Honeybees (Apis mellifera) Remember Two Near-Target Landmark Constellations

A set of experiments tested whether honeybees can remember two different landmark constellations within the same room, or whether the two constellations are mixed together in a single composite memory. Bees were trained to find a reward location with two different landmark arrays. For example, when the blue landmark was north of the yellow one, the reward was to the east; when the yellow landmark was north of the blue one, the reward was to the west. On occasional unrewarded tests, either one of the training arrays was presented (control tests), or else a training array rotated by 90° (rotated tests). A rotated array consisted of part of one training array added to a part of the other training array. Should honeybees form a single composite memory by combining the two training arrays, they should search as much at the target location on rotated tests as on control tests. Results refute this and suggest that they had two separate memories.     

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.     

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.     

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.     

Multiple landmarks, the encoding of environmental geometry and the spatial logics of a dual brain

A series of place learning experiments was carried out in young chicks (Gallus gallus) in order to investigate how the geometry of a landmark array and that of a walled enclosure compete when disoriented animals could rely on both of them to re-orient towards the centre of the enclosure. A square-shaped array (four wooden sticks) was placed in the middle of a square-shaped enclosure, the two structures being concentric. Chicks were trained to ground-scratch to search for food hidden in the centre of the enclosure (and the array). To check for effects of array degradation, one, two, three or all landmarks were removed during test trials. Chicks concentrated their searching activity in the central area of the enclosure, but their accuracy was inversely contingent on the number of landmarks removed; moreover, the landmarks still present within the enclosure appeared to influence the shape of the searching patterns. The reduction in the number of landmarks affected the searching strategy of chicks, suggesting that they had focussed mainly on local cues when landmarks were present within the enclosure. When all the landmarks were removed, chicks searched over a larger area, suggesting an absolute encoding of distances from the local cues and less reliance on the relationships provided by the geometry of the enclosure. Under conditions of monocular vision, chicks tended to rely on different strategies to localize the centre on the basis of the eye (and thus the hemisphere) in use, the left hemisphere attending to details of the environment and the right hemisphere attending to the global shape.This contribution is part of the special issue “Animal Logics” (Watanabe and Huber 2006).     

Stimulus control in the use of landmarks by pigeons in a touch-screen task

Pigeons were tested in a search task on the surface of a monitor on which their responses were registered by a touch-sensitive device. A graphic landmark array was presented consisting of a square outline (the frame) and a colored “landmark.” The unmarked goal, pecks at which produced reward, was located near the center of one edge of the frame, and the landmark was near it. The entire array was displaced without rotation on the monitor from trial to trial. On occasional no-reward tests, the following manipulations were made to the landmark array: (a) either the frame or the landmark was removed; (2) either one edge of the frame or the landmark was shifted; and (3) two landmarks were presented with or without the frame present. On these two-landmark tests, the frame, when present, defined which was the “correct” landmark. When the frame was absent, the “correct” landmark was arbitrarily determined. Results showed that pecks of 2 pigeons were controlled almost solely by the landmark, pecks of 3 were controlled primarily by the landmark but the frame could distinguish the correct landmark, and 1 bird's behavior was controlled primarily by the frame. Stimulus control in this search task is thus selective and differs across individuals. Comparisons to other search tasks and to other stimulus control experiments are made.     

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.     

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.     

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.     

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.     

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.     

Pigeons encode absolute distance but relational direction from landmarks and walls

In recent studies, researchers have examined animals' use of absolute or relational distances in finding a hidden goal. When trained with an array of landmarks, most animals use the default strategy of searching at an absolute distance from 1 or more landmarks. In contrast, when trained in enclosures, animals often use the relationship among walls. In the present study, pigeons were trained to find the center of an array of landmarks or a set of short walls that did not block external cues. Expansion tests showed that both groups of pigeons primarily used an absolute distance strategy. However, on rotational tests, pigeons continued to search in the center of the array, suggesting that direction was learned in relation to array.     

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.     

Internal sense of direction and landmark use in pigeons (Columba livia)

The relative importance of an internal sense of direction based on inertial cues and landmark piloting for small-scale navigation by White King pigeons (Columba livia) was investigated in an arena search task. Two groups of pigeons differed in whether they had access to visual cues outside the arena. In Experiment 1, pigeons were given experience with 2 different entrances and all pigeons transferred accurate searching to novel entrances. Explicit disorientation before entering did not affect accuracy. In Experiments 2-4, landmarks and inertial cues were put in conflict or tested 1 at a time. Pigeons tended to follow the landmarks in a conflict situation but could use an internal sense of direction to search when landmarks were unavailable.     

Multiple-landmark piloting in pigeons (Columba livia): landmark configuration as a discriminative cue

Using the landmark-transformation technique, researchers have shown that pigeons (Columba livia) tend to encode a goal location relative to 1 landmark, even when multiple landmarks are in the vicinity of the goal. The current experiments examined pigeons' ability to use configural information from a set of landmarks by making the arrangement of 4 landmarks a discriminative cue to the location of buried seeds. Results showed that pigeons used information from the 3 consistently placed landmarks to search accurately when 1 landmark was displaced. Findings indicate that pigeons are able to search for a goal using information from multiple landmarks instead of just 1 and that landmark use by these birds may be more flexible than previously theorized.     

Effect of absolute spatial proximity between a landmark and a goal

In two experiments rats were trained in a Morris pool to find a hidden platform in the presence of a single landmark. Circular black curtains surrounded the pool, with the single landmark inside this enclosure, so that no other room cues could provide additional information about the location of the platform. This landmark was hung from a false ceiling and rotated from trial to trial, and the position of the platform also changed on each trial, thus preserving a constant relation between the platform and the landmark. In Experiment 1, the position of the landmark was exactly above the hidden platform in Group Above and was relatively close to the hidden platform in Group Near. At the end of acquisition, test trials without the platform revealed a difference between the groups. Although a preference for searching in the correct quadrant of the pool, where the platform should have been, was found in both groups, this preference was significantly higher for Group Above. In Experiment 2, new rats in Group Near were compared to rats for which the position of the landmark was relatively far from the hidden platform in Group Far. Test trials revealed a preference for searching in the correct quadrant of the pool in both groups, but this preference was significantly higher for Group Near. The implication of these results is that the control acquired by a single landmark is different depending on its relative distance from a hidden platform: Closer landmarks acquire better control than distant ones. These results show a clear parallelism in comparison with the effect of absolute temporal proximity of the CS to the US in classical conditioning.     

Flexible learning and use of multiple-landmark information by pigeons (Columba livia) in a touch screen-based goal searching task

Three touch screen-based experiments were conducted to investigate whether pigeons would learn to use configural information about a goal's location in relation to a multiple-landmark array. In Experiment 1, 4 pigeons (Columba livia) were trained to peck a computer monitor at a location that constituted the third vertex of a hypothetical triangle defined by 2 different landmarks. The landmarks appeared in 3 orientations during the training, and the pigeons' goal-searching ability easily transferred to the landmarks presented in 3 novel orientations. Each landmark was asymmetric, so we next examined whether the pigeons used (a) the small-scale, local orientation information that could be inferred from each landmark individually, or (b) the large-scale, configural information that could be inferred from the spatial arrangement of multiple landmarks taken as a whole. Even when each single landmark appeared by itself, the pigeons were able to locate the goal accurately, suggesting that the large-scale, configural information was not essential. However, when 1 landmark locally pointed to a location that was consistent with the triangular configuration and the other landmark locally pointed to a different location, the pigeons predominantly pecked at the configurally array-consistent location. These results suggest that the pigeons redundantly learned both the large-scale, configural strategy and the local, single-landmark strategy, but they mainly used the latter information, and used the former information solely to disambiguate conflicts when the 2 landmarks pointed toward different targets. Such flexible learning and use of redundant information may reflect the pigeons' adaptation to unstable wild environments during their evolutionary history.     

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.     

Geometric rule learning by Clark's nutcrackers (Nucifraga columbiana)

Clark's nutcrackers (Nucifraga columbiana) were trained to search in a location defined by its geometric relationship to 2 landmarks. Two groups were trained to search at different points along the line connecting the landmarks, and 2 groups were trained to find the 3rd point of a triangle, on the basis of either direction or distance from the landmarks. All groups learned and transferred to new interlandmark distances. However, the constant-distance group learned more slowly, searched less accurately, and showed less transfer than the other 3 groups. When tested with new orientations of the landmarks, the birds tended to follow small but not large rotations. When tested with a single landmark, birds in the half, quarter, and constant-bearing groups searched in the appropriate direction from the landmark, but birds in the distance group did not. These results demonstrate that nutcrackers can learn a variety of geometric principles, that directional information may be weighted more heavily than distance information, and that the birds can use both absolute and relative information about spatial relationships.     

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.