Tracking and inferring spatial rotation by children and great apes

Finding hidden objects in space is a fundamental ability that has received considerable research attention from both a developmental and a comparative perspective. Tracking the rotational displacements of containers and hidden objects is a particularly challenging task. This study investigated the ability of 3-, 5-, 7-, and 9-year-old children and great apes (chimpanzees, bonobos, gorillas, and orangutans) to (a) visually track rotational displacements of a baited container on a platform and (b) infer its displacements by using the changes of position or orientation of 3 landmarks: an object on a container, the color of the containers, and the color of the platform on which the containers rested. Great apes and 5-year-old and older children successfully tracked visible rotations, but only children were able to infer the location of a correct cup (with the help of landmarks) after invisible rotations. The ability to use landmarks changed with age so that younger children solved this task only with the most explicit marker on the baited container, whereas older children, particularly 9-year-olds, were able to use landmark orientation to infer correct locations.     

Tracking the displacement of objects: a series of tasks with great apes (Pan troglodytes, Pan paniscus, Gorilla gorilla, and Pongo pygmaeus) and young children (Homo sapiens)

The authors administered a series of object displacement tasks to 24 great apes and 24 30-month-old children (Homo sapiens). Objects were placed under 1 or 2 of 3 cups by visible or invisible displacements. The series included 6 tasks: delayed response, inhibition test, A not B, rotations, transpositions, and object permanence. Apes and children solved most tasks performing at comparable levels except in the transposition task, in which apes performed better than children. Ape species performed at comparable levels in all tasks except in single transpositions, in which chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) performed better than gorillas (Gorilla gorilla) and orangutans (Pongo pygmeaus). All species found nonadjacent trials and rotations especially difficult. The number of elements that changed locations, the type of displacement, and having to inhibit predominant reaching responses were factors that negatively affected the subjects' performance.     

Do orangutans (<Emphasis Type="Italic">Pongo pygmaeus</Emphasis>) know when they do not remember?

Metacognition refers to the ability to monitor and control one’s own cognitive activities such as memory. Although recent studies have raised an interesting possibility that some species of nonhuman animals might possess such skills, subjects often required a numerous number of training trials to acquire the effective use of metacognitive responses. Here, five orangutans (Pongo pygmaeus) were tested whether they were able to escape spatial memory tests when they did not remember the location of preferred reward in a relatively small number of trials. The apes were presented with two identical cups, under one of which the experimenter hid a preferred reward (e.g., two grapes). The subjects were then presented with a third container, “escape response”, with which they could receive a less preferred but secure reward (e.g., one grape). The orangutans as a group significantly more likely selected the escape response when the baiting of the preferred reward was invisible (as compared to when it was visible) and when the hiding locations of the preferred reward were switched (as compared to when they remained unchanged). Even when the escape response was presented before the final presentation of the memory test, one orangutan successfully avoided the test in which she would likely err. These findings indicate that some orangutans appear to tell when they do not remember correct answers in memory tests.     

Invisible displacement understanding in domestic dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>): the role of visual cues in search behavior

Recently, (Collier-Baker E, Davis JM, Suddendorf T (2004) J Comp Psychol 118:421–433) suggested that domestic dogs do not understand invisible displacements. In the present study, we further investigated the hypothesis that the search behavior of domestic dogs in invisible displacements is guided by various visual cues inherent to the task rather than by mental representation of an object’s past trajectory. Specifically, we examined the role of the experimenter as a function of the final position of the displacement device in the search behavior of domestic dogs. Visible and invisible displacement problems were administered to dogs (N = 11) under two conditions. In the Visible-experimenter condition, the experimenter was visible whereas in the Concealed-experimenter condition, the experimenter was visibly occluded behind a large rigid barrier. Our data supported the conclusion that dogs do not understand invisible displacements but primarily search as a function of the final position of the displacement device and, to a lesser extent, the position of the experimenter.     

Gorillas’ use of the escape response in object choice memory tests

The ability to monitor and control one’s own cognitive states, metacognition, is crucial for effective learning and problem solving. Although the literature on animal metacognition has grown considerably during last 15 years, there have been few studies examining whether great apes share such introspective abilities with humans. Here, we tested whether four gorillas could meet two criteria of animal metacognition, the increase in escape responses as a function of task difficulty and the chosen-forced performance advantage. During testing, the subjects participated in a series of object choice memory tests in which a preferable reward (two grapes) was placed under one of two or three blue cups. The apes were required to correctly select the baited blue cup in this primary test. Importantly, the subjects also had an escape response (a yellow cup), where they could obtain a secure but smaller reward (one grape) without taking the memory test. Although the gorillas received a relatively small number of trials and thus experienced little training, three gorillas significantly declined the memory tests more often in difficult trials (e.g., when the location of the preferred reward conflicted with side bias) than in easy trials (e.g., when there was no such conflict). Moreover, even when objective cues were eliminated that corresponded to task difficulty, one of the successful gorillas showed evidence suggestive of improved memory performance with the help of escape response by selectively avoiding trials in which he would be likely to err before the memory test actually proceeded. Together, these findings demonstrate that at least some gorillas may be able to make optimal choices on the basis of their own memory trace strength about the location of the preferred reward.     

Where’s the cookie? The ability of monkeys to track object transpositions

Object permanence is the ability to represent mentally an object and follow its position even when it has disappeared from view. According to Piaget’s 6-stage scale of the sensorimotor period of development, it seems that object permanence appears in Stage 4 and fully develops in Stage 6. In this study, we investigated the ability of some species of monkeys (i.e. pig-tailed macaque, lion-tailed macaque, Celebes crested macaque, barbary macaque, De Brazza’s monkey, L’Hoest’s monkey, Allen’s swamp monkey, black crested mangabeys, collared mangabeys, Geoffroy’s spider monkey) to track the displacement of an object, which consisted of a reward hidden under one of two cups. Our findings showed that the examined subjects possess Stage 6 of object permanence. We then compared our results with data on apes and dogs participating in Rooijakkers et al. (Anim Cogn 12:789–796, 2009) experiment, where the same method was applied. The monkeys examined by us performed significantly better than the dogs but worse than the apes. In our experiment, the monkeys performed above chance level in all variants, but it should be noted that we observed significant differences in the number of correct choices according to the level of a variant’s complexity.     

Developments in infants' search for displaced objects

In an exploratory study of infants' search for displaced objects, 13-month-olds and 21-month-olds were tested on three kinds of displacement problems—visible displacements, invisible displacements, and transpositions—as well as singlehiding problems. Two aspects of performance on the displacement problems were distinguished: (1) searching within the locations involved in the displacement rather than at a control location, and (2) selecting between the displacement locations. Both the younger and the older infants were able to select the correct location on visible displacement problems and to search within the displacement locations on invisible problems. Neither age group was able to solve the transposition problems, but the older infants did at least search within the relevant locations on those problems. Age differences appeared to be due primarily to improved skill at identifying relevant locations rather than to improvements in selecting among those locations. Other factors contributing to the age differences in performance were a decrease in response biases and an increase in skills for coping with multiple possibilities.     

Great apes’ strategies to map spatial relations

We investigated reasoning about spatial relational similarity in three great ape species: chimpanzees, bonobos, and orangutans. Apes were presented with three spatial mapping tasks in which they were required to find a reward in an array of three cups, after observing a reward being hidden in a different array of three cups. To obtain a food reward, apes needed to choose the cup that was in the same relative position (i.e., on the left) as the baited cup in the other array. The three tasks differed in the constellation of the two arrays. In Experiment 1, the arrays were placed next to each other, forming a line. In Experiment 2, the positioning of the two arrays varied each trial, being placed either one behind the other in two rows, or next to each other, forming a line. Finally, in Experiment 3, the two arrays were always positioned one behind the other in two rows, but misaligned. Results suggested that apes compared the two arrays and recognized that they were similar in some way. However, we believe that instead of mapping the left–left, middle–middle, and right–right cups from each array, they mapped the cups that shared the most similar relations to nearby landmarks (table’s visual boundaries).     

Great apes use landmark cues over spatial relations to find hidden food

We investigated whether chimpanzees, bonobos, and orangutans encoded the location of a reward hidden underneath one of three identical cups in relation to (1) the other cups in the array-i.e., the relative position of the baited cup within the array; or (2) the landmarks surrounding the cups-e.g., the edge of the table. Apes witnessed the hiding of a food reward under one of three cups forming a straight line on a platform. After 30?s, they were allowed to search for the reward. In three different experiments, we varied the distance of the cups to the edge of the platform and the distance between the cups. Results showed that both manipulated variables affected apes' retrieval accuracy. Subjects' retrieval accuracy was higher for the outer cups compared with the Middle cup, especially if the outer cups were located next to the platform's edge. Additionally, the larger the distance between the cups, the better performance became.     

Gravity bias in young and adult chimpanzees (Pan troglodytes): tests with a modified opaque-tubes task

Young human children at around 2 years of age fail to predict the correct location of an object when it is dropped from the top of an S-shape opaque tube. They search in the location just below the releasing point (Hood, 1995). This type of error, called a 'gravity bias', has recently been reported in dogs and monkeys. In the present study, we investigated whether young and adult chimpanzees also show such a gravity bias in a modified version of the original opaque-tube task. The original task by Hood and colleagues required the subject to search in a location after the object had fallen, while in the task reported here, subjects were required to predict the location before the object was dropped. Thus the present procedure does not involve explicit invisible displacement operations, one of the important components of the original procedure. In Experiment 1 both young (1.5-2.5-year-old) and adult chimpanzees predicted the location of falling food items below the releasing point even when crossed tubes were used. These gravity errors remained after the extensive experience of using the tubes themselves. Experiment 2 further tested adult and 4-year-old chimpanzees under the set-up in which the straight and crossed tubes were simultaneously presented. The results were the same as those in the previous test, suggesting that developmental changes and learning effect do not affect the gravity bias in chimpanzees.     

What do dolphins (Tursiops truncatus) understand about hidden objects?

Object permanence, the ability to mentally represent and reason about objects that have disappeared from view, is a fundamental cognitive skill that has been extensively studied in human infants and terrestrial animals, but not in marine animals. A series of four experiments examined this ability in bottlenose dolphins (Tursiops truncatus). After being trained on a “find the object” game, dolphins were tested on visible and invisible displacement tasks, and transpositions. In Experiments 1 and 2, dolphins succeeded at visible displacements, but not at invisible displacements or transpositions. Experiment 3 showed that they were able to pass an invisible displacement task in which a person’s hand rather than a container was used as the displacement device. However, follow-up controls suggested they did so by learning local rules rather than via a true representation of the movement of hidden objects. Experiment 4 demonstrated that the dolphins did not rely on such local rules to pass visible displacement tasks. Thus, like many terrestrial animals, dolphins are able to succeed on visible displacement tasks, but seem unable to succeed on tasks requiring the tracking of hidden objects.     

A test of object permanence in a new-world monkey species,cotton top tamarins (<Emphasis Type="Italic">Saguinus oedipus</Emphasis>)

Cotton top tamarins were tested in visible and invisible displacement tasks in a method similar to that used elsewhere to test squirrel monkeys and orangutans. All subjects performed at levels significantly above chance on visible (n=8) and invisible (n=7) displacements, wherein the tasks included tests of the perseverance error, tests of memory in double and triple displacements, and "catch" trials that tested for the use of the experimenter's hand as a cue for the correct cup. Performance on all nine tasks was significantly higher than chance level selection of cups, and tasks using visible displacements generated more accurate performance than tasks using invisible displacements. Performance was not accounted for by a practice effect based on exposure to successive tasks. Results suggest that tamarins possess stage 6 object permanence capabilities, and that in a situation involving brief exposure to tasks and foraging opportunities, tracking objects' movements and responding more flexibly are abilities expressed readily by the tamarins. Electronic Publication     

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.     

Chimpanzees (Pan troglodytes) remember the location of a hidden food item after altering their orientation to a spatial array

Two chimpanzees (Pan troglodytes) had a direct view of an experimenter placing a food item beneath one of several cups within a horizontal spatial array. The chimpanzees then were required to move around the spatial array, shifting their orientation to the array by 180 degrees . Both chimpanzees remembered the location of the food item. In the next experiment, a visual barrier was placed between the chimpanzees and the spatial array after the food item had been hidden to prevent visual tracking of the location of the object during the chimpanzees' movement. One chimpanzee remembered the location of the hidden item in this variation. These results demonstrate another capacity for spatial memory in this species that complements data indicating chimpanzee spatial memory for invisible displacements, array rotations, and array transpositions.     

Comparing eye movements during position tracking and identity tracking: No evidence for separate systems

There is an ongoing debate as to whether people track multiple moving objects in a serial fashion or with a parallel mechanism. One recent study compared eye movements when observers tracked identical objects (Multiple Object Tracking—MOT task) versus when they tracked the identities of different objects (Multiple Identity Tracking—MIT task). Distinct eye-movement patterns were found and attributed to two separate tracking systems. However, the same results could be caused by differences in the stimuli viewed during tracking. In the present study, object identities in the MIT task were invisible during tracking, so observers performed MOT and MIT tasks with identical stimuli. Observer were able to track either position and identity depending on the task. There was no difference in eye movements between position tracking and identity tracking. This result suggests that, while observers can use different eye-movement strategies in MOT and MIT, it is not necessary.     

Attentional resources in visual tracking through occlusion: the high-beams effect

A considerable amount of research has uncovered heuristics that the visual system employs to keep track of objects through periods of occlusion. Relatively little work, by comparison, has investigated the online resources that support this processing. We explored how attention is distributed when featurally identical objects become occluded during multiple object tracking. During tracking, observers had to detect small probes that appeared sporadically on targets, distracters, occluders, or empty space. Probe detection rates for these categories were taken as indexes of the distribution of attention throughout the display and revealed two novel effects. First, probe detection on an occluder’s surface was better when either a target or distractor was currently occluded in that location, compared to when no object was behind that occluder. Thus even occluded (and therefore invisible) objects recruit object-based attention. Second, and more surprising, probe detection for both targets and distractors was always better when they were occluded, compared to when they were visible. This new attentional high-beams effect indicates that the ability to track through occlusion, though seemingly effortless, in fact requires the active allocation of special attentional resources.     

The magic cup: great apes and domestic dogs (Canis familiaris) individuate objects according to their properties

Despite current interest in dog (Canis familiaris) cognition, very little is known about how dogs represent objects and how they compare with other species, such as the great apes. Therefore, we investigated how dogs and great apes (chimpanzees [Pan troglodytes], bonobos [Pan paniscus], orangutans [Pongo pygmaeus], gorillas [Gorilla gorilla]) individuate objects in a classical violation of expectation paradigm. We used a container (magic cup) with a double bottom that allowed us to change the type of food that subjects had seen being placed in the container. Using a 2 × 2 design, we varied whether subjects received a generally preferred food and whether the food was substituted (surprise trials) or not (baseline trials). Apes showed increased begging and looking behaviors and dogs showed increased smelling behavior. Both species stayed near the experimenter more frequently in the surprise trials compared with baseline trials. Both species reacted to positive (i.e., good food substituted for bad food) and negative (i.e., bad food substituted for good food) surprises. These results suggest that apes and dogs were able to individuate objects according to their properties or type in comparable ways. In addition, we looked for frustration and elation effects, but subjects' behaviors were not influenced by the food they saw and which they received in previous trials.     

Object permanence in lemurs

Object permanence, the ability to mentally represent objects that have disappeared from view, should be advantageous to animals in their interaction with the natural world. The objective of this study was to examine whether lemurs possess object permanence. Thirteen adult subjects representing four species of diurnal lemur (Eulemur fulvus rufus, Eulemur mongoz, Lemur catta and Hapalemur griseus) were presented with seven standard Piagetian visible and invisible object displacement tests, plus one single visible test where the subject had to wait predetermined times before allowed to search, and two invisible tests where each hiding place was made visually unique. In all visible tests lemurs were able to find an object that had been in clear view before being hidden. However, when lemurs were not allowed to search for up to 25-s, performance declined with increasing time-delay. Subjects did not outperform chance on any invisible displacements regardless of whether hiding places were visually uniform or unique, therefore the upper limit of object permanence observed was Stage 5b. Lemur species in this study eat stationary foods and are not subject to stalking predators, thus Stage 5 object permanence is probably sufficient to solve most problems encountered in the wild.     

Search behaviour of cats (Felis catus) in an invisible displacement test: cognition and experience

An invisible displacement test was administered to cats in order to test the hypothesis that search behaviour in this species is influenced by their limited capacity for object permanence as well as by their previous experience with the environment. Experiment 1 compared three groups of cats in a five-choice hiding task in which the hiding places could be discriminated by their spatial positions. Two groups received a visible displacement training before the invisible displacement test and one group did not. Experiment 2 compared two groups of trained subjects in the same task, but the hiding places could be discriminated by spatial and visual cues. The results confirmed that cats are unable to solve problems with invisible displacements. The visible displacement training improved their performance, but was not sufficient to make them succeed. Experience with the hiding potential of the covers also gives more persistence to search behaviour. Finally, the distribution of search attempts is not determined by the proximity to the target and is influenced only partially by the subjects' previous experience. Like Stage 5 infants, cats rely mainly on their immediate perception. They search for an object in the last location they have seen it disappear or under the nearest cover from this location.     

Capuchin monkeys (Cebus apella) use positive, but not negative, auditory cues to infer food location

Nonhuman primates appear to capitalize more effectively on visual cues than corresponding auditory versions. For example, studies of inferential reasoning have shown that monkeys and apes readily respond to seeing that food is present (“positive” cuing) or absent (“negative” cuing). Performance is markedly less effective with auditory cues, with many subjects failing to use this input. Extending recent work, we tested eight captive tufted capuchins (Cebus apella) in locating food using positive and negative cues in visual and auditory domains. The monkeys chose between two opaque cups to receive food contained in one of them. Cup contents were either shown or shaken, providing location cues from both cups, positive cues only from the baited cup, or negative cues from the empty cup. As in previous work, subjects readily used both positive and negative visual cues to secure reward. However, auditory outcomes were both similar to and different from those of earlier studies. Specifically, all subjects came to exploit positive auditory cues, but none responded to negative versions. The animals were also clearly different in visual versus auditory performance. Results indicate that a significant proportion of capuchins may be able to use positive auditory cues, with experience and learning likely playing a critical role. These findings raise the possibility that experience may be significant in visually based performance in this task as well, and highlight that coming to grips with evident differences between visual versus auditory processing may be important for understanding primate cognition more generally.