Ordinal judgments of numerical symbols by macaques (Macaca mulatta)

Two rhesus monkeys (Macaca mulatta) learned that the arabic numerals 0 through 9 represented corresponding quantities of food pellets. By manipulating a joystick, the monkeys were able to make a selection of paired numerals presented on a computer screen. Although the monkeys received a corresponding number of pellets even if the lesser of the two numerals was selected, they learned generally to choose the numeral of greatest value even when pellet delivery was made arrhythmic. In subsequent tests, they chose the numerals of greater value when presented in novel combinations or in random arrays of up to five numerals. Thus, the monkeys made ordinal judgments of numerical symbols in accordance with their absolute or relative values.     

A primacy effect in monkeys when list position is relevant

In Experiment 1 (1a and 1b), Rhesus monkeys (Macaca mulatta) learned lists of two-choice visual discriminations in which list position was relevant to discrimination performance. For example, Stimulus A was the rewarded stimulus if it was presented at List Position 1, but was not rewarded if it was presented at any other position in the list; similarly, Stimulus B was rewarded only at List Position 2, and so on. In learning these lists, all animals showed a marked primacy effect. In Experiment 2 (2a and 2b), Rhesus monkeys and Cynomolgus monkeys (M. fascicularis) learned lists of visual discriminations in which each visual stimulus occupied a fixed position in a list, but list position was not relevant to discrimination performance. For example, Stimulus E was always rewarded, and was always presented at List Position 1. To increase the salience of list beginning as a distinctive event, successive presentations of the list were separated by 24-hr intervals. In Experiment 2 there was no primacy effect, however. These results show for the first time that a primacy effect can be obtained in visual discrimination learning by monkeys. Furthermore, they suggest that it is obtained only when list position is relevant to the discrimination learning task.     

Sequential responding and planning in capuchin monkeys (Cebus apella)

Previous experiments have assessed planning during sequential responding to computer generated stimuli by Old World nonhuman primates including chimpanzees and rhesus macaques. However, no such assessment has been made with a New World primate species. Capuchin monkeys (Cebus apella) are an interesting test case for assessing the distribution of cognitive processes in the Order Primates because they sometimes show proficiency in tasks also mastered by apes and Old World monkeys, but in other cases fail to match the proficiency of those other species. In two experiments, eight capuchin monkeys selected five arbitrary stimuli in distinct locations on a computer monitor in a learned sequence. In Experiment 1, shift trials occurred in which the second and third stimuli were transposed when the first stimulus was selected by the animal. In Experiment 2, mask trials occurred in which all remaining stimuli were masked after the monkey selected the first stimulus. Monkeys made more mistakes on trials in which the locations of the second and third stimuli were interchanged than on trials in which locations were not interchanged, suggesting they had already planned to select a location that no longer contained the correct stimulus. When mask trials occurred, monkeys performed at levels significantly better than chance, but their performance exceeded chance levels only for the first and the second selections on a trial. These data indicate that capuchin monkeys performed very similarly to chimpanzees and rhesus monkeys and appeared to plan their selection sequences during the computerized task, but only to a limited degree.     

Representation of the numerosities 1-9 by rhesus macaques (Macaca mulatta)

Three rhesus monkeys (Macaca mulatta) were trained to respond to exemplars of 1, 2, 3, and 4 in an ascending, descending, or a nonmonotonic numerical order (1-->2-->3-->4, 4-->3-->2--1, 3-->1-->4-->2). The monkeys were then tested on their ability to order pairs of the novel numerosities 5-9. In Experiment 1, all 3 monkeys ordered novel exemplars of the numerosities 1-4 in ascending or descending order. The attempt to train a nonmonotonic order (3-->1-->4-->2) failed. In Experiment 2A, the 2 monkeys who learned the ascending numerical rule ordered pairs of the novel numerosities 5-9 on unreinforced trials. The monkey who learned the descending numerical rule failed to extrapolate the descending rule to new numerosities. In Experiment 2B all 3 monkeys ordered novel exemplars of pairs of the numerosities 5-9. Accuracy and latency of responding revealed distance and magnitude effects analogous to previous findings with human participants (R. S. Moyer & T. K. Landaeur, 1967). Collectively these studies show that monkeys represent the numerosities 1-9 on at least an ordinal scale.     

Monkeys (Macaca mulatta and Cebus apella) track, enumerate, and compare multiple sets of moving items

Despite many demonstrations of numerical competence in nonhuman animals, little is known about how well animals enumerate moving stimuli. In this series of experiments, rhesus monkeys (Macaca mulatta) and capuchin monkeys (Cebus apella) performed computerized tasks in which they had to enumerate sets of stimuli. In Experiment 1, rhesus monkeys compared two sets of moving stimuli. Experiment 2 required comparisons of a moving set and a static set. Experiment 3 included human participants and capuchin monkeys to assess all 3 species' performance and to determine whether responding was to the numerical properties of the stimulus sets rather than to some other stimulus property such as cumulative area. Experiment 4 required both monkey species to enumerate subsets of each moving array. In all experiments, monkeys performed above chance levels, and their responses were controlled by the number of items in the arrays as opposed to nonnumerical stimulus dimensions. Rhesus monkeys performed comparably to adult humans when directly compared although capuchin performance was lower.     

Ordinal representation of numeric quantities by brown capuchin monkeys (Cebus apella)

Using techniques established by E. M. Brannon and H. S. Terrace (2000) with rhesus macaques (Macaca mulatta), the authors tested the ability of brown capuchins (Cebus apella) to order arrays of items ranging in quantity from 1 to 9. Three monkeys were trained on a touch screen to select the quantities 1-4 in ascending order. The monkeys exhibited successful transfer of this ability to novel representations of the quantities 1-4 and to pairs of the novel quantities 5-9. Patterns of responding with respect to numeric distance and magnitude were similar to those seen in human subjects, suggesting the use of similar psychological processes. The capuchins demonstrated an ordinal representation of quantity equivalent to that shown in Old World monkeys.     

Discrimination of word-like compound visual stimuli by monkeys

In Experiment I, two monkeys solved a successive visual discrimination in which the four positive stimuli were the visual arrays RIM, LID, RAD and LAM while the four negative stimuli were RID, LIM, RAM and LAD. In Experiment II the same monkeys first learned a discrimination where the positive stimuli were pairs of letters (e.g. OB and AK) while the negative stimulus was the letter I; in a subsequent generalization test with all four possible pairings of the stimulus elements that had been positive during training (i.e. with OB, AK, OK and AB) the monkeys responded more strongly to the pairs that had been present in initial training. These results were discussed in relation to the theoretical analysis of configurational cues in animal discrimination learning and to the mechanism underlying visual discrimination of words by people.     

Macaques’ (Macaca mulatta) use of numerical cues in maze trials

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.     

Mechanisms of same/different abstract-concept learning by rhesus monkeys (Macaca mulatta)

Experiments with 9 rhesus monkeys (Macaca mulatta) showed, for the first time, that abstract-concept learning varied with the training stimulus set size. In a same/different task, monkeys required to touch a top picture before choosing a bottom picture (same) or white rectangle (different) learned rapidly. Monkeys not required to touch the top picture or presented with the top picture for a fixed time learned slowly or not at all. No abstract-concept learning occurred after 8-item training but progressively improved with larger set sizes and was complete following 128-item training. A control monkey with a constant 8-item set ruled out repeated training and testing. Contrary to the unique-species account, it is argued that different species have quantitative, not qualitative, differences in abstract-concept learning.     

Reverse-reward learning in squirrel monkeys (Saimiri sciureus): retesting after 5 years, and assessment on qualitative transfer

Seven squirrel monkeys (Saimiri sciureus) previously trained on reverse-reward tasks were presented with the original "1-versus-4" task after a 5-year interval without reverse-reward experience (Experiment 1). None of them reliably selected the smaller food array; however, at around chance level, their performance was superior to when they were first exposed to the task almost 6 years previously, suggesting some long-term memory retention. One naive monkey consistently selected the larger array, as expected. In Experiment 2, trials consisting of 1 versus 1 piece of two qualitatively different types of food were interspersed among familiar 1-versus-4 trials. None of five monkeys tested reliably selected the less-preferred food to get the more preferred food as the reward, and one monkey scored below chance. However, when one piece of low-preference food was paired with four pieces of high-preference food (Experiment 3), all four monkeys tested avoided reaching for the latter and thereby obtained it as the reward; two monkeys obtained perfect scores on these trials. These two monkeys were trained on a specific qualitative reverse-reward pairing and then again tested on new pairings (Experiment 4), but transfer was incomplete. Compound trials that pit quantity against quality in novel ways appear taxing for squirrel monkeys, despite competence in reverse-reward on both dimensions separately.     

Rhesus monkeys (Macaca mulatta) enumerate large and small sequentially presented sets of items using analog numerical representations

Two rhesus monkeys selected the larger of two sequentially presented sets of items on a computer monitor. In Experiment 1, performance was related to the ratio of set sizes, and the monkeys discriminated between sets with up to 10 items. Performance was not disrupted when 1 set had fewer than 4 items and 1 set had more than 4 items, a critical trial type for differentiating object file and analog models of numerical representation. Experiment 2 controlled the interitem rate of presentation. Experiment 3 included some trials on which number and amount (visual surface area) offered conflicting cues. Experiment 4 varied the total duration of set presentation and the duration of item visibility. In all of the experiments, performance remained high, although total set presentation duration also acted as a partial cue for the monkeys. Overall, the data indicated that rhesus monkeys estimate the approximate number of items in sequentially presented sets and that they are not relying solely on nonnumerical cues such as rate, duration, or cumulative amount.     

Acquisition and memory of sequence order in young and adult chimpanzees (Pan troglodytes)

There is no research about age difference in the process of sequential learning in non-human primates. Is there any difference between young and adults in sequential learning process? Six chimpanzees (Pan troglodytes), 3 young and 3 adults, learned the Arabic numeral sequence 1 to 9 by touching the numerals on a touch-screen monitor in ascending order. Initially, the sequence always started with the numeral 1, i.e. ‘start-fixed task’. Training began with the sequence 1–2, 1–2–3, and continued sequentially up to 1–2–3–4–5–6–7–8–9. Later, the subjects were introduced to sequences that started with a random numeral, but always ended with 9, i.e. ‘end-fixed task’. Performance in the end-fixed task was worse relative to the familiar start-fixed task. After training with various sequences of adjacent numerals, the subjects were given a transfer test for the non-adjacent numerals. The results suggested that all chimpanzees indeed mastered sequential ordering, and although there was no fundamental difference in the acquisition process between the two age groups, there was a significant age difference in memory capacity. Based on their knowledge of sequential ordering, the subjects were then asked to perform a masking task in which once a subject touched the lowest numeral, the other numeral(s) turned to white squares. Performance of the masking task by young chimpanzees was better than that of adults in accuracy and degree of difficulty (number of numerals). Taken together, these data clearly demonstrate a similarity among subjects in the way chimpanzees acquire knowledge of sequential order regardless of age differences in sequential learning. Moreover, they reveal that once knowledge of sequential order is established, it can be a good index used to evaluate memory capacity in young and adult chimpanzees.

     

Disconnect in concept learning by rhesus monkeys (Macaca mulatta): judgment of relations and relations-between-relations

The authors investigated the role that entropy measures, discriminative cues, and symbolic knowledge play for rhesus monkeys (Macaca mulatta) in the acquisition of the concepts of same and different for use in a computerized relational matching-to-sample task. After repeatedly failing to perceive relations between pairs of stimuli in a 2-choice discrimination paradigm, monkeys rapidly learned to discriminate between 8-element arrays. Subsequent tests with smaller arrays, however, suggested that, although important for the initial acquisition of the concept, entropy is not a variable on which monkeys are dependent. Not only do monkeys choose a corresponding relational pair in the presence of a cue, but they also choose the cue itself in the presence of the relational pair--in essence, labeling those relations. Subsequent failure in the judgment of relations-between-relations, however, suggests that perhaps a qualitatively different cognitive component exists that prevents monkeys from behaving analogically.     

Mechanisms of inferential order judgments in humans (Homo sapiens) and rhesus monkeys (Macaca mulatta)

If A > B, and B > C, it follows logically that A > C. The process of reaching that conclusion is called transitive inference (TI). Several mechanisms have been offered to explain transitive performance. Scanning models claim that the list is scanned from the ends of the list inward until a match is found. Positional discrimination models claim that positional uncertainty accounts for accuracy and reaction time patterns. In Experiment 1, we trained rhesus monkeys (Macaca mulatta) and humans (Homo sapiens) on adjacent pairs (e.g., AB, BC, CD, DE, EF) and tested them with previously untrained nonadjacent pairs (e.g., BD). In Experiment 2, we trained a second list and tested with nonadjacent pairs selected between lists (e.g., B from List 1, D from List 2). We then introduced associative competition between adjacent items in Experiment 3 by training 2 items per position (e.g., B?C?, B?C?) before testing with untrained nonadjacent items. In all 3 experiments, humans and monkeys showed distance effects in which accuracy increased, and reaction time decreased, as the distance between items in each pair increased (e.g., BD vs. BE). In Experiment 4, we trained adjacent pairs with separate 9- and 5-item lists. We then tested with nonadjacent pairs selected between lists to determine whether list items were chosen according to their absolute position (e.g., D, 5-item list > E, 9-item list), or their relative position (e.g., D, 5-item list < E, 9-item list). Both monkeys' and humans' choices were most consistent with a relative positional organization.     

Rapid proportion comparison with spatial arrays of frequently used meaningful visual symbols

It has been shown that when two arrays of Arabic numerals were briefly presented, observers could accurately indicate which array contained the larger number of a target numeral. This study investigated whether this rapid proportion comparison can be extended to other meaningful symbols that share some of notable properties of Arabic numerals. We tested mainly several Japanese Kanji letters, each of which represents a meaning and can work as a word. Using physically identical stimulus sets that could be interpreted as different types of letters, Experiment 1 first confirmed the rapid proportion comparison with Arabic numerals for Japanese participants. Experiment 2 showed that the rapid proportion comparison can be extended to Kanji numerals. Experiment 3 successfully demonstrated that rapid proportion judgments can be found with non-quantitative Kanji letters that are used frequently. Experiment 4 further demonstrated the rapid proportion comparison with frequently used meaningful non-letter symbols (gender icons). The rapid processing cannot be attributed to fluent processing of familiar items, because it was not found with familiar phonograms (Japanese Kana letters). These findings suggest that the rapid proportion comparison can be commonly found with frequently used meaningful symbols, even though their meaning is not relevant to the task.     

Abstract-concept learning and list-memory processing by capuchin and rhesus monkeys

Three capuchin monkeys (Cebus apella) touched the lower of 2 pictures (same) or a white rectangle (different), increased same/different abstract-concept learning (52% to 87%) with set-size increases (8 to 128 pictures), and were better than 3 rhesus monkeys (Macaca mulatta). Three other rhesus that touched the top picture before choices learned similar to capuchins but were better at list-memory learning. Both species' serial position functions were similar in shape and changes with retention delays. Other species showed qualitatively similar shape changes but quantitatively different time-course changes. In abstract-concept learning, qualitative similarity was shown by complete concept learning, whereas a quantitative difference would have been a set-size slope difference. Qualitative similarity is discussed in relation to general-process versus modular cognitive accounts.     

Change detection by rhesus monkeys (Macaca mulatta) and pigeons (Columba livia)

Two monkeys (Macaca mulatta) learned a color change-detection task where two colored circles (selected from a 4-color set) were presented on a 4 × 4 invisible matrix. Following a delay, the correct response was to touch the changed colored circle. The monkeys' learning, color transfer, and delay transfer were compared to a similar experiment with pigeons. Monkeys, like pigeons (Columba livia), showed full transfer to four novel colors, and to delays as long as 6.4 s, suggesting they remembered the colors as opposed to perceptual based attentional capture process that may work at very short delays. The monkeys and pigeons were further tested to compare transfer with other dimensions. Monkeys transferred to shape and location changes, unlike the pigeons, but neither species transferred to size changes. Thus, monkeys were less restricted in their domain to detect change than pigeons, but both species learned the basic task and appear suitable for comparative studies of visual short-term memory. (PsycINFO Database Record (c) 2012 APA, all rights reserved).     

Memory Span for Arabic Numerals and Digit Words: Evidence for a Limited-capacity, Visuo-spatial Storage System

Six experiments examined the determinants of the numeral advantage effect: the finding that memory span for Arabic numerals (1, 2, 3, etc.) is greater than for digit words (one, two, three, etc.). The speed of item identification for numeral and digit words was unrelated to memory span for the same items and a larger memory span for numerals persisted under concurrent random generation (Experiment 1). The numeral advantage, however, was abolished when the items were presented in random locations within an invisible 3x3 grid (Experiment 2) and in locations on a horizontal plane that ran contrary to the natural direction of reading (Experiment 3). When the items were presented in the same location, a disruption of the spatial component of visuo-spatial working memory eliminated the numeral advantage (Experiment 4), whereas interference with the visual component of the system did not (Experiment 5). When the items were spatially distributed in a 3x3 matrix, however, neither visual nor spatial interference abolished the effect (Experiment 6). Taken together, these findings suggest that the numeral advantage effect is mediated by discrete components in visuo-spatial working memory dedicated to the temporary storage and renewal of visual codes and questions the assumption that the underlying mechanisms in immediate, visual serial recall are equivalent between stimulus categories.     

An assessment of memory awareness in tufted capuchin monkeys (Cebus apella)

Humans, apes, and rhesus monkeys demonstrate memory awareness by collecting information when ignorant and acting immediately when informed. In this study, five capuchin monkeys searched for food after either watching the experimenter bait one of four opaque tubes (seen trials), or not watching (unseen trials). Monkeys with memory awareness should look into the tubes before making a selection only on unseen trials because on seen trials they already know the location of the food. In Experiment 1, one of the five capuchins looked significantly more often on unseen trials. In Experiment 2, we ensured that the monkeys attended to the baiting by interleaving training and test sessions. Three of the five monkeys looked more often on unseen trials. Because monkeys looked more often than not on both trial types, potentially creating a ceiling effect, we increased the effort required to look in Experiment 3, and predicted a larger difference in the probability of looking between seen and unseen trials. None of the five monkeys looked more often on unseen trials. These findings provide equivocal evidence for memory awareness in capuchin monkeys using tests that have yielded clear evidence in humans, apes, and rhesus monkeys.     

"Constructive" enumeration by chimpanzees (Pan troglodytes) on a computerized task

Two chimpanzees used a joystick to collect dots, one at a time, on a computer monitor (see video-clip in the electronic supplementary material), and then ended a trial when the number of dots collected was equal to the Arabic numeral presented for the trial. Both chimpanzees performed substantially and reliably above chance in collecting a quantity of dots equal to the target numeral, one chimpanzee for the numerals 1–7, and the second chimpanzee for the numerals 1–6. Errors that were made were seldom discrepant from the target by more than one dot quantity, and the perceptual process subitization was ruled out as an explanation for the performance. Additionally, analyses of trial duration data indicated that the chimpanzees were responding based on the numerosity of the constructed set rather than on the basis of temporal cues. The chimpanzees' decreasing performance with successively larger target numerals, however, appeared to be based on a continuous representation of magnitude rather than a discrete representation of number. Therefore, chimpanzee counting in this type of experimental task may be a process that represents magnitudes with scalar variability in that the memory for magnitudes associated with each numeral is imperfect and the variability of responses increases as a function of the numeral's value. Accepted after revision: 11 June 2001 Electronic Publication