Search behavior in various breeds of adult dogs (Canis familiaris): object permanence and olfactory cues.

Human analog tests of object permanence were administered to various breeds of adult dogs (Canis familiaris). Experiment 1 showed that the performance of terriers, sporting, and working dogs did not differ. Dogs succeeded in solving invisible displacement problems, but performance was lower than in visible displacement tests. Familiarity with the task had some influence because invisible displacement tests were more successful if they were preceded by visible displacement tests. In Experiment 2, odor cues from the target object and the hiding screens were available or were masked. Results confirmed that success was lower in invisible than in visible displacement tests and that these problems were solved on the basis of representation of visual information rather than on the basis of olfactory cues or of local rule learning. Dogs are compared with other species that display Stage 6 object permanence.     

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.     

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     

The ecology of spatial memory in four lemur species

Evolutionary theories suggest that ecology is a major factor shaping cognition in primates. However, there have been few systematic tests of spatial memory abilities involving multiple primate species. Here, we examine spatial memory skills in four strepsirrhine primates that vary in level of frugivory: ruffed lemurs (Varecia sp.), ring-tailed lemurs (Lemur catta), mongoose lemurs (Eulemur mongoz), and Coquerel’s sifakas (Propithecus coquereli). We compare these species across three studies targeting different aspects of spatial memory: recall after a long-delay, learning mechanisms supporting memory and recall of multiple locations in a complex environment. We find that ruffed lemurs, the most frugivorous species, consistently showed more robust spatial memory than the other species across tasks—especially in comparison with sifakas, the most folivorous species. We discuss these results in terms of the importance of considering both ecological and social factors as complementary explanations for the evolution of primate cognitive skills.     

Conflict and postconflict behaviour in two lemur species with different social organizations (Eulemur fulvus and Eulemur macaco): A study on captive groups

Although the brown lemur (Eulemur fulvus) and the black lemur (Eulemur macaco) share many life parameters and are genetically closed, they show considerable difference in social organization. Dominance relationships with no systematic effect of gender characterize the former, whereas the latter is based on female dominance. The present study was done on two captive groups of brown lemurs and one semi–free‐ranging group of black lemurs. To reveal links between the specific pattern of social organization and agonistic behaviors, agonistic interactions were analyzed for each species as for their context of occurrence, symmetry, initiation, and outcome. The effect of gender in the initiation of conflicts appeared as the only notable interspecific difference, aggression being mostly initiated by females in E. macaco and by males in E. fulvus. Conflict outcome was generally in favor of the initiator, regardless of gender in both species. The analysis of postconflict behaviors revealed conciliatory processes in the brown lemur, whereas reconciliation seemed to be absent in the black lemur, a characteristic shared with the ring‐tailed lemur (Lemur catta), another lemur species with female dominance. Aggr. Behav. 28:62–74, 2002. © 2002 Wiley‐Liss, Inc.     

The effects of task and practice on search for invisibly displaced objects

The ability to successfully search for an invisibly displaced object is frequently assumed to relate to the emergence of mental representation at around 2 years of age. However, little is known about what is actually being measured in the typical “stage 6” object permanence search task. Two studies examined the effects of task factors and practice on invisible displacements and systematic search tasks to show that simple variations in ways of hiding an object change the child's ability to find it. Experiment 1 compared two common methods of presenting invisible displacements items to 36 children, approximately 2 years of age. One task was significantly easier than the other, despite the fact that they contained identical items that varied only in their order of presentation. Experiment 2 tested the effects of short-term practice on 18- and 24-month-olds' success on invisible displacements trials and on two systematic search tasks. Results indicate that too little practice inhibited performance in 18-month-olds, but that practice beyond an optimum level did not further affect their scores. In contrast, 24-month-olds continued to benefit from increased practice until they reached the maximum possible score. The findings are interpreted to mean that invisible displacements tasks may require only sensorimotor search strategies rather than representation. The findings also have implications for (a) research that attempts to relate object permanence to other variables and (b) programs that use object permanence as an assessment or screening measure.     

Mapping the contours of faith in the land of separation: spiritual geographies for children

According to Jean Piaget, children begin to develop a concept of an object, such as that it has sides that are not visible from the child's perspective or that it is likely to be where one saw it last, in early infancy. By the close of the prelinguistic phase at about 2 years old, the child has developed a mature object concept, one that comprehends the object as a continuing entity even when it is not visible. Many children's picture books demonstrate Piaget's concept of object permanence through narrative and image. This paper offers a close reading of three classics, Goodnight moon, Harold and the purple crayon and Where the wild things are, in an explanation of how children are able to develop faith in an invisible, omnipresent deity.     

THIS ARTICLE HAS BEEN RETRACTED: Enumeration of objects and substances in non‐human primates: experiments with brown lemurs (Eulemur fulvus)

Both human infants and adult non‐human primates share the capacity to track small numbers of objects across time and occlusion. The question now facing developmental and comparative psychologists is whether similar mechanisms give rise to this capacity across the two populations. Here, we explore whether non‐human primates’ object tracking abilities are subject to the same constraints as those of human infants. In particular, we examine whether one primate species, the brown lemur (Eulemur fulvus), also fails to represent and enumerate objects when they behave non‐rigidly or non‐cohesively. We presented lemurs with a series of expectancy violation studies involving simple 1 + 1 addition events in which we varied the entities to be enumerated. Like infants, lemurs successfully enumerated the two objects when those objects were rigid, cohesive individuals, but failed to enumerate similar‐looking non‐rigid piles of sand. In contrast to human infants, however, lemurs successfully enumerated non‐cohesive objects that broke into multiple pieces. These results are discussed in light of recent theories about object processing in human infants and adults.     

Object permanence in common marmosets (Callithrix jacchus)

A series of 9 search tasks corresponding to the Piagetian Stages 3-6 of object permanence were administered to 11 common marmosets (Callithrix jacchus). Success rates varied strongly among tasks and marmosets, but the performances of most subjects were above chance level on the majority of tasks of visible and invisible displacements. Although up to 24 trials were administered in the tests, subjects did not improve their performance across trials. Errors were due to preferences for specific locations or boxes, simple search strategies, and attentional deficits. The performances of at least 2 subjects that achieved very high scores up to the successive invisible displacement task suggest that this species is able to represent the existence and the movements of unperceived objects.     

Inferences about the location of food in lemurs (Eulemur macaco and Eulemur fulvus): a comparison with apes and monkeys

The ability of black and brown lemurs (Eulemur macaco and Eulemur fulvus) to make inferences about hidden food was tested using the same paradigm as in Call’s (J Comp Psycol 118:232–241, 2004) cup task experiment. When provided with either visual or auditory information about the content of two boxes (one empty, one baited), lemurs performed better in the auditory condition than in the visual one. When provided with visual or auditory information only about the empty box, one subject out of four was above chance in the auditory condition, implying inferential reasoning. No subject was successful in the visual condition. This study reveals that (1) lemurs are capable of inferential reasoning by exclusion and (2) lemurs make better use of auditory than visual information. The results are compared with the performances recorded in apes and monkeys under the same paradigm.     

Expectations about numerical events in four lemur species (Eulemur fulvus, Eulemur mongoz, Lemur catta and Varecia rubra)

Although much is known about how some primates—in particular, monkeys and apes—represent and enumerate different numbers of objects, very little is known about the numerical abilities of prosimian primates. Here, we explore how four lemur species (Eulemur fulvus, E. mongoz, Lemur catta, and Varecia rubra) represent small numbers of objects. Specifically, we presented lemurs with three expectancy violation looking time experiments aimed at exploring their expectations about a simple 1+1 addition event. In these experiments, we presented subjects with displays in which two lemons were sequentially added behind an occluder and then measured subjects duration of looking to expected and unexpected outcomes. In experiment 1, subjects looked reliably longer at an unexpected outcome of only one object than at an expected outcome of two objects. Similarly, subjects in experiment 2 looked reliably longer at an unexpected outcome of three objects than at an expected outcome of two objects. In experiment 3, subjects looked reliably longer at an unexpected outcome of one object twice the size of the original than at an expected outcome of two objects of the original size. These results suggest that some prosimian primates understand the outcome of simple arithmetic operations. These results are discussed in light of similar findings in human infants and other adult primates.This revised version was published online in March 2005 with corrections to Fig. 6.     

The role of attentiveness,mobility history,and separation of hiding sites on stage IV search behavior

The relationship between visual attentiveness, search behavior, and duration of independent mobility was examined for fifty-six 8- to 10-month-old infants presented with three versions of the Stage IV object permanence task. The number and spatial separation of hiding sites was manipulated to explore the role of these factors on visual attention and search performance. In a repeated measures design, perseverative search errors were less likely on both a two- and a six-location task with equally wide separation of the A and B hiding sites than on a two-location task with A and B close together, thus indicating that spatial separation of sites is a more important contributor to successful search than number of hiding sites alone. In addition, visual attentiveness was significantly associated with correct search at B in all three versions of the task. A significant developmental relationship was found between the length of time infants had been independently mobile and visual attentiveness during the hiding procedure. These findings are discussed in terms of the transition to self-produced mobility and expanding spatial experience in the second half of the first year and how visual attentiveness and search performance might improve as a function of such changes.     

Gaze following and gaze priming in lemurs

Although primates have often been found to co-orient visually with other individuals, members of these same species have usually failed to use co-orientation to find hidden food in object-choice experiments. This presents an evolutionary puzzle: what is the function of co-orientation if it is not used for a function as basic as locating resources? Co-orientation responses have not been systematically investigated in object-choice experiments, and requiring co-orientation with humans (as is typical in object-choice tasks) may underestimate other species’ abilities. Using an object-choice task with conspecific models depicted in photographs, we provide experimental evidence that two lemur species (Eulemur fulvus, n = 4, and Eulemur macaco, n = 2) co-orient with conspecifics. Secondly, by analysing together two measures that have traditionally been examined separately, we show that lemurs’ gaze following behaviour and ultimate choice are closely linked. Individuals were more likely to choose correctly after having looked in the same direction as the model, and thus chose objects correctly more often than chance. We propose a candidate system for the evolutionary origins of more complex gaze following: ‘gaze priming.’     

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.     

Development of object permanence in food-storing magpies (Pica pica)

The development of object permanence was investigated in black-billed magpies (Pica pica), a food-storing passerine bird. The authors tested the hypothesis that food-storing development should be correlated with object-permanence development and that specific stages of object permanence should be achieved before magpies become independent. As predicted, Piagetian Stages 4 and 5 were reached before independence was achieved, and the ability to represent a fully hidden object (Piagetian Stage 4) emerged by the age when magpies begin to retrieve food. Contrary to psittacine birds and humans, but as in dogs and cats, no "A-not-B error" occurred. Although magpies also mastered 5 of 6 invisible displacement tasks, evidence of Piagetian Stage 6 competence was ambiguous.     

Selective information use and perseveration in the search behavior of infants and young children

Two experiments examined the early development of selective information use in search. The first experiment tested 9- and 16-month-olds on a modification of Piaget's Stage IV object permanence task. It examined infants' use of information from previous experiences with an object (prior information) and from the most recent hiding (current information) to locate a hidden object. In the second experiment, 2-, $\text{2}\text{1}\text{2}$-, and 4-year-old children received these same sources of information along with new forms of prior and current information: information about the typical locations of objects (location specificity) and verbal information. No systematic perseveration was observed at 9 months, although previous findings related to perseveration were replicated. Perseveration was found at 16 months, but there was also evidence of selectivity at that age. When errors occurred, they tended to be to the prior location, but they were infrequent in comparison to correct searches at the current location. The preschoolers, while continuing to show perseveration, were more consistently selective than the infants. They also showed considerable generality in extending their selectivity to new sources of information.     

Ontogeny of object permanence in a non-storing corvid species, the jackdaw (Corvus monedula)

The aim of the present study was to investigate the ontogeny of object permanence in a non-caching corvid species, the jackdaw (Corvus monedula). Jackdaws are often presented as typical examples of non-storing corvids, as they cache either very little or not at all. We used Uzgiris and Hunt’s Scale 1 tasks to determine the age at which the certain stages set in and the final stage of this capacity that is reached. Our results show that the lack of food-storing behaviour is not associated with inferior object permanence abilities in the jackdaw, as our subjects (N = 19) have reached stage 5 competence (to follow successive visible displacements) at the average age of 61 days post-hatch and showed some evidence of stage 6 competence (to follow advanced invisible displacements) at 81 days post-hatch and thereafter. As we appreciate that object permanence abilities have a very wide ecological significance, our positive results are probably the consequence of other, more fundamental ecological pressures, such as nest-hole reproduction or prey–predator interactions.     

Brain activation during spatial updating and attentive tracking of moving targets

Keeping aware of the locations of objects while one is moving requires the updating of spatial representations. As long as the objects are visible, attentional tracking is sufficient, but knowing where objects out of view went in relation to one's own body involves an updating of spatial working memory. Here, multiple object tracking was employed to study spatial updating and its neural correlates. In a dynamic 3D-scene, targets moved among visually indistinguishable distractors. The targets and distractors either stayed visible during continuous viewpoint changes or they turned invisible. The parametric variation of tracking load revealed load-dependent activations of the intraparietal sulcus, the superior parietal lobule, and the lateral occipital cortex in response to the attentive tracking task. Viewpoint changes with invisible objects that demanded retention and updating produced load-dependent activation only in the precuneus in line with its presumed involvement in updating spatial working memory.     

Will infants search when ‘no toy’ is hidden? A study of implicit assumptions about the development of object permanence

In the Piagetian tradition, presence or absence of a sense of object permanence is inferred from whether an infant does or does not search when an object is occluded. This line of reasoning entails an implicit assumption, i.e. an infant will not search if no object is seen to disappear. This assumption was evaluated in three studies of 9- and 12-month-old infants' responses to ‘hiding’ toy and ‘no toy’ in different orders in one and two positions. The infants did not search for ‘no toy’ when it preceded toy hiding, but they did search for ‘no toy’ when it followed ‘toy’ under some conditions.     

Looking and search measures of object knowledge in preschool children

The same preschoolers were tested on an observation task and a search task involving the invisible displacement of an object. In the observation task, children watched an object roll behind a screen from which protruded the top of a solid wall. Analyses revealed significantly longer looking to impossible than to possible outcomes in all children. In search, the child was allowed to retrieve the rolled object. Most 3-year-olds but significantly fewer 2.5-year-olds completed the search successfully. An unexpected sex difference was found, with boys outperforming girls. Search performance was not associated with observation measures. The findings indicate that children visually discriminate violations of solidity but that this sensitivity is not associated with successful search performance.