Idiothetic Research Articles

Honeybees Learn Landscape Features during Exploratory Orientation Flights

Exploration is an elementary and fundamental form of learning about the structure of the world [1-3]. Little is known about what exactly is learned when an animal seeks to become familiar with the environment. Navigating animals explore the environment for safe return to an important place (e.g., a nest site) and to travel between places [4]. Flying central-place foragers like honeybees (Apis mellifera) extend their exploration into distances from which the features of the nest cannot be directly perceived [5-10]. Bees perform short-range and long-range orientations flights. Short-range flights are performed in the immediate surroundings of the hive and occur more frequently under unfavorable weather conditions, whereas long-range flights lead the bees into different sectors of the surrounding environment [11]. Applying harmonic radar technology for flight tracking, we address the question of whether bees learn landscape features during their first short-range or long-range orientation flight. The homing flights of single bees were compared after they were displaced to areas explored or not explored during the orientation flight. Bees learn the landscape features during the first orientation flight since they returned faster and along straighter flights from explored areas as compared to unexplored areas. We excluded a range of possible factors that might have guided bees back to the hive based on egocentric navigation strategies (path integration, beacon orientation, and pattern matching of the skyline). We conclude that bees localize themselves according to learned ground structures and their spatial relations to the hive.

Differences in navigation performance and postpartal striatal volume associated with pregnancy in humans

Pregnancy is accompanied by prolonged exposure to high estrogen levels. Animal studies have shown that estrogen influences navigation strategies and, hence, affects navigation performance. High estrogen levels are related to increased use of hippocampal-based allocentric strategies and decreased use of striatal-based egocentric strategies. In humans, associations between hormonal shifts and navigation strategies are less well studied. This study compared 30 peripartal women (mean age 28years) to an age-matched control group on allocentric versus egocentric navigation performance (measured in the last month of pregnancy) and gray matter volume (measured within two months after delivery). None of the women had a previous pregnancy before study participation. Relative to controls, pregnant women performed less well in the egocentric condition of the navigation task, but not the allocentric condition. A whole-brain group comparison revealed smaller left striatal volume (putamen) in the peripartal women. Across the two groups, left striatal volume was associated with superior egocentric over allocentric performance. Limited by the cross-sectional study design, the findings are a first indication that human pregnancy might be accompanied by structural brain changes in navigation-related neural systems and concomitant changes in navigation strategy.

Optimal switching between geocentric and egocentric strategies in navigation.

Animals use a combination of egocentric navigation driven by the internal integration of environmental cues, interspersed with geocentric course correction and reorientation. These processes are accompanied by uncertainty in sensory acquisition of information, planning and execution. Inspired by observations of dung beetle navigational strategies that show switching between geocentric and egocentric strategies, we consider the question of optimal reorientation rates for the navigation of an agent moving along a preferred direction in the presence of multiple sources of noise. We address this using a model that takes the form of a correlated random walk at short time scales that is punctuated by reorientation events leading to a biased random walks at long time scales. This allows us to identify optimal alternation schemes and characterize their robustness in the context of noisy sensory acquisition as well as performance errors linked with variations in environmental conditions and agent–environment interactions.

Acute stress switches spatial navigation strategy from egocentric to allocentric in a virtual Morris water maze

Stress and stress hormones are known to influence the function of the hippocampus, a brain structure critical for cognitive-map-based, allocentric spatial navigation. The caudate nucleus, a brain structure critical for stimulus–response-based, egocentric navigation, is not as sensitive to stress. Evidence for this comes from rodent studies, which show that acute stress or stress hormones impair allocentric, but not egocentric navigation. However, there have been few studies investigating the effect of acute stress on human spatial navigation, and the results of these have been equivocal. To date, no study has investigated whether acute stress can shift human navigational strategy selection between allocentric and egocentric navigation. The present study investigated this question by exposing participants to an acute psychological stressor (the Paced Auditory Serial Addition Task, PASAT), before testing navigational strategy selection in the Dual-Strategy Maze, a modified virtual Morris water maze. In the Dual-Strategy maze, participants can chose to navigate using a constellation of extra-maze cues (allocentrically) or using a single cue proximal to the goal platform (egocentrically). Surprisingly, PASAT stress biased participants to solve the maze allocentrically significantly more, rather than less, often. These findings have implications for understanding the effects of acute stress on cognitive function in general, and the function of the hippocampus in particular.

Cultural background shapes spatial reference frame proclivity

Spatial navigation is an essential human skill that is influenced by several factors. The present study investigates how gender, age, and cultural background account for differences in reference frame proclivity and performance in a virtual navigation task. Using an online navigation study, we recorded reaction times, error rates (confusion of turning axis), and reference frame proclivity (egocentric vs. allocentric reference frame) of 1823 participants. Reaction times significantly varied with gender and age, but were only marginally influenced by the cultural background of participants. Error rates were in line with these results and exhibited a significant influence of gender and culture, but not age. Participants’ cultural background significantly influenced reference frame selection; the majority of North-Americans preferred an allocentric strategy, while Latin-Americans preferred an egocentric navigation strategy. European and Asian groups were in between these two extremes. Neither the factor of age nor the factor of gender had a direct impact on participants’ navigation strategies. The strong effects of cultural background on navigation strategies without the influence of gender or age underlines the importance of socialized spatial cognitive processes and argues for socio-economic analysis in studies investigating human navigation.

Reprint of “Value of water mazes for assessing spatial and egocentric learning and memory in rodent basic research and regulatory studies”

Maneuvering safely through the environment is central to survival of all animals. The ability to do this depends on learning and remembering locations. This capacity is encoded in the brain by two systems: one using cues outside the organism (distal cues), allocentric navigation, and one using self-movement, internal cues and sometimes proximal cues, egocentric navigation. Allocentric navigation involves the hippocampus, entorhinal cortex, and surrounding structures (e.g., subiculum); in humans this system encodes declarative memory (allocentric, semantic, and episodic, i.e., memory for people, places, things, and events). This form of memory is assessed in laboratory animals by many methods, but predominantly the Morris water maze (MWM). Egocentric navigation involves the dorsal striatum and connected structures; in humans this system encodes routes and integrated paths and when over-learned becomes implicit or procedural memory. Several allocentric methods for rodents are reviewed and compared with the MWM with particular focus on the Cincinnati water maze (CWM). MWM advantages include minimal training, no food deprivation, ease of testing, reliable learning, insensitivity to differences in body weight and appetite, absence of non-performers, control methods for performance effects, repeated testing capability and other factors that make this test well-suited for regulatory studies. MWM limitations are also reviewed. Evidence-based MWM design and testing methods are presented. On balance, the MWM is arguably the preferred test for assessing learning and memory in basic research and regulatory studies and the CWM is recommended if two tests can be accommodated so that both allocentric (MWM) and egocentric (CWM) learning and memory can be effectively and efficiently assessed.

The effect of TOMM40 on spatial navigation in amnestic mild cognitive impairment

The very long (VL) poly-T variant at rs10524523 (“523”) of the TOMM40 gene may hasten the onset of late-onset Alzheimer’s disease (LOAD) and induce more profound cognitive impairment compared with the short (S) poly-T variant. We examined the influence of TOMM40 “523” polymorphism on spatial navigation and its brain structural correlates. Participants were apolipoprotein E (APOE) ε3/ε3 homozygotes with amnestic mild cognitive impairment (aMCI). The homozygotes were chosen because APOE ε3/ε3 variant is considered “neutral” with respect to LOAD risk. The participants were stratified according to poly-T length polymorphisms at “523” into homozygous for S (S/S; n = 16), homozygous for VL (VL/VL; n = 15) TOMM40 poly-T variant, and heterozygous (S/VL; n = 28) groups. Neuropsychological examination and testing in real-space human analog of the Morris Water Maze were administered. Both self-centered (egocentric) and world-centered (allocentric) spatial navigation was assessed. Brain magnetic resonance imaging scans were analyzed using FreeSurfer software. The S/S group, although similar to S/VL and VL/VL groups in demographic and neuropsychological profiles, performed better on allocentric navigation (p ≤ 0.004) and allocentric delayed recall (p ≤ 0.014), but not on egocentric navigation. Both S/VL and VL/VL groups had thinner right entorhinal cortex (p ≤ 0.043) than the S/S group, whereas only the VL/VL group had thinner left entorhinal cortex (p = 0.043) and left posterior cingulate cortex (p = 0.024) than the S/S group. In conclusion, TOMM40 “523” VL variants are related to impairment in allocentric spatial navigation and reduced cortical thickness of specific brain regions among aMCI individuals with (LOAD neutral) APOE ε3/ε3 genotype. This may reflect a specific role of TOMM40 “523” in the pathogenesis of LOAD.

Security and Spatial Searching Strategies. the Comparison of Spatial Abilities and Strategies in Anxiety-secure Dimension.

Introduction The success of orientation is influenced not only by spatial abilities but also by spatial orientation strategies. We measured the efficiency of spatial memory with traditional objetive spatial exercises. Objectives We focused on the relation between spatial abilities, strategies and security. We compared the learning time in CGA and the exploration strategies with spatial memory, as well as with the egocentric and allocentric dominant strategies in Lawton navigation strategy questionnaire. Aims We searched for egocentric and allocentric spatial navigation strategies which influence on the information processing and agoraphobic fears. Methods Computer Generated Arena (CGA), which simulates the spatial situation of Morris Water Maze (MWM). Corsi Block Task (Corsi, 1972), Block Design (Kohs, 1923), Lawton Navigation Strategy Questionnaire (Lawton, 1996). Participants: 77 university students, 54 women and 23 men, average age 22,4 years. Results The variable of the learning in CGA, in the Block Design and in the Corsi Block strongly correlated with each other. The spatial construction ability strongly correlated with the allocentric strategies, and the spatial security with the spatial memory. With linear regression in CGA 10th trial, the more complex orientation strategies explain the time of finding the target. The strategy of wayfinding had a strong explanation power too. By females the performance is more sensitive to anxiety temperament and agoraphobic level. Consequences Spatial learning was influenced not only by memory but also by learning strategies. The secure object relations correlated with the use of more developed allocentric strategies.

Sequential egocentric navigation and reliance on landmarks in Williams syndrome and typical development.

Visuospatial difficulties in Williams syndrome (WS) are well documented. Recently, research has shown that spatial difficulties in WS extend to large-scale space, particularly in coding space using an allocentric frame of reference. Typically developing (TD) children and adults predominantly rely on the use of a sequential egocentric strategy to navigate a large-scale route (retracing a sequence of left–right body turns). The aim of this study was to examine whether individuals with WS are able to employ a sequential egocentric strategy to guide learning and the retracing of a route. Forty-eight TD children, aged 5, 7, and 9 years and 18 participants with WS were examined on their ability to learn and retrace routes in two (6-turn) virtual environment mazes (with and without landmarks). The ability to successfully retrace a route following the removal of landmarks (use of sequential egocentric coding) was also examined. Although in line with TD 5-year-olds when learning a route with landmarks, individuals with WS showed significantly greater detriment when these landmarks were removed, relative to all TD groups. Moreover, the WS group made significantly more errors than all TD groups when learning a route that never contained landmarks. On a perceptual view-matching task, results revealed a high level of performance across groups, indicative of an ability to use this visual information to potentially aid navigation. These findings suggest that individuals with WS rely on landmarks to a greater extent than TD children, both for learning a route and for retracing a recently learned route. TD children, but not individuals with WS, were able to fall back on the use of a sequential egocentric strategy to navigate when landmarks were not present. Only TD children therefore coded sequential route information simultaneously with landmark information. The results are discussed in relation to known atypical cortical development and perceptual-matching abilities in WS.

Spatial memory deficit across aging: current insights of the role of 5-HT7 receptors.

Elderly persons often face biological, psychological or social changes over time that may cause discomfort or morbidity. While some cognitive domains remain stable over time, others undergo a decline. Spatial navigation is a complex cognitive function essential for independence, safety and quality of life. While egocentric (body-centered) navigation is quite preserved during aging, allocentric (externally-centered) navigation—based on a cognitive map using distant landmarks—declines with age. Recent preclinical studies showed that serotonergic 5-HT7 receptors are localized in brain regions associated with allocentric spatial navigation processing. Behavioral assessments with pharmacological or genetic tools have confirmed the role of 5-HT7 receptors in allocentric navigation. Moreover, few data suggested a selective age-related decrease in the expression of 5-HT7 receptors in pivotal brain structures implicated in allocentric navigation such as the hippocampal CA3 region. We aim to provide a short overview of the potential role of 5-HT7 receptors in spatial navigation, and to argue for their interests as therapeutic targets against age-related cognitive decline.

Dopamine depletion in either the dorsomedial or dorsolateral striatum impairs egocentric Cincinnati water maze performance while sparing allocentric Morris water maze learning

Both egocentric route-based learning and spatial learning, as assessed by the Cincinnati water maze (CWM) and Morris water maze (MWM), respectively, are impaired following an 80% dopamine (DA) loss in the neostriatum after 6-hydroxydopamine (6-OHDA) administration in rats. The dorsolateral striatum (DLS) and the dorsomedial striatum (DMS) are implicated in different navigational learning types, namely the DLS is implicated in egocentric learning while the DMS is implicated in spatial learning. This experiment tested whether selective DA loss through 6-OHDA lesions in the DMS or DLS would impair one or both types of navigation. Both DLS and DMS DA loss significantly impaired route-based CWM learning, without affecting spatial or cued MWM performance. DLS 6-OHDA lesions produced a 75% DA loss in this region, with no changes in other monoamine levels in the DLS or DMS. DMS 6-OHDA lesions produced a 62% DA loss in this region, without affecting other monoamine levels in the DMS or DLS. The results indicate a role for DA in DLS and DMS regions in route-based egocentric but not spatial learning and memory. Spatial learning deficits may require more pervasive monoamine reductions within each region before deficits are exhibited. This is the first study to implicate DLS and DMS DA in route-based egocentric navigation.

Value of water mazes for assessing spatial and egocentric learning and memory in rodent basic research and regulatory studies

Maneuvering safely through the environment is central to survival of all animals. The ability to do this depends on learning and remembering locations. This capacity is encoded in the brain by two systems: one using cues outside the organism (distal cues), allocentric navigation, and one using self-movement, internal cues and sometimes proximal cues, egocentric navigation. Allocentric navigation involves the hippocampus, entorhinal cortex, and surrounding structures (e.g., subiculum); in humans this system encodes declarative memory (allocentric, semantic, and episodic, i.e., memory for people, places, things, and events). This form of memory is assessed in laboratory animals by many methods, but predominantly the Morris water maze (MWM). Egocentric navigation involves the dorsal striatum and connected structures; in humans this system encodes routes and integrated paths and when over-learned becomes implicit or procedural memory. Several allocentric methods for rodents are reviewed and compared with the MWM with particular focus on the Cincinnati water maze (CWM). MWM advantages include minimal training, no food deprivation, ease of testing, reliable learning, insensitivity to differences in body weight and appetite, absence of non-performers, control methods for performance effects, repeated testing capability and other factors that make this test well-suited for regulatory studies. MWM limitations are also reviewed. Evidence-based MWM design and testing methods are presented. On balance, the MWM is arguably the preferred test for assessing learning and memory in basic research and regulatory studies and the CWM is recommended if two tests can be accommodated so that both allocentric (MWM) and egocentric (CWM) learning and memory can be effectively and efficiently assessed.

Neural Correlates of Spatial Navigation Changes in Mild Cognitive Impairment and Alzheimer’s Disease

Although the memory impairment is a hallmark of Alzheimer’s disease (AD), AD has also been characterized by spatial disorientation, which is present from its early stages. Spatial disorientation in AD manifests itself in getting lost in familiar and unfamiliar places and have been characterized more specifically using spatial navigation tests in both real space and virtual environments as an impairment in multiple spatial abilities, including allocentric and egocentric navigation strategies, visuo-spatial perception, or selection of relevant information for successful navigation. Patients suffering mild cognitive impairment (MCI), who are at a high risk of development of dementia, show impairment in a subset of these abilities, mainly connected with allocentric and egocentric processing. While spatial disorientation in typical AD patients probably reflects neurodegenerative changes in medial and posterior temporal, parietal, and frontal lobes, and retrosplenial cortex, the impairment of spatial navigation in MCI seem to be connected mainly with the medial temporal and also parietal brain changes. In this review, we will summarize the signs of brain disease in most MCI and AD patients showing in various tasks of spatial memory and navigation.

Assessing spatial learning and memory in rodents.

Maneuvering safely through the environment is central to survival of almost all species. The ability to do this depends on learning and remembering locations. This capacity is encoded in the brain by two systems: one using cues outside the organism (distal cues), allocentric navigation, and one using self-movement, internal cues and nearby proximal cues, egocentric navigation. Allocentric navigation involves the hippocampus, entorhinal cortex, and surrounding structures; in humans this system encodes allocentric, semantic, and episodic memory. This form of memory is assessed in laboratory animals in many ways, but the dominant form of assessment is the Morris water maze (MWM). Egocentric navigation involves the dorsal striatum and connected structures; in humans this system encodes routes and integrated paths and, when overlearned, becomes procedural memory. In this article, several allocentric assessment methods for rodents are reviewed and compared with the MWM. MWM advantages (little training required, no food deprivation, ease of testing, rapid and reliable learning, insensitivity to differences in body weight and appetite, absence of nonperformers, control methods for proximal cue learning, and performance effects) and disadvantages (concern about stress, perhaps not as sensitive for working memory) are discussed. Evidence-based design improvements and testing methods are reviewed for both rats and mice. Experimental factors that apply generally to spatial navigation and to MWM specifically are considered. It is concluded that, on balance, the MWM has more advantages than disadvantages and compares favorably with other allocentric navigation tasks.

Transcranial Direct Current Stimulation Modulates Activation and Effective Connectivity During Spatial Navigation

BackgroundAllocentric navigation declines with age and neurologic disease whereas egocentric navigation does not; differences that likely arise from maladaptive changes in brain regions mediating spatial (parietal cortex; hippocampus) but not procedural processing (caudate nucleus). Transcranial direct current stimulation (tDCS) holds promise for treating such decline given its ability to modulate neuronal excitability, but its effects have yet to be examined on spatial navigation. Objectives/hypothesesUsing healthy young adults as a model, Study 1 intended to validate a novel spatial navigation paradigm using functional magnetic resonance imaging (fMRI). Using these data to determine targets for tDCS, Study 2 aimed to determine if 1) stimulation modulates activation in a polarity-specific manner; 2) stimulation results in global and/or task-specific activation changes; 3) activation changes are accompanied by changes in effective connectivity. MethodsAll participants underwent fMRI while learning allocentric and egocentric environments. Twelve participants completed Study 1. In Study 2, 16 participants were randomized to 20 min of tDCS (2 mA) using a montage with the anode over PZ and cathode over AF4 (P+F−) or the reverse montage (P−F+). ResultsStudy 1 revealed that distinct networks preferentially mediate allocentric and egocentric navigation. Study 2 revealed polarity-dependent changes in activation and connectivity. The P+F− montage increased these measures in spatial regions, especially during allocentric navigation, and the caudate nucleus. Conversely, the P−F+ montage increased activation and connectivity in lateral prefrontal cortices and posterior hippocampus. ConclusionsThese findings support the neuromodulatory effects of tDCS in non-motor areas and demonstrate proof-of-principle for ameliorating age- and disease-related decline in navigational abilities.

Effect of Donepezil in Alzheimer Disease Can Be Measured by a Computerized Human Analog of the Morris Water Maze

Background: Drug development for Alzheimer disease (AD) is challenged by the success in animal models tested in the Morris water maze (MWM) and the subsequent failures to meet primary outcome measures in phase II or III clinical trials in patients. The human variant of MWM (hMWM) enables us to examine allocentric and egocentric navigation as in the MWM. Objective: It was the aim of this study to examine the utility of a computerized hMWM to assess the effects of donepezil in mild AD. Methods: Donepezil 5 mg/day was started after initial hMWM testing in the treated group (n = 12), and after 28 days, the dose was increased to 10 mg/day. The performance after 3 months was compared to that of a non-treated group (n = 12). Results: Donepezil stabilized or improved the spatial navigation performance after 3 months, especially in the allocentric delayed recall subtask (p = 0.014). Conclusions: The computerized hMWM has the potential to measure the effects of donepezil in mild AD. It is a sensitive cognitive outcome measure in AD clinical trials.

Egocentric virtual maze learning in adult survivors of childhood abuse with dissociative disorders: Evidence from functional magnetic resonance imaging

Present neuroimaging findings suggest two subtypes of trauma response, one characterized predominantly by hyperarousal and intrusions, and the other primarily by dissociative symptoms. The neural underpinnings of these two subtypes need to be better defined. Fourteen women with childhood abuse and the current diagnosis of dissociative amnesia or dissociative identity disorder but without posttraumatic stress disorder (PTSD) and 14 matched healthy comparison subjects underwent functional magnetic resonance imaging (fMRI) while finding their way in a virtual maze. The virtual maze presented a first-person view (egocentric), lacked any topographical landmarks and could be learned only by using egocentric navigation strategies. Participants with dissociative disorders (DD) were not impaired in learning the virtual maze when compared with controls, and showed a similar, although weaker, pattern of activity changes during egocentric learning when compared with controls. Stronger dissociative disorder severity of participants with DD was related to better virtual maze performance, and to stronger activity increase within the cingulate gyrus and the precuneus. Our results add to the present knowledge of preserved attentional and visuospatial mnemonic functioning in individuals with DD.

Spatial navigation in young versus older adults

Older age is associated with changes in the brain, including the medial temporal lobe, which may result in mild spatial navigation deficits, especially in allocentric navigation. The aim of the study was to characterize the profile of real-space allocentric (world-centered, hippocampus-dependent) and egocentric (body-centered, parietal lobe dependent) navigation and learning in young vs. older adults, and to assess a possible influence of gender. We recruited healthy participants without cognitive deficits on standard neuropsychological testing, white matter lesions or pronounced hippocampal atrophy: 24 young participants (18–26 years old) and 44 older participants stratified as participants 60–70 years old (n = 24) and participants 71–84 years old (n = 20). All underwent spatial navigation testing in the real-space human analog of the Morris Water Maze, which has the advantage of assessing separately allocentric and egocentric navigation and learning. Of the eight consecutive trials, trials 2–8 were used to reduce bias by a rebound effect (more dramatic changes in performance between trials 1 and 2 relative to subsequent trials). The participants who were 71–84 years old (p < 0.001), but not those 60–70 years old, showed deficits in allocentric navigation compared to the young participants. There were no differences in egocentric navigation. All three groups showed spatial learning effect (p’ s ≤ 0.01). There were no gender differences in spatial navigation and learning. Linear regression limited to older participants showed linear (β = 0.30, p = 0.045) and quadratic (β = 0.30, p = 0.046) effect of age on allocentric navigation. There was no effect of age on egocentric navigation. These results demonstrate that navigation deficits in older age may be limited to allocentric navigation, whereas egocentric navigation and learning may remain preserved. This specific pattern of spatial navigation impairment may help differentiate normal aging from prodromal Alzheimer’s disease.

Augmented Reality Exocentric Navigation Paradigm for Time Delayed Teleoperation

This paper describes the implementation and evaluation of a new paradigm for time delayed teleoperation - exocentric navigation and publishes experimental results of this paradigm in comparison with the more commonly used navigation algorithm which will be termed egocentric navigation. A typical example for egocentric navigation is steering of a car, where navigation is a continuous process with orientation and displacement occurring simultaneously. Exocentric navigation approaches the navigation problem as a two step process, orientation followed by displacement. It can be viewed as partial autonomy in navigation, where orientation is autonomously performed by the remote robot without assistance and displacement is determined by the human operator using, in most instances, a joystick that controls both direction and speed. Evaluation of the exocentric paradigm consists of testing it with respect to navigation, perception and manipulation. System performance is measured in terms of navigation times and number of collisions and operator performance is measured using Situational Awareness Global Assessment Technique (SAGAT). The new paradigm is tested on users both with and without prior experience with results aimed at aiding human operator training.

P.7.e.002 The effects of lisdexamfetamine dimesylate on the driving performance of young adults with ADHD

This paper describes the implementation and evaluation of a new paradigm for time delayed teleoperation - exocentric navigation and publishes experimental results of this paradigm in comparison with the more commonly used navigation algorithm which will be termed egocentric navigation. A typical example for egocentric navigation is steering of a car, where navigation is a continuous process with orientation and displacement occurring simultaneously. Exocentric navigation approaches the navigation problem as a two step process, orientation followed by displacement. It can be viewed as partial autonomy in navigation, where orientation is autonomously performed by the remote robot without assistance and displacement is determined by the human operator using, in most instances, a joystick that controls both direction and speed. Evaluation of the exocentric paradigm consists of testing it with respect to navigation, perception and manipulation. System performance is measured in terms of navigation times and number of collisions and operator performance is measured using Situational Awareness Global Assessment Technique (SAGAT). The new paradigm is tested on users both with and without prior experience with results aimed at aiding human operator training.