No evidence for overshadowing or facilitation of spatial pattern learning by visual cues

Two experiments were conducted to examine the effects of redundant and relevant visual cues on spatial pattern learning. Rats searched for hidden food items on the tops of poles that formed a square (Experiment 1) or a checkerboard (Experiment 2) pattern. The experimental groups were trained with visual cues that specified the locations of the baited poles. All groups were tested without visual cues so that any overshadowing or facilitation of spatial pattern learning by visual cues could be detected. Spatial choices were controlled by the spatial pattern and by the visual cues in both experiments. However, there was no evidence of overshadowing or facilitation of spatial pattern learning by visual cues in either experiment. The results are consistent with the idea that the representation of the spatial pattern that guides choices is not controlled by the same learning processes as those that produce associations between visual cues and food locations.     

Facilitation of learning spatial relations among locations by visual cues: generality across spatial configurations

Spatial pattern learning permits the learning of the location of objects in space relative to each other without reference to discrete visual landmarks or environmental geometry. In the present experiment, we investigated conditions that facilitate spatial pattern learning. Specifically, human participants searched in a real environment or interactive 3-D computer-generated virtual environment open-field search task for four hidden goal locations arranged in a diamond configuration located in a 5 × 5 matrix of raised bins. Participants were randomly assigned to one of three groups: Pattern Only, Landmark + Pattern, or Cues + Pattern. All participants experienced a Training phase followed by a Testing phase. Visual cues were coincident with the goal locations during Training only in the Cues + Pattern group whereas a single visual cue at a non-goal location maintained a consistent spatial relationship with the goal locations during Training only in the Landmark + Pattern group. All groups were then tested in the absence of visual cues. Results in both environments indicated that participants in all three groups learned the spatial configuration of goal locations. The presence of the visual cues during Training facilitated acquisition of the task for the Landmark + Pattern and Cues + Pattern groups compared to the Pattern Only group. During Testing the Landmark + Pattern and Cues + Pattern groups did not differ when their respective visual cues were removed. Furthermore, during Testing the performance of these two groups was superior to the Pattern Only group. Results generalize prior research to a different configuration of spatial locations, isolate spatial pattern learning as the process facilitated by visual cues, and indicate that the facilitation of learning spatial relations among locations by visual cues does not require coincident visual cues.     

Discrimination of small numerosities in young chicks

Chicks were trained to discriminate small sets of identical elements. They were then tested for choices (unrewarded) between sets of similar numerosities, when continuous physical variables such as spatial distribution, contour length, and overall surface were equalized. In all conditions chicks discriminated one versus two and two versus three stimulus sets. Similar results were obtained when elements were presented under conditions of partial occlusion. In contrast, with sets of four versus five, four versus six, and three versus four elements chicks seemed unable to discriminate on the basis of number, although nonnumerical discrimination based on perceptual cues was observed. This adds to increasing evidence for discrimination of small numerosities of up to three elements in human infants and nonhuman animals.     

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.     

The importance of "what": Infants use featural information to index events

Dynamic spatial indexing is the ability to encode, remember, and track the location of complex events. For example, in a previous study, 6-month-old infants were familiarized to a toy making a particular sound in a particular location, and later they fixated that empty location when they heard the sound presented alone (Journal of Experimental Psychology: General, 2004, Vol. 133, pp. 46-62). The basis and developmental trajectory of this ability are currently unclear. We investigated dynamic spatial indexing across the first year after birth and tested the hypothesis that the structure of visual cues supports infants' learning of sound and location associations. In our study, 3-, 6-, and 10-month-olds were tested in a dynamic spatial indexing eye movement paradigm that paired two sounds with two locations. In one condition, these were reliably paired with two sets of visual features (two toys condition), replicating the original studies. We also presented a single set of visual cues in both locations (one toy condition) and multiple sets of visual features in both locations (six toys condition). Infants from 3months of age onward showed evidence of dynamic spatial indexing in the two toys condition, but only the 10-month-olds succeeded in the one toy and six toys conditions. We argue that this may reflect a broader developmental trajectory, whereby infants first make use of multiple cue integration but with age are able to learn from a narrow set of cues.     

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.     

Visual landmarks facilitate rodent spatial navigation in virtual reality environments

Because many different sensory modalities contribute to spatial learning in rodents, it has been difficult to determine whether spatial navigation can be guided solely by visual cues. Rodents moving within physical environments with visual cues engage a variety of nonvisual sensory systems that cannot be easily inhibited without lesioning brain areas. Virtual reality offers a unique approach to ask whether visual landmark cues alone are sufficient to improve performance in a spatial task. We found that mice could learn to navigate between two water reward locations along a virtual bidirectional linear track using a spherical treadmill. Mice exposed to a virtual environment with vivid visual cues rendered on a single monitor increased their performance over a 3-d training regimen. Training significantly increased the percentage of time avatars controlled by the mice spent near reward locations in probe trials without water rewards. Neither improvement during training or spatial learning for reward locations occurred with mice operating a virtual environment without vivid landmarks or with mice deprived of all visual feedback. Mice operating the vivid environment developed stereotyped avatar turning behaviors when alternating between reward zones that were positively correlated with their performance on the probe trial. These results suggest that mice are able to learn to navigate to specific locations using only visual cues presented within a virtual environment rendered on a single computer monitor.     

Sensory information and associative cues used in food detection by wild vervet monkeys

Understanding animals’ spatial perception is a critical step toward discerning their cognitive processes. The spatial sense is multimodal and based on both the external world and mental representations of that world. Navigation in each species depends upon its evolutionary history, physiology, and ecological niche. We carried out foraging experiments on wild vervet monkeys (Chlorocebus pygerythrus) at Lake Nabugabo, Uganda, to determine the types of cues used to detect food and whether associative cues could be used to find hidden food. Our first and second set of experiments differentiated between vervets’ use of global spatial cues (including the arrangement of feeding platforms within the surrounding vegetation) and/or local layout cues (the position of platforms relative to one another), relative to the use of goal-object cues on each platform. Our third experiment provided an associative cue to the presence of food with global spatial, local layout, and goal-object cues disguised. Vervets located food above chance levels when goal-object cues and associative cues were present, and visual signals were the predominant goal-object cues that they attended to. With similar sample sizes and methods as previous studies on New World monkeys, vervets were not able to locate food using only global spatial cues and local layout cues, unlike all five species of platyrrhines thus far tested. Relative to these platyrrhines, the spatial location of food may need to stay the same for a longer time period before vervets encode this information, and goal-object cues may be more salient for them in small-scale space.     

The impact of landmark properties in shaping exploration and navigation

This study was aimed at uncovering physical and geometric properties that make a particular landmark a target of exploration and navigation. Rats were tested in a square open-field arena with additional portable corners featuring the same properties as the arena corners. It was found that the routes of progression converged upon the added corners, whether located at the arena wall or the arena center. Route convergence upon the added corners involved numerous visits to these corners. However, time spent at the added corners was relatively short compared with the arena corners, including that from which rats were introduced into the arena. There was no differential effect of testing rats in light or dark, or with a low versus a high portable corner. It is suggested that the added corners were distinct against the background of the arena enclosure, whereas the four arena corners and walls were encoded by the rats as one geometric module. This distinctness, together with the greater accessibility of the added corners, made them salient landmarks and a target of exploration. Thus, the impact of a landmark extended beyond its specific self-geometry to include accessibility and distinctness, which are contextual properties. In addition to the contextual impact on locomotor behavior there was also a temporal effect, with security initially dominating the rats’ behavior but then declining along with an increased attraction to salient landmarks. These spatiotemporal patterns characterized behavior in both lit and dark arenas, indicating that distal cues were secondary to local proximal cues in shaping routes.     

From maps to navigation: The role of cues in finding locations in a virtual environment

In two experiments, participants navigated through a large arena within a virtual environment (VE) to a location encoded in memory from a map. In both experiments, participants recalled locations by navigating through the VE, but in Experiment 2, they additionally recalled the locations on the original map. Two cues were located outside and above the walls of the arena at either north-south locations or east-west locations. The pattern of angular bias was used to infer how the cues affected the creation of spatial categories influencing memory for location in the two tasks. When participants navigated to remembered locations in the VE, two cue-based spatial categories were inferred, with cues serving to demarcate the boundaries of the categories. When participants remembered locations on the original map, two cue-based categories were again formed, but with cues serving as category prototypes. The pattern of results implies that cue-based spatial categorization schemes may be formulated differently at the memory retrieval stage depending on task constraints.     

Disconnect in concept learning by rhesus monkeys (Macaca mulatta): judgment of relations and relations-between-relations

The authors investigated the role that entropy measures, discriminative cues, and symbolic knowledge play for rhesus monkeys (Macaca mulatta) in the acquisition of the concepts of same and different for use in a computerized relational matching-to-sample task. After repeatedly failing to perceive relations between pairs of stimuli in a 2-choice discrimination paradigm, monkeys rapidly learned to discriminate between 8-element arrays. Subsequent tests with smaller arrays, however, suggested that, although important for the initial acquisition of the concept, entropy is not a variable on which monkeys are dependent. Not only do monkeys choose a corresponding relational pair in the presence of a cue, but they also choose the cue itself in the presence of the relational pair--in essence, labeling those relations. Subsequent failure in the judgment of relations-between-relations, however, suggests that perhaps a qualitatively different cognitive component exists that prevents monkeys from behaving analogically.     

Rhesus monkeys (Macaca mulatta) enumerate large and small sequentially presented sets of items using analog numerical representations

Two rhesus monkeys selected the larger of two sequentially presented sets of items on a computer monitor. In Experiment 1, performance was related to the ratio of set sizes, and the monkeys discriminated between sets with up to 10 items. Performance was not disrupted when 1 set had fewer than 4 items and 1 set had more than 4 items, a critical trial type for differentiating object file and analog models of numerical representation. Experiment 2 controlled the interitem rate of presentation. Experiment 3 included some trials on which number and amount (visual surface area) offered conflicting cues. Experiment 4 varied the total duration of set presentation and the duration of item visibility. In all of the experiments, performance remained high, although total set presentation duration also acted as a partial cue for the monkeys. Overall, the data indicated that rhesus monkeys estimate the approximate number of items in sequentially presented sets and that they are not relying solely on nonnumerical cues such as rate, duration, or cumulative amount.     

Object-goal positioning influences spatial representation in rats

Three tests investigated how the geometric relation between object/landmarks and goals influenced spatial choice behavior in rats. Two groups searched for hidden food in an object-filled circular arena containing 24 small poles. For the “Proximal” group, four distinct objects in a square configuration were placed close to four baited poles. For the “Distal” group, the identical configuration of objects was rotated 45° relative to the poles containing the hidden food. The Proximal group learned to locate the baited poles more quickly than the Distal group. Tests with removed and rearranged landmarks indicated that the two groups learned to use the objects differently. The results suggested that close proximity of objects to goals encouraged their use as beacons, while greater distance of objects from goals resulted in the global encoding of the geometric properties of the arena and the use of the objects as landmarks. Received: 22 June 1998 / Accepted after revision: 23 January 1999     

Facilitation of learning spatial relations among locations by visual cues: Implications for theoretical accounts of spatial learning

Human participants searched in a real environment or interactive 3-D virtual environment open field for four hidden goal locations arranged in a 2 × 2 square configuration in a 5 × 5 matrix of raised bins. The participants were randomly assigned to one of two groups: cues 1 pattern or pattern only. The participants experienced a training phase, followed by a testing phase. Visual cues specified the goal locations during training only for the cues 1 pattern group. Both groups were then tested in the absence of visual cues. The results in both environments indicated that the participants learned the spatial relations among goal locations. However, visual cues during training facilitated learning of the spatial relations among goal locations: In both environments, the participants trained with the visual cues made fewer errors during testing than did those trained only with the pattern. The results suggest that learning based on the spatial relations among locations may not be susceptible to cue competition effects and have implications for standard associative and dual-system accounts of spatial learning.     

Pattern vision of the honeybee (Apis mellifera): blue and green receptors in the discrimination of translocation

The visual discrimination of horizontal gratings by the honeybee (Apis mellifera) was studied in a Y-choice apparatus with fixed patterns presented vertically at a set range. Translocation in this context is the exchange of the positions of two different colored or black areas. This paper investigates what cues the bees have learned in this task. The patterns, made from combinations of calibrated colored papers, are designed to explore the parts played by the blue and green receptors when the boundary between the two colors provides contrast to only one receptor type. Horizontal translocation is not discriminated without contrast to the green receptors, but up/down translocation can be discriminated whatever the contrast at the boundary. The trained bees were tested on the same patterns made with different papers that included extreme changes in contrast. The results show that discrimination of up/down translocation involves green receptors and also blue receptors. When bees discriminate a translocation that shows contrast to only one type of receptor, they do not use the apparent brightness or the direction of the contrast to that receptor type acting alone. Instead, they discriminate the locations of colored areas irrespective of intensity differences or directions of contrasts. They use some measure of the photon flux at both receptor types and remember the difference between the colors and their locations.     

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.     

Not using the obvious: desert ants, Melophorus bagoti, learn local vectors but not beacons in an arena

Many ant species travel large distances to find food, sometimes covering distances that are up to one million times their body length. Even when these foraging trips follow convoluted paths, the ants usually find their way back to their nest with precision (Wehner et al. in J Exp Biol 199:129–140, 1996). Ants have been shown to use both compass cues in the sky (pattern of polarised light) and landmarks on Earth to return to their nest. We present two experiments conducted on a solitary foraging ant: Melophorus bagoti in their natural habitat in the central Australian desert. Ants were trained and tested in situ. We tested foragers’ ability to exit a circular arena which provided an undifferentiated panorama. Artificial visual landmarks were located near a small exit. On tests in which path integration information was not available, foragers did not use artificial landmarks as beacons. Instead, they oriented in the learned exit direction, whether or not it pointed to the nest. We suggest that M. bagoti foragers learned a context-specific local vector when cued by the context of the circular arena. Our findings present the first evidence that M. bagoti foragers learn context-specific compass directions to chart their initial path home.     

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.     

Exposure to Spatial Cues Facilitates Visual Discrimination but Not Spatial Guidance

The performance in a radial-arm maze of two groups of rats with restricted access to extra-maze visual cues was studied. One group received extensive exposure to the visual environment of the maze, whereas the second group was never exposed to the environment, aside from their experience in the maze itself. Spatial exposure resulted in a slightly improved ability to discriminate between previously visited and unvisited spatial locations, which can be explained on the basis of general perceptual learning processes. However, there was no evidence that spatial exposure resulted in control of choices by the spatial relations among the maze locations. These results are discussed in terms of theories of spatial learning which appeal to perceptual learning and cognitive mapping.     

A consistent but non-coincident visual pattern facilitates the learning of spatial relations among locations

Human participants searched in a dynamic three-dimensional computer-generated virtual-environment open-field search task for four hidden goal locations arranged in a diamond configuration located in a 5 × 5 matrix of raised bins. Participants were randomly assigned to one of two groups: visual pattern or visual random. All participants experienced 30 trials in which four goal locations maintained the same spatial relations to each other (i.e., a diamond pattern), but this diamond pattern moved to random locations within the 5 × 5 matrix from trial to trial. For participants in the visual pattern group, four locations were marked in a distinct color and arranged in a diamond pattern that moved to a random location independent of the hidden spatial pattern from trial to trial throughout the experimental session. For participants in the visual random group, four random locations were marked with a distinct color and moved to random locations independent from the hidden spatial pattern from trial to trial throughout the experimental session. As a result, the visual cues for the visual pattern group were consistent but not coincident with the hidden spatial pattern, whereas the visual cues for the visual random group were neither consistent nor coincident with the hidden spatial pattern. Results indicated that participants in both groups learned the spatial configuration of goal locations and that the presence of consistent but noncoincident visual cues facilitated the learning of spatial relations among locations.