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1.  Estimating Location without External Cues 
PLoS Computational Biology  2014;10(10):e1003927.
The ability to determine one's location is fundamental to spatial navigation. Here, it is shown that localization is theoretically possible without the use of external cues, and without knowledge of initial position or orientation. With only error-prone self-motion estimates as input, a fully disoriented agent can, in principle, determine its location in familiar spaces with 1-fold rotational symmetry. Surprisingly, localization does not require the sensing of any external cue, including the boundary. The combination of self-motion estimates and an internal map of the arena provide enough information for localization. This stands in conflict with the supposition that 2D arenas are analogous to open fields. Using a rodent error model, it is shown that the localization performance which can be achieved is enough to initiate and maintain stable firing patterns like those of grid cells, starting from full disorientation. Successful localization was achieved when the rotational asymmetry was due to the external boundary, an interior barrier or a void space within an arena. Optimal localization performance was found to depend on arena shape, arena size, local and global rotational asymmetry, and the structure of the path taken during localization. Since allothetic cues including visual and boundary contact cues were not present, localization necessarily relied on the fusion of idiothetic self-motion cues and memory of the boundary. Implications for spatial navigation mechanisms are discussed, including possible relationships with place field overdispersion and hippocampal reverse replay. Based on these results, experiments are suggested to identify if and where information fusion occurs in the mammalian spatial memory system.
Author Summary
Spatial navigation is one of the most important functions of animal brains. Multiple regions and cell types encode the current location in mammalian brains, but the underlying interactions between sensory and memory information remain unclear. Recent experimental and theoretical evidence have been found to suggest that the presence of a boundary fundamentally alters the task of navigation. In this paper, evidence is provided that it is possible to determine the location inside any familiar arena with 1-fold rotational symmetry, while completely ignoring sensory cues from the outside world. Surprisingly, the results show that the mere knowledge of the boundary's existence is enough, without requiring direct physical contact. Localization is robust despite the presence of noise modelled from the rodent head direction system, and even inaccuracies in the navigation system's memory of the boundary or internal models of noise. In circular arenas, rotational asymmetry can arise from interior structures such as barriers or voids, also without contact information. This theoretical evidence highlights the need to distinguish arena-based navigation common to most experimental studies, from open field navigation. These findings also point to novel ways to study information fusion in mammalian brains.
doi:10.1371/journal.pcbi.1003927
PMCID: PMC4214594  PMID: 25356642
2.  Geometric cues influence head direction cells only weakly in non-disoriented rats 
The influential hypothesis that environmental geometry is critical for spatial orientation has been extensively tested behaviorally, and yet findings have been conflicting. Head direction (HD) cells, the neural correlate of the “sense of direction”, offer a window into the processes underlying directional orientation, and may help clarify the issue. In the present study, HD cells were recorded as rats foraged in enclosures of varying geometry, with or without simultaneous manipulation of landmarks and self-motion cues (path integration). All geometric enclosures had single-order rotational symmetry and thus completely polarized the environment. They also had unique features, such as corners, which could, in principle, act like landmarks. Despite these strongly polarizing geometric cues, HD cells in non-disoriented rats never rotated with these shapes. By contrast, when a cue card (white or grey) was added to one wall, HD cells readily rotated with the enclosure. When path integration was disrupted by disorienting the rat, HD cells now did rotate with the enclosure even without the landmark. Collectively these findings indicate that geometry exerts little or no influence on heading computations in non-disoriented rats, but it can do so in disoriented rats. We suggest that geometric processing is only a weak influence, providing a backup system for heading calculations and being recruited only under conditions of disorientation.
doi:10.1523/JNEUROSCI.2257-11.2011
PMCID: PMC3242014  PMID: 22049411
geometry; head direction cells; navigation; orientation; path integration; landmarks
3.  Spatial Memory During Progressive Disorientation 
Human spatial representations of object locations in a room-sized environment were probed for evidence that the object locations were encoded relative not just to the observer (egocentrically) but also to each other (allocentrically). Participants learned the locations of 4 objects and then were blindfolded and either (a) underwent a succession of 70° and 200° whole-body rotations or (b) were fully disoriented and then underwent a similar sequence of 70° and 200° rotations. After each rotation, participants pointed to the objects without vision. Analyses of the pointing errors suggest that as participants lost orientation, represented object directions generally “drifted” off of their true directions as an ensemble, not in random, unrelated directions. This is interpreted as evidence that object-to-object (allocentric) relationships play a large part in the human spatial updating system. However, there was also some evidence that represented object directions occasionally drifted off of their true directions independently of one another, suggesting a lack of allocentric influence. Implications regarding the interplay of egocentric and allocentric information are considered.
doi:10.1037/0278-7393.34.3.602
PMCID: PMC2883724  PMID: 18444759
spatial representation; egocentric–allocentric frames of reference; spatial updating
4.  Retrieving Enduring Spatial Representations after Disorientation 
Cognition  2012;124(2):143-155.
Four experiments tested whether there are enduring spatial representations of objects’ locations in memory. Previous studies have shown that under certain conditions the internal consistency of pointing to objects using memory is disrupted by disorientation. This disorientation effect has been attributed to an absence of or to imprecise enduring spatial representations of objects’ locations. Experiment 1 replicated the standard disorientation effect. Participants learned locations of objects in an irregular layout and then pointed to objects after physically turning to face an object and after disorientation. The expected disorientation was observed. In Experiment 2, after disorientation, participants were asked to imagine they were facing the original learning direction and then physically turned to adopt the test orientation. In Experiment 3, after disorientation, participants turned to adopt the test orientation and then were informed of the original viewing direction by the experimenter. A disorientation effect was not observed in Experiment 2 or 3. In Experiment 4, after disorientation, participants turned to face the test orientation but were not told the original learning orientation. As in Experiment 1, a disorientation effect was observed. These results suggest that there are enduring spatial representations of objects’ locations specified in terms of a spatial reference direction parallel to the learning view, and that the disorientation effect is caused by uncertainty in recovering the spatial reference direction relative to the testing orientation following disorientation.
doi:10.1016/j.cognition.2012.05.006
PMCID: PMC3395066  PMID: 22682765
5.  The Effect of Hearing Aid Microphone Mode on Performance in an Auditory Orienting Task 
Ear and Hearing  2014;35(5):e204-e212.
Objectives:
Although directional microphones on a hearing aid provide a signal-to-noise ratio benefit in a noisy background, the amount of benefit is dependent on how close the signal of interest is to the front of the user. It is assumed that when the signal of interest is off-axis, users can reorient themselves to the signal to make use of the directional microphones to improve signal-to-noise ratio. The present study tested this assumption by measuring the head-orienting behavior of bilaterally fit hearing-impaired individuals with their microphones set to omnidirectional and directional modes. The authors hypothesized that listeners using directional microphones would have greater difficulty in rapidly and accurately orienting to off-axis signals than they would when using omnidirectional microphones.
Design:
The authors instructed hearing-impaired individuals to turn and face a female talker in simultaneous surrounding male-talker babble. Participants pressed a button when they felt they were accurately oriented in the direction of the female talker. Participants completed three blocks of trials with their hearing aids in omnidirectional mode and three blocks in directional mode, with mode order randomized. Using a Vicon motion tracking system, the authors measured head position and computed fixation error, fixation latency, trajectory complexity, and proportion of misorientations.
Results:
Results showed that for larger off-axis target angles, listeners using directional microphones took longer to reach their targets than they did when using omnidirectional microphones, although they were just as accurate. They also used more complex movements and frequently made initial turns in the wrong direction. For smaller off-axis target angles, this pattern was reversed, and listeners using directional microphones oriented more quickly and smoothly to the targets than when using omnidirectional microphones.
Conclusions:
The authors argue that an increase in movement complexity indicates a switch from a simple orienting movement to a search behavior. For the most off-axis target angles, listeners using directional microphones appear to not know which direction to turn, so they pick a direction at random and simply rotate their heads until the signal becomes more audible. The changes in fixation latency and head orientation trajectories suggest that the decrease in off-axis audibility is a primary concern in the use of directional microphones, and listeners could experience a loss of initial target speech while turning toward a new signal of interest. If hearing-aid users are to receive maximum directional benefit in noisy environments, both adaptive directionality in hearing aids and clinical advice on using directional microphones should take head movement and orientation behavior into account.
doi:10.1097/AUD.0000000000000053
PMCID: PMC4232295  PMID: 25148290
Hearing aids; Directional microphones; Head movement; Sound localization
6.  Use of Self-to-object and Object-to-object Spatial Relations in Locomotion 
Eight experiments examined the use of representations of self-to-object or object-to-object spatial relations during locomotion. Participants learned geometrically regular or irregular layouts of objects while standing at the edge or in the middle, and then pointed to objects while blindfolded in three conditions: before turning (baseline), after rotating 240 degrees (updating), and after disorientation (disorientation). The internal consistency of pointing in the disorientation condition was equivalent to that in the updating condition when participants learned the regular layout. The internal consistency of pointing was disrupted by disorientation when participants learned the irregular layout. However when participants who learned the regular layout were instructed to use self-to-object spatial relations, the effect of disorientation on pointing consistency appeared. When participants who learned the irregular layout at the periphery of the layout were instructed to use object-to-object spatial relations, the effect of disorientation disappeared. These results suggest that people represent both self-to-object and object-to-object spatial relations, and primarily use object-to-object spatial representation in a regular layout and self-to-object spatial representation in an irregular layout.
doi:10.1037/a0016273
PMCID: PMC2774707  PMID: 19686010
self-to-object spatial relations; object-to-object spatial relations; spatial updating; disorientation
7.  Precision and accuracy of the subjective haptic vertical in the roll plane 
BMC Neuroscience  2010;11:83.
Background
When roll-tilted, the subjective visual vertical (SVV) deviates up to 40° from earth-vertical and trial-to-trial variability increases with head roll. Imperfections in the central processing of visual information were postulated to explain these roll-angle dependent errors. For experimental conditions devoid of visual input, e.g. adjustments of body posture or of an object along vertical in darkness, significantly smaller errors were noted. Whereas the accuracy of verticality adjustments seems to depend strongly on the paradigm, we hypothesize that the precision, i.e. the inverse of trial-to-trial variability, is less influenced by the experimental setup and mainly reflects properties of the otoliths. Here we measured the subjective haptic vertical (SHV) and compared findings with previously reported SVV data. Twelve healthy right-handed human subjects (handedness assessed based on subjects' verbal report) adjusted a rod with the right hand along perceived earth-vertical during static head roll-tilts (0-360°, steps of 20°).
Results
SHV adjustments showed a tendency for clockwise rod rotations to deviate counter-clockwise and for counter-clockwise rod rotations to deviate clockwise, indicating hysteresis. Clockwise rod rotations resulted in counter-clockwise shifts of perceived earth-vertical up to -11.7° and an average counter-clockwise SHV shift over all roll angles of -3.3° (± 11.0°; ± 1 StdDev). Counter-clockwise rod rotations yielded peak SHV deviations in clockwise direction of 8.9° and an average clockwise SHV shift over all roll angles of 1.8° (± 11.1°). Trial-to-trial variability was minimal in upright position, increased with increasing roll (peaking around 120-140°) and decreased to intermediate values in upside-down orientation. Compared to SVV, SHV variability near upright and upside-down was non-significantly (p > 0.05) larger; both showed an m-shaped pattern of variability as a function of roll position.
Conclusions
The reduction of adjustment errors by eliminating visual input supports the notion that deviations between perceived and actual earth-vertical in roll-tilted positions arise from central processing of visual information. The shared roll-tilt dependent modulation of trial-to-trial variability for both SVV and SHV, on the other hand, indicates that the perception of earth-verticality is dominated by the same sensory signal, i.e. the otolith signal, independent of whether the line/rod setting is under visual or tactile control.
doi:10.1186/1471-2202-11-83
PMCID: PMC2912915  PMID: 20630097
8.  Pituitary apoplexy can mimic acute meningoencephalitis or subarachnoid haemorrhage 
Pituitary apoplexy is an uncommon but life-threatening condition that is often overlooked and underdiagnosed. We report a 45-year-old man who presented to our emergency department with a sudden onset headache, acute confusion, signs of meningeal irritation and ophthalmoplegia. An initial diagnosis of acute meningoencephalitis was made, which was amended to pituitary apoplexy following thorough investigation within the emergency department.
A 45-year-old man was brought to our emergency department by ambulance with a history of sudden onset of frontal headache and acute confusion. His wife provided the history. There was no significant past medical history of diabetes, hypertension, recent travel abroad, exposure to sick contacts, involvement in outdoor pursuits such as hiking/cave diving, or trauma. He worked in a bank and had been well until 24 h prior to the onset of sudden headache, which was gradually worsening in nature and associated with increasing confusion. The patient's wife reported that he had neither experienced any fevers, night sweats, or coryzal symptoms nor received any recent vaccinations. He was not on any regular medications. He was a non-smoker and occasionally consumed alcohol. There was no significant family history. On examination in the ED, his temperature was 37.6°C, his pulse was 110/min, and he was normotensive and normoglycaemic. A macular blanching rash was noted over the patient's trunk. The patient was disoriented to time and place. Neurological examination revealed reduced GCS (11/15-E3, M6, V2), marked neck stiffness, a positive Kernig's sign and a right sixth nerve palsy.
A provisional diagnosis of acute meningoencephalitis was made and the patient was started on a course of intravenous antibiotics with benzyl penicillin 1.2 g, cefotaxime 2 g and acyclovir 750 mg. Baseline blood investigations revealed hyponatraemia (122 mmol/l), a white-cell count of 11 × 109/l and a C-reactive protein > 250. Due to the sudden onset of the symptoms and lack of prodrome, an urgent CT head scan was performed to rule out a cerebrovascular event. The scan demonstrated an enlarged pituitary gland (3 cm in diameter) with impingement of the optic chiasm. The centre of the enlarged pituitary gland was noted to be hypodense in comparison to its periphery, which was consistent with a diagnosis of pituitary apoplexy. A subsequent MRI confirmed the diagnosis (Figure 1) of an enlarged sella containing abnormal soft tissue with increased signal intensity suggestive of haemorrhage (Figure 1A).
Post-MRI a lumbar puncture was performed revealing glucose 3.4 mmol/l, protein 1.0 g/l, red cells of 53/mm3 and white cells of 174/mm3 with predominant neutrophilia. No organisms were seen, and CSF cultures and HSV DNA tests were found to be negative. Endocrinological investigations demonstrated low concentrations of thyroid hormones [TSH: 0.14 mIu/l (0.35-5.5 mlU/l), FT3: 1.1 nmol/l (1.2-3.0 nmol/l), FT4: 9.6 pmol/l (8-22 pmol/l)], gonadal hormones (LH: < 1 u/l) and prolactin: 16 u/l (<450 u/l). Serum FSH was 2.9 u/l (0.8-11.5 u/L) and cortisol 575 nmol/l (450-700 nmol/l). The patient was treated for hypopituitarism based on clinical and radiological findings with intravenous fluids, hydrocortisone (100 mg) and thyroxine (50 μg) as loading doses in the ED.
Within 24 h of commencement of therapy the patient's GCS rose to 15, and within 48 h there was marked improvement in the right sixth cranial nerve palsy. Formal visual field assessment demonstrated temporal visual field loss in the left eye. The patient was discharged to his usual residence a week later and follow-up was organised with both the endocrinologists and ophthalmologists. Follow-up MRI demonstrated that there was no significant change in either size or signal characteristics of the pituitary fossa mass (Figure 1B).
doi:10.1186/1865-1380-4-63
PMCID: PMC3205010  PMID: 21975129
9.  Meperidine-induced reversible retrograde amnesia 
Introduction: Meperidine is a synthetic opioid analog that is frequently prescribed for acute pain management. Normeperidine, the only active metabolite of meperidine, is neurotoxic and can cause significant central nervous system adverse events.
Case summary: A 29-year-old woman (height, 170 cm; weight, 85 kg) presented to Marmara University Hospital Emergency Department, Istanbul, Turkey, complaining of low back pain she described as “stabbing.” Physical examination revealed impaired lower-extremity mobility and normal vital-sign findings. There was no evidence of foot drop, head or other trauma, and systemic physical examination was unremarkable. Other common causes (eg, pyelonephritis, nephrolithiasis, pancreatitis, trauma) of lower back pain were excluded. To achieve analgesia, meperidine 80 mg was administered intravenously in 100 mL of isotonic saline solution for 20 minutes. Within 20 minutes,analgesia was achieved,but the patient developed retrograde amnesia, becoming disoriented to time, location, and persons. Her speech slowed and perceptional changes developed. After the onset of amnesia, a complete physical examination was conducted.It failed to reveal focal neurologic deficit,and laboratory (sodium, potassium, magnesium, phosphorus, serum creatinine, blood urea nitrogen, albumin, bilirubin, hemoglobin,and platelet count) and subsequent vital-sign findings (blood pressure, 150/100 mm Hg; heart rate, 100 beats per minute; respiratory rate, 18 breaths per minute; body temperature, 37 ଌ and pulse oximetry,99%) were within the normal range. Noncontrast computed tomography did not reveal any abnormality. Initially, the patient's condition was attributed to medication error due to incorrect dosage or infusion rate. Despite a review of medication logs, equipment, and the vital-sign record, the etiology for the phenomenon could not be identified. Meperidine was discontinued and oxygen and intravenous isotonic saline solution were initiated as supportive treatment. Three hours after meperidine administration was discontinued, the amnesia and disorientation spontaneously resolved.
Conclusion: Meperidine was probably associated with reversible amnesia in this healthy patient after a single therapeutic dose.
doi:10.1016/j.curtheres.2008.04.006
PMCID: PMC3969970  PMID: 24692795
meperidine; amnesia; opioid analgesic; central nervous system toxicity
10.  Tracking pigeons in a magnetic anomaly and in magnetically “quiet” terrain 
Die Naturwissenschaften  2011;98(7):575-581.
Pigeons were released at two sites of equal distance from the loft, one within a magnetic anomaly, the other in magnetically quiet terrain, and their tracks were recorded with the help of GPS receivers. A comparison of the beginning of the tracks revealed striking differences: within the anomaly, the initial phase lasted longer, and the distance flown was longer, with the pigeons' headings considerably farther from the home direction. During the following departure phase, the birds were well homeward oriented at the magnetically quiet site, whereas they continued to be disoriented within the anomaly. Comparing the tracks in the anomaly with the underlying magnetic contours shows considerable differences between individuals, without a common pattern emerging. The differences in magnetic intensity along the pigeons' path do not differ from a random distribution of intensity differences around the release site, indicating that the magnetic contours do not directly affect the pigeons' routes. Within the anomaly, pigeons take longer until their flights are oriented, but 5 km from the release point, the birds, still within the anomaly, are also significantly oriented in the home direction. These findings support the assumption that magnetically anomalous conditions initially interfere with the pigeons' navigational processes, with birds showing rather individual responses in their attempts to overcome these problems.
Electronic supplementary material
The online version of this article (doi:10.1007/s00114-011-0802-3) contains supplementary material, which is available to authorized users.
doi:10.1007/s00114-011-0802-3
PMCID: PMC3128737  PMID: 21691766
Magnetic anomaly; Pigeon navigation; Homing; GPS tracking; Magnetic “map” factors; Point of Decision
11.  Directional responding of C57BL/6J mice in the Morris water maze is influenced by visual and vestibular cues and is dependent upon the anterior thalamic nuclei 
Recent findings indicate that rats navigate in spatial tasks such as the Morris water maze (MWM) using a local cue-based reference frame rather than a distal cue-based reference frame. Specifically, rats swim in a particular direction to a location relative to pool-based cues, rather than to an absolute location defined by room-based cues. Neural mechanisms supporting this bias in rodents for relative responding in spatial tasks are not yet understood. Anterior thalamic neurons discharge according to the current directional heading of the animal. The contribution of head direction (HD) cell activity to navigation has been difficult to elucidate. We found that male C57BL/6J mice trained for 4 or 7 days in the MWM exhibited an overwhelming preference for swimming in a direction relative to pool-based cues over absolute responding during a platform-less probe test. Rotation of extra-maze cues caused a corresponding rotation of the direction mice swam during probe test, suggesting that both pool- and room-based reference frames guide platform search. However, disorienting the mice before the probe test disturbed relative responding. Therefore, relative responding is guided by both internal and external cue sources. Selective inactivation of anterior thalamic nuclei (ATN) by microinfusion of muscimol or fluorophore-conjugated muscimol caused a near complete shift in preference from relative to absolute responding. Interestingly, inactivation of the dorsal CA1 region of the hippocampus did not affect relative responding. These data suggest that ATN, and HD cells therein, may guide relative responding in the MWM, a task considered by most to reflect hippocampal processing.
doi:10.1523/JNEUROSCI.4868-11.2012
PMCID: PMC3477624  PMID: 22836256
spatial navigation; directional heading; anterior thalamus; hippocampus; head direction cell; vestibular; distal cues
12.  A Chasm Between Injury and Care: Experiences of Black Male Victims of Violence 
The Journal of trauma  2010;69(6):1372-1378.
Background
Despite higher rates of stabbing and shooting violence among black men, healthcare systems have not demonstrated an efficacious response to these patients. This study describes challenges and promotive factors for engaging black male violence victims of violence with medical and mental healthcare.
Methods
Black male victims of stabbings and shootings were recruited through fliers and word of mouth, and were interviewed individually (n = 12) or in pairs (n = 4) using a semistructured guide. A racially diverse multidisciplinary team analyzed the data using Grounded Theory methods.
Results
Challenges to engagement with healthcare included the following: (1) Disconnect in the aftermath; e.g. participants reported not realizing they were seriously injured (“just a scratch” “poke”), were disoriented (“did not know where I was”), or were consumed with anger. (2) Institutional mistrust: blurred lines between healthcare and police, money-motivated care. (3) Foreshortened future: expectations they would die young. (4) Self-reliance: fix mental and substance abuse issues on their own. (5) Logistical issues: postinjury mental health symptoms, disability, and safety concerns created structural barriers to recovery and engagement with healthcare. Promotive factors included the following: (1) desire professionalism, open personality, and shared experience from clinicians; (2) turning points: injury or birth of a child serve as a “wake up call”; and (3) positive people, future-oriented friends and family.
Conclusions
For black male violence victims, medical treatment did not address circumstances of and reactions to injury. Policies delineating boundaries between medical care and law enforcement and addressing postinjury mental health symptoms, disability, and safety concerns may improve the recovery process.
doi:10.1097/TA.0b013e3181e74fcf
PMCID: PMC3005415  PMID: 20838259
Black male; Community violence; Qualitative research
13.  Vestibular prosthesis tested in rhesus monkeys 
We are studying the effectiveness of a semicircular canal prosthesis to improve postural control, perception of spatial orientation, and the VOR in rhesus monkeys with bilateral vestibular hypofunction. Balance is examined by measuring spontaneous sway of the body during quiet stance and postural responses evoked by head turns and rotation of the support surface; perception is measured with a task derived from the subjective visual vertical (SVV) test during static and dynamic rotation in the roll plane; and the angular VOR is measured during rotation about the roll, pitch, and yaw axes. After the normal responses are characterized, bilateral vestibular loss is induced with intratympanic gentamicin, and then multisite stimulating electrodes are chronically implanted into the ampullae of all three canals in one ear. The postural, perceptual, and VOR responses are then characterized in the ablated state, and then bilateral, chronic electrical stimulation is applied to the ampullary nerves using a prosthesis that senses angular head velocity in three-dimensions and uses this information to modulate the rate of current pulses provided by the implanted electrodes. We are currently characterizing two normal monkeys with these paradigms, and vestibular ablation and electrode implantation are planned for the near future. In one prior rhesus monkey tested with this approach, we found that a one-dimensional (posterior canal) prosthesis improved balance during head turns, perceived head orientation during roll tilts, and the VOR in the plane of the instrumented canal. We therefore predict that the more complete information provided by a three-dimensional prosthesis that modulates activity in bilaterally-paired canals will exceed the benefits provided by the one-dimensional, unilateral approach used in our preliminary studies.
doi:10.1109/IEMBS.2011.6090573
PMCID: PMC3594774  PMID: 22254795
14.  The influence of vision on sound localization abilities in both the horizontal and vertical planes 
Numerous recent reports have suggested that individuals deprived of vision are able to develop heightened auditory spatial abilities. However, most such studies have compared the blind to blindfolded sighted individuals, a procedure that might introduce a strong performance bias. Indeed, while blind individuals have had their whole lives to adapt to this condition, sighted individuals might be put at a severe disadvantage when having to localize sounds without visual input. To address this unknown, we compared the sound localization ability of eight sighted individuals with and without a blindfold in a hemi-anechoic chamber. Sound stimuli were broadband noise delivered via two speaker arrays: a horizontal array with 25 loudspeakers (ranging from −90° to +90°; 7.5°) and a vertical array with 16 loudspeakers (ranging from −45° to +67.5°). A factorial design was used, where we compared two vision conditions (blindfold vs. non-blindfold), two sound planes (horizontal vs. vertical) and two pointing methods (hand vs. head). Results show that all three factors significantly interact with one another with regards to the average absolute deviation error. Although blindfolding significantly affected all conditions, it did more so for head-pointing in the horizontal plane. Moreover, blindfolding was found to increase the tendency to undershoot more eccentric spatial positions for head-pointing, but not hand-pointing. Overall, these findings suggest that while proprioceptive cues appear to be sufficient for accurate hand pointing in the absence of visual feedback, head pointing relies more heavily on visual cues in order to provide a precise response. It also strongly argues against the use of head pointing methodologies with blindfolded sighted individuals, particularly in the horizontal plane, as it likely introduces a bias when comparing them to blind individuals.
doi:10.3389/fpsyg.2013.00932
PMCID: PMC3860057  PMID: 24376430
sound localization; vision; pointing methods; spatial hearing; blindness
15.  Misperception of exocentric directions in auditory space 
Acta psychologica  2008;129(1):72-82.
Previous studies have demonstrated large errors (over 30°) in visually perceived exocentric directions (the direction between two objects that are both displaced from the observer’s location; e.g., Philbeck et al., in press). Here, we investigated whether a similar pattern occurs in auditory space. Blindfolded participants either attempted to aim a pointer at auditory targets (an exocentric task) or gave a verbal estimate of the egocentric target azimuth. Targets were located at 20° to 160° azimuth in the right hemispace. For comparison, we also collected pointing and verbal judgments for visual targets. We found that exocentric pointing responses exhibited sizeable undershooting errors, for both auditory and visual targets, that tended to become more strongly negative as azimuth increased (up to −19° for visual targets at 160°). Verbal estimates of the auditory and visual target azimuths, however, showed a dramatically different pattern, with relatively small overestimations of azimuths in the rear hemispace. At least some of the differences between verbal and pointing responses appear to be due to the frames of reference underlying the responses; when participants used the pointer to reproduce the egocentric target azimuth rather than the exocentric target direction relative to the pointer, the pattern of pointing errors more closely resembled that seen in verbal reports. These results show that there are similar distortions in perceiving exocentric directions in visual and auditory space.
doi:10.1016/j.actpsy.2008.04.008
PMCID: PMC2614239  PMID: 18555205
manual pointing; auditory space perception; perception / action; perceived direction; spatial cognition
16.  The effect of traumatic brain injury on the health of homeless people 
Background
We sought to determine the lifetime prevalence of traumatic brain injury and its association with current health conditions in a representative sample of homeless people in Toronto, Ontario.
Methods
We surveyed 601 men and 303 women at homeless shelters and meal programs in 2004–2005 (response rate 76%). We defined traumatic brain injury as any self-reported head injury that left the person dazed, confused, disoriented or unconscious. Injuries resulting in unconsciousness lasting 30 minutes or longer were defined as moderate or severe. We assessed mental health, alcohol and drug problems in the past 30 days using the Addiction Severity Index. Physical and mental health status was assessed using the SF-12 health survey. We examined associations between traumatic brain injury and health conditions.
Results
The lifetime prevalence among homeless participants was 53% for any traumatic brain injury and 12% for moderate or severe traumatic brain injury. For 70% of respondents, their first traumatic brain injury occurred before the onset of homelessness. After adjustment for demographic characteristics and lifetime duration of homelessness, a history of moderate or severe traumatic brain injury was associated with significantly increased likelihood of seizures (odds ratio [OR] 3.2, 95% confidence interval [CI] 1.8 to 5.6), mental health problems (OR 2.5, 95% CI 1.5 to 4.1), drug problems (OR 1.6, 95% CI 1.1 to 2.5), poorer physical health status (–8.3 points, 95% CI –11.1 to –5.5) and poorer mental health status (–6.0 points, 95% CI –8.3 to –3.7).
Interpretation
Prior traumatic brain injury is very common among homeless people and is associated with poorer health.
doi:10.1503/cmaj.080341
PMCID: PMC2553875  PMID: 18838453
17.  Multiple Motor Learning Strategies in Visuomotor Rotation 
PLoS ONE  2010;5(2):e9399.
Background
When exposed to a continuous directional discrepancy between movements of a visible hand cursor and the actual hand (visuomotor rotation), subjects adapt their reaching movements so that the cursor is brought to the target. Abrupt removal of the discrepancy after training induces reaching error in the direction opposite to the original discrepancy, which is called an aftereffect. Previous studies have shown that training with gradually increasing visuomotor rotation results in a larger aftereffect than with a suddenly increasing one. Although the aftereffect difference implies a difference in the learning process, it is still unclear whether the learned visuomotor transformations are qualitatively different between the training conditions.
Methodology/Principal Findings
We examined the qualitative changes in the visuomotor transformation after the learning of the sudden and gradual visuomotor rotations. The learning of the sudden rotation led to a significant increase of the reaction time for arm movement initiation and then the reaching error decreased, indicating that the learning is associated with an increase of computational load in motor preparation (planning). In contrast, the learning of the gradual rotation did not change the reaction time but resulted in an increase of the gain of feedback control, suggesting that the online adjustment of the reaching contributes to the learning of the gradual rotation. When the online cursor feedback was eliminated during the learning of the gradual rotation, the reaction time increased, indicating that additional computations are involved in the learning of the gradual rotation.
Conclusions/Significance
The results suggest that the change in the motor planning and online feedback adjustment of the movement are involved in the learning of the visuomotor rotation. The contributions of those computations to the learning are flexibly modulated according to the visual environment. Such multiple learning strategies would be required for reaching adaptation within a short training period.
doi:10.1371/journal.pone.0009399
PMCID: PMC2827554  PMID: 20195373
18.  Egocentric Representation Acquired from Offline Map Learning 
PLoS ONE  2013;8(3):e60194.
It is widely accepted that people establish allocentric spatial representation after learning a map. However, it is unknown whether people can directly acquire egocentric representation after map learning. In two experiments, the participants learned a distal environment through a map and then performed the egocentric pointing tasks in that environment under three conditions: with the heading aligned with the learning perspective (baseline), after 240° rotation from the baseline (updating), and after disorientation (disorientation). Disorientation disrupted the internal consistency of pointing among objects when the participants learned the sequentially displayed map, on which only one object name was displayed at a time while the location of “self” remained on the screen all the time. However, disorientation did not affect the internal consistency of pointing among objects when the participants learned the simultaneously displayed map. These results suggest that the egocentric representation can be acquired from a sequentially presented map.
doi:10.1371/journal.pone.0060194
PMCID: PMC3610764  PMID: 23555922
19.  Incremental angular vestibulo-ocular reflex adaptation to active head rotation 
Studies on motor learning typically present a constant adaptation stimulus, corresponding to the desired final adaptive state. Studies of the auditory and optokinetic systems provide compelling evidence that neural plasticity is enhanced when the error signal driving adaptation is instead adjusted gradually throughout training. We sought to determine whether the angular vestibulo-ocular reflex (aVOR) may be adaptively increased using an incremental velocity error signal (IVE) compared with a conventional constant and large velocity-gain demand (x2). We compared the magnitude of aVOR gain change for these two paradigms across different motion contexts (active and passive). Seven individuals with normal vestibular function and six individuals with unilateral vestibular hypofunction (UVH) were exposed to the IVE and x2 (“control”) aVOR demand tasks. Each subject participated in 10 epochs of 30 active head impulses over a 15 min aVOR gain increase training session separately for the IVE and x2 paradigms, separated by either seven days (normal subjects) or 14 days (UVH subjects). For both normal and UVH subjects, both paradigms led to aVOR gain increase during the training session. For the normal subjects, the IVE paradigm led to larger aVOR gain change after training compared to the x2 paradigm, for both active (mean 17.3 ± 4% vs. mean 7.1 ± 9%, P = 0.029) and passive (mean 14.2 ± 5% vs. 4.5 ± 8%, P = 0.018) head impulses. For subjects with UVH, IVE produced a greater change in aVOR gain for active head impulses (mean 18.2 ± 9.2% vs. mean –6 ± 3.8%, P = 0.003). However, aVOR gains for passive head impulses were less consistent after IVE, with only two subjects displaying greater aVOR gain with this incremental paradigm. Some individuals generated compensatory saccades that occurred in the same direction of the deficient aVOR during either training paradigm. Our data suggest that the aVOR is modifiable when the velocity error signal is presented incrementally, and that this adaptation stimulus is particularly effective in the case of unilateral vestibular hypofunction. This has implications for programs of vestibular rehabilitation, where active head rotation is prescribed as a means to improve gaze stability.
doi:10.1007/s00221-008-1537-z
PMCID: PMC2946619  PMID: 18712370
Vestibulo-ocular reflex; Saccades; Vestibular rehabilitation; Adaptation
20.  Dancing Bees Improve Colony Foraging Success as Long-Term Benefits Outweigh Short-Term Costs 
PLoS ONE  2014;9(8):e104660.
Waggle dancing bees provide nestmates with spatial information about high quality resources. Surprisingly, attempts to quantify the benefits of this encoded spatial information have failed to find positive effects on colony foraging success under many ecological circumstances. Experimental designs have often involved measuring the foraging success of colonies that were repeatedly switched between oriented dances versus disoriented dances (i.e. communicating vectors versus not communicating vectors). However, if recruited bees continue to visit profitable food sources for more than one day, this procedure would lead to confounded results because of the long-term effects of successful recruitment events. Using agent-based simulations, we found that spatial information was beneficial in almost all ecological situations. Contrary to common belief, the benefits of recruitment increased with environmental stability because benefits can accumulate over time to outweigh the short-term costs of recruitment. Furthermore, we found that in simulations mimicking previous experiments, the benefits of communication were considerably underestimated (at low food density) or not detected at all (at medium and high densities). Our results suggest that the benefits of waggle dance communication are currently underestimated and that different experimental designs, which account for potential long-term benefits, are needed to measure empirically how spatial information affects colony foraging success.
doi:10.1371/journal.pone.0104660
PMCID: PMC4139316  PMID: 25141306
21.  Gating of neural error signals during motor learning 
eLife  2014;3:e02076.
Cerebellar climbing fiber activity encodes performance errors during many motor learning tasks, but the role of these error signals in learning has been controversial. We compared two motor learning paradigms that elicited equally robust putative error signals in the same climbing fibers: learned increases and decreases in the gain of the vestibulo-ocular reflex (VOR). During VOR-increase training, climbing fiber activity on one trial predicted changes in cerebellar output on the next trial, and optogenetic activation of climbing fibers to mimic their encoding of performance errors was sufficient to implant a motor memory. In contrast, during VOR-decrease training, there was no trial-by-trial correlation between climbing fiber activity and changes in cerebellar output, and climbing fiber activation did not induce VOR-decrease learning. Our data suggest that the ability of climbing fibers to induce plasticity can be dynamically gated in vivo, even under conditions where climbing fibers are robustly activated by performance errors.
DOI: http://dx.doi.org/10.7554/eLife.02076.001
eLife digest
The cerebellum (or ‘little brain’) is located underneath the cerebral hemispheres. Despite comprising around 10% of the brain’s volume, the cerebellum contains roughly half of the brain’s neurons. Many of the functions of the cerebellum are related to the control and fine-tuning of movement, and people whose cerebellum has been damaged have problems with balance and coordination, and with learning new motor skills.
One of the roles of the cerebellum is to control a reflex known as the vestibulo-ocular reflex, which enables us to keep our gaze fixed on an object as we turn our heads. The cerebellum relays information about head movements to the muscles that control the eyes, instructing the eyes to move in the opposite direction to the head. This keeps the image of the object we are looking at stable on the retina.
The vestibulo-ocular reflex is controlled by a circuit that includes Purkinje cells (which are the main output cells of the cerebellum) and climbing fibres (which originate in the brainstem). Any failure of the vestibulo-ocular reflex to fully compensate for head movements generates an error signal that activates the climbing fibres. These in turn modify the output of Purkinje cells, leading ultimately to adjustments in eye movements.
However, Kimpo et al. have now obtained evidence that Purkinje cells can modulate their response to the instructions they receive from climbing fibres. Monkeys sat in a rotating chair while a visual object they were trained to track with their eyes was moved to induce errors in the vestibulo-ocular reflex. When the object was moved so that a bigger reflexive eye movement was required to stabilize the image, the activation of the climbing fibres in response to the error led to a change in the response of the Purkinje cells, as expected. However, when a smaller reflexive eye movement was needed, the error-driven responses of the climbing fibres did not alter the responses of Purkinje cells. Similar results were obtained using pulses of light to artificially activate climbing fibres and thus simulate error signals.
The work of Kimpo et al. indicates that the cerebellum does not blindly follow the instructions it receives from the brainstem, but can instead modulate its responses to incoming information about performance errors. Further work is now required to identify factors that influence the responsiveness of the cerebellum: such information could ultimately be used to improve learning of motor skills and recovery from injury.
DOI: http://dx.doi.org/10.7554/eLife.02076.002
doi:10.7554/eLife.02076
PMCID: PMC3989583  PMID: 24755290
rhesus macaque; climbing fibers; cerebellum; motor learning; vestibulo-ocular reflex; supervised learning; mouse; other
22.  What is the Safest Sprint Starting Position for American Football Players? 
The main objective of this study was to perform a biomechanical analysis of three different sprint start patterns to determine the safest position in term of neck injury and Sport-Related Concussion (SRC). The second objective was to collect data on the learning process effect between football players and non-players. Three different sprint initial positions adopted by football players were studied (i.e., 4-, 3- and 2-point positions). Twenty five young healthy males, including 12 football players, participated to this study. A stereophotogrammetric system (i.e., Vicon) was used to record motion patterns and body segments positions. Various measurements related to head and trunk orientation, and player field-of-view were obtained (e.g., head height, trunk bending, time to reach upright position, head speed (vertical direction) and body speed (horizontal direction)). Learning process was found to have no influence on studied parameters. Head redress is also delayed when adopting a 4-point position leading to a reduce field-of-view during the start and increasing therefore the probability of collision. Concerning the three different positions, the 4-point position seems to be the more dangerous because leading to higher kinetic energy than the 2- and 3-point start positions. This study proposes a first biomechanical approach to understand risk/benefit balance for athletes for those three different start positions. Results suggested that the 4-point position is the most risky for football players.
Key pointsMotion analysis and biomechanical analysis of the initial start position of the sprint could be used to increase the safety of the football players.Analysis of kinematic and trajectory of the head and the time to reach the upright position could be used to determine whether or not a player can return to play after concussion.A balance needs to be found between player’s safety (2-point start) and speed (4-point start).
PMCID: PMC3990900  PMID: 24790500
Sports; sports medicine; brain concussion; biomechanics
23.  Light-dependent magnetoreception in birds: increasing intensity of monochromatic light changes the nature of the response 
Background
The Radical Pair model proposes that magnetoreception is a light-dependent process. Under low monochromatic light from the short-wavelength part of the visual spectrum, migratory birds show orientation in their migratory direction. Under monochromatic light of higher intensity, however, they showed unusual preferences for other directions or axial preferences. To determine whether or not these responses are still controlled by the respective light regimes, European robins, Erithacus rubecula, were tested under UV, Blue, Turquoise and Green light at increasing intensities, with orientation in migratory direction serving as a criterion whether or not magnetoreception works in the normal way.
Results
The birds were well oriented in their seasonally appropriate migratory direction under 424 nm Blue, 502 nm Turquoise and 565 nm Green light of low intensity with a quantal flux of 8·1015 quanta s-1 m-2, indicating unimpaired magnetoreception. Under 373 nm UV of the same quantal flux, they were not oriented in migratory direction, showing a preference for the east-west axis instead, but they were well oriented in migratory direction under UV of lower intensity. Intensities of above 36·1015 quanta s-1 m-2 of Blue, Turquoise and Green light elicited a variety of responses: disorientation, headings along the east-west axis, headings along the north-south axis or 'fixed' direction tendencies. These responses changed as the intensity was increased from 36·1015 quanta s-1 m-2 to 54 and 72·1015 quanta s-1 m-2.
Conclusion
The specific manifestation of responses in directions other than the migratory direction clearly depends on the ambient light regime. This implies that even when the mechanisms normally providing magnetic compass information seem disrupted, processes that are activated by light still control the behavior. It suggests complex interactions between different types of receptors, magnetic and visual. The nature of the receptors involved and details of their connections are not yet known; however, a role of the color cones in the processes mediating magnetic input is suggested.
doi:10.1186/1742-9994-4-5
PMCID: PMC1810254  PMID: 17302975
24.  THE EMERGENCE OF THE DISORGANIZED/DISORIENTED (D) ATTACHMENT CLASSIFICATION, 1979–1982 
History of Psychology  2015;18(1):32-46.
This article examines the emergence of the concept of infant disorganized/disoriented attachment, drawing on published and archival texts and interviews. Since this new classification was put forward by Main and Solomon (1986), “disorganized/disoriented attachment” has become an important concept in clinical and social intervention contexts. Yet whereas Main and Solomon have often been misunderstood to have introduced disorganized/disoriented attachment in order to produce an exhaustive, categorical system of infant classifications, this article will suggest quite a different account. Attention will be paid to the emergence of disorganized attachment as a classification out of results and reflections in the late 1970s regarding the limits of an alarmed infant’s capacities for maintaining behavioral and attentional avoidance. In contrasting this interpretation of Main and Solomon’s work with current, widespread misunderstandings, the article will critically examine tendencies that have supported the reification and misapplication of the concept of disorganized/disoriented attachment.
doi:10.1037/a0038524
PMCID: PMC4321742  PMID: 25664884
attachment; developmental psychology; Ainsworth Strange Situation Procedure; anomaly; classification
25.  Axis of Eye Rotation Changes with Head-Pitch Orientation during Head Impulses about Earth-Vertical 
The goal of this study was to assess how the axis of head rotation, Listing's law, and eye position influence the axis of eye rotation during brief, rapid head rotations. We specifically asked how the axis of eye rotation during the initial angular vestibuloocular reflex (VOR) changed when the pitch orientation of the head relative to Earth-vertical was varied, but the initial position of the eye in the orbit and the orientation of Listing's plane with respect to the head were fixed. We measured three-dimensional eye and head rotation axes in eight normal humans using the search coil technique during head-and-trunk (whole-body) and head-on-trunk (head-only) “impulses” about an Earth-vertical axis. The head was initially oriented at one of five pitch angles (30° nose down, 15° nose down, 0°, 15° nose up, 30° nose up). The fixation target was always aligned with the nasooccipital axis. Whole-body impulses were passive, unpredictable, manual, rotations with peak-amplitude of ∼20°, peak-velocity of ∼80°/s, and peak-acceleration of ∼1000°/s2. Head-only impulses were also passive, unpredictable, manual, rotations with peak-amplitude of ∼20°, peak-velocity of ∼150°/s, and peak-acceleration of ∼3000°/s2. During whole-body impulses, the axis of eye rotation tilted in the same direction, and by an amount proportional (0.51 ± 0.09), to the starting pitch head orientation (P < 0.05). This proportionality constant decreased slightly to 0.39 ± 0.08 (P < 0.05) during head-only impulses. Using the head-only impulse data, with the head pitched up, we showed that only 50% of the tilt in the axis of eye rotation could be predicted from vectorial summation of the gains (eye velocity/head velocity) obtained for rotations about the pure yaw and roll head axes. Thus, even when the orientation of Listing's plane and eye position in the orbit are fixed, the axis of eye rotation during the VOR reflects a compromise between the requirements of Listing's law and a perfectly compensatory VOR.
doi:10.1007/s10162-006-0029-8
PMCID: PMC2504578  PMID: 16552499
vestibuloocular reflex; axis of eye rotation; axis of head rotation; torsion; Listing's law

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