Imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation.

The 2-action method was used to examine whether imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation of the demonstrated behavior. Two groups of observers were fed before observation (satiated groups), whereas 2 other groups of observers were deprived of food before observation (hungry groups). Quail were tested either immediately following observation or after a 30-min delay. Results indicated that quail in the hungry groups imitated, whereas those in the satiated groups did not, regardless of whether their test was immediate or delayed. The results suggest that observer quail may not learn (through observation) behavior that leads to a reinforcer for which they are unmotivated at the time of test. In addition, the results show that quail are able to delay the performance of a response acquired through observation (i.e., they show deferred imitation).

The case for a cognitive approach to animal learning and behavior.

The dangers of hypothesizing about unobservable cognitive mechanisms are well known to behavior analysts. I propose, however, that carefully fashioned cognitive theories that make predictions that are inconsistent with current behavioral theories can provide useful research tools for the understanding of behavior. Furthermore, even if the results of such research may be accommodated by modifying existing behavioral theories, our understanding of behavior is often advanced by the empirical findings because it is unlikely that the research would have been conducted in the absence of such cognitive hypothesizing. Two examples of the development of emergent relations are described: The first deals with the nature of a pigeon's 'representation' of two stimuli both of which are associated with correct responding to a third in a many-to-one matching task (stimulus equivalence or common representations). The second has to do with transitive inference, the emergent relation between two stimuli mediated by their relation to a common stimulus in a simultaneous discrimination.

Episodic-like memory in pigeons.

It has been proposed that memory for personal experiences (episodic memory, rather than semantic memory) relies on the conscious review of past experience and thus is unique to humans. In an attempt to demonstrate episodic-like memory in animals, we first trained pigeons to respond to the (nonverbal) question "Did you just peck or did you just refrain from pecking?" by training them on a symbolic matching task with differential responding required to the two line-orientation samples and reinforcing the choice of a red comparison if they had pecked and the choice of a green comparison if they had not pecked. Then, in Experiment 1, after providing the conditions for (but not requiring) the pigeons to peck at one new stimulus (a yellow hue) but not at another (a blue hue), we tested them with the new hue stimuli and the red and green comparisons. In Experiment 2, we tested the pigeons with novel stimuli (a circle, which they spontaneously pecked, and a dark response key, which they did not peck) and the red and green comparisons. In both experiments, pigeons chose the comparison appropriate to the response made to the test stimulus. Thus, the pigeons demonstrated that they could remember specific details about their past experiences, a result consistent with the notion that they have the capacity for forming episodic-like memories.

Presence/absence-sample matching by pigeons: divergent retention functions may result from the similarity of behavior during the absence sample and the retention interval.

Divergent choose-absence retention functions typically found in pigeons following presence/absence-sample matching have been attributed to the development of a single-code/default coding strategy. However, such effects may result from adventitious differential responding to the samples. In Experiment 1, retention functions were divergent only when differential sample responding could serve as the basis for comparison choice. In Experiment 2, when pecking did not occur during the retention interval, a choose-absence bias was found, but when pecking occurred during the retention interval, a choose-presence bias resulted. In Experiment 3, positive transfer was found when a stimulus associated with the absence of pecking replaced the absence sample but not when a stimulus associated with pecking replaced the presence sample. Thus, presence/absence-sample matching may not encourage the development of a single-code/default coding strategy in pigeons.

"Work ethic" in pigeons: reward value is directly related to the effort or time required to obtain the reward.

Stimuli associated with less effort or with shorter delays to reinforcement are generally preferred over those associated with greater effort or longer delays to reinforcement. However, the opposite appears to be true of stimuli that follow greater effort or longer delays. In training, a simple simultaneous discrimination followed a single peck to an initial stimulus (S+FR1 S-FR1) and a different simple simultaneous discrimination followed 20 pecks to the initial stimulus (S+FR20 S-FR20). On test trials, pigeons preferred S+FR20 over S+FR1 and S-FR20 over S-FR1. These data support the view that the state of the animal immediately prior to presentation of the discrimination affects the value of the reinforcement that follows it. This contrast effect is analogous to effects that when they occur in humans have been attributed to more complex cognitive and social factors.

Selective attention in animal discrimination learning.

The traditional approach to the study of selective attention in animal discrimination learning has been to ask if animals are capable of the central selective processing of stimuli, such that certain aspects of the discriminative stimuli are partially or wholly ignored while their relationships to each other, or other relevant stimuli, are processed. A notable characteristic of this research has been that procedures involve the acquisition of discriminations, and the issue of concern is whether learning is selectively determined by the stimulus dimension defined by the discriminative stimuli. Although there is support for this kind of selective attention, in many cases, simpler nonattentional accounts are sufficient to explain the results. An alternative approach involves procedures more similar to those used in human information-processing research. When selective attention is studied in humans, it generally involves the steady state performance of tasks for which there is limited time allowed for stimulus input and a relatively large amount of relevant information to be processed; thus, attention must be selective or divided. When this approach is applied to animals and alternative accounts have been ruled out, stronger evidence for selective or divided attention in animals has been found. Similar processes are thought to be involved when animals search more natural environments for targets. Finally, an attempt is made to distinguish these top-down attentional processes from more automatic preattentional processes that have been studied in humans and other animals.

Development of a single-code/default coding strategy in pigeons.

We tested the hypothesis that pigeons could use a cognitively efficient coding strategy by training them on a conditional discrimination (delayed symbolic matching) in which one alternative was correct following the presentation of one sample (one-to-one), whereas the other alternative was correct following the presentation of any one of four other samples (many-to-one). When retention intervals of different durations were inserted between the offset of the sample and the onset of the choice stimuli, divergent retention functions were found. With increasing retention interval, matching accuracy on trials involving any of the many-to-one samples was increasingly better than matching accuracy on trials involving the one-to-one sample. Furthermore, following this test, pigeons treated a novel sample as if it had been one of the many-to-one samples. The data suggest that rather than learning each of the five sample-comparison associations independently, the pigeons developed a cognitively efficient single-code/default coding strategy.

Support for a theory of memory for event duration must distinguish between test-trial ambiguity and actual memory loss.

Staddon and Higa's (1999) trace-strength theory of timing and memory for event duration can account for pigeons' bias to "choose short" when retention intervals are introduced and to "choose long" when, following training with a fixed retention interval, retention intervals are shortened. However, it does not account for the failure of pigeons to choose short when the intertrial interval is distinct from the retention interval. That finding suggests that stimulus generalization (or ambiguity) between the intertrial interval and the retention interval may result in an effect that has been attributed to memory loss. Such artifacts must be eliminated before a theory of memory for event duration can be adequately tested.

Transfer of value from S+ to S- in simultaneous discriminations in humans.

When animals learn a simultaneous discrimination, some of the value of the positive stimulus (S+) appears to transfer to the negative stimulus (S-). The present experiments demonstrate that such value transfer can also be found in humans. In Experiment 1 humans were trained on 2 simple simultaneous discriminations, the first between a highly positive stimulus, A (1,000 points); and a negative stimulus, B (0 points); and the second between a less positive stimulus, C (100 points); and a negative stimulus, D (0 points). On test trials, most participants preferred B over D. In Experiments 2 and 3 the value of the 2 original discriminations was equated in training (A[100]B[0] and C[100]D[0]). In Experiment 2 the values of the positive stimuli were then altered (A[1,000]C[0]); again, most participants preferred B over D. In Experiment 3, however, when the values of B and D were altered (B[1,000]D[0]), participants were indifferent to A and C. Thus, the mechanism that underlies value transfer in humans appears to be related to Pavlovian second-order conditioning. Similar mechanisms may be involved in assimilation processes in social contexts.

Simultaneous discrimination learning in pigeons: value of S- affects the relative value of its associated S+.

In a simple simultaneous discrimination involving a positive stimulus (S+) and a negative stimulus (S-), it has been hypothesized that positive value can transfer from the S+ to the S- (thus increasing the relative value of the S-) and also that negative value can transfer from the S- to the S+ (thus diminishing the relative value of the S+; Fersen, Wynne, Delius, & Staddon, 1991). Evidence for positive value transfer has been reported in pigeons (e.g. Zentall & Sherburne, 1994). The purpose of the present experiments was to determine, in a simultaneous discrimination, whether the S- diminishes the value of the S+ or the S- is contrasted with the S+ (thus enhancing the value of the S+). In two experiments, we found evidence for contrast, rather than value transfer, attributable to simultaneous discrimination training. Thus, not only does the S+ appear to enhance the value of the S-, but the S- appears to enhance rather than reduce the value of the S+.

Transfer across delayed discriminations: II. Differences in the substitutability of initial versus test stimuli.

In 2 experiments, pigeons were trained on, and then transferred to, delayed simple discriminations in which the initial stimuli signalled reinforcement versus extinction following a retention interval. Experiment 1 showed that discriminative responding on the retention test transferred to novel test stimuli that had appeared in another delayed simple discrimination but not to stimuli having the same reinforcement history off-baseline. By contrast, Experiment 2 showed that performances transferred to novel initial stimuli whether they had been trained on-baseline or off-baseline. These results suggest that the test stimuli in delayed simple discriminations acquire control over responding only in the memory task itself. On the other hand, control by the initial stimuli, if coded as outcome expectancies, does not require such task-specific training.

Delayed matching in pigeons: can apparent memory loss be attributed to the delay of reinforcement of sample-orienting behavior?

In earlier research using constant-delay matching with pigeons, there is evidence that delay of reinforcement of sample-orienting behavior may contribute to the decline in matching accuracy with increasing delay between sample and comparison stimuli. In the present research using this procedure, we found that a significant decline in matching accuracy between the first and second session can occur when delays are relatively long. This effect cannot be accounted for in terms of either additional memory loss or surprise (generalization decrement) associated with the increase in delay. Furthermore, the decline in matching accuracy occurred regardless of whether the delay was inserted between samples and comparisons (where it would be expected to affect the use of sample memory in making the comparison choice response) or between comparisons and reinforcement (where it would not be expected to affect the use of sample memory in making the comparison choice response). Thus, the decrease in matching accuracy between Session 1 and 2 following an increase in delay appears to be unrelated to sample memory at the time of choice. Instead, the results suggest that delay of reinforcement of sample-orienting behavior may play an important role in the negative slope of the retention functions obtained when constant- or mixed-delay matching procedures are used to assess animal memory.

Win-stay/lose-shift and win-shift/lose-stay learning by pigeons in the absence of overt response mediation.

Win-stay/lose-shift and win-shift/lose-stay behavior in pigeons was compared using a two-alternative conditional discrimination for which the number of trials involving each of the task components could be precisely controlled. One group was rewarded for pecking the location just pecked if those pecks were followed by food and for pecking the other location if those pecks were not followed by food (win-stay/lose-shift). Another group was rewarded for pecking the location just pecked if those pecks were not followed by food and for pecking the other location if those pecks were followed by food (win-shift/lose-stay). With increasing delay to comparison choice, pigeons were more accurate on trials when initial pecking was followed by the absence of food than by food (Experiment 1). However, when hypothesized overt response mediation was discouraged (Experiment 2), a win-stay superiority effect emerged with increasing delay to comparison choice. Thus, unlike rats, pigeons may be somewhat predisposed to repeat a response to a location to which responses have been previously rewarded.

Imitative learning in male Japanese quail (Coturnix japonica) using the two-action method.

The study of imitative learning in animals has suffered from the presence of a number of confounding motivational and attentional factors (e.g., social facilitation and stimulus enhancement). The two-action method avoids these problems by exposing observers to demonstrators performing a response (e.g., operating a treadle) using 1 of 2 distinctive topographies (e.g., by pecking or by stepping). Japanese quail (Coturnix japonica) observers exposed to conspecific demonstrators showed a high correlation between the topography of the response they observed and the response they performed. These data provide strong evidence for the existence of true imitative learning in an active, precocious bird under conditions that control for alternative accounts.

Value transfer in a simultaneous discrimination appears to result from within-event pavlovian conditioning.

When pigeons acquire a simple simultaneous discrimination, some of the value acquired by the S+ transfers to the S-. The mechanism underlying this transfer of value was examined in three experiments. In Experiment 1, pigeons trained on two simultaneous discriminations (A + B- and C +/- D-) showed a preference for B over D. This preference was reduced, however, following the devaluation of A. In Experiment 2, when after the same original training, value was given to D, the pigeons' preference for C did not significantly increase. In Experiment 3, when both discriminations involved partial reinforcement (S +/-), A + C- training resulted in a preference for B over D, whereas B + D- training resulted in a preference for A over C. Thus, simultaneous discrimination training appears to result in bidirectional within-event conditioning involving the S+ and S-.

Most directed forgetting in pigeons can be attributed to the absence of reinforcement on forget trials during training or to other procedural artifacts.

In research on directed forgetting in pigeons using delayed matching procedures, remember cues, presented in the delay interval between sample and comparisons, have been followed by comparisons (i.e., a memory test), whereas forget cues have been followed by one of a number of different sample-independent events. The source of directed forgetting in delayed matching to sample in pigeons was examined in a 2 x 2 design by independently manipulating whether or not forget-cue trials in training ended with reinforcement and whether or not forget-cue trials in training included a simultaneous discrimination (involving stimuli other than those used in the matching task). Results were consistent with the hypothesis that reinforced responding following forget cues is sufficient to eliminate performance deficits on forget-cue probe trials. Only when reinforcement was omitted on forget-cue trials in training (whether a discrimination was required or not) was there a decrement in accuracy on forget-cue probe trials. When reinforcement is present, however, the pattern of responding established during and following a forget cue in training may also play a role in the directed forgetting effect. These findings support the view that much of the evidence for directed forgetting using matching procedures may result from motivational and behavioral artifacts rather than the loss of memory.

Role of differential sample responding in the differential outcomes effect involving delayed matching by pigeons.

The role of differential sample responding in the differential outcomes effect was examined. In Experiment 1, we trained pigeons on a one-to-many matching task with differential sample responding required. Differential outcomes were associated with samples and comparisons, with comparisons only, or with neither samples nor comparisons. Slopes of delay functions for trials with pecked versus nonpecked samples suggested use of a single-code-default strategy in the nondifferential-outcomes group but not in the differential-outcomes groups. In Experiment 2, differential sample responding and differential outcomes were manipulated independently. Again, there were significant differences in the relative slopes of the delay functions. Results suggest that differential outcomes exert their effect independently of differential sample responding.

Transfer of value from S+ to S- in a simultaneous discrimination.

Value transfer theory has been proposed to account for transitive inference effects (L. V. Fersen, C. D. L. Wynne, J. D. Delius, & J. E. R. Staddon, 1991), in which following training on 4 simultaneous discriminations (A+B-, B+C-, C+D-, D+E-) pigeons show a preference for B over D. According to this theory, some of the value of reinforcement acquired by each S+ transfers to the S-. In the transitive inference experiment, C (associated with both reward and nonreward) can transfer less value to D than A (associated only with reward) can transfer to B. Support for value transfer theory was demonstrated in 2 experiments in which an S- presented in the context of a stimulus to which responses were always reinforced (S+) was preferred over an S- presented in the context of a stimulus to which responses were sometimes reinforced (S +/-).

Directed forgetting in animals.

Directed-forgetting research with animals suggests that animals show disrupted test performance only under certain conditions. Important variables are (a) whether during training, the cue to forget (F cue) signals nonreward (i.e., that the trial is over) versus reward (i.e., that reinforcement can be obtained) and (b) given that reinforcement can be obtained on F-cue trials, whether the post-F-cue response pattern is compatible with the baseline memory task. It is proposed that some findings of directed forgetting can be attributed to trained response biases, whereas others may be attributable perhaps to frustration-produced interference. It is suggested that directed forgetting in animals should be studied using procedures similar to those used to study directed forgetting in humans. This can be accomplished by presenting, within a trial, both to-be-remembered and to-be-forgotten material.