Strategies in landmark use by children, adults, and marmoset monkeys

Common marmosets (Callithrix jacchus jacchus), human children, and human adults learned to find a goal that was located in the center of a square array of four identical landmarks. The location of the landmark array and corresponding goal varied across trials, so the task could not be solved without using the landmark array. In Experiment 1, a matrix of discrete goal locations was presented and the landmarks surrounded and were adjacent to the correct location during training. After training, an expansion test was given in which the distance between landmarks was increased. Marmosets, children (ages 5–9), and adults all readily learned to use the landmarks to search accurately during training. On the expansion test, adults uniformly searched in the center of the array. Monkeys and children concentrated their searching near the landmarks rather than in the center. The monkeys, but not the children, searched more often on the directionally appropriate side of the landmarks than on other sides of the landmarks. In Experiment 2, children (ages 3–5) were trained with a continuous search space and with the goal farther from the landmarks so that a beaconing strategy rule could not be used. Several of the children failed to acquire the training task. Of those who learned to find the goal, three searched in the middle on expansion tests but most searched nearer to the landmarks. The “middle rule” strategy that is uniformly used by adult humans does not appear to be a preferred strategy for children or non-human primates.     

Hippocampus lesions impair landmark array spatial learning in homing pigeons: a laboratory study

Hippocampal (HF)-lesioned pigeons display impaired homing ability when flying over familiar terrain, where they are presumably relying on a map-like representation of familiar landmarks to navigate. However, research carried out in the field precludes a direct test of whether hippocampal lesions compromise the ability of homing pigeons to navigate by familiar landmarks. To examine more thoroughly the relationship between hippocampus and landmark spatial learning, control, neostriatum-lesioned, and HF-lesioned homing pigeons were trained on two open field, laboratory, conditional discrimination tasks. One was a visual landmark array task, and the other was a room color discrimination task. For the tasks, the correct of three differently colored food bowls was determined by the spatial relationship among a group of five landmarks and room color, respectively. Intact control birds successfully learned both tasks, while neostriatum-lesioned birds successfully learned the landmark array task-the only task on which they were trained. By contrast, HF-lesioned birds successfully learned the room color task but were unable to learn the landmark array task. The data support the hypothesis that homing performance deficits observed in the field following hippocampal lesions are in part a consequence of an impairment in the ability of lesioned pigeons to use familiar visual landmarks for navigation.     

Age-related differences in the use of background layout in visual search

The effect of background layout on visual search performance, and more specifically on the tendency to refixate previously inspected locations and objects, was investigated. Older and younger adults performed a search task in which a background layout or landmark was present or absent in a gaze contingent visual search paradigm. Regardless of age, participants demonstrated fewer refixations when landmarks were present, with older adults showing a larger landmark advantage. This visual search advantage did not come at the cost of saccadic latency. Furthermore, the visual search performance advantage obtained in the presence of a background layout or landmark was observed both for individuals with small and large memory spans.     

Children's use of geometry and landmarks to reorient in an open space

Eight experiments tested the abilities of 3-4-year-old children to reorient themselves and locate a hidden object in an open circular space furnished with three or four landmark objects. Reorientation was tested by hiding a target object inside one of the landmarks, disorienting the child, observing the child's search for the target, and comparing the child's performance to otherwise similar trials in which the child remained oriented. On oriented trials, children located the target successfully in every experiment. On disoriented trials, in contrast, children failed to locate the object when the landmarks were indistinguishable from one another but formed a distinctive geometric configuration (a triangle with sides of unequal length or a rectangle). This finding provides evidence that the children failed to use the geometric configuration of objects to reorient themselves. As in past research, children also did not appear to reorient themselves in accord with non-geometric properties of the layout. In contrast to these findings, children successfully located the object in relation to a geometric configuration of walls. Moreover, adults, who were tested in two further experiments, located the object by using both geometric and non-geometric information. Together, these ten experiments provide evidence that early-developing navigational abilities depend on a mechanism that is sensitive to the shape of the permanent, extended surface layout, but that is not sensitive to geometric or non-geometric properties of objects in the layout.     

Solving small spaces: investigating the use of landmark cues in brown capuchins (Cebus apella)

Some researchers have recently argued that humans may be unusual among primates in preferring to use landmark information when reasoning about some kinds of spatial problems. Some have explained this phenomenon by positing that our species’ tendency to prefer landmarks stems from a human-unique trait: language. Here, we test this hypothesis—that preferring to use landmarks to solve such tasks is related to language ability—by exploring landmark use in a spatial task in one non-human primate, the brown capuchin monkey (Cebus apella). We presented our subjects with the rotational displacement task, in which subjects attempt to relocate a reward hidden within an array of hiding locations which are subsequently rotated to a new position. Over several experiments, we varied the availability and the salience of a landmark cue within the array. Specifically, we varied (1) visual access to the array during rotation, (2) the type of landmark, (3) the consistency of the landmark qualities, and (4) the amount of exposure to the landmark. Across Experiments 1 through 4, capuchins did not successfully use landmarks cues, suggesting that non-linguistic primates may not spontaneously use landmarks to solve some spatial problems, as in this case of a small-scale dynamic spatial task. Importantly, we also observed that capuchins demonstrated some capacity to learn to use landmarks in Experiment 4, suggesting that non-linguistic creatures may be able to use some landmarks cues in similar spatial tasks with extensive training.     

Reorientation in the real world: the development of landmark use and integration in a natural environment

An influential series of studies have argued that young children are unable to use landmark information to reorient. However, these studies have used artificial experimental environments that may lead to an underestimation of the children’s ability. We tested whether young children could reorient using landmarks in an ecologically valid setting. Children aged between 3 and 7 years completed a reorientation task in open parkland, and the properties of the search array (size and distinctiveness) were manipulated in a within-subjects design. Responses were recorded using Global Positioning Systems technology. All age groups performed above chance level, demonstrating that young children can reorient using natural landmarks. This behaviour was modulated by the nature of the search array: children were more accurate when the locations were spaced in a large array, and when the search locations were distinctively coloured. This suggests that the integration between landmarks and search locations, at different spatial scales, is a key factor in characterising human reorientation in the real world.     

Rapid discrimination of visual patterns in children and adults

Rapid visual discrimination in children (9 to 11 yr.) and adults was investigated using two tasks in which the subject had to search for a target pattern embedded in 35 background patterns. The time available for inspecting the search arrays was altered by varying the stimulus duration. In one task, there was a large difference in the feature or 'texton' content of patterns between the target and the background; in the other, this difference was small. In the first task, the children could detect the target pattern in a briefly flashed search array with high accuracy, because the target 'popped out' perceptually from the array, but in the second task the same detection rate was not reached until the stimulus duration was much longer, since a serial searching strategy was required. Results achieved by children and adults were similar. It seems that serial visual search is as efficient in children as in adults.     

Strategies in landmark use by orangutans and human children

Landmark use has been demonstrated in a variety of organisms, yet the manner in which landmarks are encoded and subsequently used appears to vary between and sometimes within species, even when faced with identical landmark arrays. In the present experiments, orangutans and human children were shown a square array of identical landmarks and were trained to locate a hidden goal in the centre of the array. In Experiments 1 and 2, the search space appeared to be discrete, with white gridlines dividing up the space, and in Experiments 3a and 3b, the search space was uniformly coloured, making it appear continuous. In all experiments, following training, subjects were given a single expansion test, to determine their landmark strategy use, based on peak search activity. The orangutans appeared to use absolute directional vectors from individual landmarks, with peak search activities on the inner corners of the square array, and they used this strategy persistently. In contrast, human children showed two landmark-based strategies, absolute directional vectors and a relational or “middle” strategy, with the majority of children starting their search in the middle region. Although some children, especially young children, persistently used one strategy like the orangutans, many changed strategies when the original one failed to yield the hidden goal.     

The development of landmarks in spatial memory: the role of differential experience

To assess how differential experience with objects in a spatial array might serve to establish relative landmarks within the array, first and fifth graders learned models of a town and farm under two conditions. In a homogeneous condition, all elements in the array were visited an equal number of times. In a landmark condition, the relative landmark status of an element was established by distributing the same total number of visits unequally across the elements. The effects of such landmarks on spatial memory were assessed both in reconstructions of the entire array and by pairwise distance estimations. The landmark condition led to a general improvement in spatial recall accuracy as well as providing a relative landmark within the array to help organize the space. The results suggest that different levels of experience (controlling for overall experience and object salience) can establish elements as relative landmarks in spatial memory. Although there was clear developmental improvement in spatial memory, the specific landmark effects were similar for both first and fifth graders.     

Wayfinding behavior and spatial knowledge of adults and children in a virtual environment: The role of landmarks

This study investigated the effect of different organizations of landmark-location pairings as fine-space information on wayfinding behavior and spatial knowledge on a total of 90 participants: 30 second graders, 30 sixth graders, and 30 adults. All participants had to find their way to a goal in a virtual environment with either randomized or categorical landmarks, or without any landmarks. Thereafter, they had to find the shortest way from the start position to the goal in two consecutive trials (wayfinding performance), and they had to solve a number of spatial knowledge tasks. The results showed that independent of their categorical function, the existence of landmarks influenced the way-finding performance of adults and children in the same way. Whereas the presence of landmarks had no effect on spatial survey knowledge, landmark knowledge itself was influenced by the categorical function of the landmarks presented. Moreover, second graders showed limited achievement compared to adults independent of the existence of landmarks. The main results implicate firstly that children at school age indeed are able to use landmark-location pairings as fine-space information like adults during learning an unknown environmental space, and secondly that a dissociation between wayfinding behavior and spatial knowledge might exist.     

Transfer of spatial behaviour controlled by a landmark array with a distinctive shape

In two experiments, rats swam to a submerged platform in one corner of a rectangular or kite-shaped array created by four identical landmarks attached to the walls of a circular pool. After training in the rectangular array, rats expressed a preference for the corner in the kite-shaped array that was geometrically equivalent to where the platform was located previously. After training in either array, the removal of two landmarks from the rectangular array, or the landmark at the apex of the kite-shaped array, did not affect the control over searching exerted by the remaining landmarks. The results imply that rats use local rather than global spatial representations when searching for a hidden goal with reference to an array of landmarks.     

The role of landmarks and boundaries in the development of spatial memory

It has been suggested that learning an object's location relative to (1) intramaze landmarks and (2) local boundaries is supported by parallel striatal and hippocampal systems, both of which rely upon input from a third system for orientation. However, little is known about the developmental trajectories of these systems' contributions to spatial learning. The present study tested 5- and 7-year-old children and adults on a water maze-like task in which all three types of cue were available. Participants had to remember the location of an object hidden in a circular bounded environment containing a moveable intramaze landmark and surrounded by distal cues. Children performed less accurately than adults, and showed a different pattern of error. While adults relied most on the stable cue provided by the boundary, children relied on both landmark and boundary cues similarly, suggesting a developmental increase in the weighting given to boundary cues. Further, adults were most accurate in coding angular information (dependent on distal cues), whereas children were most accurate in coding distance, suggesting a developing ability to use distal cues to orient. These results indicate that children as young as 5 years use boundary, intramaze landmark, and distal visual cues in parallel, but that the basic accuracy and relative weighting of these cues changes during subsequent development.     

Landmark Stability: Further Studies Pointing to a Role in Spatial Learning

Two experiments were conducted to investigate the possible role of landmark stability in spatial learning. Rats were trained to search in a large arena for food hidden at a consistent distance and direction from either a single radially symmetric landmark or an array of two landmarks. We varied the relative degree to which the landmark array and/or the cues of the training context predicted the location of food, without varying the conditional probability of food being available given either cue. Experiment 1 used vestibular disorientation to ensure control of search location by experimenter-controlled cues. The results showed that making either a single landmark or a cluster of two adjacent landmarks the sole spatial predictor of reward location reduced the accuracy of search compared to a condition where both the landmark array and context cues were reliable spatial predictors. Varying global landmark stability had no effect when training was conducted using an array of two landmarks located some distance from each other. Context cues, when tested alone, triggered very little searching in appropriate locations, and the absolute magnitude of control over search was insufficient to account for the superiority of stable landmarks. The better learning with a stable landmark, and the dependence of this effect on the geometrical arrangement of landmarks, points to the conditions of spatial learning involving additional principles to those of simple associative conditioning. Experiment 2 examined landmark stability using a single landmark and fixed directional cues in the absence of vestibular disorientation. This also revealed a relative advantage of landmark stability, but animals with a landmark that moved from trial to trial did show some evidence of learning. Context cues when tested alone had minimal influence. Parametric manipulation of landmark stability offers a novel way of influencing spatial learning and thus understanding better the process through which egocentric representations of perceived space are transformed into allocentric representations of the real world. The purpose of this paper is to describe two experiments concerned with identifying the psychological processes of allocentric spatial learning. The results point to the idea that landmark stability is an important factor in spatial learning. Specifically, they reveal that whether or not a landmark will be used for the purpose of representing the location of another object (including hidden objects) is influenced by whether it is perceived as geometrically stable with respect to at least one other landmark and/or certain geometric features of the environment. This phenomenon is relevant to the application of associative learning principles to the spatial domain.     

Immediate spatial distortions of pointing movements induced by visual landmarks

We tested the influence of two horizontally aligned visual landmarks on pointing movements to memorized targets, to investigate whether the visuomotor system can make use of an egocentric representation unaffected by visual context. The endpoints of pointing movements were systematically distorted toward the nearest visual landmark, indicating that spatial representations included both target and nontarget information. These distortions were not due to the presence of the landmarks during the movement but, rather, to their presence in the encoding phase. Qualitatively similar distortions were present even with the shortest possible retention phase, when the target was extinguished at movement onset. Finally, we found the same pattern of distortion when participants were forced to remember the target within an allocentric frame of reference. We argue that even early memory representations for pointing movements are influenced by visual information in the surrounding visual field.     

Blocking in the spatial domain with arrays of discrete landmarks.

A characteristic feature of associative conditioning is that learning a predictive relationship between two events can block later learning about an added event. It is not yet well established whether blocking occurs in the spatial domain or the circumstances in which it does. We now report, using rats trained to search for hidden food near landmarks in an open field arena, that blocking can occur in spatial learning. The animals noticed the added landmark at the start of the blocking phase and explored it, but either failed to incorporate it into their spatial map or developed a representation in which only some landmarks actually control behavior. Additionally, performance at asymptote was controlled by the shape of the landmark array rather than the individual landmarks comprising it, indicating that blocking in the spatial domain may represent a failure to alter the encoded geometry of a learned array.     

Absence of an interaction between navigational strategies based on local and distal landmarks

In 3 experiments, rats were required to escape from a Morris pool by swimming to a submerged platform that was located at the apex of a notional, equilateral triangle with 2 different landmarks occupying the corners at the base. Training for 1 group was always conducted in view of the landmarks surrounding the pool and with the triangular array in a fixed orientation. Subjects could therefore identify the direction of the platform from a single landmark within the pool by reference to cues outside the pool or to the other landmark within the pool. Both strategies were used, and the results from additional groups revealed that the first of these strategies did not affect the acquisition of the second one.     

Clark's nutcrackers (Nucifraga columbiana) and the effects of goal--landmark distance on overshadowing

Three groups of Clark's nutcrackers (Nucifraga columbiana) were trained to find a goal location defined by an array of 4 landmarks that varied in goal--landmark distance. The arrays for each group differed in the distance of the closest landmark and contained goal--landmark distances that were common across groups, allowing for the examination of the effects of both relative and absolute goal--landmark distance on encoding of a landmark array. All 3 groups readily learned the task and were subsequently tested in probe tests with only single landmarks from the array available. Search error in tests with single landmarks was compared both within and across groups. Results demonstrated that both relative and absolute goal-landmark distances are important in spatial search.     

Global and local spatial landmarks: their role during foraging by Columbian ground squirrels (Spermophilus columbianus)

Locating food and refuge is essential for an animal's survival. However, little is known how mammals navigate under natural conditions and cope with given environmental constraints. In a series of six experiments, I investigated landmark-based navigation in free-ranging Columbian ground squirrels (Spermophilus columbianus). Squirrels were trained individually to find a baited platform within an array of nine identical platforms and artificial landmarks set up on their territories. After animals learned the location of the food platform in the array, the position of the latter with respect to local artificial, local natural, and global landmarks was manipulated, and the animal's ability to find the food platform was tested. When only positions of local artificial landmarks were changed, squirrels located food with high accuracy. When the location of the array relative to global landmarks was altered, food-finding accuracy decreased but remained significant. In the absence of known global landmarks, the presence of a familiar route and natural local landmarks resulted in significant but not highly accurate performance. Squirrels likely relied on multiple types of cues when orienting towards a food platform. Local landmarks were used only as a secondary mechanism of navigation, and were not attended to when a familiar route and known global landmarks were present. This study provided insights into landmark use by a wild mammal in a natural situation, and it demonstrated that an array of platforms can be employed to investigate landmark-based navigation under such conditions.     

Landmark-based spatial search in honeybees. I.

Honeybees were trained to find sugar water in the middle of an array of two landmarks of different colours. Unrewarded tests compared searching on the training array with searching on rotated arrays. On rotated tests, a system using the angles between landmarks would continue to search in the middle. A system using vectors to individual elements would search at locations outside the rotated array at which the distances and compass directions to a subset of landmarks matched. Results indicated that bees used both elements and interlandmark angles, but they relied most on one favourite landmark element. Results support the template model of landmark use in honeybees, with the minor parametric modification that weights given to different elements may be unequal. Received:6 July 1998 / Accepted after revision:7 December 1998