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Brannon, E. M., & Terrace, H. S. (1998). Ordering of the numerosities 1 to 9 by monkeys. Science, 282(5389), 746–749.
Abstract: A fundamental question in cognitive science is whether animals can represent numerosity (a property of a stimulus that is defined by the number of discriminable elements it contains) and use numerical representations computationally. Here, it was shown that rhesus monkeys represent the numerosity of visual stimuli and detect their ordinal disparity. Two monkeys were first trained to respond to exemplars of the numerosities 1 to 4 in an ascending numerical order (1 --> 2 --> 3 --> 4). As a control for non-numerical cues, exemplars were varied with respect to size, shape, and color. The monkeys were later tested, without reward, on their ability to order stimulus pairs composed of the novel numerosities 5 to 9. Both monkeys responded in an ascending order to the novel numerosities. These results show that rhesus monkeys represent the numerosities 1 to 9 on an ordinal scale.
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Washburn, D. A., Smith, J. D., & Shields, W. E. (2006). Rhesus monkeys (Macaca mulatta) immediately generalize the uncertain response. J Exp Psychol Anim Behav Process, 32(2), 185–189.
Abstract: Rhesus monkeys (Macaca mulatta) have learned, like humans, to use an uncertain response adaptively under test conditions that create uncertainty, suggesting a metacognitive process by which human and nonhuman primates may monitor their confidence and alter their behavior accordingly. In this study, 4 rhesus monkeys generalized their use of the uncertain response, without additional training, to 2 familiar tasks (2-choice discrimination learning and mirror-image matching to sample) that predictably and demonstrably produce uncertainty. The monkeys were significantly less likely to use the uncertain response on trials in which the answer might be known. These results indicate that monkeys, like humans, know when they do not know and that they can learn to use a symbol as a generalized means for indicating their uncertainty.
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Brannon, E. M., Cantlon, J. F., & Terrace, H. S. (2006). The role of reference points in ordinal numerical comparisons by rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process, 32(2), 120–134.
Abstract: Two experiments examined ordinal numerical knowledge in rhesus macaques (Macaca mulatta). Experiment 1 replicated the finding (E. M. Brannon & H. S. Terrace, 2000) that monkeys trained to respond in descending numerical order (4-->3-->2-->1) did not generalize the descending rule to the novel values 5-9 in contrast to monkeys trained to respond in ascending order. Experiment 2 examined whether the failure to generalize a descending rule was due to the direction of the training sequence or to the specific values used in the training sequence. Results implicated 3 factors that characterize a monkey's numerical comparison process: Weber's law, knowledge of ordinal direction, and a comparison of each value in a test pair with the reference point established by the first value of the training sequence.
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Beran, M. J., Smith, J. D., Redford, J. S., & Washburn, D. A. (2006). Rhesus macaques (Macaca mulatta) monitor uncertainty during numerosity judgments. J Exp Psychol Anim Behav Process, 32(2), 111–119.
Abstract: Two rhesus macaques (Macaca mulatta) judged arrays of dots on a computer screen as having more or fewer dots than a center value that was never presented in trials. After learning a center value, monkeys were given an uncertainty response that let them decline to make the numerosity judgment on that trial. Across center values (3-7), errors occurred most often for sets adjacent in numerosity to the center value. The monkeys also used the uncertainty response most frequently on these difficult trials. A 2nd experiment showed that monkeys' responses reflected numerical magnitude and not the surface-area illumination of the displays. This research shows that monkeys' uncertainty-monitoring capacity extends to the domain of numerical cognition. It also shows monkeys' use of the purest uncertainty response possible, uncontaminated by any secondary motivator.
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Beran, M. J., Pate, J. L., Washburn, D. A., & Rumbaugh, D. M. (2004). Sequential responding and planning in chimpanzees (Pan troglodytes) and rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process, 30(3), 203–212.
Abstract: Chimpanzees (Pan troglodytes) and rhesus macaques (Macaca mulatta) selected either Arabic numerals or colored squares on a computer monitor in a learned sequence. On shift trials, the locations of 2 stimuli were interchanged at some point. More errors were made when this interchange occurred for the next 2 stimuli to be selected than when the interchange was for stimuli later in the sequence. On mask trials, all remaining stimuli were occluded after the 1st selection. Performance exceeded chance levels for only 1 selection after these masks were applied. There was no difference in performance for either stimulus type (numerals or colors). The data indicated that the animals planned only the next selection during these computerized tasks as opposed to planning the entire response sequence.
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Beran, M. J. (2007). Rhesus monkeys (Macaca mulatta) succeed on a computerized test designed to assess conservation of discrete quantity. Anim. Cogn., 10(1), 37–45.
Abstract: Conservation of quantity occurs through recognition that changes in the physical arrangement of a set of items do not change the quantity of items in that set. Rhesus monkeys (Macaca mulatta) were presented with a computerized quantity judgment task. Monkeys were rewarded for selecting the greater quantity of items in one of two horizontal arrays of items on the screen. On some trials, after a correct selection, no reward was given but one of the arrays was manipulated. In some cases, this manipulation involved moving items closer together or farther apart to change the physical arrangement of the array without changing the quantity of items in the array. In other cases, additional items were added to the initially smaller array so that it became quantitatively larger. Monkeys then made another selection from the two rows of items. Monkeys were sensitive to these manipulations, changing their selections when the number of items in the rows changed but not when the arrangement only was changed. Therefore, monkeys responded on the basis of the quantity of items, and they were not distracted by non-quantitative manipulations of the sets.
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Jordan, K. E., & Brannon, E. M. (2006). Weber's Law influences numerical representations in rhesus macaques (Macaca mulatta). Anim. Cogn., 9(3), 159–172.
Abstract: We present the results of two experiments that probe the ability of rhesus macaques to match visual arrays based on number. Three monkeys were first trained on a delayed match-to-sample paradigm (DMTS) to match stimuli on the basis of number and ignore continuous dimensions such as element size, cumulative surface area, and density. Monkeys were then tested in a numerical bisection experiment that required them to indicate whether a sample numerosity was closer to a small or large anchor value. Results indicated that, for two sets of anchor values with the same ratio, the probability of choosing the larger anchor value systematically increased with the sample number and the psychometric functions superimposed. A second experiment employed a numerical DMTS task in which the choice values contained an exact numerical match to the sample and a distracter that varied in number. Both accuracy and reaction time were modulated by the ratio between the correct numerical match and the distracter, as predicted by Weber's Law.
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Harris, E. H., & Washburn, D. A. (2005). Macaques' (Macaca mulatta) use of numerical cues in maze trials. Anim. Cogn., 8(3), 190–199.
Abstract: We tested the ability of number-trained rhesus monkeys to use Arabic numeral cues to discriminate between different series of maze trials and anticipate the final trial in each series. The monkeys' prior experience with numerals also allowed us to investigate spontaneous transfer between series. A total of four monkeys were tested in two experiments. In both experiments, the monkeys were trained on a computerized task consisting of three reinforced maze trials followed by one nonreinforced trial. The goal of the maze was an Arabic numeral 3, which corresponded to the number of reinforced maze trials in the series. In experiment 1 (n=2), the monkeys were given probe trials of the numerals 2 and 4 and in experiment 2 (n=2), they were given probe trials of the numerals 2-8. The monkeys receiving the probe trials 2 and 4 showed some generalization to the new numerals and developed a pattern of performing more slowly on the nonreinforced trial than the reinforced trial before it for most series, indicating the use of the changing numeral cues to anticipate the nonreinforced trial. The monkeys receiving probe trials of the numerals 2-8 did not predict precisely when the nonreinforced trial would occur in each series, but they did incorporate the changing numerals into their strategy for performing the task. This study provides the first evidence that number-trained monkeys can use Arabic numerals to perform a task involving sequential presentations.
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Treichler, F. R. (2005). Successive reversal of concurrent discriminations by macaques (Macaca mulatta): proactive interference effects. Anim. Cogn., 8(2), 75–83.
Abstract: Rhesus monkeys received concurrent within-session training on eight, two-choice object pairs and then underwent successive reversals of these problems. Initially, reversals required about six times more training than acquisition with no improvement over seven successive reversals. Surprisingly, performance on these eight problems was unimpaired if they were embedded in different eight-problem tasks, thereby indicating a release from proactive interference. When the original eight problems again underwent successive reversal, no improvement was seen over seven reversals, although there was significantly less error-per-reversal than in the initial test. Subsequently, monkeys appeared to be developing a learning set for successive reversal because performance on successive reversal of eight novel problems was not different from that seen with the old familiar task. Set acquisition was confirmed when proficient reversal was eventually achieved on both old and new concurrent tasks. Thus, “concurrent reversal set” did develop, but it required arduous training to overcome proactive interference effects on memory. The ubiquitous influence of measurement context on organization of monkey memory was noted.
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Hampton, R. R., Zivin, A., & Murray, E. A. (2004). Rhesus monkeys (Macaca mulatta) discriminate between knowing and not knowing and collect information as needed before acting. Anim. Cogn., 7(4), 239–246.
Abstract: Humans use memory awareness to determine whether relevant knowledge is available before acting, as when we determine whether we know a phone number before dialing. Such metacognition, or thinking about thinking, can improve selection of appropriate behavior. We investigated whether rhesus monkeys ( Macaca mulatta) are capable of a simple form of metacognitive access to the contents of short-term memory. Monkeys chose among four opaque tubes, one of which concealed food. The tube containing the reward varied randomly from trial to trial. On half the trials the monkeys observed the experimenter baiting the tube, whereas on the remaining trials their view of the baiting was blocked. On each trial, monkeys were allowed a single chance to select the tube containing the reward. During the choice period the monkeys had the opportunity to look down the length of each tube, to determine if it contained food. When they knew the location of the reward, most monkeys chose without looking. In contrast, when ignorant, monkeys often made the effort required to look, thereby learning the location of the reward before choosing. Looking improved accuracy on trials on which monkeys had not observed the baiting. The difference in looking behavior between trials on which the monkeys knew, and trials on which they were ignorant, suggests that rhesus monkeys discriminate between knowing and not knowing. This result extends similar observations made of children and apes to a species of Old World monkey, suggesting that the underlying cognitive capacities may be widely distributed among primates.
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