Robust estimation of tear film surface quality in lateral shearing interferometry

Interferometry is a sensitive technique for recording tear film surface irregularities in a noninvasive manner. At the same time, the technique is hindered by natural eye movements resulting in measurement noise. Estimating tear film surface quality from interferograms can be reduced to a spatial-average-localized weighted estimate of the first harmonic of the interference fringes. However, previously reported estimation techniques proved to perform poorly in cases where the pattern fringes were significantly disturbed. This can occur in cases of measuring tear film surface quality on a contact lens on the eye or in a dry eye. We present a new estimation technique for extracting the first harmonic from the interference fringes that combines the traditional spectral estimation techniques with morphological image processing techniques. The proposed technique proves to be more robust to changes in interference fringes caused by natural eye movements and the degree of dryness of the contact lens and corneal surfaces than its predecessors, resulting in tear film surface quality estimates that are less noisy

The impact of presbyopic spectacles and contact lenses on driving performance

Presbyopia affects individuals from the age of 45 years onwards, resulting in difficulty in accurately focusing on near objects. There are many optical corrections available including spectacles or contact lenses that are designed to enable presbyopes to see clearly at both far and near distances. However, presbyopic vision corrections also disturb aspects of visual function under certain circumstances. The impact of these changes on activities of daily living such as driving are, however, poorly understood. Therefore, the aim of this study was to determine which aspects of driving performance might be affected by wearing different types of presbyopic vision corrections. In order to achieve this aim, three experiments were undertaken. The first experiment involved administration of a questionnaire to compare the subjective driving difficulties experienced when wearing a range of common presbyopic contact lens and spectacle corrections. The questionnaire was developed and piloted, and included a series of items regarding difficulties experienced while driving under day and night-time conditions. Two hundred and fifty five presbyopic patients responded to the questionnaire and were categorised into five groups, including those wearing no vision correction for driving (n = 50), bifocal spectacles (BIF, n = 54), progressive addition lenses spectacles (PAL, n = 50), monovision (MV, n = 53) and multifocal contact lenses (MTF CL, n = 48). Overall, ratings of satisfaction during daytime driving were relatively high for all correction types. However, MV and MTF CL wearers were significantly less satisfied with aspects of their vision during night-time than daytime driving, particularly with regard to disturbances from glare and haloes. Progressive addition lens wearers noticed more distortion of peripheral vision, while BIF wearers reported more difficulties with tasks requiring changes in focus and those who wore no vision correction for driving reported problems with intermediate and near tasks. Overall, the mean level of satisfaction for daytime driving was quite high for all of the groups (over 80%), with the BIF wearers being the least satisfied with their vision for driving. Conversely, at night, MTF CL wearers expressed the least satisfaction. Research into eye and head movements has become increasingly of interest in driving research as it provides a means of understanding how the driver responds to visual stimuli in traffic. Previous studies have found that wearing PAL can affect eye and head movement performance resulting in slower eye movement velocities and longer times to stabilize the gaze for fixation. These changes in eye and head movement patterns may have implications for driving safety, given that the visual tasks for driving include a range of dynamic search tasks. Therefore, the second study was designed to investigate the influence of different presbyopic corrections on driving-related eye and head movements under standardized laboratory-based conditions. Twenty presbyopes (mean age: 56.1 ± 5.7 years) who had no experience of wearing presbyopic vision corrections, apart from single vision reading spectacles, were recruited. Each participant wore five different types of vision correction: single vision distance lenses (SV), PAL, BIF, MV and MTF CL. For each visual condition, participants were required to view videotape recordings of traffic scenes, track a reference vehicle and identify a series of peripherally presented targets while their eye and head movements were recorded using the faceLAB® eye and head tracking system. Digital numerical display panels were also included as near visual stimuli (simulating the visual displays of a vehicle speedometer and radio). The results demonstrated that the path length of eye movements while viewing and responding to driving-related traffic scenes was significantly longer when wearing BIF and PAL than MV and MTF CL. The path length of head movements was greater with SV, BIF and PAL than MV and MTF CL. Target recognition was less accurate when the near stimulus was located at eccentricities inferiorly and to the left, rather than directly below the primary position of gaze, regardless of vision correction type. The third experiment aimed to investigate the real world driving performance of presbyopes while wearing different vision corrections measured on a closed-road circuit at night-time. Eye movements were recorded using the ASL Mobile Eye, eye tracking system (as the faceLAB® system proved to be impractical for use outside of the laboratory). Eleven participants (mean age: 57.25 ± 5.78 years) were fitted with four types of prescribed vision corrections (SV, PAL, MV and MTF CL). The measures of driving performance on the closed-road circuit included distance to sign recognition, near target recognition, peripheral light-emitting-diode (LED) recognition, low contrast road hazards recognition and avoidance, recognition of all the road signs, time to complete the course, and driving behaviours such as braking, accelerating, and cornering. The results demonstrated that driving performance at night was most affected by MTF CL compared to PAL, resulting in shorter distances to read signs, slower driving speeds, and longer times spent fixating road signs. Monovision resulted in worse performance in the task of distance to read a signs compared to SV and PAL. The SV condition resulted in significantly more errors made in interpreting information from in-vehicle devices, despite spending longer time fixating on these devices. Progressive addition lenses were ranked as the most preferred vision correction, while MTF CL were the least preferred vision correction for night-time driving. This thesis addressed the research question of how presbyopic vision corrections affect driving performance and the results of the three experiments demonstrated that the different types of presbyopic vision corrections (e.g. BIF, PAL, MV and MTF CL) can affect driving performance in different ways. Distance-related driving tasks showed reduced performance with MV and MTF CL, while tasks which involved viewing in-vehicle devices were significantly hampered by wearing SV corrections. Wearing spectacles such as SV, BIF and PAL induced greater eye and head movements in the simulated driving condition, however this did not directly translate to impaired performance on the closed- road circuit tasks. These findings are important for understanding the influence of presbyopic vision corrections on vision under real world driving conditions. They will also assist the eye care practitioner to understand and convey to patients the potential driving difficulties associated with wearing certain types of presbyopic vision corrections and accordingly to support them in the process of matching patients to optical corrections which meet their visual needs.

The effect of simulated astigmatic refractive error on reading performance

Background: Few studies have specifically investigated the functional effects of uncorrected astigmatism on measures of reading fluency. This information is important to provide evidence for the development of clinical guidelines for the correction of astigmatism. Methods: Participants included 30 visually normal, young adults (mean age 21.7 ± 3.4 years). Distance and near visual acuity and reading fluency were assessed with optimal spectacle correction (baseline) and for two levels of astigmatism, 1.00DC and 2.00DC, at two axes (90° and 180°) to induce both against-the-rule (ATR) and with-the-rule (WTR) astigmatism. Reading and eye movement fluency were assessed using standardized clinical measures including the test of Discrete Reading Rate (DRR), the Developmental Eye Movement (DEM) test and by recording eye movement patterns with the Visagraph (III) during reading for comprehension. Results: Both distance and near acuity were significantly decreased compared to baseline for all of the astigmatic lens conditions (p < 0.001). Reading speed with the DRR for N16 print size was significantly reduced for the 2.00DC ATR condition (a reduction of 10%), while for smaller text sizes reading speed was reduced by up to 24% for the 1.00DC ATR and 2.00DC condition in both axis directions (p<0.05). For the DEM, sub-test completion speeds were significantly impaired, with the 2.00DC condition affecting both vertical and horizontal times and the 1.00DC ATR condition affecting only horizontal times (p<0.05). Visagraph reading eye movements were not significantly affected by the induced astigmatism. Conclusions: Induced astigmatism impaired performance on selected tests of reading fluency, with ATR astigmatism having significantly greater effects on performance than did WTR, even for relatively small amounts of astigmatic blur of 1.00DC. These findings have implications for the minimal prescribing criteria for astigmatic refractive errors.

The relationship between CFH and ARMS2 genotypes and neuroretinal function in persons without age-related macular degeneration

Purpose: To determine whether neuroretinal function differs in healthy persons with and without common risk gene variants for age- related macular degeneration (AMD) and no ophthalmoscopic signs of AMD, and to compare those findings in persons with manifest early AMD. Methods and Participants: Neuroretinal function was assessed with the multifocal electroretinogram (mfERG) (VERIS, Redwood City, CA,) in 32 participants (22 healthy persons with no clinical signs of AMD and 10 early AMD patients). The 22 healthy participants with no AMD were risk genotypes for either the CFH (rs380390) and/or ARMS2 (rs10490920). We used a slow flash mfERG paradigm (3 inserted frames) and a 103 hexagon stimulus array. Recordings were made with DTL electrodes; fixation and eye movements were monitored online. Trough N1 to peak P1 (N1P1) response densities and P1-implicit times (IT) were analysed in 5 concentric rings. Results: N1P1 response densities (mean ± SD) for concentric rings 1-3 were on average significantly higher in at-risk genotypes (ring 1: 17.97 nV/deg2 ± 1.9, ring 2: 11.7 nV/deg2 ±1.3, ring 3: 8.7 nV/deg2 ± 0.7) compared to those without risk (ring 1: 13.7 nV/deg2 ± 1.9, ring 2: 9.2 nV/deg2 ±0.8, ring 3: 7.3 nV/deg2 ± 1.1) and compared to persons with early AMD (ring 1: 15.3 nV/deg2 ± 4.8, ring 2: 9.1 nV/deg2 ±2.3, ring 3 nV/deg2: 7.3± 1.3) (p<0.5). The group implicit times, P1-ITs for ring 1 were on average delayed in the early AMD patients (36.4 ms ± 1.0) compared to healthy participants with (35.1 ms ± 1.1) or without risk genotypes (34.8 ms ±1.3), although these differences were not significant. Conclusion: Neuroretinal function in persons with normal fundi can be differentiated into subgroups based on their genetics. Increased neuroretinal activity in persons who carry AMD risk genotypes may be due to genetically determined subclinical inflammatory and/or histological changes in the retina. Assessment of neuroretinal function in healthy persons genetically susceptible to AMD may be a useful early biomarker before there is clinical manifestation of AMD.

The impact of interface modality on police officers' visual behaviour when using an in-vehicle system

Background Standard operating procedures state that police officers should not drive while interacting with their mobile data terminal (MDT) which provides in-vehicle information essential to police work. Such interactions do however occur in practice and represent a potential source of driver distraction. The MDT comprises visual output with manual input via touch screen and keyboard. This study investigated the potential for alternative input and output methods to mitigate driver distraction with specific focus on eye movements. Method Nineteen experienced drivers of police vehicles (one female) from the NSW Police Force completed four simulated urban drives. Three drives included a concurrent secondary task: imitation licence plate search using an emulated MDT. Three different interface methods were examined: Visual-Manual, Visual-Voice, and Audio-Voice (“Visual” and “Audio” = output modality; “Manual” and “Voice” = input modality). During each drive, eye movements were recorded using FaceLAB™ (Seeing Machines Ltd, Canberra, ACT). Gaze direction and glances on the MDT were assessed. Results The Visual-Voice and Visual-Manual interfaces resulted in a significantly greater number of glances towards the MDT than Audio-Voice or Baseline. The Visual-Manual and Visual-Voice interfaces resulted in significantly more glances to the display than Audio-Voice or Baseline. For longer duration glances (>2s and 1-2s) the Visual-Manual interface resulted in significantly more fixations than Baseline or Audio-Voice. The short duration glances (<1s) were significantly greater for both Visual-Voice and Visual-Manual compared with Baseline and Audio-Voice. There were no significant differences between Baseline and Audio-Voice. Conclusion An Audio-Voice interface has the greatest potential to decrease visual distraction to police drivers. However, it is acknowledged that an audio output may have limitations for information presentation compared with visual output. The Visual-Voice interface offers an environment where the capacity to present information is sustained, whilst distraction to the driver is reduced (compared to Visual-Manual) by enabling adaptation of fixation behaviour.

Did you see that? Dissociating advanced visual information and ball flight constrains perception and action processes during one-handed catching

The integration of separate, yet complimentary, cortical pathways appears to play a role in visual perception and action when intercepting objects. The ventral system is responsible for object recognition and identification, while the dorsal system facilitates continuous regulation of action. This dual-system model implies that empirically manipulating different visual information sources during performance of an interceptive action might lead to the emergence of distinct gaze and movement pattern profiles. To test this idea, we recorded hand kinematics and eye movements of participants as they attempted to catch balls projected from a novel apparatus that synchronised or de-synchronised accompanying video images of a throwing action and ball trajectory. Results revealed that ball catching performance was less successful when patterns of hand movements and gaze behaviours were constrained by the absence of advanced perceptual information from the thrower's actions. Under these task constraints, participants began tracking the ball later, followed less of its trajectory, and adapted their actions by initiating movements later and moving the hand faster. There were no performance differences when the throwing action image and ball speed were synchronised or de-synchronised since hand movements were closely linked to information from ball trajectory. Results are interpreted relative to the two-visual system hypothesis, demonstrating that accurate interception requires integration of advanced visual information from kinematics of the throwing action and from ball flight trajectory.

Cerebellar grey matter deficits in first-episode schizophrenia mapped using cortical pattern matching

Cerebellar dysfunction has been proposed to lead to “cognitive dysmetria” in schizophrenia via the cortico-cerebellar-thalamic-cortical circuit, contributing to a range of cognitive and clinical symptoms of the disorder. Here we investigated total cerebellar grey and white matter volumes and cerebellar regional grey matter abnormalities in 13 remitted first-episode schizophrenia patients with less than 2 years’ duration of illness. Patient data were compared to 13 pair-wise age, gender, and handedness-matched healthy volunteers using cortical pattern averaging on high-resolution magnetic resonance images. Total cerebellar volume and total grey matter volumes in first-episode schizophrenia patients did not differ from healthy control subjects, but total cerebellar white matter was increased and total grey to white matter ratios were reduced in patients. Four clusters of cerebellar grey matter reduction were identified: (i) in superior vermis; (ii) in the left lobuli VI; (iii) in right-inferior lobule IX, extending into left lobule IX; and (iv) bilaterally in the areas of lobuli III, peduncle and left flocculus. Grey matter deficits were particularly prominent in right lobuli III and IX, left flocculus and bilateral pedunculi. These cerebellar areas have been implicated in attention control, emotional regulation, social functioning, initiation of smooth pursuit eye movements, eye-blink conditioning, language processing, verbal memory, executive function and the processing of spatial and emotional information. Consistent with common clinical, cognitive, and pathophysiological signs of established illness, our findings demonstrate cerebellar pathology as early as in first-episode schizophrenia.

The impact of simulated astigmatism on functional measures of visual performance

Uncorrected refractive error, including astigmatism, is a leading cause of reversible visual impairment. While the ability to perform vision-related daily activities is reduced when people are not optimally corrected, only limited research has investigated the impact of uncorrected astigmatism. Given the capacity to perform vision-related daily activities involves integration of a range of visual and cognitive cues, this research examined the impact of simulated astigmatism on visual tasks that also involved cognitive input. The research also examined whether the higher levels of complexity inherent in Chinese characters makes them more susceptible to the effects of astigmatism. The effects of different powers of astigmatism, as well as astigmatism at different axes were investigated in order to determine the minimum level of astigmatism that resulted in a decrement in visual performance.

Functional imaging of peripheral vision and dorsal stream function in the human cerebral cortex

Visual information processing in brain proceeds in both serial and parallel fashion throughout various functionally distinct hierarchically organised cortical areas. Feedforward signals from retina and hierarchically lower cortical levels are the major activators of visual neurons, but top-down and feedback signals from higher level cortical areas have a modulating effect on neural processing. My work concentrates on visual encoding in hierarchically low level cortical visual areas in human brain and examines neural processing especially in cortical representation of visual field periphery. I use magnetoencephalography and functional magnetic resonance imaging to measure neuromagnetic and hemodynamic responses during visual stimulation and oculomotor and cognitive tasks from healthy volunteers. My thesis comprises six publications. Visual cortex forms a great challenge for modeling of neuromagnetic sources. My work shows that a priori information of source locations are needed for modeling of neuromagnetic sources in visual cortex. In addition, my work examines other potential confounding factors in vision studies such as light scatter inside the eye which may result in erroneous responses in cortex outside the representation of stimulated region, and eye movements and attention. I mapped cortical representations of peripheral visual field and identified a putative human homologue of functional area V6 of the macaque in the posterior bank of parieto-occipital sulcus. My work shows that human V6 activates during eye-movements and that it responds to visual motion at short latencies. These findings suggest that human V6, like its monkey homologue, is related to fast processing of visual stimuli and visually guided movements. I demonstrate that peripheral vision is functionally related to eye-movements and connected to rapid stream of functional areas that process visual motion. In addition, my work shows two different forms of top-down modulation of neural processing in the hierachically lowest cortical levels; one that is related to dorsal stream activation and may reflect motor processing or resetting signals that prepare visual cortex for change in the environment and another local signal enhancement at the attended region that reflects local feed-back signal and may perceptionally increase the stimulus saliency.

Correction of Ocular Artifacts in EEG Recordings using Empirical Mode Decomposition

EEG recordings are often contaminated with ocular artifacts such as eye blinks and eye movements. These artifacts may obscure underlying brain activity in the electroencephalogram (EEG) data and make the analysis of the data difficult. In this paper, we explore the use of empirical mode decomposition (EMD) based filtering technique to correct the eye blinks and eye movementartifacts in single channel EEG data. In this method, the single channel EEG data containing ocular artifact is segmented such that the artifact in each of the segment is considered as some type of slowly varying trend in the dataand the EMD is used to remove the trend. The filtering is done using partial reconstruction from components of the decomposition. The method is completely data dependent and hence adaptive and nonlinear. Experimental results are provided to check the applicability of the method on real EEG data and the results are quantified using power spectral density (PSD) as a measure. The method has given fairlygood results and does not make use of any preknowledge of artifacts or the EEG data used.

Online Estimation of Evolving Human Visual Interest

Regions in video streams attracting human interest contribute significantly to human understanding of the video. Being able to predict salient and informative Regions of Interest (ROIs) through a sequence of eye movements is a challenging problem. Applications such as content-aware retargeting of videos to different aspect ratios while preserving informative regions and smart insertion of dialog (closed-caption text) into the video stream can significantly be improved using the predicted ROIs. We propose an interactive human-in-the-loop framework to model eye movements and predict visual saliency into yet-unseen frames. Eye tracking and video content are used to model visual attention in a manner that accounts for important eye-gaze characteristics such as temporal discontinuities due to sudden eye movements, noise, and behavioral artifacts. A novel statistical-and algorithm-based method gaze buffering is proposed for eye-gaze analysis and its fusion with content-based features. Our robust saliency prediction is instantiated for two challenging and exciting applications. The first application alters video aspect ratios on-the-fly using content-aware video retargeting, thus making them suitable for a variety of display sizes. The second application dynamically localizes active speakers and places dialog captions on-the-fly in the video stream. Our method ensures that dialogs are faithful to active speaker locations and do not interfere with salient content in the video stream. Our framework naturally accommodates personalisation of the application to suit biases and preferences of individual users.

Contributions of parietal cortex to reach planning

Sensory-motor circuits course through the parietal cortex of the human and monkey brain. How parietal cortex manipulates these signals has been an important question in behavioral neuroscience. This thesis presents experiments that explore the contributions of monkey parietal cortex to sensory-motor processing, with an emphasis on the area's contributions to reaching. First, it is shown that parietal cortex is organized into subregions devoted to specific movements. Area LIP encodes plans to make saccadic eye movements. A nearby area, the parietal reach region (PRR), plans reaches. A series of experiments are then described which explore the contributions of PRR to reach planning. Reach plans are represented in an eye-centered reference frame in PRR. This representation is shown to be stable across eye movements. When a sequence of reaches is planned, only the impending movement is represented in PRR, showing that the area is more related to movement planning than to storing the memory of reach targets. PRR resembles area LIP in each of these properties: the two areas may provide a substrate for hand-eye coordination. These findings yield new perspectives on the functions of the parietal cortex and on the organization of sensory-motor processing in primate brains.

Analog VLSI-based, neuromorphic sensorimotor systems: modeling the primate oculomotor system

Using neuromorphic analog VLSI techniques for modeling large neural systems has several advantages over software techniques. By designing massively-parallel analog circuit arrays which are ubiquitous in neural systems, analog VLSI models are extremely fast, particularly when local interactions are important in the computation. While analog VLSI circuits are not as flexible as software methods, the constraints posed by this approach are often very similar to the constraints faced by biological systems. As a result, these constraints can offer many insights into the solutions found by evolution. This dissertation describes a hardware modeling effort to mimic the primate oculomotor system which requires both fast sensory processing and fast motor control. A one-dimensional hardware model of the primate eye has been built which simulates the physical dynamics of the biological system. It is driven by analog VLSI circuits mimicking brainstem and cortical circuits that control eye movements. In this framework, a visually-triggered saccadic system is demonstrated which generates averaging saccades. In addition, an auditory localization system, based on the neural circuits of the barn owl, is used to trigger saccades to acoustic targets in parallel with visual targets. Two different types of learning are also demonstrated on the saccadic system using floating-gate technology allowing the non-volatile storage of analog parameters directly on the chip. Finally, a model of visual attention is used to select and track moving targets against textured backgrounds, driving both saccadic and smooth pursuit eye movements to maintain the image of the target in the center of the field of view. This system represents one of the few efforts in this field to integrate both neuromorphic sensory processing and motor control in a closed-loop fashion.

Responses to auditory stimuli in macaque lateral intraparietal area

The lateral intraparietal area (LIP) of macaque posterior parietal cortex participates in the sensorimotor transformations underlying visually guided eye movements. Area LIP has long been considered unresponsive to auditory stimulation. However, recent studies have shown that neurons in LIP respond to auditory stimuli during an auditory-saccade task, suggesting possible involvement of this area in auditory-to-oculomotor as well as visual-to-oculomotor processing. This dissertation describes investigations which clarify the role of area LIP in auditory-to-oculomotor processing.

Extracellular recordings were obtained from a total of 332 LIP neurons in two macaque monkeys, while the animals performed fixation and saccade tasks involving auditory and visual stimuli. No auditory activity was observed in area LIP before animals were trained to make saccades to auditory stimuli, but responses to auditory stimuli did emerge after auditory-saccade training. Auditory responses in area LIP after auditory-saccade training were significantly stronger in the context of an auditory-saccade task than in the context of a fixation task. Compared to visual responses, auditory responses were also significantly more predictive of movement-related activity in the saccade task. Moreover, while visual responses often had a fast transient component, responses to auditory stimuli in area LIP tended to be gradual in onset and relatively prolonged in duration.

Overall, the analyses demonstrate that responses to auditory stimuli in area LIP are dependent on auditory-saccade training, modulated by behavioral context, and characterized by slow-onset, sustained response profiles. These findings suggest that responses to auditory stimuli are best interpreted as supramodal (cognitive or motor) responses, rather than as modality-specific sensory responses. Auditory responses in area LIP seem to reflect the significance of auditory stimuli as potential targets for eye movements, and may differ from most visual responses in the extent to which they arc abstracted from the sensory parameters of the stimulus.

Macaque lateral intraparietal area and oculomotor behaviors

Neurons in the primate lateral intraparietal area (area LIP) carry visual, saccade-related and eye position activities. The visual and saccade activities are anchored in a retinotopic framework and the overall response magnitude is modulated by eye position. It was proposed that the modulation by eye position might be the basis of a distributed coding of target locations in a head-centered space. Other recording studies demonstrated that area LIP is involved in oculomotor planning. These results overall suggest that area LIP transforms sensory information for motor functions. In this thesis I further explore the role of area LIP in processing saccadic eye movements by observing the effects of reversible inactivation of this area. Macaque monkeys were trained to do visually guided and memory saccades and a double saccade task to examine the use of eye position signal. Finally, by intermixing visual saccades with trials in which two targets were presented at opposite sides of the fixation point, I examined the behavior of visual extinction.

In chapter 2, I will show that lesion of area LIP results in increased latency of contralesional visual and memory saccades. Contralesional memory saccades are also hypometric and slower in velocity. Moreover, the impairment of memory saccades does not vary with the duration of the delay period. This suggests that the oculomotor deficits observed after inactivation of area LIP is not due to the disruption of spatial memory.

In chapter 3, I will show that lesion of area LIP does not severely affect the processing of spontaneous eye movement. However, the monkeys made fewer contralesional saccades and tended to confine their gaze to the ipsilesional field after inactivation of area LIP. On the other hand, lesion of area LIP results in extinction of the contralesional stimulus. When the initial fixation position was varied so that the retinal and spatial locations of the targets could be dissociated, it was found that the extinction behavior could best be described in a head-centered coordinate.

In chapter 4, I will show that inactivation of area LIP disrupts the use of eye position signal to compute the second movement correctly in the double saccade task. If the first saccade steps into the contralesional field, the error rate and latency of the second saccade are both increased. Furthermore, the direction of the first eye movement largely does not have any effect on the impairment of the second saccade. I will argue that this study provides important evidence that the extraretinal signal used for saccadic localization is eye position rather than a displacement vector.

In chapter 5, I will demonstrate that in parietal monkeys the eye drifts toward the lesion side at the end of the memory saccade in darkness. This result suggests that the eye position activity in the posterior parietal cortex is active in nature and subserves gaze holding.

Overall, these results further support the view that area LIP neurons encode spatial locations in a craniotopic framework and is involved in processing voluntary eye movements.