant aversion to “odor of nonreward” appeared after the donor odorants had received 12 exposures to reward; (2) production of odor of nonreward by odorant subjects changed as a function of training experience with reward; (3) after repeated exposure to odor of nonreward, the escape response habituated; (4) greater or different odor excretion in extinction resulted from subjects trained on a continuous reward schedule than on a partial reward schedule. Relationships of the data to frustration theory were discussed, assuming that inferred differences in production of odor reflect differences in frustration reaction.     

Effects of varied and partial reward on discrete-trial patterning of rats

Rats were trained in the discrete-trial operant apparatus with single alternation of large reward and small reward or large reward and nonreward. followed by either transfer or extinction. The results showed that both groups acquired appropriate response patterning, that patterning is conditioned, and that the stimuli which control patterning are derived from reward conditions on immediately preceding trials. A modification of the sequential hypothesis of instrumental learning was proposed to account for the results.     

Taste + odor interactions in compound aversion conditioning

In three experiments with rats, taste + odor interactions in compound aversion conditioning were investigated. In Experiment 1, two odors (0.02% almond and 0.02% orange) were compared on single-element odor aversions, taste (denatonium) potentiated odor aversions, and potentiated odor aversions following taste extinction. Although no odor differences were seen following single-element conditioning, both types of potentiated orange odor aversions were stronger than their almond odor counterparts. These data show that odors of similar conditionability are differentially potentiated by the same taste. To determine whether these differences were due to unique perceptual representations, the effects of elemental extinction or compound extinction on aversions to the compound were investigated in Experiments 2 and 3. In Experiment 2, orange odor extinction weakened responding to the compound significantly more than taste extinction did. In contrast, almond odor extinction and taste extinction produced similar decrements in responding to the compound in Experiment 3. These results suggest that the perceptual representation of these specific taste + odor compounds are different, and they are discussed in regard to configural and within-compound association accounts of potentiation.     

Nonreward anticipated: Effects on extinction runway performance in the rat

Following a 47-day extinction procedure, the reinstatement of the cue previously associated with reward produced an immediate improvement in performance. The cue was the opportunity to traverse the alley following an “anticipated” nonrewarded runway trial. Moreover, the animals trained in this matter exhibited daily increments in performance during the initial phase of extinction testing. The results were interpreted as consistent with the notion that the difference in extinction performance of one group as compared to another does not necessarily reflect the relative “strengths” of the instrumentally acquired habits. Instead, it probably indicates the degree of similarity of the extinction testing procedure to the acquisition training condition previously associated with reinforcement for the two individual groups.     

Imitative learning in Japanese quail (<Emphasis Type="Italic">Coturnix japonica</Emphasis>) using the bidirectional control procedure

In the bidirectional control procedure, observers are exposed to a conspecific demonstrator responding to a manipulandum in one of two directions (e.g., left vs. right). This procedure controls for socially mediated effects (the mere presence of a conspecific) and stimulus enhancement (attention drawn to a manipulandum by its movement), and it has the added advantage of being symmetrical (the two different responses are similar in topography). Imitative learning is demonstrated when the observers make the response in the direction that they observed it being made. Recently, however, it has been suggested that when such evidence is found with a predominantly olfactory animal, such as the rat, it may result artifactually from odor cues left on one side of the manipulandum by the demonstrator. In the present experiment, we found that Japanese quail, for which odor cues are not likely to play a role, also showed significant correspondence between the direction in which the demonstrator and the observer push a screen to gain access to reward. Furthermore, control quail that observed the screen move, when the movement of the screen was not produced by the demonstrator, did not show similar correspondence between the direction of screen movement observed and that performed by the observer. Thus, with the appropriate control, the bidirectional procedure appears to be useful for studying imitation in avian species.     

A Pavlovian analysis of food-attraction conditioning in the snailHelix aspersa

Following brief pairing of an odor with a feeding experience (food-attraction conditioning), snails will lower their tentacles when subsequently presented with that odor alone. Three experiments investigated the possible behavioral mechanism mediating food-attraction conditioning in the snailHelix aspersa. It is suggested that food-attraction conditioning is an example of Pavlovian conditioning. In this case, the odor (conditioned stimulus) is paired with oral stimulation (unconditioned stimulus), which elicits lowering of the tentacles (unconditioned response). Following conditioning, the odor comes to elicit lowering of the tentacles (conditioned response). Experiment 1 ruled out nonassociative effects, such as habituation and sensitization, using an unpaired control group. Experiment 2 provided further evidence against a role for habituation of neophobia, through the demonstration of extinction following conditioning. In Experiment 3, an omission procedure was used to rule out the possible role of instrumental contingencies.     

A nonverbal false belief task: the performance of children and great apes

A nonverbal task of false belief understanding was given to 4- and 5-year-old children (N = 28) and to two species of great ape: chimpanzees and orangutans (N = 7). The task was embedded in a series of finding games in which an adult (the hider) hid a reward in one of two identical containers, and another adult (the communicator) observed the hiding process and attempted to help the participant by placing a marker on the container that she believed to hold the reward. An initial series of control trials ensured that participants were able to use the marker to locate the reward, follow the reward in both visible and invisible displacements, and ignore the marker when they knew it to be incorrect. In the crucial false belief trials, the communicator watched the hiding process and then left the area, at which time the hider switched the locations of the containers. When the communicator returned, she marked the container at the location where she had seen the reward hidden, which was incorrect. The hider then gave the subject the opportunity to find the sticker. Successful performance required participants to reason as follows: the communicator placed the marker where she saw the reward hidden; the container that was at that location is now at the other location; so the reward is at the other location. Children were also given a verbal false belief task in the context of this same hiding game. The two main results of the study were: (1) children's performance on the verbal and nonverbal false belief tasks were highly correlated (and both fit very closely with age norms from previous studies), and (2) no ape succeeded in the nonverbal false belief task even though they succeeded in all of the control trials indicating mastery of the general task demands.     

Social working memory: Memory for another rat's spatial choices can increase or decrease choice tendencies

In two experiments using a radial-arm maze, pairs of rats made choices among eight maze locations, each containing a large quantity of one of two food types. The choices made by 1 rat affected the choices made by the other rat. Under most conditions, visits by 1 rat increased the tendency of the other rat to subsequently choose that maze location. However, the effect depended on the quality of the food available in a particular location. When it was possible for the rats to observe each other on the maze arms and a rat had experienced that a location contained the less preferred food type, a previous visit to that location by the foraging partner decreased the tendency to visit that location. These effects are attributed to working memory for the spatial choices of another rat, and they indicate that memory produced by a rat’s own visit to a maze location is integrated with memory for the behavior of another rat to determine spatial choice     

Reciprocal overshadowing in the discrimination of color-odor compounds by honeybees: Further tests of a continuity model

Honeybees were trained to discriminate between simultaneously presented color-odor compounds, one group with color and odor confounded and a control group with color relevant and odor irrelevant; in subsequent differentially reinforced training with the colors in the absence of the odors, the performance of the two groups was the same (Experiment 1). When, however, response to the colors was measured in a 10-min extinction test, discrimination was found to be poorer after confounded training (Experiment 2), and like results were obtained in an extinction test with the odors after control animals had been trained with odor relevant and color irrelevant, the confounded animals showing poorer discrimination of the odors than the controls (Experiment 3). The results of the first two experiments, in which overshadowing of color by odor was found only with an extinction test, require us to take seriously the possibility that our previous modeling experiments (with probability of correct choice in differentially reinforced training as the measure of performance) may have been insufficiently sensitive to noncontinuity effects. Our first efforts to model extinction suggest, however, that all the results of the present experiments can be understood without sacrifice of the parsimonious independence principle.     

Emergence of choice strategies in the rat: Shift-stay differentiation precedes win-lose differentiation

The choice strategies of rat pups between 12 and 21 days of age were studied in a two-choice discrimination procedure that used access to a nonlactating anesthetized dam as a reward. After receiving a reward (win) or not receiving a reward (lose), a pup repeated a previous choice (stay) or made an alternative response (shift). At 12 and 15 days of age, pups demonstrated neither a stay nor a shift response strategy; at 18 and 21 days of age, pups had a robust tendency to shift. Winning or losing did not alter behavior until 21 days of age, at which point winning produced a greater tendency to return to the initially visited location for rats in the stay condition.     

Spatial localization of a goal: Beacon homing and landmark piloting by rats on a radial maze

We performed six experiments in order to examine the ability of rats to use moving beacons and landmarks as cues to the location of reward on an eight-arm radial maze. In Experiments 1–4, the cues and goals were moved before each trial, and groups in which a single beacon was placed on the rewarded arm, a single landmark indicated that reward was on the arm immediately to the left of a landmark, or two landmarks were placed on each side of the reward arm were compared. The rats rapidly learned to track the reward in the beacon condition, failed to find the reward sooner than chance expectation with a single landmark, and did only slightly better than chance with two landmarks. In Experiments 5 and 6, the rats were trained in five trials per day, with the landmark and goal locations constant over daily rewarded trials, and in two extinction trials that were inserted among the rewarded trials. The rats found the goal arm at substantially better than chance expectancy with both one and two landmarks. Our results, in agreement with data from recent swimming pool experiments (A. D. L. Roberts & Pearce, 1998), show that rats will use the relationship between moving landmarks and a goal in order to find reward.     

Reward and learning in honeybees: analysis of an overshadowing effect

Previous experiments have shown that honeybees trained with colored targets baited with 5- versus 20-µl drops of sucrose solution fail to develop a preference for the 20-µl color when the location of the drop on each target is marked by a white dot (dot-color overshadowing) but that discrimination is not impaired by dots when the targets differ in odor rather than in color. In Experiments 1–3, dot-color overshadowing failed to appear with differences in concentration rather than amount of sucrose (50% vs. 20% or 0%), but it did appear in Experiments 4 and 5 with a difference in probability of reward (consistent vs. partial). Experiment 6 showed no dot-odor overshadowing with a difference in probability of reward. The results are not generally predictable from the Rescorla-Wagner principle of shared associative strength, but point instead (in conjunction with those of earlier experiments) to competition for visual attention.     

Social effects on spatial choice in the radial arm maze

Social memory was investigated in the context of a spatial working memory task. Pairs of rats were tested in an eight-arm radial maze. Under most conditions, there was a tendency to choose maze locations that had been visited earlier by the other rat. The possibility that this tendency is produced by common preferences for particular maze locations was ruled out. An opposite tendency to avoid visits to locations that had been visited earlier during the trial by another rat was found only when the maze location contained two pellets (rather than an undepletable supply), the rats’ ability to see each other in the maze was restricted to the central arena, and the maze location had been previously visited by the focal rat. The amount of food available in maze locations did not otherwise modulate social influences on spatial choice. The results indicate that memory for a rat’s own previous choices is combined with memory for the choices made by another rat.     

Extinction responding following partial reinforcement: The effects of number of rewarded trials and magnitude of reward

Rats were trained on a daily partial reward schedule of small magnitude of reward (S), nonreward (N), and large magnitude of reward (L), which began with SN or SSNN for all animals. The remainder of the daily schedule was defined by the factorial combination of the number of rewards (1 vs. 3) and the magnitude of reward (S vs. L). Following 18 days of such training, 20 trials of extinction were administered. It was found that increasing the number of rewarded trials in a partial reinforcement schedule decreased resistance to extinction. Furthermore, increased number of large-magnitude rewards reduced resistance to extinction to a greater extent than increased number of small-magnitude rewards.     

Partial reinforcement delayed extinction effect after regularly alternating reward training

When extinction is delayed very long, the superior resistance to extinction of the random schedule group relative to the alternating schedule group disappears (partial reinforcement delayed extinction effect, PRDE). Two experiments assessed the effects of reinforcement/nonreinforcement on Trial 1 on the PRDE. Following extended partial reinforcement acquisition training in a runway, rats received extinction training after a short (1-day) or long (23-day) retention interval. The schedules used in Experiment 1 were: a single-alternation (SA) schedule beginning each day with a rewarded (r) trial, for Group r-SA; an SA schedule beginning with a nonrewarded (n) trial, for Group n-SA; and a random (Rd) schedule, for Group Rd. The schedules and group names used in Experiment 2 were r-SA, Rd, and r-Rd. The results were that (1) rats given r-SA schedules yielded considerable resistance under delayed extinction, (2) those given Rd and r-Rd schedules showed a decline in resistance to extinction over a long retention interval, (3) those given the n-SA schedule showed relatively low resistance at both retention intervals, although retention deficit was not greater than in the case of the Rd schedule, and thus, (4) the PRDE was found in both experiments, although only weakly in Experiment 1. The results indicated that a regularly alternating reward pattern was a more important determinant than was type of reward on Trial 1 for the PRDE. The PRDE due to differential retention deficits among schedules is discussed on the basis of dual-process associative sequential mechanisms and cognitive rule-encoding mechanisms.     

Reward sequence and reinforcement level as determinants of S− behavior in differential conditioning

In two differential conditioning experiments, groups of 10 rats each differed with respect to average reward and schedule of reward received in S+. Nonreward (N) occurred on all S? trials. In both experiments, extinction of responding to S? (resistance to discrimination) was extensively regulated by reward sequence and was largely independent of average reward. In Experiment 1, resistance to discrimination was a function of transitions from N to rewarded (R) trials (N-R transitions). In Experiment 2, resistance to discrimination was increased by large reward on the R trial of N-R transitions and decreased by large reward on the R trial of R-N transitions. These schedule effects on resistance to discrimination parallel the effects of comparable schedules on resistance to extinction following partial reinforcement. The results are discussed in terms of sequential theory, reinforcement level theory, and their implications for various schedule manipulations that have previously shown S? behavior to be inversely related to average reward in S+.     

Instrumental escape learning in the rat at 24-hour intertriai interval: Effects of magnitude and schedule of reinforcement

Three experiments investigated the effects of magnitude and schedule of reinforcement and level of training in instrumental escape learning at a 24-h intertriai interval. In Experiment I, two magnitudes of reinforcement were factorially combined with two schedules of reinforcement (CRF and PRF). Under PRF, large reward produced greater resistance to extinction than did small reward, while the reverse was true under CRF. In Experiment II, two levels of acquisition training were factorially combined with three schedules of reinforcement (CRF, single-alternation, and nonalternated PRF). Patterned running was observed late in acquisition in the single-alternation extended-training condition. Resistance to extinction was greater for the nonalternated PRF condition than for the single-alternation condition following extended acquisition, and the reverse was true following limited acquisition. Experiment III confirmed the extinction findings of Experiment II. The results of all three experiments supported an analysis of escape learning at spaced trials in terms of Capaldi’s (1967) sequential theory.     

Frustration and sexual behavior in male rats

Frustration is an emotional state produced by the surprising omission in quantity and/or quality of an appetitive reinforcer. The aversive properties of stressors, such as electric shocks, produce responses similar to those elicited by a state of frustration. In this set of three experiments, we assessed the effects of water immersion (WIM, in Experiment 1)—that is, a physical stressor—and first (in Experiments 1 and 2) and second trials of a consummatory extinction (cE; i.e., a surprising reward omission; in Experiment 3) on the sexual behavior of male rats, as compared with nonstressed animals. The results showed a sexual deficit in the animals subjected to either WIM or cE, relative to control subjects, although these experimental conditions differed in the component of the male sexual response that was affected. The present results accord with the fear = frustration hypothesis, and with Amsel’s frustration theory.     

Partial reinforcement effect: The expectancy of reward on nonreward trials

In order to determine the importance of the development of expectancy of reward prior to partial reward trials; rats were given 20 continuously reinforced trials prior to 20 partially reinforced trials (CRF-PRF) and compared to Ss given only 20 partially reinforced trials (PRF). Control groups received 20 or 40 continuously reinforced trials (CRF-20, CRF-40) to determine the effect of differing numbers of acquisition trials. Results showed that terminal acquisition differences were minimal in the run segment of the alley and that Group CRF-PRF was more resistant to extinction than Group PRF, and both were more resistant to extinction than the CRF-20 and CRF-40 groups, which did not differ from each other. These results were interpreted as supporting the notion that the expectancy of reward on nonreward trials during partial reinforcement acquisition is a determiner of the magnitude of the partial reinforcement extinction effect.     

Aversive conditioning of naturally produced reward and nonreward odors in rats

Past experiments have reported that rats encountering reward (R) or nonreward (N) goal events emit odors that can be utilized as discriminative stimuli for instrumental behavior by conspecifics. In the present study, thirsty male rats were aversively conditioned by ingestional toxicosis to R and N odors, and their suppression of water consumption in the presence of these odors was measured. Thirsty trained donors were placed into chambers containing R or N goal events to generate, respectively, the R or N odors. Test animals were given eight differential conditioning trials (four with one odor as CS+; four with the other as CS?), involving placement into an odorcontaining chamber with water available, followed by a LiCl injection on CS+ trials. Animals tested in their CS+ odor consumed significantly less water than did CS? and control subjects. Both R and N odors were conditioned by aversive means and readily discriminable from each other. This represents the first laboratory demonstration of aversive conditioning of such naturally produced odors, and it suggests that aversive conditioning may be useful in the study of odorous emissions generally. Implications for innate meanings of R and N odors are discussed.