Rhesus monkeys (Macaca mulatta) spontaneously compute addition operations over large numbers

Mathematics is a uniquely human capacity. Studies of animals and human infants reveal, however, that this capacity builds on language-independent mechanisms for quantifying small numbers (<4) precisely and large numbers approximately. It is unclear whether animals and human infants can spontaneously tap mechanisms for quantifying large numbers to compute mathematical operations. Moreover, all available work on addition operations in non-human animals has confounded number with continuous perceptual properties (e.g. volume, contour length) that correlate with number. This study shows that rhesus monkeys spontaneously compute addition operations over large numbers, as opposed to continuous extents, and that the limit on this ability is set by the ratio difference between two numbers as opposed to their absolute difference.     

Shared system for ordering small and large numbers in monkeys and humans

There is increasing evidence that animals share with adult humans and perhaps human infants a system for representing objective number as psychological magnitudes that are an analogue of the quantities they represent. Here we show that rhesus monkeys can extend a numerical rule learned with the values 1 through 9 to the values 10, 15, 20, and 30, which suggests that there is no upper limit on a monkey's numerical capacity. Instead, throughout the numerical range tested, both accuracy and latency in ordering two numerical values were systematically controlled by the ratio of the values compared. In a second experiment, we directly compared humans' and monkeys' performance in the same ordinal comparison task. The qualitative and quantitative similarity in their performance provides the strongest evidence to date of a single nonverbal, evolutionarily primitive mechanism for representing and comparing numerical values.     

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.     

Probability-reward preferences of rhesus monkeys

The relative preferences of rhesus monkeys for reward probability versus amount were investigated with procedures which contrasted general experience with specific instructions, and evaluated the relationship between probability-amount combinations and preference strength. Four stimulus objects, each signifying a different combination of reward frequency and amount (100% with one unit; 50% with two units; 33% with three units; or 25% with four units), were presented in pairs, one pair per daily session, with trial schedules providing the same amount of reward within each set of 12 trials. In Phase A, 4 monkeys (Group 1) were tested on the six choice-pairs with no preliminary training. In Phase B, Group 1 was joined by an additional 4 monkeys (Group 2), and each of the tasks was preceded by a demonstration of the relevant stimuli, one at a time, together with their associated probabilities and amounts. Group 1 animals developed preferences during Phase A for the more frequently rewarded objects, which persisted into Phase B, whereas Group 2 animals showed no preferences. This result indicates that preliminary instructions concerning the reward combinations associated with stimulus objects can prevent the development of a preference for greater probability over greater amount of reward, but cannot extinguish it once it has been formed or reestablished within the context of a particular task.     

Spontaneous representations of small numbers of objects by rhesus macaques: examinations of content and format

The project of comparative cognition benefits from common measures across species. We report here on five experiments using the violation of expectancy looking time measure with free-ranging rhesus macaques (Macaca mulatta), each designed to build on current knowledge concerning spontaneous representations of number. Each subject, tested in only one experimental condition, watched as eggplants were placed behind a screen one at a time, after which the screen was removed revealing an outcome that either matched or did not match the number placed there. Subjects looked longer at impossible than possible outcomes in 1+1=2 or 3, 1 small+1 small=1 big or 2 small, 2+1=2 or 3, and 2+1=3 or 4 conditions. They failed in 2+1+1=4 or 3 or 5 and in 1+1+1=2 or 3 conditions. This pattern of results closely matches that observed across several previous studies of human infants. The data allow us to test among four different proposals concerning the format and content of the mental representations underlying looking in these experiments. Object file representations are favored over: (i) low-level perceptual representations, (ii) representations of continuous variables such as volume or surface area, and (iii) analog magnitude representations of number. We conclude by considering exactly how the object tracking system revealed in these and other related experiments does and does not represent number, and how it might be one evolutionary precursor of the human specific system of number representations.     

Confidence judgments by humans and rhesus monkeys

Researchers have begun to evaluate animals' metacognitive capacities. Continuing this evaluation, the present authors asked whether monkeys could use the analog of a confidence-rating scale to judge retrospectively their accuracy in a psychophysical discrimination. Monkeys and human participants classified stimuli as dense or sparse without feedback. Then, they made a secondary confidence judgment by choosing responses that risked different levels of timeout but could earn different levels of reward. Human participants were instructed to use these responses to express levels of confidence. They used 2- and 3-level confidence-rating scales appropriately. Monkeys used the 2-level scale appropriately--especially one who performed almost identically to humans. Neither monkey used the 3-level scale as the humans did. These studies place humans and animals for the first time in the identical risk-taking situation that collects declarative confidence judgments from humans. They demonstrate the first retrospective reports of uncertainty by non-human primates while also suggesting their limits in this area.     

Music perception and octave generalization in rhesus monkeys

Two rhesus monkeys were tested for octave generalization in 8 experiments by transposing 6- and 7-note musical passages by an octave and requiring same or different judgments. The monkeys showed no octave generalization to random-synthetic melodies, atonal melodies, or individual notes. They did show complete octave generalization to childhood songs (e.g., "Happy Birthday") and tonal melodies (from a tonality algorithm). Octave generalization was equally strong for 2-octave transpositions but not for 0.5- or 1.5-octave transpositions of childhood songs. These results combine to show that tonal melodies form musical gestalts for monkeys, as they do for humans, and retain their identity when transposed with whole octaves so that chroma (key) is preserved. This conclusion implicates similar transduction, storage, processing, and relational memory of musical passages in monkeys and humans and has implications for nature-nurture origins of music perception.     

Behavioral and pharmacological modulation of respiration in rhesus monkeys.

Changes in respiration associated with schedule-controlled behavior were determined in seated rhesus monkeys prepared with a pressure-displacement head plethysmograph for monitoring ventilation continuously during behavioral experiments. Subjects were trained to press a lever under fixed-ratio 40 and fixed-interval 300-s schedules of stimulus termination. Episodic increases in ventilation were closely associated with periods of responding under both schedules. Recurring episodes of increased ventilation occurred during fixed-ratio responding, and were separated by brief 10-s timeouts during which ventilation decreased. Under the fixed-interval schedule, both ventilation and response rate typically increased as the 300-s interval elapsed. The effects of cocaine, caffeine, and two adenosine agonists, 5'-N-ethylcarboxamidadenosine (NECA) and 2-(carboxyethylphenylamino)adenosine-5'-carboxamide (CGS 21680), on behavior and respiration were determined using a cumulative-dosing procedure. Drug-induced suppression of behavior eliminated the episodic increases in ventilation during the performance components of both schedules. Schedule-related increases in ventilation were compared to those produced by elevated levels of CO2 in inspired air. Exposure to 4% CO2 mixed in air increased ventilation in all subjects, and the combined effects of CO2 exposure and schedule-controlled responding on respiration appeared to be additive. The results suggest that behavioral activities may increase ventilation through increased metabolic demand and increased CO2 production.     

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.