Fish can infer social rank by observation alone.

Transitive inference (TI) involves using known relationships to deduce unknown ones (for example, using A > B and B > C to infer A > C), and is thus essential to logical reasoning. First described as a developmental milestone in children, TI has since been reported in nonhuman primates, rats and birds. Still, how animals acquire and represent transitive relationships and why such abilities might have evolved remain open problems. Here we show that male fish (Astatotilapia burtoni) can successfully make inferences on a hierarchy implied by pairwise fights between rival males. These fish learned the implied hierarchy vicariously (as 'bystanders'), by watching fights between rivals arranged around them in separate tank units. Our findings show that fish use TI when trained on socially relevant stimuli, and that they can make such inferences by using indirect information alone. Further, these bystanders seem to have both spatial and featural representations related to rival abilities, which they can use to make correct inferences depending on what kind of information is available to them. Beyond extending TI to fish and experimentally demonstrating indirect TI learning in animals, these results indicate that a universal mechanism underlying TI is unlikely. Rather, animals probably use multiple domain-specific representations adapted to different social and ecological pressures that they encounter during the course of their natural lives.

Androgen level and male social status in the African cichlid, Astatotilapia burtoni.

In vertebrates, circulating androgen levels are regulated by the hypothalamic-pituitary-gonadal (HPG) axis through which the brain controls the gonads via the pituitary. Androgen levels ultimately depend on factors including season, temperature, social circumstance, age, and other variables related to reproductive capacity and opportunity. Previous studies with an African cichlid fish, Astatotilapia burtoni, suggested that changes in both testosterone and 11-ketotestosterone (11-KT), an androgen specific to teleost fish, depend on male social status. Here we characterize circulating plasma concentrations of testosterone and 11-KT in socially dominant (territorial) and socially subordinate (non-territorial) males. Territorial males have significantly higher circulating levels of both forms of androgen, which is another defining difference between dominant and subordinate males in this species. These results underscore how internal and external cues related to reproduction are integrated at the level of the HPG axis.

Required pecking alters judgments of the passage of time by pigeons.

There is evidence that how humans perceive time is affected by the activity in which they are engaged when they are judging time. In humans, typically, the more demanding the task, the faster time seems to pass. We asked whether a similar effect could be found in pigeons. Pigeons were trained to discriminate between a short- (2-sec) and a long- (10-sec) duration stimulus. Depending on the color of the stimulus (white or blue), the pigeons were required to peck (at least once per second or the trial was aborted) or to refrain from pecking (pecks aborted the trial). Once these tasks had been acquired to a high degree, probe trials involving white and blue stimuli were presented at durations between 2 and 10 sec. On trials in which the pigeons were required to peck, the point of subjective equality (i.e., the point at which pigeons are equally likely to choose the stimulus associated with long stimuli as the stimulus associated with short stimuli) was almost 1 sec longer than on trials in which th epigeons were required to refrain from pecking. In other words, on trials that required pecking, more time passed before the pigeons indicated that the probe duration was at the subjective midpoint between 2 and 10 sec than on trials that did not require pecking. This result suggests that like humans, the pigeons underestimated the passage of time when they were active or when attention to time-related cues had to be shared with attention to satisfying the response rate requirement.

Memory mechanisms in pigeons: evidence of base-rate neglect.

In delayed matching to sample, once acquired, pigeons presumably choose comparisons according to their memory for (the strength of) the sample. When memory for the sample is sufficiently weak, comparison choice should depend on the history of reinforcement associated with each of the comparison stimuli. In the present research, pigeons acquired two matching tasks in which Sample S1 was associated with one comparison from each task, C1 and C3, whereas Sample S2 was associated with Comparison C2, and Sample S3 was associated with Comparison C4. As the retention interval increased, the pigeons showed a bias to choose the comparison (C1 or C3) associated with the more frequently occurring sample (S1). Thus, pigeons were sensitive also to the (irrelevant) likelihood that each of the samples was presented. The results suggest that pigeons may allow their reference memory for the overall sample frequency to influence comparison choice, independent of the comparison stimuli present.

Second-order contrast based on the expectation of effort and reinforcement.

Pigeons prefer signals for reinforcement that require greater effort (or time) to obtain over those that require less effort to obtain (T. S. Clement, J. Feltus, D. H. Kaiser, & T. R. Zentall, 2000). Preference was attributed to contrast (or to the relatively greater improvement in conditions) produced by the appearance of the signal when it was preceded by greater effort. In Experiment 1, the authors of the present study demonstrated that the expectation of greater effort was sufficient to produce such a preference (a second-order contrast effect). In Experiments 2 and 3, low versus high probability of reinforcement was substituted for high versus low effort, respectively, with similar results. In Experiment 3, the authors found that the stimulus preference could be attributed to positive contrast (when the discriminative stimuli represented an improvement in the probability of reinforcement) and perhaps also negative contrast (when the discriminative stimuli represented reduction in the probability of reinforcement).

Choice based on exclusion in pigeons.

When humans acquire a conditional discrimination and are given a novel-sample-comparison choice, they often reject a comparison known to be associated with a different sample and choose the alternative comparison by default (or by exclusion). In Experiment 1, we found that if, following matching training, we replaced both of the samples, acquisition took five times longer than if we replaced only one of the samples. Apparently, the opportunity to reject one of the comparisons facilitated the association of the other sample with the remaining comparison. In Experiment 2, we first trained pigeons to treat two samples differently (to associate Sample A with Comparison 1 and Sample B with Comparison 2) and then trained them to associate one of those samples with a new comparison (e.g., Sample A with Comparison 3) and to associate a novel sample (Sample C) with a different, new comparison (Comparison 4). When Sample B then replaced Sample C, the pigeons showed a significant tendency to choose Comparison 4 over Comparison 3. Thus, when given the opportunity, pigeons will choose by exclusion.

Symmetry training in pigeons can produce functional equivalences.

Functional stimulus equivalence has been demonstrated using a transfer of training design with matching-to-sample training in which two sample stimuli are associated with the same comparison stimulus (A-B, C-B; many-to-one matching). Equivalence is shown by training a new association (A-D) and demonstrating the presence of an emergent relation (C-D). In the present experiment, we show that symmetry training, in which a bidirectional association is trained between two stimuli (A-B, B-A, using successive stimulus presentations followed by reinforcement), can also produce functional equivalence using a transfer of training design (i.e., train B-C, test A-C). The results suggest that training pigeons in the substitutability of two stimuli may be sufficient to produce functional stimulus equivalence between them. The results also have implications for the development of an emergent transitive relation, because training on A-B and B-C relations results in the emergence of an untrained A-C relation, if B-A training also is provided.

Pigeons group time intervals according to their relative duration.

In the present research, we asked whether pigeons tended to judge time intervals not only in terms of their absolute value but also relative to a duration from which they must be discriminated (i.e., longer or shorter). Pigeons were trained on two independent temporal discriminations. In one discrimination, sample durations of 2 and 8 sec were associated with, for example, red and green hue comparisons, respectively, and in the other discrimination, sample durations of 4 and 16 sec were associated with vertical and horizontal line comparisons, respectively. If pigeons are trained on a temporal discrimination and tested with intermediate durations, the subjective midpoint typically occurs close to the geometric mean of the two trained values. The 4- and 8-sec values were selected to be the geometric mean of the two values in the other discrimination. When a 4-sec test sample was presented with the comparisons from the 2- and 8-sec discrimination, the pigeons preferred the comparison associated with the shorter sample. Similarly, when an 8-sec test sample was presented with the comparisons from the 4- and 16-sec discrimination, the pigeons preferred the comparison associated with the longer sample. Thus, a relative grouping effect was found. That is, durations that should have produced indifferent choice were influenced by their relative durations (shorter than or longer than the alternative) during training.

Functional equivalence in pigeons involving a four-member class.

Research suggests that animals are capable of forming functional equivalence relations or stimulus classes of the kind usually demonstrated by humans (e.g., the class defined by an object and the word for that object). In pigeons, such functional equivalences are typically established using many-to-one matching-to-sample in which two samples are associated with one comparison stimulus and two different samples are associated with the other. Evidence for the establishment of functional equivalences between samples associated with the same comparison comes from transfer tests. In Experiment 1, we found that pigeons can form a single class consisting of four members (many-to-one matching) when the alternative class has only one member (one-to-one matching). In Experiment 2, we ruled out the possibility that the pigeons acquired the hybrid one-to-one/many-to-one task by developing a single-code/default coding strategy as earlier research suggested that it might. Thus, pigeons can develop a functional class consisting of as many as four members, with the alternative class consisting of a single member.

Discriminative stimuli that follow the absence of reinforcement are preferred by pigeons over those that follow reinforcement.

Clement, Feltus, Kaiser, and Zentall (2000) found that when pigeons have to work to obtain a discriminative stimulus that is followed by reinforcement, they prefer a discriminative stimulus that requires greater effort over one that requires less effort. The authors suggested that such a preference results from the greater change in hedonic value that occurs between the more aversive event and the onset of the stimulus that signals reinforcement, a contrast effect. It was hypothesized that any stimulus that follows a relatively more aversive event would be preferred over a stimulus that follows a relatively less aversive event. In the present experiment, the authors tested the counterintuitive prediction of that theory, that pigeons should prefer a discriminative stimulus that follows the absence of reinforcement over a discriminative stimulus that follows reinforcement. Results supported the theory.

Behavioral coping strategies in a cichlid fish: the role of social status and acute stress response in direct and displaced aggression.

The African cichlid fish, Astatotilapia burtoni, has a complex social system with a sophisticated social hierarchy that offers unique opportunities to understand how social rank and its physiological substrates relate to behavioral strategies. In A. burtoni, a small fraction of the males are dominant (T, territorial), as distinguished by being large, brightly colored, reproductively active, and aggressively defending territories. In contrast, the majority of males are non-dominant (NT, non-territorial), being smaller, drably colored, sexually immature, and typically schooling with females. The social system is regulated by aggressive interactions between males and behavioral responses to aggression can be direct or displaced with respect to the animal that acts. To determine whether direct and displaced behaviors are differentially exhibited by T and NT males, individuals were shown a video presentation of a dominant male displaying aggressively. Analysis of aggressive acts toward the video display and displaced activity toward a tank mate revealed that T males exhibited more direct behavior (toward the video display), while NT males engaged in more displaced behavior (toward tank mates). Because similar experiments with primates suggest that shifts in behavioral strategies are linked to changes in the stress response (as measured by circulating cortisol levels), we measured cortisol levels of T and NT males following exposure to the aggressive stimulus. Although in some animals subordinate males are reported to have higher cortisol levels, here we show that in A. burtoni the endocrine response to specific situations can vary considerably even among animals of the same status. Interestingly, NT males with intermediate cortisol levels showed more directed behavior while NT males with both high and low cortisol levels showed more displaced. This suggests an optimal physiological stress response in NT males that predisposes them to challenge aggressors perhaps making it more likely for them to ascend in status.