Reorienting in Virtual 3D Environments: Do Adult Humans Use Principal Axes,Medial Axes or Local Geometry?

Studies have shown that animals, including humans, use the geometric properties of environments to orient. It has been proposed that orientation is accomplished primarily by encoding the principal axes (i.e., global geometry) of an environment. However, recent research has shown that animals use local information such as wall length and corner angles as well as local shape parameters (i.e., medial axes) to orient. The goal of the current study was to determine whether adult humans reorient according to global geometry based on principal axes or whether reliance is on local geometry such as wall length and sense information or medial axes. Using a virtual environment task, participants were trained to select a response box located at one of two geometrically identical corners within a featureless rectangular-shaped environment. Participants were subsequently tested in a transformed L-shaped environment that allowed for a dissociation of strategies based on principal axes, medial axes and local geometry. Results showed that participants relied primarily on a medial axes strategy to reorient in the L-shaped test environment. Importantly, the search behaviour of participants could not be explained by a principal axes-based strategy.     

Not all locations are created equal: exploring how adults hide and search for objects

Little is known about the strategies people use to effectively hide objects from others, or to search for objects others have hidden. The present research extends a recent investigation of people's hiding and searching strategies in a simple room with 9 cache location. In the present studies, people hid and searched for three objects under more than 70 floor tiles in complex real and virtual rooms. Experiment 1 replicated several finding of Talbot et al within the more complex real and virtual environments. Specifically, people traveled further from origin and selected more dispersed locations when hiding than when searching. Experiments 2 and 3 showed that: 1) people were attracted to an area of darkness when searching and avoided locations close to a window when hiding, 2) when search attempts were limited to three choices, people searched farther from origin and dispersed their locations more when hiding than when searching, and 3) informing people that they would need to recover their hidden objects altered their hiding behavior and increased recovery accuracy. Across all experiments, consistencies in location preferences emerged, with more preference for the middle of the room during hiding and more preference for corners of the room during searching. Even though the same people participated in both the hiding and searching tasks, it appears that people use different strategies to select hiding places than to search for objects hidden by others.     

Control of rodent and human spatial navigation by room and apparatus cues

A growing body of literature indicates that rats prefer to navigate in the direction of a goal in the environment (directional responding) rather than to the precise location of the goal (place navigation). This paper provides a brief review of this literature with an emphasis on recent findings in the Morris water task. Four experiments designed to extend this work to humans in a computerized, virtual Morris water task are also described. Special emphasis is devoted to how directional responding and place navigation are influenced by room and apparatus cues, and how these cues control distinct components of navigation to a goal. Experiments 1 and 2 demonstrate that humans, like rats, perform directional responses when cues from the apparatus are present, while Experiment 3 demonstrates that place navigation predominates when apparatus cues are eliminated. In Experiment 4, an eyetracking system measured gaze location in the virtual environment dynamically as participants navigated from a start point to the goal. Participants primarily looked at room cues during the early segment of each trial, but primarily focused on the apparatus as the trial progressed, suggesting distinct, sequential stimulus functions. Implications for computational modeling of navigation in the Morris water task and related tasks are discussed.     

Visual cue learning and odometry in guiding the search behavior of desert ants, Melophorus bagoti, in artificial channels

Terrestrial panoramic cues, path integration and search behavior are the main navigational strategies used by ants to locate food and find their way back to the nest. Searching becomes important when the other navigational cues are either not available or cannot provide sufficient information to pinpoint the goal. When searching in one-dimensional channels Melophorus bagoti ants exhibit a systematic drift in the starting-point-to-goal direction as they turn back and forth, sometimes past the goal location ( Narendra et al., 2008). Here we show that this drift in channels is not a stereotypical part of the search behavior in these ants. It rather depends on the conditions of training. In experiments in which the nest entrance is located not at the end but at the side of the channel, forward drift is not always part of the nest search. Experiments on food searches showed that with the food source at the end of the channel, ants performed a linear drift in the starting-point-to-food direction. With food at the side of the channel, they showed a less pronounced drift toward the food source. In this constrained environment, especially with the goal at the end of the channel, ants seem to learn a routine such as ‘run along the channel’, and mix this routine with their usual strategy of turning back and forth in search.     

Rats’ (Rattus norvegicus) temporal discrimination of brief auditory stimuli: How does it compare with that of humans (Homo sapiens) and zebra finches (Taeniopygia guttata)?

The present study with rats replicated an experiment on the ability of zebra finches and humans to discriminate among brief auditory stimuli (see Weisman et al., 1999, Experiment 2). We trained rats with 27 3-kHz tones that varied in duration from 10 ms to 1420 ms. Reinforcement was contingent on responding (approaching the food well) to the nine medium-durations range tones (56-255 ms) but not to the nine short-durations range (10-46 ms) or long-durations range tones (309-1420 ms). Rats also received post-discrimination transfer tests with 2 kHz and 4 kHz tones that varied over the same durations as the 3 kHz tones. Rats acquired the temporal discrimination to a slightly lower level of accuracy than seen in finches or humans by Weisman et al. (1999). We tested for transfer of the temporal discrimination to find that rats, similar to humans (data from Weisman et al., 1999), transferred to untrained 2-kHz and 4-kHz tones at levels approaching accuracy to that achieved to the trained 3-kHz tone. By contrast, zebra finches (data from Weisman et al., 1999) failed to transfer their discrimination to the trained tone. We conclude that (a) rats discriminate among tone durations at least as well as they do among auditory frequencies and (b) rats like humans, but unlike finches, are insensitive to absolute pitch in their temporal discrimination.     

Adaptive Patch Searching Strategies in Fragmented Landscapes

The search strategies dispersers employ to search for new habitat patches affect individuals’ search success and subsequently landscape connectivity and metapopulation viability. Some evidence indicates that individuals within the same species may display a variety of behavioural patch searching strategies rather than one species-specific strategy. This may result from landscape heterogeneity. We modelled the evolution of individual patch searching strategies in different landscapes. Specifically, we analysed whether evolution can favour different, co-existing, behavioural search strategies within one population and to what extent this coexistence of multiple strategies was dependent on landscape configuration. Using an individual-based simulation model, we studied the evolution of patch searching strategies in three different landscape configurations: uniform, random and clumped. We found that landscape configuration strongly influenced the evolved search strategy. In uniform landscapes, one fixed search strategy evolved for the entire spatially structured population, while in random and clumped landscapes, a set of different search strategies emerged. The coexistence of several search strategies also strongly depended on the dispersal mortality. We show that our result can affect landscape connectivity and metapopulation dynamics. Co-ordinating editor: N. Yamamura     

Intramaze cue utilization in the water maze: effects of sex and estrous cycle in rats

Rats can use a wide spectrum of intra- and extramaze information while navigating through the environment. The current study examined the relative contribution of an intramaze cue with regard to its proximity to the goal. Three experiments were conducted and the impact of intramaze cue removal or rotation on water maze search was examined. In males, the effect of the intramaze cue declined monotonically in relation to the proximity of the cue to the goal. A more complex relationship between cue location and utilization was found in estrous and proestrus females. Estrous females showed a strong effect of the cue only when it was near the goal, ignoring it when it was situated further away. Conversely proestrus females were affected by the cue under all conditions. It is concluded that previous reports of behavioral differences may stem from the fact that proestrus females are affected by and attend to a wider range of stimuli, while estrous females are more affected by salient stimuli.     

Identifying Stride-To-Stride Control Strategies in Human Treadmill Walking

Variability is ubiquitous in human movement, arising from internal and external noise, inherent biological redundancy, and from the neurophysiological control actions that help regulate movement fluctuations. Increased walking variability can lead to increased energetic cost and/or increased fall risk. Conversely, biological noise may be beneficial, even necessary, to enhance motor performance. Indeed, encouraging more variability actually facilitates greater improvements in some forms of locomotor rehabilitation. Thus, it is critical to identify the fundamental principles humans use to regulate stride-to-stride fluctuations in walking. This study sought to determine how humans regulate stride-to-stride fluctuations in stepping movements during treadmill walking. We developed computational models based on pre-defined goal functions to compare if subjects, from each stride to the next, tried to maintain the same speed as the treadmill, or instead stay in the same position on the treadmill. Both strategies predicted average behaviors empirically indistinguishable from each other and from that of humans. These strategies, however, predicted very different stride-to-stride fluctuation dynamics. Comparisons to experimental data showed that human stepping movements were generally well-predicted by the speed-control model, but not by the position-control model. Human subjects also exhibited no indications they corrected deviations in absolute position only intermittently: i.e., closer to the boundaries of the treadmill. Thus, humans clearly do not adopt a control strategy whose primary goal is to maintain some constant absolute position on the treadmill. Instead, humans appear to regulate their stepping movements in a way most consistent with a strategy whose primary goal is to try to maintain the same speed as the treadmill at each consecutive stride. These findings have important implications both for understanding how biological systems regulate walking in general and for being able to harness these mechanisms to develop more effective rehabilitation interventions to improve locomotor performance.     

Spatial context and top-down strategies in visual search

Marvin M. Chun and Yuhong Jiang (1998) investigated the role of spatial context on visual search. They used two display conditions. In the Old Display condition, the spatial arrangement of items in the search display was kept constant throughout the experiment. In the New Display condition, the spatial arrangement of items was always novel from trial to trial. The results showed better performance with Old Displays than with New Displays. The authors proposed that repeated spatial context help guiding attention to the target location, thus they termed this effect Contextual Cueing. We present three attempts to reproduce this effect. Experiments 1 and 2 were near exact replications of experiments in Chun and Jiang's report, where we failed to obtain Contextual Cueing. Post-experimental interviews revealed that participants used different search strategies when performing the task: an 'active' strategy (an active effort to find the target), or a 'passive' strategy (intuitive search). In Experiment 3, we manipulated task instructions to bias participants into using active or passive strategies. A robust Contextual Cueing Effect was obtained only in the passive instruction condition.     

Food searching decisions in four cyprinid species

Synopsis Four species of cyprinids (vimba, bream, Danubian bleak and roach) which had been incubated and reared under identical conditions differed qualitatively as well as quantitatively in the use of prey search strategies during comparative experiments. While bream and vimba prefer to use sampling techniques under conditions of reduced visibility of benthic prey, roach favor almost exclusively directed search methods. Vimba and Danubian bleak, in contrast to bream and roach, rapidly increase their total search activity in response to successive presentations of novel prey. These results have several ecological implications which are confirmed by field data. It is concluded that food searching cyprinids follow decision rules which are species specific both in the amount of sampling and in the responsiveness to rapidly changing conditions.     

Reactive Searching and Infotaxis in Odor Source Localization

Male moths aiming to locate pheromone-releasing females rely on stimulus-adapted search maneuvers complicated by a discontinuous distribution of pheromone patches. They alternate sequences of upwind surge when perceiving the pheromone and cross- or downwind casting when the odor is lost. We compare four search strategies: three reactive versus one cognitive. The former consist of pre-programmed movement sequences triggered by pheromone detections while the latter uses Bayesian inference to build spatial probability maps. Based on the analysis of triphasic responses of antennal lobe neurons (On, inhibition, Off), we propose three reactive strategies. One combines upwind surge (representing the On response to a pheromone detection) and spiral casting, only. The other two additionally include crosswind (zigzag) casting representing the Off phase. As cognitive strategy we use the infotaxis algorithm which was developed for searching in a turbulent medium. Detection events in the electroantennogram of a moth attached to a robot indirectly control this cyborg, depending on the strategy in use. The recorded trajectories are analyzed with regard to success rates, efficiency, and other features. In addition, we qualitatively compare our robotic trajectories to behavioral search paths. Reactive searching is more efficient (yielding shorter trajectories) for higher pheromone doses whereas cognitive searching works better for lower doses. With respect to our experimental conditions (2 m from starting position to pheromone source), reactive searching with crosswind zigzag yields the shortest trajectories (for comparable success rates). Assuming that the neuronal Off response represents a short-term memory, zigzagging is an efficient movement to relocate a recently lost pheromone plume. Accordingly, such reactive strategies offer an interesting alternative to complex cognitive searching.     

Cognitive performance in rhesus monkeys varies by sex and prenatal androgen exposure

Men and women differ on performance and strategy on several spatial tasks. Rodents display similar sex differences, and manipulations of early hormone exposure alter the direction of these differences. However, most cognitive testing of nonhuman primates has utilized sample sizes too small to investigate sexually differentiated behaviors. This study presents an investigation of sex differences and the effects of prenatal androgen on spatial memory and strategy use in rhesus monkeys. Monkeys prenatally exposed to vehicle, testosterone, or the androgen receptor blocker flutamide performed a search task in which 5 of 12 goal boxes contained food rewards. Spatial consistency and the presence of local landmarks were varied. Performance when both spatial and marker cues were available did not differ by sex or prenatal treatment. Contrary to predictions, females easily solved the task when local markers were removed, and their performance outscored males. Although eliminating spatial consistency and requiring subjects to use local markers impaired performance by all monkeys, females continued to locate correct goal boxes at higher than chance levels and scored better than males. Blocking prenatal androgen exposure in males improved use of local markers. These findings suggest that the tendency to attend to landmarks and to use them in solving spatial problems is typical of females across many species, including rodents, humans, and rhesus monkeys. In rhesus monkeys and rodents, developmental androgen eliminates this specialization. However, these results are the only known example of better performance of females than males when salient markers are removed.     

Why a Confirmation Strategy Dominates Psychological Science

Our research explored the incidence and appropriateness of the much-maligned confirmatory approach to testing scientific hypotheses. Psychological scientists completed a survey about their research goals and strategies. The most frequently reported goal is to test the non-absolute hypothesis that a particular relation exists in some conditions. As expected, few scientists reported testing universal hypotheses. Most indicated an inclination to use a confirmation strategy to test the non-absolute hypotheses that a particular relation sometimes occurs or sometimes does not occur, and a disconfirmation strategy to test the absolute hypotheses that a particular relation always occurs or never occurs. The confirmatory search that dominates the field was found to be associated with the testing of non-absolute hypotheses. Our analysis indicates that a confirmatory approach is the normatively correct test of the non-absolute hypotheses that are the starting point of most studies. It also suggests that the strategy of falsification that was once proposed by Popper is generally incorrect given the infrequency of tests of universal hypotheses.     

Hippocampal and striatal dependent navigation: Sex differences are limited to acquisition

Estrogen has been demonstrated to enhance the use of hippocampal-based place learning while reducing the use of striatal-based motor-response strategy (Korol, D.L., Malin, E.L., Borden, K.A., Busby, R.A., & Couper-Leo, J. (2004). Shifts in preferred learning strategy across the estrous cycle in female rats. Horm. Behav. 45, 330–338). Previous research has focused on task acquisition and the switch from a place to motor-response navigation with training. The current paradigm allowed an examination of the interplay between these two systems by having well-trained animals switch strategies “on demand.” Female and male Sprague–Dawley rats were taught a motor-response task on a plus maze. The rats were then introduced to a place task and taught to switch, by cue, from the motor-response to place strategy. Finally, the rats were trained to continuously alternate between place and motor-responses strategies. The maze configuration allowed for an analysis of cooperative choices (both strategies result in the same goal arm), competitive choices (both strategies result in different goal arms), and single strategy choices (can only use the motor-response strategy). The results indicate that sex and estrogen-related effects on navigation strategy are limited to the initial stages of learning a task. The role of sex and estrogen is diminished once the task is well learned, and presumably, the relative involvement of the hippocampal and striatal systems is established.     

Zigzagging: theoretical insights on climbing strategies

Human and animal trails on steep hillsides often exhibit dramatic switchbacks and shortcuts. Helbing et al. have recently examined the emergence of human trail systems on flat terrains while Minetti and Margaria established the effect of gradients on human metabolic efficiency. In this paper we use these ideas to develop a semi-quantitative theoretical model of the behaviour of humans moving on a terrain with relief. The model determines the direction of movement by minimising metabolic cost per unit of distance in a desired direction. The structure of the theory resembles the Landau Theory of Phase Transitions, much used in theoretical physics. We find that both hairpin bends (switchbacks) and shortcuts appear as efficient strategies for downhill walkers, while uphill walkers retain switchbacks. For weakly inclined slopes, the best strategy involves walking directly uphill or downhill. For sufficiently steep slopes, however, we find that the best strategy should undergo a transition to a broken symmetry solution corresponding to the switchback trail patterns typical of rugged environments. The critical slope at which this transition takes place should be less steep for uphill and downhill walkers. The theory should be amenable to empirical investigation. Amongst other applications, this model will enable us to generalize the work of previous authors to real landscapes, eventually permitting the reconstruction of ancient patterns of movement in archaeological landscapes.     

Cycling around a Curve: The Effect of Cycling Speed on Steering and Gaze Behavior

Although it is generally accepted that visual information guides steering, it is still unclear whether a curvature matching strategy or a ‘look where you are going’ strategy is used while steering through a curved road. The current experiment investigated to what extent the existing models for curve driving also apply to cycling around a curve, and tested the influence of cycling speed on steering and gaze behavior. Twenty-five participants were asked to cycle through a semicircular lane three consecutive times at three different speeds while staying in the center of the lane. The observed steering behavior suggests that an anticipatory steering strategy was used at curve entrance and a compensatory strategy was used to steer through the actual bend of the curve. A shift of gaze from the center to the inside edge of the lane indicates that at low cycling speed, the ‘look where you are going’ strategy was preferred, while at higher cycling speeds participants seemed to prefer the curvature matching strategy. Authors suggest that visual information from both steering strategies contributes to the steering system and can be used in a flexible way. Based on a familiarization effect, it can be assumed that steering is not only guided by vision but that a short-term learning component should also be taken into account.     

No evidence that consistent auditory cues facilitate 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: Consistent or Inconsistent. 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 Consistent group, each goal location within the pattern always provided a unique and consistent auditory cue throughout the experimental session. For participants in the Inconsistent group, the same distinct auditory cues were provided for each goal location; however, the locations of these auditory cues within the pattern itself were randomized from trial-to-trial throughout the experimental session. Results indicated that participants in both groups learned the spatial configuration of goal locations, but the presence of consistent auditory cues did not facilitate the learning of spatial relations among locations.     

Mice recognize the center of an enclosure

In recent years, it has been shown that animals can localize the geometric center of an area by reference to the shape of the environment. We trained a group of mice (experimental group) to search for a pellet hidden under sand in the center of a square-shaped dry maze. Three weeks later, they were tested in a triangular enclosure half the size of the training area and a circular enclosure double the size of the training area to see transfer to these enclosures. We compared their searching behavior with that of subjects that had received no training. The results show that the experimental group searched the geometric center of each enclosure in both transfer tests, while the untrained control group walked along the walls. This indicates that the experimental group localized the center not by reference to the absolute distance from the corners but by equal distances from all walls (geometric center).     

Reorienting when cues conflict: A role for information content in spatial learning?

In two experiments, human participants searched in dynamic three-dimensional virtual-environment rectangular enclosures. Unlike previous studies involving learning of features and geometry, we trained features and geometry separately before placing them in conflict. Specifically, participants learned to respond to rewarded features located along the principle axis of a rectangular search space and to respond to rewarded geometry of a rectangular search space in separate training phases followed by a single test trial. During the test trial, features and geometry were placed in conflict by situating rewarded bins during feature training in unrewarded geometric corners from geometry training and unrewarded bins during feature training in rewarded geometric corners from geometry training. Results of Experiment 1 indicated that although all participants learned features and geometry at an equivalent rate and to an equivalent level, performance during the test trial indicated no preferential responding to features or geometry. However, choice reaction time was significantly longer during the test trial compared to that of last feature and last geometry training trials. Experiment 2 attempted to dissociate information content of features and geometry from their acquired associative strength by rewarding only one geometric corner during geometry training. Results of Experiment 2 indicated that although features had presumably acquired greater associative strength relative to that of geometry by the end of training, performance during the test trial indicated no preferential responding to features or geometry. As in Experiment 1, choice reaction time was significantly longer during the test trial compared to that of last feature and last geometry training trials. Collectively, results seem to provide converging evidence against a view-based matching account of spatial learning, appear inconsistent with standard associative-based accounts of spatial learning, and suggest that information content of spatial cues may play an important role in spatial learning.     

Sex differences in search and gathering skills

The hunter–gatherer theory of sex differences states that female cognition has evolutionarily adapted to gathering and male cognition to hunting. Existing studies corroborate that men excel in hunting-related skills, but there is only indirect support for women excelling in gathering tasks. This study tested if women would outperform men in laboratory-based computer tests of search and gathering skills. In Experiment 1, men found target objects faster and made fewer mistakes than women in a classic visual search study. In Experiment 2, participants gathered items (fruits or letters presented on screen), and again, men performed significantly better. In Experiment 3, participants' incidental learning of object locations in a search experiment was studied, but no statistically significant sex differences were observed. These findings found the opposite of what was expected based on the hypothesis that female cognition has adapted to gathering. Alternative interpretations of the role of object location memory, female gathering roles and the division of labor between the sexes are discussed.