Erratum to: Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities

Although many studies have shown that nonhuman animals can choose the larger of two discrete quantities of items, less emphasis has been given to discrimination of continuous quantity. These studies are necessary to discern the similarities and differences in discrimination performance as a function of the type of quantities that are compared. Chimpanzees made judgments between continuous quantities (liquids) in a series of three experiments. In the first experiment, chimpanzees first chose between two clear containers holding differing amounts of juice. Next, they watched as two liquid quantities were dispensed from opaque syringes held above opaque containers. In the second experiment, one liquid amount was presented by pouring it into an opaque container from an opaque syringe, whereas the other quantity was visible the entire time in a clear container. In the third experiment, the heights at which the opaque syringes were held above opaque containers differed for each set, so that sometimes sets with smaller amounts of juice were dropped from a greater height, providing a possible visual illusion as to the total amount. Chimpanzees succeeded in all tasks and showed many similarities in their continuous quantity estimation to how they performed previously in similar tasks with discrete quantities (for example, performance was constrained by the ratio between sets). Chimpanzees could compare visible sets to nonvisible sets, and they were not distracted by perceptual illusions created through various presentation styles that were not relevant to the actual amount of juice dispensed. This performance demonstrated a similarity in the quantitative discrimination skills of chimpanzees for continuous quantities that matches that previously shown for discrete quantities.     

Quantity judgments of auditory and visual stimuli by chimpanzees (Pan troglodytes)

Many species can choose between two visual sets of stimuli on the basis of quantity. This is true when sets are both visible, or are presented one set at a time or even one item at a time. However, we know comparatively little about how well nonhuman animals can compare auditory quantities. Here, three chimpanzees (Pan troglodytes) chose between two sets of food items when they only heard each item fall into different containers rather than seeing those items. This method prevented the chimpanzees from summing the amount of visible food they saw because there were no visual cues. Chimpanzees performed well, and their performance matched that of previous experiments with regard to obeying Weber's law. They also performed well with comparisons between a sequentially presented auditory set and a fully visible set, demonstrating that duration of presentation was not being used as a cue. In addition, they accommodated empty sets into these judgments, although not perfectly. Thus, chimpanzees can judge auditory quantities in flexible ways that show many similarities to how they compare visual quantities.     

Numerical judgments by chimpanzees (Pan troglodytes) in a token economy

We presented four chimpanzees with a series of tasks that involved comparing two token sets or comparing a token set to a quantity of food. Selected tokens could be exchanged for food items on a one-to-one basis. Chimpanzees successfully selected the larger numerical set for comparisons of 1 to 5 items when both sets were visible and when sets were presented through one-by-one addition of tokens into two opaque containers. Two of four chimpanzees used the number of tokens and food items to guide responding in all conditions, rather than relying on token color, size, total amount, or duration of set presentation. These results demonstrate that judgments of simultaneous and sequential sets of stimuli are made by some chimpanzees on the basis of the numerousness of sets rather than other non-numerical dimensions. The tokens were treated as equivalent to food items on the basis of their numerousness, and the chimpanzees maximized reward by choosing the larger number of items in all situations.     

Ordinal judgments and summation of nonvisible sets of food items by two chimpanzees and a rhesus macaque

Two chimpanzees and a rhesus macaque rapidly learned the ordinal relations between 5 colors of containers (plastic eggs) when all containers of a given color contained a specific number of identical food items. All 3 animals also performed at high levels when comparing sets of containers with sets of visible food items. This indicates that the animals learned the approximate quantity of food items in containers of a given color. However, all animals failed in a summation task, in which a single container was compared with a set of 2 containers of a lesser individual quantity but a greater combined quantity. This difficulty was not overcome by sequential presentation of containers into opaque receptacles, but performance improved if the quantitative difference between sizes was very large.     

Quantity judgments of sequentially presented food items by capuchin monkeys (Cebus apella)

Recent assessments have shown that capuchin monkeys, like chimpanzees and other Old World primate species, are sensitive to quantitative differences between sets of visible stimuli. In the present study, we examined capuchins’ performance in a more sophisticated quantity judgment task that required the ability to form representations of food quantities while viewing the quantities only one piece at a time. In three experiments, we presented monkeys with the choice between two sets of discrete homogeneous food items and allowed the monkeys to consume the set of their choice. In Experiments 1 and 2, monkeys compared an entirely visible food set to a second set, presented item-by-item into an opaque container. All monkeys exhibited high accuracy in choosing the larger set, even when the entirely visible set was presented last, preventing the use of one-to-one item correspondence to compare quantities. In Experiment 3, monkeys compared two sets that were each presented item-by-item into opaque containers, but at different rates to control for temporal cues. Some monkeys performed well in this experiment, though others exhibited near-chance performance, suggesting that this species’ ability to form representations of food quantities may be limited compared to previously tested species such as chimpanzees. Overall, these findings support the analog magnitude model of quantity representation as an explanation for capuchin monkeys’ quantification of sequentially presented food items.

Chimpanzees remember the results of one-by-one addition of food items to sets over extended time periods

Four chimpanzees were highly accurate in selecting the larger of two concurrent accumulations of bananas in two opaque containers over a span of 20 min. One at a time, bananas were placed into the containers, which were outside the chimpanzees' cages. The chimpanzees never saw more than one banana at a time, and there were no cues indicating the locations of the bananas after they were placed into the containers. The performance of these animals matched that of human infants and young children in similar tests. The chimpanzees were successful even when the sets to be compared were sufficiently large (5 vs. 8, 5 vs. 10, and 6 vs. 10) to cast doubt on the possibility that the chimpanzees were using an object file mechanism. These chimpanzees are the first nonhuman animals to demonstrate extended memory for accumulated quantity.     

Discrete quantity judgments in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus): the effect of presenting whole sets versus item-by-item

The authors examined quantity-based judgments for up to 10 items for simultaneous and sequential whole sets as well as for sequentially dropped items in chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), bonobos (Pan paniscus), and orangutans (Pongo pygmaeus). In Experiment 1, subjects had to choose the larger of 2 quantities presented in 2 separate dishes either simultaneously or 1 dish after the other. Representatives of all species were capable of selecting the larger of 2 quantities in both conditions, even when the quantities were large and the numerical distance between them was small. In Experiment 2, subjects had to select between the same food quantities sequentially dropped into 2 opaque cups so that none of the quantities were ever viewed as a whole. The authors found some evidence (albeit weaker) that subjects were able to select the larger quantity of items. Furthermore, the authors found no performance breakdown with the inclusion of certain quantities. Instead, the ratio between quantities was the best performance predictor. The authors conclude that quantity-based judgments rely on an analogical system, not a discrete object file model or perceptual estimation mechanism, such as subitizing.     

Spontaneous logicomathematical constructions by chimpanzees (Pan troglodytes, P. paniscus)

Two experiments investigated the spontaneous construction of precursory logicomathematical operations by human-enculturated and language-reared chimpanzees (Pan troglodytes, Pan paniscus) when they were interacting freely with objects. In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with sets of six objects. Chimpanzees constructed equivalence, order and reversibility relations within single sets of objects as well as between two or three contemporaneous sets of objects. The chimpanzees’ logicomathematical operations were more advanced, including infrequent and minimal operations on three sets, than those of some previously investigated younger nonenculturated common chimpanzees. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with sets of 12 objects. Chimpanzees constructed more advanced operations on single sets, but not on contemporaneous sets. The results suggest partial convergence and partial divergence between development of logicomathematical cognition in chimpanzees and humans. Received: 20 January 1999 / Accepted after revision: 9 August 1999     

Piagetian liquid conservation in the great apes (Pan paniscus, Pan troglodytes, and Pongo pygmaeus)

An understanding of Piagetian liquid conservation was investigated in 4 bonobos (Pan paniscus), 5 chimpanzees (Pan troglodytes), and 5 orangutans (Pongo pygmaeus). The apes were tested in the ability to track the larger of 2 quantities of juice that had undergone various kinds of transformations. The accuracy of the apes' judgment depended on the shape or number of containers into which the larger quantity was transferred. The apes made their choice mainly on the basis of visual estimation but showed modest success when the quantities were occluded. The results suggest that the apes rely to a greater extent on visual information, although they might have some appreciation of the constancy of liquid quantities.     

Prospective memory in children and chimpanzees

Prospective memory (PM) involves remembering to do something at a specific time in the future. Here, we investigate the beginnings of this ability in young children (3-year-olds; Homo sapiens) and chimpanzees (Pan troglodytes) using an analogous task. Subjects were given a choice between two toys (children) or two food items (chimpanzees). The selected item was delivered immediately, whereas the unselected item was hidden in an opaque container. After completing an ongoing quantity discrimination task, subjects could request the hidden item by asking for it (children) or by pointing to the container and identifying the item on a symbol board (chimpanzees). Children and chimpanzees showed evidence of prospective-like memory in this task, as evidenced by successful retrieval of the item at the end of the task, sometimes spontaneously with no prompting from the experimenter. These findings contribute to our understanding of PM from an ontogenetic and comparative perspective.     

Inferences by exclusion in the great apes: the effect of age and species

This study investigated the ability of chimpanzees, gorillas, orangutans, and bonobos to make inferences by exclusion using the procedure pioneered by Premack and Premack (Cognition 50:347–362, 1994) with chimpanzees. Thirty apes were presented with two different food items (banana vs. grape) on a platform and covered with identical containers. One of the items was removed from the container and placed between the two containers so that subjects could see it. After discarding this item, subjects could select between the two containers. In Experiment 1, apes preferentially selected the container that held the item that the experimenter had not discarded, especially if subjects saw the experimenter remove the item from the container (but without seeing the container empty). Experiment 3 in which the food was removed from one of the containers behind a barrier confirmed these results. In contrast, subjects performed at chance levels when a stimulus (colored plastic chip: Exp. 1; food item: Exp. 2 and Exp. 3) designated the item that had been removed. These results indicated that apes made inferences, not just learned to use a discriminative cue to avoid the empty container. Apes perceived and treated the item discarded by the experimenter as if it were the very one that had been hidden under the container. Results suggested a positive relationship between age and inferential ability independent of memory ability but no species differences.This contribution is part of the special issue “Animal Logics” (Watanabe and Huber 2006).     

An olfactory discrimination procedure for mice

This paper describes an olfactory discrimination procedure for mice that is inexpensively implemented and leads to rapid discrimination learning. Mice were first trained to dig in small containers of sand to retrieve bits of buried chocolate. For discrimination training, two containers were presented simultaneously for eight trials per session. One container held sand mixed with cinnamon, and the other held sand mixed with nutmeg. Both containers were baited with chocolate buried in the sand. One odor was designated S+, and mice were allowed to dig and retrieve the chocolate from this container. The other odor was S-, and both containers were removed immediately if subjects began to dig in an S- container. After meeting a two-session acquisition criterion, subjects were given a series of discrimination reversals. In Experiment 1, 12 Swiss-Webster mice (6 male and 6 female) acquired the olfactory discrimination in three to five sessions and completed 3 to 10 successive discrimination reversals within a 50-session testing limit. In Experiment 2, subjects were 14 Pah(enu2) mice, the mouse mutant for phenylketonuria; 7 were homozygotes in which the disorder was expressed (PKU), and 7 were heterozygotes with normal metabolism (non-PKU). Thirteen mice completed pretraining in four to seven sessions, acquisition required 3 to 12 sessions, and all mice completed at least three reversals. Learning rates were similar in PKU and non-PKU mice. We discuss issues related to implementation and several potentially useful procedural variations.     

Object permanence in orangutans (Pongo pygmaeus), chimpanzees (Pan troglodytes), and children (Homo sapiens).

Juvenile and adult orangutans (n = 5; Pongo pygmaeus), chimpanzees (n = 7; Pan troglodytes), and 19- and 26-month-old children (n = 24; Homo sapiens) received visible and invisible displacements. Three containers were presented forming a straight line, and a small box was used to displace a reward under them. Subjects received 3 types of displacement: single (the box visited 1 container), double adjacent (the box visited 2 contiguous containers), and double nonadjacent (the box visited 2 noncontiguous containers). All species performed at comparable levels, solving all problems except the invisible nonadjacent displacements. Visible displacements were easier than invisible, and single were easier than double displacements. In a 2nd experiment, subjects saw the baiting of either 2 adjacent or 2 nonadjacent containers with no displacements. All species selected the empty container more often when the baited containers were nonadjacent than when they were adjacent. It is hypothesized that a response bias and inhibition problem were responsible for the poor performance in nonadjacent displacements.     

Apes' and children's understanding of cooperative and competitive motives in a communicative situation

Chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) (Study 1) and 18- and 24-month-old human children (Study 2) participated in a novel communicative task. A human experimenter (E) hid food or a toy in one of two opaque containers before gesturing towards the reward's location in one of two ways. In the Informing condition, she attempted to help the subject find the hidden object by simply pointing to the correct container. In the Prohibiting condition, E held out her arm toward the correct container (palm out) and told the subject firmly 'Don't take this one.' As in previous studies, the apes were at chance in the Informing condition. However, they were above chance in the new Prohibiting condition. Human 18-month-olds showed this same pattern of results, whereas 24-month-olds showed the opposite pattern: they were better in the Informing condition than in the Prohibiting condition. In our interpretation, success in the Prohibiting condition requires subjects to understand E's goal toward them and their behavior, and then to make an inference (she would only prohibit if there were something good in there). Success in the Informing condition requires subjects to understand a cooperative communicative motive - which apparently apes and young infants find difficult.     

Spontaneous use of tools as straws in great apes

Great apes can use multiple tools to extract food embedded in substrates and can invent new ways to exploit those resources. We tested five bonobos, five chimpanzees, and six orangutans in a task in which they had to use (and modify) a tool as a straw to drink the juice located inside a container. Experiment 1 showed that four orangutans and one chimpanzee invented the use of a piece of electric cable to get the juice. Experiment 2 investigated whether subjects could transform a non-functional hose into a functional one by removing blockages that impeded the free flow of juice. Orangutans outperformed chimpanzees and bonobos by differentially removing those blockages that prevented the flow of juice, often doing so before attempting to extract the juice. In Experiment 3, we presented chimpanzees and orangutans with four 3-tool sets (each tool set contained a single straw-like tool) and allowed them to select one tool. Unlike chimpanzees, orangutans succeeded in selecting the straw-like tool above chance levels without having to physically manipulate it. We suggest that orangutans’ superior performance is related to their greater reliance on mouth actions during foraging. Experiment 4 investigated whether orangutans were also capable of selecting the suitable tool not by its appearance, but by the effects that it produced. After witnessing the experimenter blow bubbles or absorb liquid with a functional tool but fail to accomplish the same thing with the non-functional tool, orangutans failed to select the functional tool above chance levels.     

Chimpanzee handedness revisited: 55 years since Finch (1941)

Chimpanzees and other great apes have long held the fascination of psychologists because of their morphological and behavioral similarities to humans. This paper describes the historical interest in studies on chimpanzee handedness and reviews current findings. Data are presented which suggest that transient changes in posture result in the transient expression of right-handedness in chimpanzees. The role of tool use as an evolutionary mechanism underlying the expression of right-handedness is challenged. Rather, emphasis is placed on the role of bimanual feeding as a behavioral adaptation for the expression of handedness. Suggestions for further research on the nature of nonhuman primate handedness are made in light of these findings.     

Generalising principles in spite of procedural differences: Children's understanding of division

Between ages 5 and 7, children are known to be quite good at sharing discrete quantities but very bad at sharing continuous quantities. Our aim was to find whether they can transfer their understanding of logical relations from discrete to continuous quantities though the procedures used in sharing these quantities are markedly different.Two samples of 5- to 7-year-olds participated in two studies. In the first study, the items involved partitive division; in the second, quotitive division tasks. In both studies, the children solved tasks with discrete and continuous quantities.Performance varied significantly across age level and logical principle (equivalence between different rounds of sharing versus inverse relation between the divisor and the quotient) but not across type of quantity (discrete versus continuous). There was a very strong relation between performance across type of quantity. We conclude that children can generalise reasoning principles in division across type of quantity in spite of the difference in sharing procedures.     

Spontaneous class grouping behavior by bonobos (Pan paniscus) and common chimpanzees (P. troglodytes)

Two experiments investigated spontaneous class grouping behavior by human-enculturated and language-reared bonobos (Pan paniscus) and common chimpanzees (P. troglodytes). In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with six objects. The objects embodied three conditions: additive, multiplicative and disjoint classes. All chimpanzees spontaneously produced single- and two-category classifying. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with 12 objects in the same class conditions. Chimpanzees mainly produced single-category classifying. Their two-category classifying was more rudimentary than that found in experiment 1. Chimpanzees did not produce any three-category classifying which would be necessary to construct the hierarchies that humans begin to construct during early childhood. Received: 1 February 1999 / Accepted after revision: 9 August 1999     

The discrimination of discrete and continuous amounts in African grey parrots (Psittacus erithacus)

A wealth of research in infants and animals demonstrates discrimination of quantities, in some cases nonverbal numerical perception, and even elementary calculation capacities. We investigated the ability of three African grey parrots (Psittacus erithacus) to select the largest amount of food between two sets, either discrete food items (experiment 1) or as volume of a food substance (experiment 2). The two amounts were presented simultaneously and were visible at the time of choice. Parrots were tested several times for all possible combinations between 1 and 5 seeds or 0.2 and 1 ml of food substance. In both conditions, subjects performed above chance for almost all combinations. Accuracy was negatively correlated with the ratio, that is performance improved with greater differences between amounts. Therefore, these results with both individual items and volume discrimination suggest that parrots use an analogue of magnitude, rather than object-file mechanisms to quantify items and substances.     

Chimpanzees (Pan troglodytes) know what can be seen from where

Visual perspective taking research has established that chimpanzees recognize what others can or cannot see in the presence of occluding barriers. Less is known about chimpanzees’ appreciation of what they themselves can or cannot see in similar situations. Additionally, it is unclear whether chimpanzees must rely on others’ gaze cues to solve such tasks or whether they have a more general appreciation of what can be seen from where. Hence, we investigated chimpanzees’ ability to anticipate what they would or would not be able to see from different visual perspectives. Food was hidden among arrays of open containers, with different containers providing visual access from unique viewing perspectives. Chimpanzees immediately adopted the correct perspective for each container type. Follow-up experiments showed that they were not simply moving to align themselves with visible openings. Our study thus suggests that chimpanzees have good visual perspective taking abilities with regard to themselves as well as others, and that both likely reflect a more general knowledge, at least implicit, of what can be seen from where.