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Sex differences in a virtual water maze: An eye tracking and pupillometry study
Abstract
Sex differences in human spatial navigation are well known. However, the exact strategies that males and females employ in order to navigate successfully around the environment are unclear. While some researchers propose that males prefer environment-centred (allocentric) and females prefer self-centred (egocentric) navigation, these findings have proved difficult to replicate. In the present study we examined eye movements and physiological measures of memory (pupillometry) in order to compare visual scanning of spatial orientation using a human virtual analogue of the Morris Water Maze task. Twelve women and twelve men (average age=24 years) were trained on a visible platform and had to locate an invisible platform over a series of trials. On all but the first trial, participants' eye movements were recorded for 3s and they were asked to orient themselves in the environment. While the behavioural data replicated previous findings of improved spatial performance for males relative to females, distinct sex differences in eye movements were found. Males tended to explore consistently more space early on while females demonstrated initially longer fixation durations and increases in pupil diameter usually associated with memory processing. The eye movement data provides novel insight into differences in navigational strategies between the sexes.
... Many rodent tasks were adapted for human research using virtual reality, including the Morris Water Maze task (MWM), the radial arm maze, and the T-maze. A male advantage is usually found in the MWM (Astur et al., 1998;Astur et al., 2004;Newhouse et al., 2007;Mueller et al., 2008;Woolley et al., 2010) and was also found in one study that investigated wayfinding (Iaria et al., 2008). Studies examining performance in radial mazes did not find sex differences (Astur et al., 2004;Levy et al., 2005). ...
... Consistent with this idea, the absence of landmarks can disadvantage women as they often rely on landmarks when navigating and are impaired when there are none (Sandstrom et al., 1998;Andersen et al., 2012), while men's navigation performance will remain unaffected because they use other sources of information such as cardinal directions and Euclidean coordinates (Dabbs Jr et al., 1998;Sandstrom et al., 1998;Saucier et al., 2002;Postma et al., 2004;Ruggiero et al., 2008;Andersen et al., 2012). Additionally, men are better at estimating precise angles and distances (Holding and Holding, 1988;Postma et al., 1998;Lawton and Morrin, 1999;Postma et al., 2004;Iachini et al., 2005), which could explain why they perform better than women in navigation paradigms in which these skills are required, such as the Morris Water Maze (Astur et al., 1998;Astur et al., 2004;Rizk-Jackson et al., 2006;Mueller et al., 2008;Nowak and Moffat, 2011;Dahmani et al., 2012). ...
... The aim of experiment 4 was to compare spatial and response learners in latencies to find specific targets using a Morris Water Maze, a task that can only be solved using a spatial strategy (Morris, 1987)32. Considering several studies have found men to outperform women on this task, we hypothesized that response women would have higher latencies to find the target compared to other groups (Astur et al., 1998;Astur et al., 2004;Rizk-Jackson et al., 2006;Mueller et al., 2008;Chai and Jacobs, 2009;Nowak and Moffat, 2011). In experiment 5 we aimed to test the use of landmarks using an Invisible Sensor Task (IST), which is a real-life spatial task that was modelled after the Morris Water Maze. ...
Sex differences in navigation have been a topic of investigation for decades and has been subjected to various contradictory findings and debates. The aim of this work was to compare the spatial memory of men and women tested in various different types of spatial tasks, while controlling for navigation strategies and aging. It is generally thought that men outperform women in navigation and that women have higher scores on object location tasks. However, many studies fail to control for different factors that may bias one sex or the other. We aggregated the data of 465 participants (349 young adults, 127 older adults) who took part in various studies conducted in our laboratory, which include both published and original unpublished data, in order to investigate sex differences. In these studies, we used a number of different paradigms: virtual radial arm mazes, a virtual wayfinding task, an object location task, a virtual Morris Water Maze, and the invisible sensor task which is a real-life model of the Morris Water Maze. While our results may seem discordant at first glance, they demonstrate that several factors can impact the performance of men and women on spatial tasks, including spontaneous navigation strategies, environmental characteristics, and age. We replicated findings showing that women favor proximal landmarks compared to men who favor distal landmarks, women have better memory than men for the position of objects in the absence of reference frames, but they will have poorer scores when navigation requires specific angles, distances and polar coordinates. Moreover, we found that in aging, women who avoid the use of landmarks when navigating a radial maze show stronger reliance on these non-spatial strategies than men. On the other hand, women who rely on landmarks, do so to the same extent as men. Our findings highlight the need to carefully take into consideration these factors in order to produce a more harmonious understanding of sex differences in navigation. Finally, the interaction between spontaneous navigation strategies, sex, and age is discussed in terms of its implications for risk of Alzheimer’s disease.
... Overall, the reported results support essential differences between men and women regarding attitudes, product searching efficiency, and in-store shopping time (Bogomolova et al. 2016;Hasan 2010;Sommer et al. 1992). Some research exploring gender-based differences in virtual spatial Navigation has confirmed that, on average, males and females perform differently in their employment of specific spatial Navigation and goal-orientated strategies (Mueller et al. 2008;Spiers et al. 2008;Tlauka et al. 2005). Using motion-tracking immersive VR systems, one recent study demonstrated that women spend significantly more time instore (mostly interacting with products) than men (Schnack et al. 2020). ...
... Findings have consistently suggested that women's visual attention to products tends to be greater than men's, likely indicating their different buying styles and strategies (Hwang and Lee 2018). Also, studies based on spatial cognition have reported evidence of different cognitive strategies adopted by men and women when navigating VR environments (Andersen et al. 2012;Mueller et al. 2008). For instance, in a virtual maze setting, relatively longer fixation durations and greater increases in pupil diameter have been observed in women compared to men (e.g., Mueller et al. 2008). ...
... Also, studies based on spatial cognition have reported evidence of different cognitive strategies adopted by men and women when navigating VR environments (Andersen et al. 2012;Mueller et al. 2008). For instance, in a virtual maze setting, relatively longer fixation durations and greater increases in pupil diameter have been observed in women compared to men (e.g., Mueller et al. 2008). ...
The use of virtual reality (VR) technology in the context of retail is a signifcant trend in current consumer research, as it
ofers market researchers a unique opportunity to measure purchase behavior more realistically. Yet, efective methods for
assessing the virtual shopping experience based on consumer’s demographic characteristics are still lacking. In this study, we
examine the validity of behavioral biometrics for recognizing the gender and age of customers in an immersive VR environment. We used behavior measures collected from eye-tracking, body posture (head and hand), and spatial navigation sources.
Participants (n=57) performed three tasks involving two diferent purchase situations. Specifcally, one task focused on free
browsing through the virtual store, and two other tasks focused on product search. A set of behavioral features categorized
as kinematic, temporal, and spatial domains was processed based on two strategies. First, the relevance of such features in
recognizing age and gender with and without including the spatial segmentation of the virtual space was statistically analyzed.
Second, a set of implicit behavioral features was processed and demographic characteristics were recognized using a statistical
supervised machine learning classifer algorithm via a support vector machine. The results confrmed that both approaches
were signifcantly insightful for determining the gender and age of buyers. Also, the accuracy achieved when applying the
machine learning classifer (>70%) indicated that the combination of all metrics and tasks was the best classifcation strategy.
The contributions of this work include characterizing consumers in v-commerce spaces according to the shopper’s profile.
... There are two recognized spatial navigation strategies: (1) local landmark usage and (2) distant cues (eg, cardinal directions) utilization [11]. All neurotypical humans are capable of utilizing either strategy when required by a given situation; however, individuals have a reliable tendency to default to one strategy over the other, and sex is a strong predictor of one's default strategy [11][12][13][14][15]. In these spatial tasks, females tend to default to using local landmarks while males default to utilizing distal cues. ...
... Our finding that sex is predictive of differences in physician GI during EHR usage is consistent with work in other navigatory domains-both spatial and electronic [11][12][13][14][15][20][21][22]25]. When viewed as a whole, the work done in this area supports that sex-linked navigatorial differences are global and have at least part of their roots in biology as opposed to being solely linked to socialization or environmental cues [17][18][19]. ...
... This is not a completely unexpected finding. While differences in default navigatory strategies are associated with significant performance differences in spatial navigation tasks, there is more variance in the types and the extent of performance differences exhibited in various types of electronic navigation as in this study [12]. This is likely in part due to the fact that biologic differences would have evolved due to evolutionary pressure from spatial orientation tasks. ...
Background
Use of electronic health records (EHRs) has increased dramatically over the past decade. Their widespread adoption has been plagued with numerous complaints about usability, with subsequent impacts on patient safety and provider well-being. Data in other fields suggest biological sex impacts basic patterns of navigation in electronic media.
Objective
This study aimed to determine whether biological sex impacted physicians’ navigational strategies while using EHRs.
Methods
This is a secondary analysis of a prior study where physicians were given verbal and written signout, and then, while being monitored with an eye tracker, were asked to review a simulated record in our institution’s EHR system, which contained 14 patient safety items. Afterward, the number of safety items recognized was recorded.
Results
A total of 93 physicians (female: n=46, male: n=47) participated in the study. Two gaze patterns were identified: one characterized more so by saccadic (“scanning”) eye movements and the other characterized more so by longer fixations (“staring”). Female physicians were more likely to use the scanning pattern; they had a shorter mean fixation duration (P=.005), traveled more distance per minute of screen time (P=.03), had more saccades per minute of screen time (P=.02), and had longer periods of saccadic movement (P=.03). The average proportion of time spent staring compared to scanning (the Gaze Index [GI]) across all participants was approximately 3:1. Females were more likely than males to have a GI value <3.0 (P=.003). At the extremes, males were more likely to have a GI value >5, while females were more likely to have a GI value <1. Differences in navigational strategy had no impact on task performance.
Conclusions
Females and males demonstrate fundamentally different navigational strategies while navigating the EHR. This has potentially significant impacts for usability testing in EHR training and design. Further studies are needed to determine if the detected differences in gaze patterns produce meaningful differences in cognitive load while using EHRs.
... McGivern et al., 1997;Andersen et al., 2011). Accordingly, sex differences in eye-movements during spatial tasks have previously been described (Andersen et al., 2011;Mueller et al., 2008;Hussein et al., 2016). However, up to date no study has investigated sex differences in eye movements during number processing. ...
... It has been demonstrated that participants tendency towards global or local processing relates to cognitive processing styles in various tasks including spatial tasks (Basso and Lowery, 2004; 7 (Andersen et al., 2011;Mueller et al., 2008;Hussein et al., 2016), no study so far has explored sex differences in the number comparison using eye-tracking. ...
... This is in line with our hypothesis and the assumption of a more decomposed processing style in women. Longer fixation durations in women have also been reported for other stimulus material (Andersen et al., 2011;Mueller et al., 2008), supporting the assumption that women take in their visual environment in more detail than men. However, the stronger compatibility effect in fixation durations in women was mostly attributable to decade digits rather than unit digits, which is contrary to our expectation and previous results by Moeller et al. (2009). ...
Multi-digit numbers are of a hierarchical nature with whole number magnitudes depending on digit magnitudes. Processing of multi-digit numbers can occur in a holistic or decomposed fashion. The unit-decade compatibility effect during number comparison is often used as a measure of decomposed processing. It refers to the fact that performance is reduced when the larger number contains the smaller unit digit (e.g. 73 vs. 26). It has been demonstrated that women show a larger compatibility effect than men, which is in accordance with their general tendency towards focusing on stimulus details during processing of visual hierarchical stimuli (local processing style). Such a local processing style has been related to higher progesterone and lower testosterone levels. One method to study individual processing styles is eye-tracking. The aim of the present study was to examine whether sex and sex hormones (estradiol, progesterone, testosterone) relate to eye movement behavior in the number comparison task. Unlike previous studies we found no evidence for sex differences in the behavioral compatibility effect. Nevertheless, women look more often and longer at individual digits and show a stronger compatibility effect in fixation durations compared to men, while men show more saccades between numbers than women. Estradiol and progesterone were related to fewer fixations and shorter fixation durations and more saccades between numbers in men, but not in women. Furthermore, the compatibility effect in the number of fixations and fixation durations was negatively related to testosterone in women. In summary, this is the first study to demonstrate sex differences and sex hormone influences on eye gaze behavior during number comparison.
... Behavioural and psychometric tests have repeatedly demonstrated gender differences in spatial ability and performance, with males showing an advantage over females in psychometric measures of spatial ability and in performance of spatial tasks (e.g., Astur et al., 1998;Astur et al., 2016;Hao et al., 2016;Hegarty et al., 2006;Moffat et al., 1998;Mueller et al., 2008;Persson et al., 2013;Ross et al., 2006;Wegman et al., 2014, see Maguire et al., 1999 for a review). However, the neurophysiological mechanisms that may underlie such prominent group differences in performance are still unclear. ...
... Overall, the behavioural results showed clear gender differences in spatial navigation performance/ability: males had higher scores on the sense of direction scale and faster performance on the maze task; males also showed significantly improved performance in the familiar vs. new environments, while females did not. These results are in line with the idea that males and females differ in how they process environmental cues (e.g., Sandstrom et al., 1998;Saucier et al., 2002;Chai & Jacobs, 2009;Iachini et al., 2005;Kong et al., 2017;Lawton, 1994;Mueller et al., 2008). ...
... The behavioural data agree with previous studies (e.g., Astur et al., 1998;Mueller et al., 2008, see a recent meta-analysis Nazareth et al., 2019) showing that males were faster than females to find the hidden platform in a virtual Morris water maze task. We further showed that males, but not females, significantly improved performance from the new (training set one) to familiar (training set two) environment. ...
Hippocampal rhythms are important for spatial navigation. This study examined whether gender differences in human navigation performance are associated with differences in hippocampal rhythms. We measured brain activities in males and females with whole-head magnetoencephalography (MEG), while they performed a virtual Morris water maze task. Behavioural results showed clear gender differences: males were significantly faster than females; unlike males, females did not show improved navigation performance in a familiar vs. new environment. MEG results showed that the magnitudes of right hippocampal/parahippocampal theta rhythm were similar between the two groups during navigation in a new environment; however, unlike males who exhibited a significant decrease in right hippocampal/parahippocampal theta power in the familiar environment shown before, females showed no change. This result may suggest faster environmental learning in males vs. females. After navigating in the new environment during the inter-trial (ITI) rest periods, males showed significantly higher right hippocampal/parahippocampal high-gamma power than females, suggesting greater consolidation in males. Moreover, right hippocampal/parahippocampal theta power during navigation correlated with navigation performance in both genders; high-gamma power during the ITI was correlated with navigation performance only in males. These associations may provide further support for the functional importance of gamma and theta rhythms in navigation. Overall, this study provides a new perspective for understanding the neurophysiological mechanisms underlying gender differences in spatial navigation.
... Research with rodents has found male rats tend to rely on configurational cues when completing orientation tasks in a maze (Foreman, 1985;Margueles & Gallistel, 1988;Suzuki, Augerinos, & Black, 1980;Williams, Barnett, & Meck, 1990;Williams & Meck, 1991). Extending this research paradigm to humans, sex differences have also been found using virtual analogues of the Morris Water Maze (Astur, Ortiz, & Sutherland, 1998;Mueller, Jackson, & Skelton, 2008;Newhouse, Newhouse, & Astur, 2007;Sandstrom, Kaufman, & Huettel, 1998). Originally, the Morris Water Maze (MWM) was developed to study spatial learning and memory in rodents (Morris, 1984). ...
... It was surprising to find little explicit description of the specifications of the VMWM used in each study, and little attention to the value of developing a 'standard' model and practices. After much deliberation, we built our own maze (described below) from the specifications provided by Mueller et al. (2008). We sought to determine the utility of the VMWM in investigating how males and females with varying degrees of videogame expertise orient themselves in the presence and absence of proximal cues. ...
... We expected to find sex differences in VMWM performance favouring males and videogame experts. Previous VMWM research has found a gender difference in uptake and use of distal cues (Mueller et al., 2008;Sandstrom et al., 1998), but it is not clear how proximal landmarks affect navigation performance in this measure of spatial learning and memory. To see how these elements in a virtual environment affected performance, we developed two versions of the VMWM, one containing proximal information consisting of five landmark-type cues around the pool wall, and the other lacking proximal landmarks. ...
Spatial abilities associated with success in educational and occupational fields of Science, Technology, Engineering, and Mathematics (STEM) have been repeatedly shown to be gendered, with males demonstrating measurably better spatial abilities than females. Less is known about why this is, or about how experience with spatial systems (videogames, for example) affects these abilities. We conducted two experiments with 82 participants with varying degrees of videogame experience on measures of mental rotation, spatial learning, and spatial memory. Spatial learning and memory were tested in a Virtual Morris Water Maze. In the first experiment, the maze lacked proximal landmarks. Males proved faster and more accurate than females in learning the location of the hidden platform. As predicted males also outperformed females in mental rotation abilities. Mental rotation correlated with performance in the virtual maze, indicating that in the absence of proximal landmarks, participants relied on strategies requiring mental rotation. Experienced 3D videogame players did not demonstrate superior spatial learning and memory, but performed better than novices in mental rotation. In the second experiment, the maze had proximal cues, in the form of landmarks on the circumference of the virtual pool, and gender-based differences in navigational performance significantly diminished. Under these changed environmental conditions, mental rotation ability did not correlate with performance in the VMWM, suggesting that given proximal cues, the need for mental rotation diminishes. Differences between videogame novices and experts also decreased when proximal cues were provided. Females in particular obtained more discernible benefits from videogame experience. Together, these experiments reveal how the spatial abilities and strategies used to solve the Morris maze task vary with environmental design. Given the structural similarities between the virtual maze and videogame environments, these results offer insight into how spatial experience gained through videogame playing can affect aspects of spatial cognition, and can help identify design elements that contribute to their improvement.
... Given that eye fixations are correlated to the focus of attention (Groner and Groner, 1989), the duration and frequency spent fixating on certain aspects of the environment, reflect the amount of cognitive resources allocated to processing this type of information (Greef et al., 2009). Several studies assess sex differences in gaze behavior during navigation (Miyahira et al., 2000a,b;Campagne et al., 2005;Mueller et al., 2008a;Cazzato et al., 2010;Andersen et al., 2012). An increased fixation duration, pupil dilation and decreased performance during a Morris water maze has been demonstrated by Mueller et al. (2008a) in women compared to men, who explored the environment more and therefore had decreased fixation periods. ...
... Several studies assess sex differences in gaze behavior during navigation (Miyahira et al., 2000a,b;Campagne et al., 2005;Mueller et al., 2008a;Cazzato et al., 2010;Andersen et al., 2012). An increased fixation duration, pupil dilation and decreased performance during a Morris water maze has been demonstrated by Mueller et al. (2008a) in women compared to men, who explored the environment more and therefore had decreased fixation periods. The preferred landmark strategy for women led investigators to believe that women would look at landmarks more frequently and longer than men (Andersen et al., 2012), but as the number of landmarks in an environment increases, so does the time spent looking at landmarks equally for men and women (Andersen et al., 2012). ...
Sex differences in spatial navigation have been related to different navigation strategies. For example, women are more likely to utilize local landmark-information in the environment compared to men. Furthermore, sex differences appear to be more pronounced when distances need to be judged in Euclidian terms and an allocentric representation of the environment is necessary. This suggests differential attentional processes during spatial navigation in men and women. However, eye-tracking studies on spatial navigation exploring these attentional processes are rare. The present study (39 men and 36 women) set out to investigate sex differences in eye-movements during spatial navigation in a 3D environment using virtual reality goggles. While we observed the expected sex differences in overall navigation performance, women did not benefit from the landmark-based instructions. Gaze fixations were in accordance with the preferred Euclidian strategy in men, but did not confirm the expected landmark-based strategy in women. However, high estradiol levels where related to an increased focus on landmark information. Surprisingly, women showed longer gaze distances than men, although the utilization of distal landmarks has been related to allocentric representations preferred by men. In fact, larger gaze distances related to slower navigation, even though previous studies suggest that the utilization of distal landmarks is beneficial for navigation. The findings are discussed with respect to the utility of virtual reality presentation for studies on sex differences in navigation. While virtual reality allows a full first-person immersion in the environment, proprioceptive and vestibular information is lacking.
... We did not observe any sex difference in the ability to select and use of specific navigation strategy. This result seems in contrast with results from a recent survey in healthy participants [96], results from navigation tasks [4,69] and data from neuroimaging studies suggesting that men and women use different strategies and activate different brain areas during navigation [36,47]. However, our results are consistent with other studies that did not observe any sex-related difference in the use of navigation strategies in virtual radial mazes and T-mazes [57,85]. ...
... As previously demonstrated, females generally spend more time looking at surrounding landmarks and show impaired performance when no landmarks are available [1,84], while males typically show an advantage in time performance as they are more prone to use additional information from the environment, such as room geometry or Euclidian coordinates. Although this is not correlated to the use of a spontaneous spatial strategy during navigation [1], most of the previous findings that revealed an effect of sex resulted from the analysis of the time and accuracy to navigate [36,69] and not from a direct comparison of behavioural data and verbal reports. Indeed, a third point concerns the methodology used to determine the strategies. ...
During navigation, humans mainly rely on egocentric and allocentric spatial strategies, two different frames of reference working together to build a coherent representation of the environment. Spatial memory deficits during navigation have been repeatedly reported in patients with vestibular disorders. However, little is known about how vestibular disorders can change the use of spatial navigation strategies. Here, we used a new reverse T-maze paradigm in virtual reality to explore whether vestibular loss specifically modifies the use of egocentric or allocentric spatial strategies in patients with unilateral (n = 23) and bilateral (n = 23) vestibular loss compared to healthy volunteers (n = 23) matched for age, sex and education level. Results showed that the odds of selecting and using a specific strategy in the T-maze were significantly reduced in both unilateral and bilateral vestibular loss. An exploratory analysis suggests that only right vestibular loss decreased the odds of adopting a spatial strategy, indicating an asymmetry of vestibular functions. When considering patients who used strategies to navigate, we observed that a bilateral vestibular loss reduced the odds to use an allocentric strategy, whereas a unilateral vestibular loss decreased the odds to use an egocentric strategy. Age was significantly associated with an overall lower chance to adopt a navigation strategy and, more specifically, with a decrease in the odds of using an allocentric strategy. We did not observe any sex difference in the ability to select and use a specific navigation strategy. Findings are discussed in light of previous studies on visuo-spatial abilities and studies of vestibulo-hippocampal interactions in peripheral vestibular disorders. We discuss the potential impact of the history of the disease (chronic stage in patients with a bilateral vestibulopathy vs. subacute stage in patients with a unilateral vestibular loss), of hearing impairment and non-specific attentional deficits in patients with vestibular disorders.
... Forty-two experiments with a total of 96 effects were included for eye-related outcome measure changes during learning over time (Table 2.2e), with average effect sizes of 0.39 (SD = 0.24) and 1.53 (SD = 1.04) for ηp 2 and Hedge's g respectively. Effects have been reported on relatively low-level tasks examining processes such as discrimination learning (DeLoss et al., 2014) and visual search (Hout & Goldinger, 2012;Neider et al., 2010) and on higher-level tasks such as route learning or spatial navigation (de Condappa & Wiener, 2016;Mueller et al., 2008). Most effects were reported for gaze, for a total of 84 effects. ...
... Only one of the current meta-study's included experiments specifically looked at the effect of gender. The study by Mueller et al. (2008) examined gender differences in spatial navigation learning and demonstrated that eye movements developed differently over time for males and females, while pupil size did not. Gender effects may therefore be modulated by task or measurement. ...
Up till now, there has been an unresolved discussion in the literature regarding the validity of non-invasive neurophysiological measures in learning. On the one hand studies have shown promise for these measures in learning (Krigolson et al., 2015; Lai et al., 2013; Leff et al., 2011), while on the other hand there has been caution for the use of such measures (Ansari et al., 2011; Brouwer et al., 2014; Cowley, 2015; Dahlstrom-Hakki et al., 2019). This dissertation was aimed to address this discussion. To this aim, this dissertation focused on experimentally examining non-invasive neurophysiological changes during learning and factors that influence these changes. Additionally, this dissertation focused on providing insight into how to move towards applying these measures validly and effectively in a wide range of settings, not only in the laboratory but also in real-world contexts.
Considering all of these findings together, it becomes clear that understanding the assessment of learning through neurophysiology requires an understanding of the interplay between learning, neurophysiology, behavior, individual differences, and task-related aspects. Comprehending this complex interaction is key to resolving the discussion regarding the validity of non-invasive neurophysiological measures in learning. As the reported findings demonstrate non-invasive neurophysiology to be able to provide insight into learning, the discussion should not be focussed on whether neurophysiological measures are able to assess learning, but on how to obtain valid assessments across different learning tasks and across different trainees.Although it is clear that further development and research are needed for largescale application of neurophysiology in learning and training, the potential of neurophysiology is expected to increase as the field advances (see Chapter 8 for a more in-depth discussion). Industry could benefit from being involved in future endeavors to move the field forward. Vice versa, development and research can move forward in promising directions when taking into account the needs and experiences from the industry.
The embedding of the work presented in this dissertation within the CAMPIONE project highlights how fundamental research can provide valuable contributions to application. Even though fundamental research may sometimes seem to be far removed from application, understanding the fundamentals will ultimately lead to the most valid and reliable application. I am looking forward to seeing future research contribute to our knowledge about assessment of learning through non-invasive neurophysiological measures and to seeing application of neurophysiology in training and education advance. This dissertation has paved the way and I hope many scholars and other professionals will follow up on the presented work.
... We only included males in the study to avoid differences in HPA activation caused by hormonal changes associated with the menstrual cycle or oral contraceptives. To control for prior experience with computer games and joysticks, participants completed a questionnaire [46] before starting the vMWM task. All participants provided written consent and received monetary compensation (60€). ...
... The vMWM task is a validated analogue of the original animal research tool for human studies [46]. Similar to our previous work [13], we used the vMWM task software version 1.2 developed by Derek Hamilton and colleagues [52]. ...
... We only included males in the study to avoid differences in HPA activation caused by hormonal changes associated with the menstrual cycle or oral contraceptives. To control for prior experience with computer games and joysticks, participants completed a questionnaire [46] before starting the vMWM task. All participants provided written consent and received monetary compensation (60€). ...
... The vMWM task is a validated analogue of the original animal research tool for human studies [46]. Similar to our previous work [13], we used the vMWM task software version 1.2 developed by Derek Hamilton and colleagues [52]. ...
... We only included males in the study to avoid differences in HPA activation caused by hormonal changes associated with the menstrual cycle or oral contraceptives. To control for prior experience with computer games and joysticks, participants completed a questionnaire [46] before starting the vMWM task. All participants provided written consent and received monetary compensation (60€). ...
... The vMWM task is a validated analogue of the original animal research tool for human studies [46]. Similar to our previous work [13], we used the vMWM task software version 1.2 developed by Derek Hamilton and colleagues [52]. ...
Background:
Acute stress leads to a rapid release of noradrenaline and glucocorticoids, which in turn influence cognitive functions such as spatial learning and memory. However, few studies have investigated noradrenergic and glucocorticoid effects on spatial learning and memory in humans. Therefore, we examined the separate and combined effects of noradrenergic and glucocorticoid stimulation on spatial learning and memory.
Methods:
One hundred and four healthy men (mean age = 24.1 years ±SD 3.5) underwent the virtual Morris Water Maze (vMWM) task to test spatial learning and spatial memory retrieval after receiving either 10 mg hydrocortisone or 10 mg yohimbine (an alpha 2-adrenergic receptor antagonist that increases noradrenergic activity), 10 mg hydrocortisone and 10 mg yohimbine combined, or placebo. The vMWM task took place 90 min after yohimbine was administered and 75 min after hydrocortisone was administered. Placebo was given at the same times. Salivary cortisol and alpha amylase levels were measured to check pharmacological stimulation.
Results:
Hydrocortisone and yohimbine increased salivary cortisol and alpha amylase levels. Participants' task performance improved over time, suggesting successful spatial learning. However, separate and combined noradrenergic and glucocorticoid stimulation had no effect on spatial learning and spatial memory retrieval compared with placebo.
Conclusions:
In healthy young men, hydrocortisone and/or yohimbine did not alter spatial learning or spatial memory retrieval. Importantly, pharmacological stimulation took place prior to learning. Further studies should examine the effects of glucocorticoid and noradrenergic stimulation during encoding, consolidation, and retrieval.
... Gaps, missing samples in the gaze data that were between 50 and 500 ms in duration, were considered blinks. Fixations were only included if they were greater than 100 ms in duration [67][68][69]. For each fixation, the fixated object in the VE was defined as the object that had the highest percentage of collisions of the 3D gaze vector. ...
U.S. service members maintain constant situational awareness (SA) due to training and experience operating in dynamic and complex environments. Work examining how military experience impacts SA during visual search of a complex naturalistic environment, is limited. Here, we compare Active Duty service members and Civilians’ physiological behavior during a navigational visual search task in an open-world virtual environment (VE) while cognitive load was manipulated. We measured eye-tracking and electroencephalogram (EEG) outcomes from Active Duty (N = 21) and Civilians (N = 15) while they navigated a desktop VE at a self-regulated pace. Participants searched and counted targets (N = 15) presented among distractors, while cognitive load was manipulated with an auditory Math Task. Results showed Active Duty participants reported significantly greater/closer to the correct number of targets compared to Civilians. Overall, Active Duty participants scanned the VE with faster peak saccade velocities and greater average saccade magnitudes compared to Civilians. Convolutional Neural Network (CNN) response (EEG P-300) was significantly weighted more to initial fixations for the Active Duty group, showing reduced attentional resources on object refixations compared to Civilians. There were no group differences in fixation outcomes or overall CNN response when comparing targets versus distractor objects. When cognitive load was manipulated, only Civilians significantly decreased their average dwell time on each object and the Active Duty group had significantly fewer numbers of correct answers on the Math Task. Overall, the Active Duty group explored the VE with increased scanning speed and distance and reduced cognitive re-processing on objects, employing a different, perhaps expert, visual search strategy indicative of increased SA. The Active Duty group maintained SA in the main visual search task and did not appear to shift focus to the secondary Math Task. Future work could compare how a stress inducing environment impacts these groups’ physiological or cognitive markers and performance for these groups.
... Table S1), as has been found in animal studies in the MWM (female meadow voles, Galea et al. 2002). We only found men to be less idle, corroborating the many reports that men simply tend to spatially cover more ground in mazes (Mueller et al. 2008). ...
Rationale
Animal studies suggest that the so-called “female” hormone estrogen enhances spatial navigation and memory. This contradicts the observation that males generally out-perform females in spatial navigation and tasks involving spatial memory. A closer look at the vast number of studies actually reveals that performance differences are not so clear.
Objectives
To help clarify the unclear performance differences between men and women and the role of estrogen, we attempted to isolate organizational from activational effects of estrogen on spatial navigation and memory.
Methods
In a double-blind, placebo-controlled study, we tested the effects of orally administered estradiol valerate (E2V) in healthy, young women in their low-hormone menstrual cycle phase, compared to healthy, young men. Participants performed several first-person, environmentally rich, 3-D computer games inspired by spatial navigation and memory paradigms in animal research.
Results
We found navigation behavior suggesting that sex effects dominated any E2 effects with men performing better with allocentric strategies and women with egocentric strategies. Increased E2 levels did not lead to general improvements in spatial ability in either sex but to behavioral changes reflecting navigation flexibility.
Conclusion
Estrogen-driven differences in spatial cognition might be better characterized on a spectrum of navigation flexibility rather than by categorical performance measures or skills.
... Furthermore, measures of electrodermal activity are employed in game user studies as indicators of emotional changes, however, obtained results are mixed, as some studies did not detect any changes in measurements due to gameplay conditions or could not identify a correspondence between subjective measures. Eye-Related measures, such as fixations and movements of the eye were found to be related to cognitive activity (e.g Jennet et al., 2008;Mueller, Jackson & Skelton., 2008;Alkan & Çağıltay, 2007) and pupil size indicated the emotional changes (Mojzisch et al., 2006). ...
Games User Research (GUR) has become an important subdomain of the Human- Computer Interaction field within the last few years, as gaming has becomes a daily entertainment for many people rather than being in the interest of a few game enthusiasts. Researchers require specific tools to measure the users’ responses and attitudes towards the games. Game User Experience Satisfaction Scale is one of the recent additions to GUR tools, which had already been adapted into Turkish as GUESSTR. Through this study, we aimed to verify a shorter form of GUESS-TR which is compatible with a currently existing English shorter version with 18 items that measure game user experience through 9 factors. Data revealed that a 20-item version resembling 10 factors, namely GUESS-20-TR, is a valid and reliable measure of game user experience. We provided evidence for construct validity through confirmatory factor analysis. Spearman-Brown prophecy coefficients indicate that the 2-item subscales are reliable. Heterotrait - monotrait ratios show that items indicate different constructs, i.e. discriminant validity. Based on Pearson correlation, the mean scores obtained with the short form GUESS-20-TR are highly consistent with the 51-item Turkish version.
... The primary dependent measure for the VMT (as established in previous studies) is In a secondary exploratory analysis, we will evaluate the impact of cybersickness symptoms and sense of presence in performance in both VMT versions. Furthermore, we will assess the effect of gender and perspective taking on our participants' performance on the VMT, as both measures have shown to have an impact in maze and spatial learning (Coutrot et al., 2018;Mueller et al., 2008;Piber et al., 2018;Woolley et al., 2010). ...
... Some studies have shown that men have an advantage in deciphering and remembering spaces. When exposed to new environments, men visually explore more spaces than women to understand the overall character and layout of the space [50]. Women generally rely more heavily on landmarks and exhibit sustained landmark-directed gaze during eye tracking, whereas landmark-directed gaze decreases over time among men [51]. ...
Improving the inclusiveness of urban green spaces and enabling various groups to equally enjoy their benefits is the basis of sustainable urban development. Urban park design generally starts from a gender-neutral perspective, ignoring differences in needs related to gender, particularly women’s sensitivity to the environment. This study focused on visual perception and explored gender differences and proposed causes of visual-behavior differences while viewing landscapes. We used photo data from Mizumoto Park in Tokyo and recruited 16 master’s students living nearby to participate in an eye-tracking experiment. The results indicate that men and women have different eye-movement patterns and that elemental ratios affect eye movement behavior more among women than men. Moreover, this study found that men gaze longer at trees and more briefly at shrubs, flowers, and artificial elements than women. Attention-grabbing paths/grounds had a negative effect on the aesthetic evaluation of the scene among women but not men. Based on these findings, suggestions for optimizing women’s experiences at the visual level are proposed for aspects of vegetation density, visual focus, and road design. This study informs park design and improvement with the premise that gender alters the perception of these environments.
... For example, studies found gender-related differences, as women seem to follow more often a systematic strategy and look at the landscape more intensively and with longer fixation times than men [37]. Accordingly, the increase in pupil diameter indicates a more intensive processing of memories of women, while men tend to get a quick overview of the landscape [101]. Another study related eye movements to human behavior, indicating that the level of nature relatedness could explain whether individuals looked more likely at trees than buildings [102]. ...
Profound knowledge about landscape preferences is of high importance to support decision making, in particular, in the context of emerging socioeconomic developments to foster a sustainable spatial development and the maintenance of attractive landscapes. Eye-tracking experiments are increasingly used to examine how respondents observe landscapes, but such studies are very time-consuming and costly. For the first time, this study explored the potential of using eye-tracking simulation software in a mountain landscape by (1) identifying the type of information that can be obtained through eye-tracking simulation and (2) examining how this information contributes to the explanation of landscape preferences. Based on 78 panoramic landscape photographs, representing major landscape types of the Central European Alps, this study collected 19 indicators describing the characteristics of the hotspots that were identified by the Visual Attention Software by 3M (3M-VAS). Indicators included quantitative and spatial information (e.g., number of hotspots, probabilities of initially viewing the hotspots) as well variables indicating natural and artificial features within the hotspots (e.g., clouds, lighting conditions, natural and anthropogenic features). In addition, we estimated 18 variables describing the photo content and calculated 12 landscape metrics to quantify spatial patterns. Our results indicate that on average 3.3 hotspots were identified per photograph, mostly containing single trees and tree trunks, buildings and horizon transitions. Using backward stepwise linear regression models, the hotspot indicators increased the model explanatory power by 24%. Thus, our findings indicate that the analysis of eye-tracking hotspots can support the identification of important elements and areas of a landscape, but it is limited in explaining preferences across different landscape types. Future research should therefore focus on specific landscape characteristics such as complexity, structure or visual appearance of specific elements to increase the depth of information obtained from eye-tracking simulation software.
... A nivel de percepción visual, se ha podido conocer que las mujeres suelen realizar fijaciones más largas que los hombres, mientras que estos últimos exploran más el espacio (Mueller et al., 2008;Piber et al., 2018). Además, las mujeres son más proclives a observar los objetos y sus características, y en cambio, los hombres tienden a prestar más atención a las relaciones espaciales entre los elementos que componen una escena (McGivern et al., 2019). ...
Hand-eye coordination and visuospatial integration are among the numerous skills that define human beings. These two properties are the result of a shared evolution of the visual and tactile senses, and are directly related to manipulation. The use of tools, for example, requires the combination of these two characteristics. From an evolutionary point of view, the relationship between manipulation, hand anatomy and tools has mainly been studied. However, the other source of information, vision, has not been explored yet. Therefore, the main aim of this doctoral thesis is to find out visuospatial changes in the archaeological record through the analysis of the visual behaviour in relation to Lower Palaeolithic stone tools.
Although cognitive abilities cannot be studied directly, the analysis of eye movements provides indirect evidence of the amount of visuospatial attention allocated to exploring a given scene. Likewise, the archaeological record can provide information on the visuospatial relationship between the individual and the technology. Therefore, in this research work, a series of visual attention studies are conducted in different scanning conditions, using different eye-tracking devices to analyse eye movements and visual attention. The exploration situations analysed include passive observation of photographs of tools, observation of tools in peripersonal space, manipulation and tactile exploration of tools, and finally, tool-making. For this purpose, a total of 215 participants have collaborated in six experimental studies, carried out at the National Centre for Research on Human Evolution (CENIEH, Burgos) and the University of Lincoln (UK). Apart from two studies focused on experience with lithic technology, all experimental processes are carried out on individuals with no archaeological knowledge in order to avoid biases that influence primary visual exploration. Experimental studies used examples of the first stone tool technologies, like choppers and handaxes.
The different studies of visual perception reveal that attention is not affected by the most prominent properties of the tools, instead, the tools show a characteristic pattern of observation. Furthermore, the distribution of visuospatial attention is similar in all the scanning states analysed. Stone tools trigger the same visual response regardless of the shown mode (image or replica) or location (peripersonal or personal space). In this sense, the functional areas of the tools are more observed than those areas associated with manipulation, despite the widely known tendency to observe the centre of the artefacts. The results obtained suggest that visual attention is affected by the processing of affordances or action possibilities of the tools. Despite the fact that the gaze is generally directed to the functional regions of prehistoric tools, there are differences between the examples of the technological modes analysed. Choppers are more observed in the function-associated zones, while handaxes need more exploration in the manipulative regions. We propose that the differences in the distribution of visuospatial attention according to the technological element observed correspond to the predominance of one type of affordance over another. Choppers require more executive processing while handaxes require more reflection on grasping affordances. In addition, functional areas attract a higher degree of visuospatial attention in participants with lithic industry knowledge. This behaviour is a consequence of previous experience about potential uses and modes of manipulation.
Another aspect to consider concerns the influence of the morphology of the tool on its perception. From the quantification of the shape, as well as other structural and functional variables, it has been found that the characteristics that most affect exploration are the size, weight and morphology of the tool base. However, shape does not have the same influence on choppers and handaxes. We suggest that the simpler morphology of choppers allows the focus to be on the functional area of the tool, as a complex manipulation strategy is not necessary. In this case, the scanning pattern of the worked pebbles is exclusively affected by weight. On the other hand, the shape and size of the handaxes directly influence the visual scanning pattern. Small and elongated handaxes allow for easier handling of the tool. Therefore, we propose that functional affordances condition the first tools, while the affordances associated with manipulation become more important as potential uses and morphological complexity increase. Finally, the role of vision during lithic tool manufacture is considered. Preliminary results indicate that visual behaviour is different depending on the tool manufactured.
This research work constitutes the first study on the visual response triggered by lithic technology. Several conclusions can be considered on the basis of the studies developed. At the methodological level, eye tracking is established as a useful technology to study one of the main human cognitive activities, visual perception. On the other hand, the evidence found can be related to different visuospatial processing, and therefore, to different cognitive capacities required according to the lithic technology explored.
... The subject under consideration was investigated by such authors as Brugger A. et al. [11], Kiefer P. et al. [12], Müller S. et al. [13], Franchak J. et al. [14], showing that eye-tracking is an extremely useful tool for assessing the effectiveness of existing navigation elements, as well as for proving the need for their placement. Instead of using assumptions based on generalized models, eye-tracking technology provides researchers with the exact data representing where the visual attention of specific users is directed. ...
Navigation and wayfinding are well-known issues that people face in their life every day. In many situations, and especially in the case of medical facilities, it is vital that users find the desired destination quickly and successfully. This is not only about comfort, but also about life and health safety. In this regard, the development of an effective navigation system becomes an extremely important mission for health care institutions. Therefore, the eye-tracking technology, which makes it feasible to look at the problem through visitor’s eyes, appears to be a pertinent tool for an objective assessment of the existing navigation, as well as finding ways to optimize it. This article considers present approaches to the analysis of navigation systems dealing with a large number of clients and describes our visual navigation study conducted in a health center. The first part of our study included the current navigation environment evaluation and the decision-making points detection, serving as a starting point for the implementation of a new, user-oriented navigation in the medical center. In the second part of the study the effectiveness of the applied system was confirmed. The results obtained show that navigation environment improvement can significantly reduce users’ time to reach the goal, as well as length of their route. Thus, we managed to apply the eye-tracking method to the improve user experience in a medical establishment.
... For example, the GVS-induced ocular vestibular-evoked myogenic potential (oVEMP) amplitude was reported higher in males than in females (57). Males were shown to normally perform better in vestibular function-related tasks, in particular, visuospatial cognition such as 3D figures, spatial orientation, and maze navigation (58,59). ...
Objectives
In this study, the specific threshold intensities and response characteristics of galvanic vestibular stimulation (GVS) on vestibular (conscious) and cutaneous (detrimental) perception as well as oculomotor nystagmus (reflex) were determined.
Methods
The threshold intensities for vestibular and cutaneous perception and oculomotor response induced by GVS were determined in 25 right-handed healthy subjects (32.6 ± 7.2 years of age; 56% female). The subjects were seated upright, and eye movements were recorded while a direct GVS current was applied with paradigms of cathode on the right and anode on the left (CRAL) and also cathode on the left and anode on the right (CLAR).
Results
Subjects experienced dizziness, sense of spinning, or fall tendency, which was more frequently directed to the cathode (76%) than the anode (24%, p < 0.001, chi-square one-variable test) at mean current greater than 0.98 ± 0.29 mA (mean vestibular threshold). The current also triggered a more frequent mild tingling sensation at the cathode (56%) than the anode (30%) or on both sides (14%; p = 0.001, chi-square one-variable test) when above the mean cutaneous threshold of 0.9 ± 0.29 mA. Above the mean oculomotor threshold of 1.61 ± 0.35 mA, combined horizontal and torsional nystagmus was more frequent toward the cathode (86%) than toward the anode (p < 0.001, chi-square one-variable test). The mean oculomotor threshold was significantly higher than both the vestibular (p < 0.001, Mann–Whitney U-test) and cutaneous (p < 0.001, Mann–Whitney U-test) thresholds, which were comparable (p = 0.317, Mann–Whitney U-test). There was no significant disparity in these specific thresholds between the two GVS paradigms. The vestibular threshold was significantly higher in males than in females [1 (0.5–1.25) mA vs. 0.75 (0.625–1.125) mA, Z = −2.241, p = 0.025, Mann–Whitney U-test]. However, the thresholds of cutaneous perception and oculomotor response did not differ by sex.
Conclusion
The findings indicate that thresholds for vestibular and somatosensory perception are lower than the oculomotor threshold. Therefore, a strategy to reduce GVS current intensity to the level of vestibular or somatosensory perception threshold could elicit beneficial vestibular effects while avoiding undesirable effects such as oculomotor consequences.
... It has been shown that spatial memory abilities present gender differences, with males excelling at the behavioral level of spatial memory ability [9][10][11], as evidenced by shorter reaction times and higher accuracy [12]. Some researchers have found through a watermaze experimental-research paradigm that males are considered to be more gifted than females in spatial memory [13,14] and this may influence task performance [15]. However, it has also been shown that there are no differences between males and females in spatial memory abilities [16,17]. ...
Objective:
To investigate the intervention effect of orienteering exercises on the spatial memory ability of college students of different genders and its underlying mechanism.
Methods:
Forty-eight college students were randomly screened into experimental and control groups, 12 each of male and female, by SBSOD scale. The effects of 12 weeks of orienteering exercises on the behavioral performance and brain activation patterns during the spatial memory tasks of college students of different genders were explored by behavioral tests and the fNIRS technique.
Results:
After the orienteering exercise intervention in the experimental group, the male students had significantly greater correct rates and significantly lower reaction times than the female students; left and right dorsolateral prefrontal activation was significantly reduced in the experimental group, and the male students had a significantly greater reduction in the left dorsolateral prefrontal than the female students. The degree of activation in the left and right dorsolateral prefrontals of the male students and the right dorsolateral prefrontals of the female students correlated significantly with behavioral performance, and the functional coupling between the brain regions showed an enhanced performance.
Discussion:
Orienteering exercises improve the spatial memory ability of college students, more significantly in male students. The degree of activation of different brain regions correlated with behavioral performance and showed some gender differences.
... For example, the Morris water-maze is a widely used task to study the physiological mechanisms of spatial learning in rodents [12][13][14]. Similarly, adaptations of the Morris water-maze in virtual environments [15][16][17] and other maze tasks [18,19] have been used to investigate spatial navigation in humans. However, in recent years experimental designs of spatial navigation research have shifted to make use of more naturalistic settings [20][21][22][23][24][25][26][27]. ...
Vision provides the most important sensory information for spatial navigation. Recent technical advances allow new options to conduct more naturalistic experiments in virtual reality (VR) while additionally gathering data of the viewing behavior with eye tracking investigations. Here, we propose a method that allows one to quantify characteristics of visual behavior by using graph-theoretical measures to abstract eye tracking data recorded in a 3D virtual urban environment. The analysis is based on eye tracking data of 20 participants, who freely explored the virtual city Seahaven for 90 minutes with an immersive VR headset with an inbuild eye tracker. To extract what participants looked at, we defined "gaze" events, from which we created gaze graphs. On these, we applied graph-theoretical measures to reveal the underlying structure of visual attention. Applying graph partitioning, we found that our virtual environment could be treated as one coherent city. To investigate the importance of houses in the city, we applied the node degree centrality measure. Our results revealed that 10 houses had a node degree that exceeded consistently two-sigma distance from the mean node degree of all other houses. The importance of these houses was supported by the hierarchy index, which showed a clear hierarchical structure of the gaze graphs. As these high node degree houses fulfilled several characteristics of landmarks, we named them "gaze-graph-defined landmarks". Applying the rich club coefficient, we found that these gaze-graph-defined landmarks were preferentially connected to each other and that participants spend the majority of their experiment time in areas where at least two of those houses were visible. Our findings do not only provide new experimental evidence for the development of spatial knowledge, but also establish a new methodology to identify and assess the function of landmarks in spatial navigation based on eye tracking data.
... Survey knowledge is based on cognitive maps, which are mental representations of the environment that reflect the spatial configuration of landmarks relative to one another (O'Keefe & Nadel, 1978;Taylor et al., 1999;Thorndyke & Hayes-Roth, 1982;Tolman, 1948;Zimmer, 2004). Some work has shown that distal landmarks best support survey knowledge because they provide global orientation information about the environment (De Condappa, 2016;Hamilton et al., 2002;Jacobs et al., 1997;Livingstone-Lee et al., 2011;Mueller et al., 2008). ...
We used a driving simulator to investigate landmark-based route navigation in young adults. Previous research has examined how proximal and distal landmarks influence route navigation, however, these effects have not been extensively tested in ecologically-relevant settings. We used a virtual town in which participants learned various routes while simultaneously driving. We first examined the effect of four different landmark conditions on navigation performance, such that each driver experienced one of four versions of the town with either proximal landmarks only, distal landmarks only, both proximal and distal landmarks, or no landmarks. Drivers were given real-time navigation directions along a route to a target destination, and were then tested on their ability to navigate to the same destination without directions. We found that the presence of proximal landmarks significantly improved route navigation. We then examined the effect of prior exposure to proximal vs. distal landmarks by testing the same drivers in the same environment they previously encountered, but with the landmarks removed. In this case, we found that prior exposure to distal landmarks significantly improved route navigation. The present results are in line with existing research on route navigation and landmarks, suggesting that these findings can be extended to ecologically-relevant settings.
... In some approaches, eye movements and physiological measures of memory (pupillometry) are analysed in order to compare visual exploration of spatial orientation. However, this experimentation is limited due to non-immersive tasks just using the computer screen (Mueller et al., 2008). ...
The Morris Water Maze (MWM) has become one of the most widely used laboratory tools in behavioural neuroscience. It has been used in some of the most sophisticated experiments in the study of spatial learning and memory with animals. However, human based studies have been very limited due to the use of unrealistic scenarios, usually presented on a computer screen where participants’ attention is poorly controlled. Recent advances in virtual reality (VR) enable the generation of 3D environments with a high level of realism and user’s immersion. The user’s attention plays a key role in spatial learning. Current VR systems integrate eyetracking devices to measure the user’s attention over virtual entities. In this paper, we present an easy-to-use game-based simulator of the MWM, using eye-tracking VR technology to extract information about the user’s attention. This research still in progress has achieved important hints according to the design of the virtual scenario, user interaction and experimentation. The study conducted in this paper validates the technology as a novel way to perform MWM focused on spatial learning and memory with human participants
... Alguns estudos mostram que mulheres e fêmeas têm melhor desempenho em tarefas que usam preferencialmente navegação alocêntrica (por exemplo, FERNANDEZ-BAIZAN et al., 2018) e outros mostram melhor desempenho deste sexo em tarefas que usam navegação egocêntrica (por exemplo, VORHEES e WILIAMS, 2016). Por outro lado, Mueller e colaboradores (2008) mostraram que mulheres e homens começam a explorar um novo espaço com estratégias egocêntricas e gradualmente mudavam para estratégias alocêntricas conforme se familiarizavam com o espaço(MUELLER et al., 2008). Outros estudos também não conseguiram demonstrar diferenças sexuais na preferência de navegação(BURGESS et al., 2004;FERGUSON et al., 2019;SCHEURINGER et al., 2017). ...
The female sex has been neglected in neuroscience and behavioral pharmacology research, because of the alleged increased variability in results caused by hormonal fluctuations. In studies with rodents, the vaginal lavage procedure (VLP) is a common method of smear collection to determine the stage of the estrous cycle by cytological analysis. However, little is known about the consequences of this procedure. In neuroscience, spatial memory has been a relevant issue in terms of differences between the sexes. In general, a worse performance is pointed out in females compared to males. However, due to the aforementioned negligence, spatial memory tests were standardized on males. Among the tests that evaluate spatial memory, the task of object location recognition (OLR) allows the evaluation of spatial memory with a character of recognition; the T-maze (TM) task allows the assessment of spatial memory with a referential character and the use of reward; and the plus-maze discriminative avoidance task (PMDAT) allows the concomitant assessment of spatial memory (using aversive stimulation), anxiety-like behavior and locomotion. The aim of the present study was to investigate sex differences and the effects of VLP on the performance of rats in behavioral tasks above. Adult male Wistar rats, adult female Wistar rats, and adult female Wistar rats submitted to 14 days of VLP were evaluated in behavioral tasks. In addition, another set of female rats were submitted to VLP to determine plasma corticosterone levels. Females not submitted to VLP and males had similar performance in the PMDAT task, while females were slightly better in the OLR and TM tasks. Female rats submitted to VLP had impaired performance in the tasks of PMDAT and OLR, but there were no alterations on anxiety-like behavior. The vaginal secretion collection procedure also resulted in a higher level of plasma corticosterone, suggesting that VLP is a stressful manipulation. In conclusion, females do not seem to be naturally worse at spatial memory tasks, and comparative performance in relation to males depends on the type of task and the parameters used in the comparison. Still, the use of VLP to control hormonal variation promotes behavioral and physiological changes per see.
https://repositorio.unifesp.br/handle/11600/61457
... After initial saccade detection, fixations of <100 ms were discarded and not used in any subsequent analysis (Ouerhani et al., 2004;Mueller et al., 2008;Andersen et al., 2012). In addition to standard metrics associated with fixations (e.g., duration), each fixation was assigned a virtual environment object label using the following approach. ...
Eye tracking has been an essential tool within the vision science community for many years. However, the majority of studies involving eye-tracking technology employ a relatively passive approach through the use of static imagery, prescribed motion, or video stimuli. This is in contrast to our everyday interaction with the natural world where we navigate our environment while actively seeking and using task-relevant visual information. For this reason, an increasing number of vision researchers are employing virtual environment platforms, which offer interactive, realistic visual environments while maintaining a substantial level of experimental control. Here, we recorded eye movement behavior while subjects freely navigated through a rich, open-world virtual environment. Within this environment, subjects completed a visual search task where they were asked to find and count occurrence of specific targets among numerous distractor items. We assigned each participant into one of four target conditions: Humvees, motorcycles, aircraft, or furniture. Our results show a statistically significant relationship between gaze behavior and target objects across Target Conditions with increased visual attention toward assigned targets. Specifically, we see an increase in the number of fixations and an increase in dwell time on target relative to distractor objects. In addition, we included a divided attention task to investigate how search changed with the addition of a secondary task. With increased cognitive load, subjects slowed their speed, decreased gaze on objects, and increased the number of objects scanned in the environment. Overall, our results confirm previous findings and support that complex virtual environments can be used for active visual search experimentation, maintaining a high level of precision in the quantification of gaze information and visual attention. This study contributes to our understanding of how individuals search for information in a naturalistic (open-world) virtual environment. Likewise, our paradigm provides an intriguing look into the heterogeneity of individual behaviors when completing an un-timed visual search task while actively navigating.
... Many wayfinding tests in VR are specifically designed to be sensitive to hippocampal function, thus requiring subjects to find a hidden platform (virtual water maze) or location, using external environmental cues (Moffat and Resnick, 2002;Morganti et al., 2008Morganti et al., , 2013Davis et al., 2017). Many studies have shown that VR can be used to test spatial navigation, especially using specific maze tasks, in older adults with and without dementia (Moffat and Resnick, 2002;Shukitt-Hale et al., 2004;Vembar et al., 2004;Mueller et al., 2008;Jheng and Pai, 2009;Schoenfeld et al., 2010;Tangen et al., 2015;Parizkova et al., 2018). Using VR for spatial navigation testing has led to a rich understanding of strategies and abilities in a wide range of age groups, among individuals with varying types of abilities (Diersch and Wolbers, 2019). ...
Aim: To examine the feasibility of using large scale spatial, self-mobile, virtual reality, and eye tracking in older adults with and without Alzheimer's disease (AD).
Methods: Older adults with early stage AD ( n = 38) and a control group without AD ( n = 50) were asked to find their way in a large, projected VR simulation of a retirement community repeatedly over 10 trials for each of 2 days, while wearing eye tracking glasses. Feasibility measures, including tolerance, side effects, and ability to complete the VR and eye tracking were collected. This study reports the analysis of the feasibility data for the VR and eye tracking and comparison of findings between the groups.
Results: Over 80% of the subjects were able to complete the VR portion of the study. Only four subjects, all in the AD group, could not use the joystick and were excluded. Withdrawal rate (18%) was similar between the groups [ X ² (2) = 2.82, N = 88, p = 0.245] with most withdrawals occurring after the fourth trial. Simulation sickness was not significantly different between the groups. Only 60% of the subjects had completed eye tracking videos; more subjects in the AD group had complete eye tracking videos than the control group; X ² (1) = 7.411, N = 88, p = 0.006. Eye tracking incompletion was primarily due to inability to calibration issues.
Conclusion: Virtual reality testing and eye tracking can be used in older adults with and without AD in a large-scale way-finding task, but that there are some limitations.
... Particularly, this ITI should be taken when the participant has located the goal, causing them to become stationary but capable of examining the environment visually (as rats would do in the MWM). The reason shorter ITI times are used may be due to human participants possessing full control over the task (see Mueller et al. 2008). Some participants may rush to get to the next trial, whereas others may wish to have extended breaks (particularly if using VR gear). ...
The ability to accurately recall locations and navigate our environment relies on multiple cognitive mechanisms. The behavioural and neural correlates of spatial navigation have been repeatedly examined using different types of mazes and tasks with animals. Accurate performances of many of these tasks have proven to depend on specific circuits and brain structures and some have become the standard test of memory in many disease models. With the introduction of virtual reality (VR) to neuroscience research, VR tasks have become a popular method of examining human spatial memory and navigation. However, the types of VR tasks used to examine navigation across laboratories appears to greatly differ, from open arena mazes and virtual towns to driving simulators. Here, we examined over 200 VR navigation papers, and found that the most popular task used is the virtual analogue of the Morris water maze (VWM). Although we highlight the many advantages of using the VWM task, there are also some major difficulties related to the widespread use of this behavioural method. Despite the task's popularity, we demonstrate an inconsistency of use - particularly with respect to the environmental setup and procedures. Using different versions of the virtual water maze makes replication of findings and comparison of results across researchers very difficult. We suggest the need for protocol and design standardisation, alongside other difficulties that need to be addressed, if the virtual water maze is to become the 'gold standard' for human spatial research similar to its animal counterpart.
... Firstly, there are studies that demonstrated that eye movements act as a function of culture (Chua et al., 2005;Kelly et al., 2010). Secondly, gender is also believed to influence eye movement strategies (Mueller et al., 2008). Another study showed that eye movements differ by gender only in the adult phase (Miyahira et al., 2000). ...
... These experimental tasks provide rich data sets with a wide range of measures that allow assessing the precision with which spatial locations can be memorized, such as distances and angular differences between the estimated position of the target or own location and the correct locations, time spent searching in the vicinity of the correct location, and path trajectories amongst others. These tasks have also been used to investigate spatial encoding strategies (e.g., Mueller et al., 2008) and reference frames (e.g., King et al., 2002;King et al., 2004) used during navigation as well as effects of (a)typical aging on spatial navigation (e.g., Moffat et al., 2007). More recently, the vMWM has also been applied to investigate the precision of spatial representations in patients with hippocampal lesions (Kolarik et al., 2018;Kolarik et al., 2016). ...
The aim of the current study was to develop a novel task that allows for the quick assessment of spatial memory precision with minimal technical and training requirements. In this task, participants memorized the position of an object in a virtual room and then judged from a different perspective, whether the object has moved to the left or to the right. Results revealed that participants exhibited a systematic bias in their responses that we termed the reversed congruency effect. Specifically, they performed worse when the camera and the object moved in the same direction than when they moved in opposite directions. Notably, participants responded correctly in almost 100% of the incongruent trials, regardless of the distance by which the object was displaced. In Experiment 2, we showed that this effect cannot be explained by the movement of the object on the screen, but that it relates to the perspective shift and the movement of the object in the virtual world. We also showed that the presence of additional objects in the environment reduces the reversed congruency effect such that it no longer predicts performance. In Experiment 3, we showed that the reversed congruency effect is greater in older adults, suggesting that the quality of spatial memory and perspective-taking abilities are critical. Overall, our results suggest that this effect is driven by difficulties in the precise encoding of object locations in the environment and in understanding how perspective shifts affect the projected positions of the objects in the two-dimensional image.
... For instance, research demonstrates large individual differences in accuracy at using distal (stable far away landmarks, such as mountains) or proximal (nearby landmarks, such as a building or street sign) cues. This is shown with variations on the Morris Water Maze task (Daugherty et al., 2015;Mueller, Jackson, & Skelton, 2008;Padilla et al., 2017;Woolley et al., 2010) in which individuals learn a target location in a virtual environment (VE) that contains only distal or only proximal cues, and then must return to the location from a novel position. Successful performance requires accurate encoding of the location relative to the available cues. ...
Individuals differ in preferences to use route versus survey strategies or distal versus proximal cues for navigation. The current study aimed to examine the effects of environmental structure experience in environment representations. Two groups of participants from Salt Lake City (Utah, USA) and Padua (Veneto, Italy) completed a series of navigation tasks in familiar and novel virtual environments as well as navigation strategy questionnaires. The results showed that Padua participants – compared to Utah participants - had more accurate survey knowledge of locations in their city and country, were more accurate at using proximal cues to remember target locations, and were more likely to use navigation strategies that involved shortcuts. Utah participants did not use distal cues more accurately or use more survey-based strategies despite their higher reported sense of direction and cardinal knowledge compared to Padua participants. Overall the results support that environmental demands shape environment strategies and performance.
... Moreover, this replay of eye movements is accompanied by image-specific patterns of brain activity during recall (Bone et al., 2019). Indeed, eye-tracking has been used to investigate strategies in many cognitive domains, such as learning (for a review, see Lai et al., 2013), reading (for a review, see Rayner, 1998), memory (for a review, see Hannula et al., 2010), face recognition (Chaby et al., 2017), and navigation (Mueller et al., 2008;Livingstone-Lee et al., 2011;Andersen et al., 2012). This link between eye movements and cognition extends to the solving of spatial tasks (Thomas and Lleras, 2007). ...
The ability to recognise places is known to deteriorate with advancing age. In this study, we investigated the contribution of age-related changes in spatial encoding strategies to declining place recognition ability. We recorded eye movements while younger and older adults completed a place recognition task first described by Muffato et al. (2019). Participants first learned places, which were defined by an array of four objects, and then decided whether the next place they were shown was the same or different to the one they learned. Places could be shown from the same spatial perspective as during learning or from a shifted perspective (30° or 60°). Places that were different to those during learning were changed either by substituting an object in the place with a novel object or by swapping the locations of two objects. We replicated the findings of Muffato et al. (2019) showing that sensitivity to detect changes in a place declined with advancing age and declined when the spatial perspective was shifted. Additionally, older adults were particularly impaired on trials in which object locations were swapped; however, they were not differentially affected by perspective changes compared to younger adults. During place encoding, older adults produced more fixations and saccades, shorter fixation durations, and spent less time looking at objects compared to younger adults. Further, we present an analysis of gaze chaining, designed to capture spatio-temporal aspects of gaze behaviour. The chaining measure was a significant predictor of place recognition performance. We found significant differences between age groups on the chaining measure and argue that these differences in gaze behaviour are indicative of differences in encoding strategy between age groups. In summary, we report a direct replication of Muffato et al. (2019) and provide evidence for age-related differences in spatial encoding strategies, which are related to place recognition performance.
... Given the reports of sex differences in navigation and spatial cognition (Coutrot et al., 2019;Mueller, Jackson, & Skelton, 2008), we first ran an exploratory analysis focusing on sex, but did not find any performance differences between sexes (see Online Supplementary Materials). As the current study was not designed to investigate sex differences, we did not include sex as a factor in any further analyses. ...
Successful navigation requires memorising and recognising the locations of objects across different perspectives. Although these abilities rely on hippocampal functioning, which is susceptible to degeneration in older adults, little is known about the effects of ageing on encoding and response strategies that are used to recognise spatial configurations. To investigate this, we asked young and older participants to encode the locations of objects in a virtual room shown as a picture on a computer screen. Participants were then shown a second picture of the same room taken from the same (0°) or a different perspective (45° or 135°) and had to judge whether the objects occupied the same or different locations. Overall, older adults had greater difficulty with the task than younger adults although the introduction of a perspective shift between encoding and testing impaired performance in both age groups. Diffusion modelling revealed that older adults adopted a more conservative response strategy, while the analysis of gaze patterns showed an age-related shift in visual-encoding strategies with older adults attending to more information when memorising the positions of objects in space. Overall, results suggest that ageing is associated with declines in spatial processing abilities, with older individuals shifting towards a more conservative decision style and relying more on encoding target object positions using room-based cues compared to younger adults, who focus more on encoding the spatial relationships among object clusters.
... Our findings are in line with previous experimental studies that reported gender differences in visual-spatial skills-i.e., it has been observed that boys/men exhibit better spatial perception, spatial visualization, mental turn, and spatial-temporal performance than girls/women, as well as fewer errors when searching for and interpreting stimuli from the environment [48][49][50]. Moreover, during a visual-spatial task, women prefer strategies based on memory function, while men analyze current spatial relationships [51]. Interestingly, it has been reported that sport activity may reduce gender differences in visual-spatial ability. ...
Reactive agility (RA) directly refers to athletes’ visuomotor processing of the specific conditions for team sports. The aim of the study was to identify the factors among age, gender, sport discipline, time participation in a sports activity, reaction time, and visual field which could have an impact on visual-motor performance in RA tasks in young, competitive team sports players. The study included boys (n = 149) and girls (n = 157) aged 13–15 participating in basketball, volleyball and handball. Anthropometric measurements were carried out, and the Peripheral Perception (PP) test was used to evaluate the visual-motor performance under laboratory conditions. The Five-Time Shuttle Run to Gates test was used to determine the RA. A multiple regression analysis was performed to identify the relationships between the visual-motor performance in an RA task (dependent variable) and the remaining independent variables (continuous and categorical). The findings of the current study indicate that the main predictive factors of visual-motor performance in RA among young athletes are gender (ß = −0.46, p < 0.000) and age (ß = −0.30, p < 0.000). Moreover, peripheral perception positively affected the achievements in the RA task in boys (ß = −0.25, p = 0.020). The sport discipline does not differentiate the visual-motor performance in RA in team sports players in the puberty period.
... In the MWM, male subjects were more efficient than females to navigate to the platform position (which may be an indicator better allocentric processes in males). However, females and males spent the same amount of time gazing at the "allocentric space", which the authors define as encompassing distal landmarks and the room unpolarized walls (Mueller et al., 2008). In the more complex maze, males and females selected allocentric and egocentric strategy in the same proportion, but the pattern of eye fixations was more clustered on landmarks in females (Andersen et al., 2012). ...
This doctoral work investigates the behavioural consequences of visual and cognitive aging within a spatial cognition framework. The key novelty is the use of ecological environments, which allows to test navigation in natural, yet controlled conditions. We show an age-related spatial reorganization, in which mental representations are preferentially anchored on geometric cues in older adults and on landmarks in young adults. Recording of gaze and body dynamics allowed us to identify oculomotor signatures that were predictive of the spatial coding preference. These results challenge the current view that focuses on egocentric versus allocentric strategies in aging, and question whether it is a matter of availability of spatial cues rather than strategy per se. We verified this prediction by showing that older adults, similarly to children, are as efficient as young adults at using allocentric strategies, given that their preferred cue (i.e. geometry) is available at the time of navigation decision. This work highlights the need for enriching the allocentric/egocentric framework, classically used to interpret the impact of healthy aging on spatial cognition, by integrating the spatial cue processing dimension. This enriched framework may be useful to develop innovative solutions to prevent/counteract the autonomy loss induced by visuo-spatial cognitive aging.
... Finally, it may be worthwhile to consider the effects of gender in learning with realistic visualizations in future studies. Previous research suggests gender-dependent differences in the cognitive processing of spatial stimuli (Mueller, Jackson, & Skelton, 2008; see Castro-Alonso, Wong, Adesope, Ayres, & Paas, 2019, for a metaanalysis in the context of learning media) and may thus be a factor to be considered when applying our results to the design of visualizations. ...
Realistic graphics have been found to result in a worse learning performance than schematic visualizations. These negative results of realism are usually explained by assuming that realistic visualizations induce more cognitive load. Therefore, realism has been linked to the facet of extraneous cognitive load in the model of cognitive load theory. However, recent results indicate that realism may be used to good advantage when it is utilized only in selected parts of visualizations. We tested the hypothesis that realism can be used as a form of signaling. In an experimental study (N = 50), we presented each participant with one realistic and one schematic anatomical visualization. As predicted, retention performance was highest for the realistic components. Furthermore, we found that extraneous load ratings were overall higher when a particularly detailed visualization was shown to participants (although no main effect of realism was found). Our results can be explained in terms of the disfluency effect, which holds that more visually demanding learning materials can in some cases enhance learning. An important implication of the study is that educators may use the degree of realism to focus learners' attention.
... A spatial navigation study examining sex differences (Mueller et al. 2008) employed eye-tracking and pupillometry. Twenty-four participants (M = 24, Male = 12) played The Arena Maze. ...
The subjective experience of emotions which is usually caused by a specific stimulus either real or virtual typically gets accompanied by physiological and behavioral changes in the body. An essential part of the gaming experience is formed by emotional responses, as physiological responses are not affected by subjective tendencies, considering those physiological responses may contribute to the insights of game user research. In this chapter, the origins of psychophysiological responses will be examined on a theoretical basis along with a review of the psychophysiological measurement methods previously employed in the game studies. We will explore their relationship with emotional responses as well as their correspondence with the self-report based evaluations.
... There are also several studies discussing sex differences in eye-tracking indices during the performance of wayfinding tasks; visual attention on landmarks has been discussed substantially, and other eye-tracking indices also show differences between males and females. For example, Sven [30] found sex differences in fixation duration and pupil diameter when performing a spatial navigation task. Valentina [31] found that when planning a route, the number of fixations and fixation ratio between males and females is significantly different, but these studies are limited to pure eyetracking data without considering semantic information on the environment. ...
Spatial orientation is an important task in human wayfinding. Existing research indicates sex-related similarities and differences in performance and strategies when executing spatial orientation behaviors, but few studies have investigated the similarities and differences in visual behaviors between males and females. To address this research gap, we explored visual behavior similarities and differences between males and females using an eye-tracking method. We recruited 40 participants to perform spatial orientation tasks in a desktop environment and recorded their eye-tracking data during these tasks. The results indicate that there are no significant differences between sexes in efficiency and accuracy of spatial orientation. In terms of visual behaviors, we found that males fixated significantly longer than females on roads. Males and females had similar fixation counts in building, signpost, map, and other objects. Males and females performed similarly in fixation duration for all five classes. Moreover, fixation duration was well fitted to an exponential function for both males and females. The base of the exponential function fitted by males’ fixation duration was significantly lower than that of females, and the coefficient difference of exponential function was not found. Females were more effective in switching from maps to signposts, but differences of switches from map to other classes were not found. The newfound similarities and differences between males and females in visual behavior may aid in the design of better human-centered outdoor navigation applications.
... The maze adopted was of medium difficulty, and the addition of a concurrent secondary task would undoubtedly require a heavy load of working memory. That's probably why male participants performed faster than female participants in Experiment 1. Besides, different ways of exploring (Lin et al., 2012;Mueller, Jackson, & Skelton, 2008) and wayfinding strates (Lawton & Janos, 2002;Sandstrom, Kaufman, & Huettel, 1998;Lawton, 1996;Chen, Chang, & Chang, 2009;Nowak et al., 2015) could also be possible attributions. As mentioned above, operations in the maze was programmed in a egocentric perspective, while the general layout map offered an allocentric perspective. ...
The role of phonological loop and visuospatial sketchpad in wayfinding was tested in a virtual maze employing dual-task paradigm through two experiments. Experiment 1 was designed to detect the effects of experimental conditions (control/phonological dual-task/visuospatial dual-task group) and gender on wayfinding performance. The results showed that it took more time in dual-task groups than the control group to complete the primary task, and male participants travelled significantly faster than female participants. The differences between dual-task and control group mainly came from the female participants, while the male participants were relatively not affected by the secondary tasks. In order to further explore the effect of dual-task conditions on male participants, Experiment 2 was conducted to explore how the two dual-task conditions might affect male participants’ wayfinding as the difficulty of secondary tasks increased. It was found that wayfinding performance in dual-task groups was significantly deteriorated in comparison with the control group. There was also no significant difference between the two types of dual tasks. Our study demonstrated that both the phonological loop and the visuospatial sketchpad mattered in wayfinding and to break the advantage in men required more challenging secondary tasks.
... Analyses of pupil size have been used to study navigation strategies as well as measures of fixations and gaze position, providing information about the allocation of perceptual attention and integration of information (Condappa & Wiener, 2014;Mueller et al., 2008; for a review see Kiefer et al., 2017). Interestingly, in the past, research was restricted to laboratories, but recent technologies provide now the possibility to study spatial navigation in real situations and in real time (Kiefer et al., 2014;Wenczel et al.,2017). ...
Eye tracking (ET) studies are becoming increasingly popular due to rapid methodological and technological advances as well as the development of cost efficient and portable eye trackers. Although historically ET has been mostly employed in psychophysics or developmental cognition studies, there is also promising scope to use ET for movement disorders and measuring cognitive processes in neurodegeneration. Particularly, ET can be a powerful tool for cognitive and neuropsychological assessments of patients with pathologies affecting motor and verbal abilities, as tasks can be adapted without requiring motor (except eye movements) or verbal responses. In this review, we will examine the existing evidence of ET methods in neurodegenerative conditions and its potential clinical impact for cognitive assessment. We highlight that current evidence for ET is mostly focused on diagnostics of cognitive impairments in neurodegenerative disorders, where it is debatable whether it has any more sensitivity or specificity than existing cognitive assessments. By contrast, there is currently a lack of ET studies in more advanced disease stages, when patients’ motor and verbal functions can be significantly affected, and standard cognitive assessments are challenging or often not possible. We conclude that ET is a promising method not only for cognitive diagnostics but more importantly, for potential cognitive disease tracking in progressive neurodegenerative conditions.
There are various styles to visually represent relational data in the form of node-link diagrams. In particular, for planar graphs we can find orthogonal node-link diagrams consisting of links bending only at ninety degrees a successful and prominent variant. One of the benefits of such drawings is the tracking of longer paths through a network with the eyes due to their limited number of link orientations, changes, and variations, but on the negative side the links can have arbitrary bending shapes. In this article we developed a novel way to visualize such orthogonal planar drawings by making use of mazes that look more natural to the human eye due to the street-like visual metaphor that many people are familiar with. Tracking paths is one of the major tasks in such graph visualizations, similar to orthogonal node-link diagrams, however, we argue that mazes are a more natural way to find paths. To get insights in the visual scanning behavior when reading graph mazes we conducted a comparative eye tracking study with 26 male versus female participants of different experience levels while also alternating between orthogonal node-link drawings and graph mazes as well as different graph size levels. The major result of this comparative study is that the participants can track paths in both representation styles, including a geodesic path tendency in their visual search behavior, but typically have a longer fixation duration at branching nodes and locations in the mazes that lead to opposite directions to the geodesic path tendency, maybe the viewers had to start a reorientation phase in their visual scanning behavior. We also found out that the size, that is the number of graph vertices has an impact on the visual scanning behavior for both orthogonal node-link diagrams as well as street-like maze representations, but for the mazes we found this impact to be less strong (in terms of the eye movement data metrics fixation durations and saccade lengths) compared to the node-link diagrams. To conclude the article, we discuss limitations and scalability issues of our approach. Moreover, we give an outlook and future work for possible extensions.
A psychophysiological eye-tracking experiment was conducted to investigate map navigation (including tasks of familiarization with the map, route memorization and recall of the route) as a function of landmark type (no landmarks, object landmarks, verbal landmarks, both types of landmarks) and gender. It was found that the number of fixations decreased and the fixation duration increased during recall compared to map familiarization and route memorization. The effect of task type on eye movements varied between route proximity areas. In the proximal areas, the number of fixations decreased during recall in comparison to memorization, while in the distal (“off route”) area the effect was opposite. These changes were pronounced in the male participants and did not reach statistical significance in the females. Men switched more frequently from the proximal to the distal area during memorization, thus forming a connection between the route and the remote environment. In contrast, women’s gaze was more often directed to the distal area during route recall. The map with object landmarks differed from the other maps in the lower number and longer fixation duration, indicating the advantage of object landmarks and, possibly, a preference for the object visual cognitive style among our participants.
Zukünftig können Fahrzeuge automatisiert fahren und Reisende die Fahrzeit zum Video schauen nutzen. Adrian Brietzke untersucht dabei realitätsnah das Auftreten von Kinetose (Reisekrankheit) bei den Reisenden. Es wird ein ganzheitlicher Ansatz beginnend bei der Identifikation der Betroffenengruppe über die empirische Analyse der Ist-Situation im Stop-and-Go-Verkehr bis zu einer Entwicklung und Bewertung von technischen Maßnahmen vorgestellt. Systematisch erhobene Nutzererfahrungen werden ausgewertet sowie das akute Auftreten von Kinetose in Realfahrstudien untersucht. Die gewonnenen Erkenntnisse erlauben eine Überprüfung der Ursachen für Kinetose und liefern Ansätze zur Weiteerentwicklung von Maßnahmen zur Reduzierung von Kinetose.
Background
Major depressive disorder (MDD) is associated with impairments in spatial learning and memory and with altered functioning of central mineralocorticoid receptors (MR) and glutamatergic N-methyl-D-aspartate receptors (NMDA-R). Both receptors are highly expressed in the hippocampus and prefrontal cortex – brain areas that are critical for spatial learning and memory. Here, we examined the effects of separate and combined MR and NMDA-R stimulation on spatial learning and memory in individuals with MDD and healthy controls.
Methods
We used a randomized, double-blind, placebo-controlled between-group study design to examine the effects of separate and combined stimulation of the MR (with 0.4 mg fludrocortisone) and NMDA-R (with 250 mg D-cycloserine) in 116 unmedicated individuals with MDD (mean age: 34.7 ± 13.3 years; 78.4% women) and 116 age-, sex-, and education-matched healthy controls. Participants were randomly assigned to one of four conditions: 1) placebo; 2) MR stimulation; 3) NMDA-R stimulation; and 4) combined MR/NMDA-R stimulation. Three hours after drug administration, spatial learning and memory were assessed using a virtual Morris Water Maze task.
Results
Individuals with MDD and healthy controls did not differ in spatial learning and memory performance. Neither separate nor combined MR or NMDA-R stimulation altered measures of spatial performance.
Conclusion
In this study of relatively young, predominantly female, and unmedicated individuals, we found no effect of MDD and no effect of separate or combined MR and NMDA-R stimulation on spatial learning and memory.
Vision provides the most important sensory information for spatial navigation. Recent technical advances allow new options to conduct more naturalistic experiments in virtual reality (VR) while additionally gather data of the viewing behavior with eye tracking investigations. Here, we propose a method that allows to quantify characteristics of visual behavior by using graph-theoretical measures to abstract eye tracking data recorded in a 3D virtual urban environment.
The analysis is based on eye tracking data of 20 participants, who freely explored the virtual city Seahaven for 90 minutes with an immersive VR headset with an inbuild eye tracker. To extract what participants looked at, we defined “gaze” events, from which we created gaze graphs. On these, we applied graph-theoretical measures to reveal the underlying structure of visual attention.
Applying graph partitioning, we found that our virtual environment could be treated as one coherent city. To investigate the importance of houses in the city, we applied the node degree centrality measure. Our results revealed that 10 houses had a node degree that exceeded consistently two-sigma distance from the mean node degree of all other houses. The importance of these houses was supported by the hierarchy index, which showed a clear hierarchical structure of the gaze graphs. As these high node degree houses fulfilled several characteristics of landmarks, we named them “gaze-graph-defined landmarks”. Applying the rich club coefficient, we found that these gaze-graph-defined landmarks were preferentially connected to each other and that participants spend the majority of their experiment time in areas where at least two of those houses were visible.
Our findings do not only provide new experimental evidence for the development of spatial knowledge, but also establish a new methodology to identify and assess the function of landmarks in spatial navigation based on eye tracking data.
Author Summary
The ability to navigate and orient ourselves in an unknown environment is important in everyday life. To better understand how we are able to learn about a new environment, it is important to study our behavior during the process of spatial navigation. New technical advances allow us to conduct studies in naturalistic virtual environments with participants wearing immersive VR-headsets. In addition, we can use eye trackers to observe the participant’s eye movements. This is interesting, because observing eye movements allows us to observe visual attention and therefore important cognitive processes. But, it can be difficult to analyze eye tracking data that was measured in a VR environment, as there is no established algorithm yet. Therefore, we propose a new method to analyze such eye tracking data. In addition, our method allows us to transform the eye tracking data into graphs which we can use to find new patterns in behavior that were not accessible before. Using this methodology, we found that participants who spend 90 min exploring a new virtual town used some houses as orientation anchors which we call gaze-graph-defined landmarks. Our further analysis revealed also new characteristics of those houses that were not yet associated with landmarks.
The development of effective system of path searching in space is becoming an important task for organizations that render consumer services, and the use of neuromarketing, eye tracking in particular, makes it possible to solve it. The article reviews the approaches of scientists regarding navigation systems research; the research results using the method of trajectory tracking and eye movement for improvement of the search of necessary objects by service consumers in healthcare institution are outlined. The method testing proved to be successful. The use of eye tracking in the marketing research boosted the quality and made it possible to perform necessary improvement with clear proof of its effect, as well as demonstrate the significance of eye tracking use for improvement of the spatial consumer orientation.
Adults, ranging in age from 20 to 78 years, were required to give directions to a hypothetical stranger while looking at a map. The direction giving was scored according to the strategies employed by the direction giver (e.g., landmarks, relational turns, road names, and cardinality). The "map present" direction-giving paradigm was employed to reduce the influence of memory. The results suggest that the aging decline in spatial abilities does not influence direction-giving strategies when memory demands are minimal. Older adults are as proficient as young adults when employing direction-giving strategies. Middle-age females employed a significantly higher frequency of strategies relative to young males, young females, middle-age males, and older females. When accuracy was examined, gender-related differences favoring males were obtained for the relational strategy.
What attracts attention when we inspect a scene? Two experiments recorded eye movements while viewers inspected pictures of natural office scenes in which two objects of interest were placed. One object had low contour density and uniform colouring (a piece of fruit), relative to another that was visually complex (for example, coffee mugs and commercial packages). In each picture the visually complex object had the highest visual saliency according to the Itti and Koch algorithm. Two experiments modified the task while the pictures were inspected, to determine whether visual saliency is invariably dominant in determining the pattern of fixations, or whether the purpose of inspection can provide a cognitive override that renders saliency secondary. In the first experiment viewers inspected the scene in preparation for a memory task, and the more complex objects were potent in attracting early fixations, in support of a saliency map model of scene inspection. In the second experiment viewers were set the task of detecting the presence of a low saliency target, and the effect of a high saliency distractor was negligible, supporting a model in which the saliency map can be built with cognitive influences that override low-level visual features.
The generality of the place-learning impairment associated with hippocampal system damage was challenged using methods of training that permitted subjects to form an individual association between the place of escape and a particular navigational route in an open-field water maze. Both normal rats and rats with fornix lesions (FX rats) acquired this task rapidly, although FX rats were slightly slower in achieving minimum escape latencies. In postcriterion testing, FX rats occasionally made near misses but, more often, their escape performance was indistinguishable from that of intact rats. Results from a variety of probe tests indicated that FX rats, like normal rats, had based their performance on a representation of multiple distal cues but their representation, unlike that of normal rats, was inflexible in that it could not be used to guide performance when the cues or starting position were altered. These results parallel those from other studies of hippocampal function in animals and humans: The learning deficit consequent to hippocampal system damage (1) is not specific to a particular category of learning materials, but is dependent on the representational demands of the task; (2) is observed when task demands encourage a representation based on relations among multiple cues, but not when the task encourages adaptation to an individual (or compound) stimulus; (3) spares acquisition of fundamental procedures needed to perform the task; and (4) impairs the flexible use of learned information in tests other than repetition of the learning experience.
The generality of the place-learning impairment associated with hippocampal system damage was challenged using methods of training that permitted subjects to form an individual association between the place of escape and a particular navigational route in an open-field water maze. Both normal rats and rats with fornix lesions (FX rats) acquired this task rapidly, although FX rats were slightly slower in achieving minimum escape latencies. In postcriterion testing, FX rats occasionally made near misses but, more often, their escape performance was indistinguishable from that of intact rats. Results from a variety of probe tests indicated that FX rats, like normal rats, had based their performance on a representation of multiple distal cues but their representation, unlike that of normal rats, was inflexible in that it could not be used to guide performance when the cues or starting position were altered. These results parallel those from other studies of hippocampal function in animals and humans: The learning deficit consequent to hippocampal system damage (1) is not specific to a particular category of learning materials, but is dependent on the representational demands of the task; (2) is observed when task demands encourage a representation based on relations among multiple cues, but not when the task encourages adaptation to an individual (or compound) stimulus; (3) spares acquisition of fundamental procedures needed to perform the task; and (4) impairs the flexible use of learned information in tests other than repetition of the learning experience.
In recent years, the magnitude, consistency, and stability across time of cognitive sex differences have been questioned. The present study examined these issues in the context of spatial abilities. A meta-analysis of 286 effect sizes from a variety of spatial ability measures was conducted. Effect sizes were partitioned by the specific test used and by a number of variables related to the experimental procedure in order to achieve homogeneity. Results showed that sex differences are significant in several tests but that some intertest differences exist. Partial support was found for the notion that the magnitude of sex differences has decreased in recent years. Finally, it was found that the age of emergence of sex differences depends on the test used. Results are discussed with regard to their implications for the study of sex differences in spatial abilities.
The space around us is represented not once but many times in parietal cortex. These multiple representations encode locations and objects of interest in several egocentric reference frames. Stimulus representations are transformed from the coordinates of receptor surfaces, such as the retina or the cochlea, into the coordinates of effectors, such as the eye, head, or hand. The transformation is accomplished by dynamic updating of spatial representations in conjunction with voluntary movements. This direct sensory-to-motor coordinate transformation obviates the need for a single representation of space in environmental coordinates. In addition to representing object locations in motoric coordinates, parietal neurons exhibit strong modulation by attention. Both top-down and bottom-up mechanisms of attention contribute to the enhancement of visual responses. The saliance of a stimulus is the primary factor in determining the neural response to it. Although parietal neurons represent objects in motor coordinates, visual responses are independent of the intention to perform specific motor acts.
Visuospatial navigation in animals and human subjects is generally studied using maze exploration. We used functional MRI to observe brain activation in male and female subjects as they searched for the way out of a complex, three-dimensional, virtual-reality maze. Navigation activated the medial occipital gyri, lateral and medial parietal regions, posterior cingulate and parahippocampal gyri as well as the right hippocampus proper. Gender-specific group analysis revealed distinct activation of the left hippocampus in males, whereas females consistently recruited right parietal and right prefrontal cortex. Thus we demonstrate a neural substrate of well established human gender differences in spatial-cognition performance.
Spatial learning and memory has been linked to the hippocampus and temporal lobes and though these areas are often damaged in traumatic brain injury (TBI), spatial learning deficits after TBI have not received much attention. In the present study, a virtual environment was used to challenge people with TBI to solve a task comparable to the Morris water maze, which in turn has been shown to be highly sensitive to hippocampal and frontal lobe dysfunction in laboratory animals. A regular computer monitor was used to present 12 participants with TBI and 12 age- and sex-matched comparison participants with a computer-generated, three-dimensional "virtual arena maze," consisting of a large round arena within a very large square room. Participants were required to learn the place of an invisible target on the floor of the room based solely on distal cues on the walls of the room. Eight of the 12 participants with moderate to severe TBI showed substantial place-learning deficits in comparison to the uninjured participants. Performance in the virtual environment correlated with self-reported frequency of wayfinding problems in everyday life and with scores on a test of episodic memory, the Rivermead Behavioural Memory Task. These data confirm that deficits in spatial learning and memory follow TBI, and suggest that the virtual arena maze may provide a new method for objectively assessing them.
During natural vision, the brain efficiently processes views of the external world as the eyes actively scan the environment. To better understand the neural mechanisms underlying this process, we recorded the activity of individual temporal cortical neurons while monkeys looked for and identified familiar targets embedded in natural scenes. We found a group of visual neurons that exhibited stimulus-selective neuronal bursts just before the monkey's response. Most of these cells showed similar selectivity whether effective targets were viewed in isolation or encountered in the course of exploring complex scenes. In addition, by embedding target stimuli in natural scenes, we could examine the activity of these stimulus-selective cells during visual search and at the time targets were fixated and identified. We found that, during exploration, neuronal activation sometimes began shortly before effective targets were fixated, but only if the target was the goal of the next fixation. Furthermore, we found that the magnitude of this early activation varied inversely with reaction time, indicating that perceptual information was integrated across fixations to facilitate recognition. The behavior of these visually selective cells suggests that they contribute to the process of noticing familiar objects in the real world.
The reported research extends classic findings that after briefly viewing structured, but not random, chess positions, chess masters reproduce these positions much more accurately than less-skilled players. Using a combination of the gaze-contingent window paradigm and the change blindness flicker paradigm, we documented dramatically larger visual spans for experts while processing structured, but not random, chess positions. In addition, in a check-detection task, a minimized 3 x 3 chessboard containing a King and potentially checking pieces was displayed. In this task, experts made fewer fixations per trial than less-skilled players, and had a greater proportion of fixations between individual pieces, rather than on pieces. Our results provide strong evidence for a perceptual encoding advantage for experts attributable to chess experience, rather than to a general perceptual or memory superiority.
The use of virtual environment (VE) technology to assess spatial navigation in humans has become increasingly common and provides an opportunity to quantify age-related deficits in human spatial navigation and promote a comparative approach to the neuroscience of cognitive aging. The purpose of the present study was to assess age differences in navigational behavior in a VE and to examine the relationship between this navigational measure and other more traditional measures of cognitive aging. Following pre-training, participants were confronted with a VE spatial learning task and completed a battery of cognitive tests. The VE consisted of a richly textured series of interconnected hallways, some leading to dead ends and others leading to a designated goal location in the environment. Compared to younger participants, older volunteers took longer to solve each trial, traversed a longer distance, and made significantly more spatial memory errors. After 5 learning trials, 86% of young and 24% of elderly volunteers were able to locate the goal without error. Performance on the VE navigation task was positively correlated with measures of mental rotation and verbal and visual memory.
The recording and analysis of eye movements are fundamental to a diverse set of research applications, including studies in which reading, visual search, and both overt and covert visuospatial attention are examined. Software tools supplied with commonly available eye-tracking equipment have generally been limited in functionality and nonextensible. Because of this dearth of available software, ELAB was created to provide an extensible framework for analyzing various aspects of eye movements. The program consists of a series of open-source MATLAB functions. The program's data structures keep raw data, analysis preferences, and analyzed data separate, thus maintaining data fidelity and promoting extensibility.
It is widely believed that male mammals have better spatial ability than females. A large number of evolutionary hypotheses have been proposed to explain these differences, but few species have been tested. The authors critically review the proposed evolutionary explanations for sex differences in spatial cognition and conclude that most of the hypotheses are either logically flawed or, as yet, have no substantial support. Few of the data exclusively support or exclude any current hypotheses. The hypothesis with the strongest support proposes that range size was the selection pressure that acted to increase spatial ability. The authors suggest ways in which these hypotheses could be tested by presenting explicit predictions and suggesting suitable test species or conditions.
Survivors of traumatic brain injury (TBI) often have spatial navigation deficits. This study examined such deficits and conducted a detailed analysis of navigational behaviour in a virtual environment.
TBI survivors were tested in a computer simulation of the Morris water maze task that required them to find and remember the location of an invisible platform that was always in the same location. A follow-up questionnaire assessed everyday spatial ability.
Fourteen survivors of moderate-to-severe TBI were compared to 12 non-injured participants.
TBI survivors navigated to a visible platform but could not learn the location of the invisible platform. The difference between TBI survivors and uninjured participants was best indicated by two new dependent variables, path efficacy and spatial scores.
This study confirms the capacity of virtual environments to reveal spatial navigation deficits after TBI and establishes the best way to identify such deficits.
Motivated by the fact that previous visual memory paradigms have imposed encoding and retrieval constraints, the present article presents two experiments that address how observers allocate eye movements in memory and comparison processes in the absence of constraints. A comparative visual search design (Pomplun, Sichelschmidt, et al., 2001) was utilized in which observers searched for a difference between two images presented side by side. Robust time course effects were obtained, whereby search was characterized by brief fixations and a high proportion of comparative saccades. Then, upon target detection, fixations were extended, more comparative saccades were elicited, and the search focus was narrowed. The saliency and presence of differences did not guide attention, and detection was contingent upon direct fixation of the targets. The results indicate that, when full control is given, observers adopt a strategy that cuts down on memory usage in favor of restricted encoding and active scanning.
The hippocampus (HC) and associated neural structures are hypothesized to contribute to individual differences in human spatial navigation. However, functional imaging studies and theoretical models underscore the importance of extrahippocampal regions as well. The purpose of the present study was to examine age differences in virtual environment navigation and to assess possible relationships between navigation and structural integrity of hippocampal and extrahippocampal brain regions. Healthy adult volunteers completed a virtual navigation task and underwent magnetic resonance imaging to assess volumes of the caudate nucleus (CN), cerebellum, HC, prefrontal, and primary visual cortices. Results demonstrated robust age-related differences in place learning. Moreover, individual differences in regional brain volumes as well as performance on the tests of memory and executive functions contributed to age differences in human place learning. High performance in a virtual navigation task was associated with larger volume of the CN and prefrontal gray and white matter. Larger hippocampal volume was associated with improved performance in the young but not old participants. We conclude that human navigation requires both hippocampal and extrahippocampal brain systems and draws on executive resources for successful performance.
In two sessions with free scanning and memory instructions, eye-movement patterns from nine artists were compared with those of nine artistically untrained participants viewing 16 pictures representing a selection of categories from ordinary scenes to abstraction: 12 pictures were made to accommodate an object-oriented viewing mode (selection of recognisable objects), and a pictorial viewing mode (selection of more structural features), and 4 were abstract. The artistically untrained participants showed preference for viewing human features and objects, while the artists spent more scanning time on structural/abstract features. A group by session interaction showed a change of viewing strategy in the artists, who viewed more objects and human features in the memory task session. A verbal test of recall memory showed no overall difference in the number of pictures remembered, but the number of correctly remembered pictorial features was significantly higher for artists than for the artistically untrained viewers, irrespective of picture type. No differences in fixation frequencies/durations were found between groups across sessions, but a significant task-dependent-group by session interaction of fixation frequency/duration showed that the artistically untrained participants demonstrated repetition effects in fewer, longer fixations with repeated viewing, while the opposite pattern obtained for the artists.
We investigated the processing effort during visual search and counting tasks using a pupil dilation measure. Search difficulty was manipulated by varying the number of distractors as well as the heterogeneity of the distractors. More difficult visual search resulted in more pupil dilation than did less difficult search. These results confirm a link between effort and increased pupil dilation.
The pupil dilated more during the counting task than during target-absent search, even though the displays were identical, and the two tasks were matched for reaction time. The moment-to-moment dilation pattern during search suggests little effort in the early stages, but increasingly more effort towards response, whereas the counting task involved an increased initial effort, which was sustained throughout the trial.
These patterns can be interpreted in terms of the differential memory load for item locations in each task. In an additional experiment, increasing the spatial memory requirements of the search evoked a corresponding increase in pupil dilation. These results support the view that search tasks involve some, but limited, memory for item locations, and the effort associated with this memory load increases during the trials. In contrast, counting involves a heavy locational memory component from the start.
Previous work has shown that eye movement behaviour is affected by previous experience, such that alterations in viewing patterns can be observed to previously viewed compared to novel displays. The current work addresses the extent to which such effects of memory on eye movement behaviour are obligatory; that is, we examined whether prior experience could alter subsequent eye movement behaviour under a variety of testing conditions, for stimuli that varied on the nature of the prior exposure. While task demands influenced whether viewing was predominantly directed to the novel versus familiar faces, viewing of the familiar faces was distinguished from viewing of the novel faces, regardless of whether the task required incidental encoding or intentional retrieval. Changes in scanning of previously viewed over novel faces emerged early in viewing; in particular, viewing duration of the first fixation to the familiar faces was often significantly different from the duration of the first fixation directed to the novel faces, regardless of whether prior exposure was solely in the context of the experiment or due to real-world exposure. These findings suggest that representations maintained in memory may be retrieved and compared with presented information obligatorily.
We present evidence for an evolved sexually dimorphic adaptation that activates spatial memory and navigation skills in response to fruits, vegetables and other traditionally gatherable sessile food resources. In spite of extensive evidence for a male advantage on a wide variety of navigational tasks, we demonstrate that a simple but ecologically important shift in content can reverse this sex difference. This effect is predicted by and consistent with the theory that a sexual division in ancestral foraging labour selected for gathering-specific spatial mechanisms, some of which are sexually differentiated. The hypothesis that gathering-specific spatial adaptations exist in the human mind is further supported by our finding that spatial memory is preferentially engaged for resources with higher nutritional quality (e.g. caloric density). This result strongly suggests that the underlying mechanisms evolved in part as adaptations for efficient foraging. Together, these results demonstrate that human spatial cognition is content sensitive, domain specific and designed by natural selection to mesh with important regularities of the ancestral world.
The strongest sex differences on any cognitive task, favoring men, are found for tasks that require the mental rotation of three-dimensional objects. A number of studies have explored functional brain activation during mental rotation tasks, and sex differences have been noted in some. However, in these studies there was a substantial confounding factor because male and female subjects differed in overall performance levels. In contrast, our functional brain activation study examined cortical activation patterns for males and females who did not differ in overall level of performance on three mental rotation tasks. This allowed us to eliminate any confounding influences of overall performance levels. Women exhibited significant bilateral activations in the intraparietal sulcus (IPS) and the superior and inferior parietal lobule, as well as in the inferior temporal gyrus (ITG) and the premotor areas. Men showed significant activation in the right parieto-occitpital sulcus (POS), the left intraparietal sulcus and the left superior parietal lobule (SPL). Both men and women showed activation of the premotor areas but men also showed an additional significant activation of the left motor cortex. No significant activation was found in the inferior temporal gyrus. Our results suggest that there are genuine between-sex differences in cerebral activation patterns during mental rotation activities even when performances are similar. Such differences suggest that the sexes use different strategies in solving mental rotation tasks.
In a study of sex differences in navigation strategy and geographic knowledge, 90 men and 104 women completed cognitive spatial tests, gave directions from local maps, and identified places on a world map. On the spatial tests, men were better than women in mental rotation skill, but men and women were similar in object location memory. In giving directions, men were more abstract and Euclidian, using miles and north–south– east–west terms, whereas women were more concrete and personal, using landmarks and left–right terms. Older subjects of both sexes gave more abstract Euclidian direc-tions than younger subjects did. On the world map, men identified more places than women did. The data fit a causal model in which sex predicts world map knowledge and the use of Euclidian directions, both directly and indirectly through a sex difference in spatial skills. The age effect, which was independent of sex, supports a developmental view of spatial cognition. © 1998 Elsevier Science Inc. his article begins with two findings about the spatial activities and skills of men and women. The first is that women use more landmarks than men do in navigating (Choi and Silverman 1997). The second is that men know more than women do about world geography (Liben 1995). These findings are puzzling, and understanding them may help us understand other differ-ences between men and women. In this article we examine factors that may relate to both navigation strategy and geographic knowledge.
This study used computer-generated (“virtual”) mazes to investigate sex differences in the efficiency of spatial route learning in humans. Correlations between maze performance and traditional psychometric measures of spatial ability also were examined. Male and female subjects completed a total of five learning trials on each of two spatial mazes and completed a battery of spatial and verbal cognitive tests. As well as demonstrating the typical male advantage on psychometric measures of spatial performance, robust sex differences favoring males were found for both the time required to solve the mazes (d = 1.59) and the number of spatial memory errors committed ( d = 1.40). Highly significant positive correlations were obtained between scores on the paper and pencil tests and performance on the maze task. The results of the present study are consistent with results from studies in other mammalian species suggesting a male advantage for spatial navigation through a novel environment.
The current study used a variety of behavioral and self-report measures to examine gender differences in way finding behavior following incidental leaming in an unfamiliar indoor environment. College students were led along a circuitous route and then asked to find their way back to the start. Choice of route back, comments made during the task, directional accuracy, and self-report after the task indicated three relatively efficient patterns of way finding and a distinctly inefficient pattern, characterized by frequent back-trailing, poor directional accuracy, and uncertainty about the task. Gender differences showed a discrepancy between measures of way finding efficiency and directional accuracy. Although there was no gender difference in choice of route back, men were significantly more accurate than women in locating the direction of the destination. Also, women reported significantly more uncertainty about carrying out the task.
The use of augmented reality (AR) in formal education could prove a key component in future learning environments that are richly populated with a blend of hardware and software applications. However, relatively little is known about the potential of ...
Major questions remain about the exact role of hormones in cognition. Furthermore, the extent to which early perturbation in steroid function affects human brain development continues to be a wide open area of research. Congenital adrenal hyperplasia (CAH), a genetic disorder of steroid dysfunction characterized in part by in utero over-production of testosterone, was used as a natural model for addressing this question. Here, CAH (n=54, mean age=17.53, 31 female) patients were compared to healthy age- and sex-matched individuals (n=55, mean age=19.02, 22 female) on a virtual equivalent of the Morris Water Maze task [Morris, R., 1984. Developments of a water-maze procedure for studying spatial learning in the rat. J. Neurosci. Methods 11, 47-60], an established measure of sex differences in spatial cognition in rodents. Findings revealed that females with CAH with the most severe form of the disease and expected highest level of in utero exposure to androgens were found to perform similarly to both healthy males and CAH males, whereas strong sex differences were apparent in milder forms of the disorder and in controls. Moreover, advanced bone age, an indicator of long-term childhood exposure to testosterone was correlated with improved performance. The results indicate that individuals exposed to both excess androgens prenatally and prolonged exposure during childhood may manifest long-lasting changes in cognitive function. Such finding suggests a pivotal role of hormonal function on brain development in humans, mirroring results from the animal literature.
In a combined behavioral and morphometric study, we observed a testosterone-related sexual dimorphism in the granule cell layer of the hippocampus of Sprague-Dawley rats that appears to be related to sex differences in spatial performance. This cell layer was larger and laterally asymmetrical in males. Neonatal testosterone treatment of females resulted in a more male-like hippocampus. These treated females also performed better than controls, and as well as males, on a spatial navigation task. In addition, a strong correlation between the size of granule cell layers and maze performance was detected.
Sex differences in spatial ability are widely acknowledged, yet considerable dispute surrounds the magnitude, nature, and age of first occurrence of these differences. This article focuses on 3 questions about sex differences in spatial ability: What is the magnitude of sex differences in spatial ability? On which aspects of spatial ability are sex differences found? and When, in the life span, are sex differences in spatial ability first detected? Implications for clarifying the linkage between sex differences in spatial ability and other differences between males and females are discussed. We use meta-analysis, a method for synthesizing empirical studies, to investigate these questions. Results of the meta-analysis suggest that sex differences arise on some types of spatial ability but not others, that large sex differences are found only on measures of mental rotation, that smaller sex differences are found on measures of spatial perception, and that, when sex differences are found, they can be detected across the life span.
During a short-term memory task, pupil diameter is a measure of the amount of material which is under active processing at any time. The pupil dilates as the material is presented and constricts during report. The rate of change of these functions is related to task difficulty.
Developments of an open-field water-maze procedure in which rats learn to escape from opaque water onto a hidden platform are described. These include a procedure (A) for automatically tracking the spatial location of a hooded rat without the use of attached light-emitting diodes; (B) for studying different aspects of spatial memory (e.g. working memory); and (C) for studying non-spatial discrimination learning. The speed with which rats learn these tasks suggests that they may lend themselves to a variety of behavioural investigations, including pharmacological work and studies of cerebral function.
In this study 12 depressed outpatients were compared to 12 healthy controls with respect to their performance on a number of cognitive tasks, including a recognition-memory task, and their eye movements and pupil size were recorded while watching a traffic film. The recognition-memory task consisted of words with intermediate hedonic tone (neutral words), words with high hedonic tone ('good' words) and words with low hedonic tone ('bad' words). Patients performed slower on perceptual-motor tasks which could be characterized as effort-demanding, while no difference between groups was found on effortless tasks. In addition, the range of horizontal eye movements, an indication of visual span, was found to be less in patients. Signal-detection analysis on the recognition-memory data showed an impairment of 'pure' memory in depressives. Analyses on response bias indicated that patients had more false alarms, but only with respect to good words. It is concluded that patients exhibit cognitive deficits, including memory impairment, a narrower visual span and a risky response strategy on good words, which may be an indication of the trouble patients have in processing emotionally toned words.
The study of navigational ability in humans is often limited by the restricted availability and inconvenience of using large novel environments. In the present study we use a computer-generated virtual environment to study sex differences in human spatial navigation. Adult male and female participants navigated through a virtual water maze where both landmarks and room geometry were available as distal cues. Manipulation of environmental characteristics revealed that females rely predominantly on landmark information, while males more readily use both landmark and geometric information. We discuss these results as a possible link between recent human research reporting hippocampal activation in spatial tasks and animal work showing sex differences in both spatial ability and hippocampal development.
In many mammalian species, it is known that males and females differ in place learning ability. The performance by men and women is commonly reported to also differ, despite a large amount of variability and ambiguity in measuring spatial abilities. In the non-human literature, the gold standard for measuring place learning ability in mammals is the Morris water task. This task requires subjects to use the spatial arrangement of cues outside of a circular pool to swim to a hidden goal platform located in a fixed location. We used a computerized version of the Morris water task to assess whether this task will generalize into the human domain and to examine whether sex differences exist in this domain of topographical learning and memory. Across three separate experiments, varying in attempts to maximize spatial performance, we consistently found males navigate to the hidden platform better than females across a variety of measures. The effect sizes of these differences are some of the largest ever reported and are robust and replicable across experiments. These results are the first to demonstrate the effectiveness and utility of the virtual Morris water task for humans and show a robust sex difference in virtual place learning.
The Morris water maze (MWM) has been used to assess cognitive function in rats after a variety of lesions designed to model brain damage and to assess the effects of drugs, growth factors, and neural transplants on post-operative deficits. The present study examined recovery of spatial navigation in the MWM over time in order to model the spontaneous recovery of cognitive function seen in humans. Diffuse axonal injury, a neuropathology commonly associated with traumatic brain injury (TBI), was modelled by transecting the perforant path (PP) bilaterally, either caudal to the hippocampus or dorsal to it at the decussation of the dorsal hippocampal commissure. Both groups with PP cuts showed substantial deficits initially, but spatial performance recovered with time and training. Recovery of platform finding was nearly complete within 14 days of testing, but recovery of platform searching did not occur for 2 or 3 more weeks. When the platform was moved to a new location, a continuing deficit in learning rate was revealed. When the platform was moved to a new position every day, this deficit was even more evident. These results illustrate the multi-faceted nature of recovery after brain injury and provide a new model for assessing the effects of manipulations designed to modulate recovery.
The Morris water maze (MWM) was described 20 years ago as a device to investigate spatial learning and memory in laboratory rats. In the meanwhile, it has become one of the most frequently used laboratory tools in behavioral neuroscience. Many methodological variations of the MWM task have been and are being used by research groups in many different applications. However, researchers have become increasingly aware that MWM performance is influenced by factors such as apparatus or training procedure as well as by the characteristics of the experimental animals (sex, species/strain, age, nutritional state, exposure to stress or infection). Lesions in distinct brain regions like hippocampus, striatum, basal forebrain, cerebellum and cerebral cortex were shown to impair MWM performance, but disconnecting rather than destroying brain regions relevant for spatial learning may impair MWM performance as well. Spatial learning in general and MWM performance in particular appear to depend upon the coordinated action of different brain regions and neurotransmitter systems constituting a functionally integrated neural network. Finally, the MWM task has often been used in the validation of rodent models for neurocognitive disorders and the evaluation of possible neurocognitive treatments. Through its many applications, MWM testing gained a position at the very core of contemporary neuroscience research.
For nonhumans, it has been shown that the hippocampus (HPC) is critical for spatial memory. We tested patients with unilateral HPC resections on a virtual analogue of a classic spatial task to assess HPC functioning in nonhumans: the Morris water task. We found that when humans are required to use spatial cues to navigate to a hidden escape platform in a pool, patients with HPC resections display severe impairments in spatial navigation relative to age-matched controls and age-matched patients who have had extra-HPC resections. This effect occurred for every patient tested and was evident regardless of side of surgery. Hence, it is apparent across species and irrespective of which hemisphere is damaged that the human HPC is critical for spatial/relational memory.
We examined individual differences in working memory appearing in the effective visual field size while reading Japanese text. Working memory capacity was measured by a Japanese reading span test, and the subjects were divided into high- and low-score groups. Reading performance was measured by reading time, comprehension, and eye movements using a variable moving window through which the subject could read areas of the Japanese text. As the window size decreased, the reading time increased significantly. High-span subjects showed better performance in reading time, comprehension, and fixation duration than low-span subjects even in small visual fields. Interestingly, high-span subjects appear to show better information integration during reading, whereas low-span subjects showed less integration without parafoveal vision. These findings suggest that reading performance was better for subjects with larger working memory resources in a parafoveal restriction condition.
Subjects were required to navigate through a virtual 3D labyrinth presented on a screen while fMRI images were obtained. Contrasting the fMRI images obtained during the navigation trials with appropriate control conditions revealed a bilateral network comprising the parietal lobe (including the intraparietal sulcus) and various lateral and medial premotor areas. The subjects using an allocentric strategy showed stronger activation in the medial temporal areas including the parahippocampal region, the hippocampus, and the thalamus. In addition, the cerebellum was also active in those subjects. We believe that this activation pattern is related to visually guided memory retrieval based on generalized spatial maps. The stronger activation in the thalamic-basal ganglia-cerebellar-loop points to a more automatic support of memory and attentional processes possibly supporting memorization of spatial maps.
In this preliminary study we investigate gender differences in object location memory. Our purpose is to extend the results about object location memory obtained in laboratory settings to a real 3-D environment and to further distinguish the specific components involved in this kind of memory by considering the strategies adopted to perform the task. To do this, we join the three-level model of spatial representations (landmark, route, and survey) proposed by Siegel and White (1975) with the three subcomponents of spatial memory (what, where, and what + where) identified by Postma and De Haan (1996). We adopted the object relocation task devised by Postma and De Haan (1996), adapted to a real environment. Seven common objects were placed on the floor of a cylindrical room. Sixty-four males and 64 females were asked to memorize the spatial layout. Next, the experimenter moved the objects to a different position along with seven new objects and the participants had to relocate the original objects to their initial positions. In line with Postma, Izendoorn, and De Haan (1998), we found no gender difference in object recognition, and in recalling absolute distance and categorical spatial relations; however males were better than females in recalling the distance between objects and the size of the layout. Overall, the data show a male advantage in some components of spatial cognition closely linked to the encoding of the metric structure of the spatial relationships at both route and survey level.
Object descriptions are extracted and retained across saccades when observers view natural scenes. We investigated whether particular object properties are encoded and the stability of the resulting memories. We tested immediate recall of multiple types of information from real-world scenes and from computer-presented images of the same scenes. The relationship between fixations and properties of object memory was investigated. Position information was encoded and accumulated from multiple fixations. In contrast, identity and colour were encoded but did not require direct fixation and did not accumulate. In the current experiments, participants were unable to recall any information about shape or relative distances between objects. In addition, where information was encoded we found differential patterns of stability. Data from viewing real scenes and images were highly consistent, with stronger effects in the real-world conditions. Our findings imply that object files are not dependent upon the encoding of any particular object property and so are robust to dynamic visual environments.
In the present study we investigated spatial navigation in male and female Wistar rats in the Morris water escape task. Rats were subjected to procedures which required the use of a place (PLACE), cue (CUE) and egocentric (EGO) response to learn the task efficiently. In a first experiment rats were successively tested in the PLACE, CUE and EGO condition and in a second experiment the order of tasks was reversed. The first experiment showed that female rats swam longer distances and took more time to find the platform in the PLACE task. Further, the female rats spent less time near the previous platform position than the male rats during probe trial. No sex difference was found in the CUE and EGO task. In the second experiment, the female rats took longer to find the platform than the male rats in the EGO task. In the CUE and PLACE task no differences between the sexes was found during acquisition. However, the male rats spent more time near the previous platform position than the female rats during the probe trial of the PLACE task. On basis of present data it is concluded that the use of a PLACE-based strategy is better in male Wistar rats. CUE learning is not sex-dependent. The ability to use EGO strategies appears not to be different between male and female Wistar rats, but appears dependent on pre-exposure to the task.
Visual-spatial navigation in familiar and unfamiliar environments is an essential requirement of daily life. Animal studies indicated the importance of the hippocampus for navigation. Neuroimaging studies demonstrated gender difference or strategies dependent difference of neural substrates for navigation. Using functional magnetic resonance imaging, we measured brain activity related to navigation in four groups of normal volunteers: good navigators (males and females) and poor navigators (males and females). In a whole group analysis, task related activity was noted in the hippocampus, parahippocampal gyrus, posterior cingulate cortex, precuneus, parietal association areas, and the visual association areas. In group comparisons, good navigators showed a stronger activation in the medial temporal area and precuneus than poor navigators. There was neither sex effect nor interaction effect between sex and navigation ability. The activity in the left medial temporal areas was positively correlated with task performance, whereas activity in the right parietal area was negatively correlated with task performance. Furthermore, the activity in the bilateral medial temporal areas was positively correlated with scores reflecting preferred navigation strategies, whereas activity in the bilateral superior parietal lobules was negatively correlated with them. Our data suggest that different brain activities related to navigation should reflect navigation skill and strategies.
How do bottom-up and top-down guidance signals combine to guide search behavior? Observers searched for a target either with or without a preview (top-down manipulation) or a color singleton (bottom-up manipulation) among the display objects. With a preview, reaction times were faster and more initial eye movements were guided to the target; the singleton failed to attract initial saccades under these conditions. Only in the absence of a preview did subjects preferentially fixate the color singleton. We conclude that the search for realistic objects is guided primarily by top-down control. Implications for saliency map models of visual search are discussed.
We examined differences between 10-year-olds and young adults in resource recruitment and regulation during tasks of sustained attention and spatial working memory. We administered participants spatial 0- and 1-back tasks and used pupillary dilation as a measure of resource recruitment. Repeated administration of 0-back led to smaller pupillary dilations and greater response time (RT) variability, revealing a vigilance decrement. Effects of repeated administration of 0-back and differences between 0- and 1-back in d' and RTs were similar between ages. Results further suggested that the children may not have been as effective as adults in extracting frequency information. Thus, on simple tasks of sustained attention and working memory, children recruit resources in a manner similar to adults. Finally, d' was correlated with RT variability on both tasks at both ages, highlighting the role of attentional fluctuations on both tasks.
Gonadal steroid effects during puberty are often hypothesized to account for the male advantage seen in certain spatial tasks. One spatial task where males consistently show better performance than females is the Morris Water Task in which subjects must navigate to a goal location in a pool. We examined whether sex differences exist in pre-pubertal children completing a Virtual Morris Water Task, which has previously shown strong sex differences in adults. Pre-pubertal boys show superior performance to similar-aged girls, as evidenced by shorter latencies to find the platform and stronger preferences for the platform location during a probe trial. These results suggest that sex differences in spatial learning and memory exist prior to puberty and do not appear to require the effects of sex hormones at puberty. Rather, these differences may reflect early-life hormonal effects on hippocampal-dependent processes and may suggest different preferential learning strategies by boys and girls.
Navigation in real environments is often impaired by traumatic brain injury (TBI). These deficits in wayfinding appear to be due to disruption of cognitive processes underlying navigation and may in turn be due to damage to the hippocampus and frontal lobes. These wayfinding problems after TBI were investigated using a virtual simulation of a Morris Water Maze (MWM), a standard test of hippocampal function in laboratory animals. The virtual environment consisted of a large virtual arena in a very large virtual room whose walls provided views of a naturalistic landscape. Eleven community-dwelling TBI survivors and 12 comparison participants, matched for gender, age and education were tested to see if they could find a location in the arena marked by one of the following: (a) a visible platform, (b) a single proximal object, (c) a single proximal object among seven other distracter objects, or (d) distal features inside and outside the room. The proximal objects allowed participants to use egocentric (body-centered) navigational strategies that rely on relatively simple stimulus-response associations. The absence of proximal cues forced the participants to rely on distal features of the environment (room walls, landscape elements) and tested their ability to use allocentric (world-based) navigational strategies requiring cognitive mapping. Results indicated that the navigation of TBI survivors was not impaired when the proximal cues were present but was impaired when proximal cues were absent. These results provide more evidence that the navigational deficit after TBI is due to an inability to form, remember or use cognitive maps.