Configural representations in spatial working memory

Two studies investigate the nature of representations in spatial working memory, directly addressing the question of whether people represent the configuration information above and beyond independent positional information when representing multiple sequentially presented locations. In Experiment 1, participants performed a location memory task in which they recalled the location of objects that were presented sequentially on a screen. Comparison of participants' data with simulated data modelled to represent independent positional representation of spatial locations revealed that participants represented the configural properties of shorter sequences (3 and 4 locations) but not of longer ones (5 and 7 locations). Experiment 2 employed a sequential recognition task in which participants were asked to judge whether two consequently presented spatial sequences were identical. These experiments confirmed sensitivity to configural properties of spatial sequences.     

Testing visual short-term memory of pigeons (Columba livia) and a rhesus monkey (Macaca mulatta) with a location change detection task

Change detection is commonly used to assess capacity (number of objects) of human visual short-term memory (VSTM). Comparisons with the performance of non-human animals completing similar tasks have shown similarities and differences in object-based VSTM, which is only one aspect (“what”) of memory. Another important aspect of memory, which has received less attention, is spatial short-term memory for “where” an object is in space. In this article, we show for the first time that a monkey and pigeons can be accurately trained to identify location changes, much as humans do, in change detection tasks similar to those used to test object capacity of VSTM. The subject’s task was to identify (touch/peck) an item that changed location across a brief delay. Both the monkey and pigeons showed transfer to delays longer than the training delay, to greater and smaller distance changes than in training, and to novel colors. These results are the first to demonstrate location-change detection in any non-human species and encourage comparative investigations into the nature of spatial and visual short-term memory.     

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.     

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.     

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.     

Does terrain slope really dominate goal searching?

If you can locate a target by using one reliable source of information, why would you use an unreliable one? A similar question has been faced in a recent study on homing pigeons, in which, despite the presence of better predictors of the goal location, the slope of the floor in an arena dominated the searching process. This piece of evidence seems to contradict straightforward accounts of associative learning, according to which behavior should be controlled by the stimulus that best predicts the reward, and has fueled interest toward one question that, to date, has received scarce attention in the field of spatial cognition: how are vertical spaces represented? The purpose of this communication is to briefly review the studies on this issue, trying to determine whether slope is a special cue--driving behavior irrespective of other cues--or simply a very salient one.     

Individual strategies and release site features determine the extent of deviation in clock-shifted pigeons at familiar sites

When homing from familiar areas, homing pigeons are able to exploit previously acquired topographical information, but the mechanisms behind this ability are still poorly understood. One possibility is that they recall the familiar release site topographical features in association with the home direction (site-specific compass orientation strategy), another that the spatial relationships among landmarks guide their route home (piloting strategy), without relying on the compass mechanism. The two strategies can be put in conflict by releasing clock-shifted birds at familiar locations, in order to highlight which is preferred. We analysed GPS tracks of clock-shifted pigeons, with familiarity controlled at each of three different release sites, and we observed that pigeons can display individual preferences for one of the two orientation strategies and that some characteristic features of the release site have an important role in determining the level of landmark-based homeward orientation.     

No sex differences in spatial location memory for abstract designs

Previous research has demonstrated a female advantage, albeit imperfectly, on tests of object location memory where object identity information is readily available. However, spatial and visual elements are often confounded in the experimental tasks used. Here spatial and visual memory performance was compared in 30 men and 30 women by presenting 12 abstract designs in a spatial array for recall and recognition (visual memory) and spatial location ("object" location memory). Object location memory was measured via a sensitive absolute displacement score defined as the distance in mms between the position assigned to the object during recall and the actual position it originally occupied. There were no sex differences in either the visual or spatial location tests. Controlling for age and estimated IQ scores made no impact on the results. These data suggest an absence of a sex difference in purely visual and spatial aspects of object location memory.     

Do binding deficits account for age-related decline in visual working memory?

Remembering visual material, such as objects, faces, and spatial locations, over a short period of time (seconds) becomes more difficult as we age. We investigated whether these deficits could be explained by a simple reduction in visual working memory capacity or by an impairment in one's ability to form or maintain appropriate associations among pieces of related information. In three experiments, we used recognition and recall tests to address the efficacy with which older adults can create bound object representations by varying the number of features of each object that had to be remembered for a subsequent memory test. Results demonstrated that whereas older adults exhibited reduced memory capacity as compared with that of younger adults, both groups stored integrated object representations in visual working memory. These results are contrasted with other work that suggests that age-related memory decline is due, at least in part, to associative deficits.     

No evidence for binding of items to task-irrelevant backgrounds in visual working memory

When representing visual features such as color and shape in visual working memory (VWM), participants also represent the locations of those features as a spatial configuration of the locations of those features in the display. In everyday life, we encounter objects against some background, yet it is unclear whether the configural representation in memory obligatorily constitutes the entire display, including that (often task-irrelevant) background information. In three experiments, participants completed a change detection task on color and shape; the memoranda were presented in front of uniform gray backgrounds, a textured background (Exp. 1), or a background containing location placeholders (Exps. 2 and 3). When whole-display probes were presented, changes to the objects’ locations or feature bindings impacted memory performance—implying that the spatial configuration of the probes influenced participants’ change decisions. Furthermore, when only a single item was probed, the effect of changing its location or feature bindings was either diminished or completely extinguished, implying that single probes do not necessarily elicit the entire spatial configuration. Critically, when task-irrelevant backgrounds were also presented that may have provided a spatial configuration for the single probes, the effect of location or bindings was not moderated. These findings suggest that although the spatial configuration of a display guides VWM-based recognition, this information does not necessarily always influence the decision process during change detection.     

Finding the best angle: pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>) weight angular information more heavily than relative wall length in an open-field geometry task

Pigeons were trained to locate food in two geometrically equivalent corners of a parallelogram-shaped enclosure. Both the angular amplitude of the corners and the length of the walls alone were sufficient for successfully completing the task. Following training, birds were tested in three separate conditions that manipulated the geometric information available. During tests in both a rectangular-shaped enclosure that preserved the wall length information but not the angular amplitude, and a rhombus-shaped enclosure that did the opposite, pigeons located their goal corners with a high degree of accuracy, indicating an ability to use both types of geometric information in isolation. This result is consistent with prior research with domestic chicks. However, in a conflict test in a reverse parallelogram-shaped enclosure, in which the correct angular location was paired with an incorrect wall length location, birds showed a preference for the correct angular location. This suggests that pigeons weight angles more heavily than wall lengths in this type of navigation task, which differs from findings in a similar task conducted with the domestic chick. Results in the conflict test also suggest that pigeons did not use the principal axis as their main strategy of small-scale navigation.     

Time-space learning in homing pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>): orientation to an artificial light source

Time-space learning reflects an ability to represent in memory event-stimulus properties together with the place and time of the event; a capacity well developed in birds. Homing pigeons were trained in an indoor octagonal arena to locate one food goal in the morning and a different food goal in the late afternoon. The goals differed with respect to their angular/directional relationship to an artificial light source located outside the arena. Further, the angular difference in reward position approximated the displacement of the sun's azimuth that would occur during the same time period. The experimental birds quickly learned the task, demonstrating the apparent ease with which birds can adopt an artificial light source to discriminate among alternative spatial responses at different times of the day. However, a novel midday probe session following successful learning revealed that the light source was interpreted as a stable landmark and not as a surrogate sun that would support compass orientation. Probe sessions following a phase shift of the light–dark cycle revealed that the mechanism employed to make the temporal discrimination was prevailingly based on an endogenous circadian rhythm and not an interval timing mechanism.     

Recognition memory and the recall of spatial location

It is argued that memory for events should depend on the ability to recall their attributes. Two experiments suggest that this is so for the attribute of spatial location. When words presented in spatial arrays were later re-presented in a joint test of recognition memory and location recall, it was found that the surer one’s recognition, the greater the accuracy of location recall. Instructions to attend to a word’s location resulted in poorer recognition memory and marginally poorer location recall than were obtained when location was “incidentally” learned. Finally, when words were presented in color as well as in space, recognition memory was best when both attributes were recalled, was better when only location, was recalled than when only color was recalled, and was poorest when neither attribute was recalled.     

Integration of spatial maps in pigeons

The integration of spatial maps in pigeons was investigated using a spatial analog to sensory preconditioning. The pigeons were tested in an open-field arena in which they had to locate hidden food among a 4×4 grid of gravel-filled cups. In phase 1, the pigeons were exposed to a consistent spatial relationship (vector) between landmark L (a red L-shaped block of wood), landmark T (a blue T-shaped block of wood) and the hidden food goal. In phase 2, the pigeons were then exposed to landmark T with a different spatial vector to the hidden food goal. Following phase 2, pigeons were tested with trials on which they were presented with only landmark L to examine the potential integration of the phase 1 and 2 vectors via their shared common elements. When these test trials were preceded by phase 1 and phase 2 reminder trials, pigeons searched for the goal most often at a location consistent with their integration of the LT phase 1 and Tphase 2 goal vectors. This result indicates that integration of spatial vectors acquired during phases 1 and 2 allowed the pigeons to compute a novel Lgoal vector. This suggests that spatial maps may be enlarged by successively integrating additional spatial information through the linkage of common elements.     

Septal area lesions impair spatial working memory in homing pigeons (Columba livia)

The septo-hippocampal system in birds resembles that of mammals, motivating research into the function of the avian hippocampus while surprisingly little attention has been given to the septum. To investigate a possible role of the avian septum in memory, the effects of septal area lesions on a spatial working memory (SpWM) task was tested in homing pigeons. After preoperative training on an analogue eight-arm (feeders) radial maze, now sham-operated control and septal lesioned pigeons were then trained again on the same task of locating the four feeders on the test phase of a trial that were not baited during the sample phase of a trial. During the test phase of a working memory trial, septal lesioned pigeons, compared to both their own preoperative performance and the performance of the controls, required significantly more choices to locate the four baited feeders not baited during the sample phase of a trial, and they made significantly fewer correct responses to the now baited feeders on their first four choices. The results demonstrate that, like its mammalian counterpart, the avian septum plays an important role in SpWM, suggesting that at least some functional properties of the septum are evolutionarily conserved in birds and mammals.     

Feature memory and binding in young and older adults

Intact memory for complex events requires not only memory for particular features (e.g., item, location, color, size), but also intact cognitive processes for binding the features together. Binding provides the memorial experience that certain features belong together. The experiments presented here were designed to explicate these as potentially separable sources of age-associated changes in complex memory—namely, to investigate the possibility that age-related changes in memory for complex events arise from deficits in (1) memory for the kinds of information that comprise complex memories, (2) the processes necessary for binding this information into complex memories, or (3) both of these components. Young and older adults were presented with colored items located within an array. Relative to young adults, older adults had a specific and disproportionate deficit in recognition memory for location, but not for item or for color. Also, older adults consistently demonstrated poorer recognition memory for bound information, especially when all features were acquired intentionally. These feature and binding deficits separately contribute to what have been described as older adults’ context and source memory impairments.     

Out of mind, but not out of sight: Intentional control of visual memory

Does visual information enjoy automatic, obligatory entry into memory, or, after such information has been seen, can it still be actively excluded? To characterize the process by which visual information could be excluded from memory, we used Sternberg’s (1966, 1975) recognition paradigm, measuring visual episodic memory for compound grating stimuli. Because recognition declines as additional study items enter memory, episodic recognition performance provides a sensitive index of memory’s contents. Three experiments showed that an item occupying a fixed serial position in a series of study items could be intentionally excluded from memory. In addition, exclusion does not depend on lowlevel information, such as the stimulus’s spatial location, orientation, or spatial frequency, and does not depend on the precise timing of irrelevant information, which suggests that the exclusion process is triggered by some event during a trial. The results, interpreted within the framework of a summed similarity model for visual recognition, suggest that exclusion operates after considerable visual processing of the to-be-excluded item.     

Learning of magnetic compass directions in pigeons

A proof of magnetic compass learning by pigeons under laboratory conditions has been attempted for decades, but all experiments have failed so far. The aim of the present study was to test whether pigeons can learn magnetic compass directions in an operant chamber if magnetic cues are presented as true spatial cues. Experimental sessions were carried out in the local geomagnetic field and in magnetic fields with matched total intensity and inclination, but different directions generated with Helmholtz-coils. Birds demonstrated successful learning with a performance level comparable to that in learning studies with magnetic anomalies. In addition, we compared the data from magnetic learning in the laboratory with performance from homing experiments in the field. The birds that were more successful in the learning experiment had vanishing bearings farther away from the home direction than the group mean at unfamiliar, but not at familiar sites. This might suggest that better learners explore unknown locations in a different way. Our findings represent the first evidence for operant magnetic compass learning in pigeons and also provide a link between behavioural data from the field and the laboratory.     

Multiple cue use and integration in pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>)

Encoding multiple cues can improve the accuracy and reliability of navigation and goal localization. Problems may arise, however, if one cue is displaced and provides information which conflicts with other cues. Here we investigated how pigeons cope with cue conflict by training them to locate a goal relative to two landmarks and then varying the amount of conflict between the landmarks. When the amount of conflict was small, pigeons tended to integrate both cues in their search patterns. When the amount of conflict was large, however, pigeons used information from both cues independently. This context-dependent strategy for resolving spatial cue conflict agrees with Bayes optimal calculations for using information from multiple sources.     

Working memory for visual features and conjunctions in schizophrenia

The visual working memory (WM) storage capacity of patients with schizophrenia was investigated using a change detection paradigm. Participants were presented with 2, 3, 4, or 6 colored bars with testing of both single feature (color, orientation) and feature conjunction conditions. Patients performed significantly worse than controls at all set sizes but demonstrated normal feature binding. Unlike controls, patient WM capacity declined at set size 6 relative to set size 4. Impairments with subcapacity arrays suggest a deficit in task set maintenance: Greater impairment for supercapacity set sizes suggests a deficit in the ability to selectively encode information for WM storage. Thus, the WM impairment in schizophrenia appears to be a consequence of attentional deficits rather than a reduction in storage capacity.