Lateral intraparietal area (LIP) neurons are thought to compute the decision of where to look. Specifically, their firing rate is thought to represent accumulated evidence for a decision by ramping up to a high level, the putative decision bound, before an eye movement to a given location. However, LIP neurons are also highly responsive to visual stimuli. Because all previous research put a visual stimulus (a saccadic target) in the response field (RF) location of a neuron during decision formation, it is unknown if LIP neurons can still show decision computation without this visual drive. We therefore recorded the spiking activity of LIP neurons in a conventional decision task where a monkey decides the direction of a noisy motion stimulus and indicates his decision with a saccade. On half the trials, the Choice Targets remained on for the whole trial, as is conventionally done. On the remaining trials, targets were flashed at the beginning of the trial and absent during motion-viewing. Furthermore, we recorded the activity of any neuron with an RF, instead of only neurons exhibiting persistent memory activity before an instructed saccade. This enabled us to also test the long-held assumption that only cells with persistent memory activity show decision signals. Our results show that 1) cells without persistent activity indeed show decision signals, 2) population response drops without RF stimulation (although individual neurons were affected in diverse ways), 3) distinct, repeating response “motifs” exist among cells, 4) a signal exists where neural response is lower for stronger motion strength stimuli, regardless of direction. These results prove that contrary to dogma, a neuron’s ability to show a long “memory” response is not related to an ability to accumulate evidence over time for a decision. Also, LIP firing rate cannot be interpreted as a pure decision variable because it simultaneously represents decision-irrelevant, visual stimuli. Finally, diverse, but repeating responses among cells suggest the existence of cell types in LIP. These results demonstrate that LIP acts as a bank of potentially useful signals, and raises the question of how they might be used for a decision. / text

In many everyday activities, we must visually process multiple objects embedded in complex real world scenes. Our visual system can flexibly extract behaviorally relevant visual information from such scenes, even though it has a severely limited processing capacity. This dissertation proposes that human superior intra-parietal sulcus (IPS) plays a central role in this flexible visual information processing. In Chapter 1, using functional magnetic resonance imaging (fMRI) with univariate analysis, I found that distractor processing in superior IPS was attenuated when target locations were known in advance. In Chapter 2, using multi-voxel pattern analysis (MVPA), I showed that superior IPS encoded object shapes, but only when such information was required by task. In Chapter 3, I showed that, given a set of perceptually distinct, but semantically grouped visual inputs, superior IPS could represent abstract object identity. The neural similarity of identities in superior IPS significantly correlated with perceived similarity between identities, confirming the representation in this region indeed reflected identity. Taken together, these results suggest that human superior IPS encodes a wide range of visual information, from simple features to abstract identities, in a task-dependent manner, enabling flexible goal-directed visual information processing in the human brain. / Psychology

Recent functional neuroimaging research has highlighted the role of the posterior parietal cortex (PPC) during episodic memory retrieval (Vilberg & Rugg, 2007; Vincent et al., 2006; Wagner et al., 2005; Wheeler & Buckner, 2003; Yonelinas et al., 2005). It has been shown that the PPC is active during the retrieval of old information, when subjects incorrectly believe that new information has been encountered before (false alarms), and when retrieval is accompanied by recollection (Wagner et al., 2005). A preliminary positron emission tomography (PET) study suggests that the PPC is also active when retrieval occurs unintentionally or incidentally, prompted by the presence of an old stimulus item. Collectively these studies suggest that activity in the PPC may be driven by familiarity, irrespective of task demands and independent of its veracity. The main goal of this thesis was to explore, with functional magnetic resonance imaging (fMRI), the conditions under which activity in the PPC may be modulated by the familiarity of veridical and perceived memories. Subjects were presented with scene/word pairs that were repeated 1, 3, or 7 times on Day 1. On Day 2, while MRI scans were recorded, subjects studied either, new items, old pairs (same-form) one additional time (2nd,, 4th, or 8th presentation), or pairs in which the word associated with each picture was changed (different-form) one additional time (2nd,, 4th, or 8th presentation). Same form pairs activated bilateral inferior parietal lobule compared to novel pairs replicating the findings from the PET study on incidental retrieval. Same and different form pairs did not differ on amount of parietal activity. Additionally, same form pairs that were repeated 8 times displayed more PPC activity than same form pairs repeated two and four times while there was no difference across different form repetitions. Together these results indicate that items that are old and that those that are new but are perceived as old activate the inferior parietal lobule.

Superior parietal cortex has been implicated in visuomotor guidance and is proposed to be specialised for action in the lower visual field and peripersonal space. Two patients, one with a right superior parietal lesion leading to optic ataxia (ME), and one with a lesion affecting right inferior parietal cortex (LH), were compared to elderly controls (n=8) and young controls (n=8) on a reciprocal pointing task with movements made in the near-far direction (i.e., sagittal plane) or right-left direction (i.e., fronto-parallel plane). In contrast to both control groups, who demonstrated a speed-accuracy trade-off in movement time and peak velocity, neither of the patients did. When the time spent post-peak velocity (represented as a percentage of total movement time) was examined, both patients demonstrated larger times post-peak velocity than controls for all movement directions. Furthermore, while rightward movements of the right hand had higher times post-peak velocity than leftward movements there were no directional patterns for near-far movements which contrasted with controls who had larger times post-peak velocity for near movements. The patient with the more superior lesion (ME) had the greatest difficulty with movements made back toward the body suggestive of a role for superior parietal cortex in the fine tuning of movements made in this region of space (i.e., personal or peripersonal space). In contrast, all directions of movement seemed to be equally affected in the patient with a more inferior lesion. These results are discussed in terms of the different roles played by inferior and superior parietal cortex in the control of visually guided movements.

Superior parietal cortex has been implicated in visuomotor guidance and is proposed to be specialised for action in the lower visual field and peripersonal space. Two patients, one with a right superior parietal lesion leading to optic ataxia (ME), and one with a lesion affecting right inferior parietal cortex (LH), were compared to elderly controls (n=8) and young controls (n=8) on a reciprocal pointing task with movements made in the near-far direction (i.e., sagittal plane) or right-left direction (i.e., fronto-parallel plane). In contrast to both control groups, who demonstrated a speed-accuracy trade-off in movement time and peak velocity, neither of the patients did. When the time spent post-peak velocity (represented as a percentage of total movement time) was examined, both patients demonstrated larger times post-peak velocity than controls for all movement directions. Furthermore, while rightward movements of the right hand had higher times post-peak velocity than leftward movements there were no directional patterns for near-far movements which contrasted with controls who had larger times post-peak velocity for near movements. The patient with the more superior lesion (ME) had the greatest difficulty with movements made back toward the body suggestive of a role for superior parietal cortex in the fine tuning of movements made in this region of space (i.e., personal or peripersonal space). In contrast, all directions of movement seemed to be equally affected in the patient with a more inferior lesion. These results are discussed in terms of the different roles played by inferior and superior parietal cortex in the control of visually guided movements.

Successful recognition memory decisions depend on mnemonic and decision making processes that are computed by multiple, distributed brain areas. However, little is known about what computations these areas perform or how these areas are connected. Here, I collected behavioral and functional magnetic resonance imaging data from humans during the performance of an old-new recognition memory task with retrospective confidence judgments. Across runs, choice bias was successfully manipulated by providing rewards for correct responses that were either symmetric (equal reward for hits and correct rejections) or asymmetric (one response worth more than the other). Successful recognition memory was associated with activation in anterior prefrontal, parahippocampal, posterior cingulate, and parietal cortex. Resting state functional connectivity demonstrated that these brain areas are organized into two distinct networks. The first network includes parahippocampal cortex and angular gyrus. The second network includes lateral prefrontal cortex and intraparietal sulcus. The hippocampal-cortical network was most active during old vs. new decisions, did not differentiate hits from false alarms, and was differentially active during low confidence old and new judgments. In contrast, while the frontoparietal network was robustly activated by hits, it was not activated during either false alarms or low confidence old judgments. Thus, these two distinct networks can be distinguished by their relative connectivity to the medial temporal lobe vs. lateral prefrontal cortex and their responses during uncertain old judgments and errors. The choice bias manipulation had opposing effects on the parietal components of these networks, which further suggests these networks make distinct contributions to mnemonic decision making. / Psychology

The posterior parietal cortex (PPC) has been reliably implicated in visuospatial attention, such that the dorsal regions (dPPC) are associated with voluntary ‘top-down’ attention, whereas the ventral regions (vPPC) are associated with automatic ‘bottom-up’ attentional processes. The Attention-to-Memory model (AtoM: Ciaramelli, Grady, & Moscovitch, 2008) has suggested that it also plays a similar role in memory retrieval, suggesting that the PPC mediates a domain-general attentional process. Furthermore, domain-generality of attentional processes may account for differences in perception and memory function accompanying old age. This study examined domain-generality by determining the shared variance in performance of tasks thought to recruit top-down and bottom-up attentional processes mediated across both domains. Results clearly suggested generality across domains in top-down processing; and in bottom-up processing, depending on its operationalization. Ageing was characterized by an absence of shared variance across domains and slower reaction times during bottom-up attentional reorienting only in perception.

The posterior parietal cortex (PPC) has been reliably implicated in visuospatial attention, such that the dorsal regions (dPPC) are associated with voluntary ‘top-down’ attention, whereas the ventral regions (vPPC) are associated with automatic ‘bottom-up’ attentional processes. The Attention-to-Memory model (AtoM: Ciaramelli, Grady, & Moscovitch, 2008) has suggested that it also plays a similar role in memory retrieval, suggesting that the PPC mediates a domain-general attentional process. Furthermore, domain-generality of attentional processes may account for differences in perception and memory function accompanying old age. This study examined domain-generality by determining the shared variance in performance of tasks thought to recruit top-down and bottom-up attentional processes mediated across both domains. Results clearly suggested generality across domains in top-down processing; and in bottom-up processing, depending on its operationalization. Ageing was characterized by an absence of shared variance across domains and slower reaction times during bottom-up attentional reorienting only in perception.

abstract: Understanding where our bodies are in space is imperative for motor control, particularly for actions such as goal-directed reaching. Multisensory integration is crucial for reducing uncertainty in arm position estimates. This dissertation examines time and frequency-domain correlates of visual-proprioceptive integration during an arm-position maintenance task. Neural recordings were obtained from two different cortical areas as non-human primates performed a center-out reaching task in a virtual reality environment. Following a reach, animals maintained the end-point position of their arm under unimodal (proprioception only) and bimodal (proprioception and vision) conditions. In both areas, time domain and multi-taper spectral analysis methods were used to quantify changes in the spiking, local field potential (LFP), and spike-field coherence during arm-position maintenance. In both areas, individual neurons were classified based on the spectrum of their spiking patterns. A large proportion of cells in the SPL that exhibited sensory condition-specific oscillatory spiking in the beta (13-30Hz) frequency band. Cells in the IPL typically had a more diverse mix of oscillatory and refractory spiking patterns during the task in response to changing sensory condition. Contrary to the assumptions made in many modelling studies, none of the cells exhibited Poisson-spiking statistics in SPL or IPL. Evoked LFPs in both areas exhibited greater effects of target location than visual condition, though the evoked responses in the preferred reach direction were generally suppressed in the bimodal condition relative to the unimodal condition. Significant effects of target location on evoked responses were observed during the movement period of the task well. In the frequency domain, LFP power in both cortical areas was enhanced in the beta band during the position estimation epoch of the task, indicating that LFP beta oscillations may be important for maintaining the ongoing state. This was particularly evident at the population level, with clear increase in alpha and beta power. Differences in spectral power between conditions also became apparent at the population level, with power during bimodal trials being suppressed relative to unimodal. The spike-field coherence showed confounding results in both the SPL and IPL, with no clear correlation between incidence of beta oscillations and significant beta coherence. / Dissertation/Thesis / Doctoral Dissertation Biomedical Engineering 2017

The purpose of this project was to compare the Attention to Memory (AtoM) and the Episodic Buffer (EB) Models. The AtoM model proposes that the ventral parietal cortex (VPC) and dorsal parietal cortex (DPC) are responsive to bottom-up and top- down attention to memory, respectively, (Cabeza, 2008; Cabeza et al., 2008; Ciaramelli et al., 2008). The EB model, on the other hand, proposes that the VPC is involved in the episodic buffer component of Baddeley's working memory model (Vilberg & Rugg, 2008). Using objective (source) and subjective (Remember/Know) retrieval tasks, specific patterns of PPC activity were posited based on the propositions of the AtoM model. These expectations included greater VPC activity for Remember and False Alarms compared to Correct Rejections and Subjective Know, greater DPC activity for Know and Objective Remember compared to Subjective Remember and correct rejections, and no difference in VPC activity for remembering both font and color compared to remembering only one contextual detail. During encoding participants saw words in one of two colors, red or yellow, and in one of two fonts, curvy or straight, and were required to indicate the color the word was presented in. Following each encoding scan participants performed either an Objective or Subjective retrieval task. During Objective retrieval task, participants performed a forced-choice source memory test choosing the word with the correct fontand color or the "new" option. During Subjective retrieval participants were presented with the word in a neutral font and white color and performed a Remember/Know test. On the Subjective retrieval task both VPC and DPC were active for recollection compared to familiar items and Correct Rejections. On the Objective retrieval task the DPC was active for all correct old responses. Neither the VPC nor the DPC were significantly active for False Alarms on both the Subjective and Objective tasks. Both VPC and DPC were more active for Subjective Remember compared to Objective Remember response. Neither PPC region was more active for remembering font and color compared to remembering only font or color. Memory load effects for retrieval of information from long-term memory were only seen in the hippocampus on the Subjective retrieval task. These patterns of activity support the role of the VPC in recollection, as seen on the Subjective task, and the role of the DPC in familiarity, as shown in both the Subjective and Objective tasks. The role of the VPC and DPC during recollection and familiarity processing supports both the AtoM and the EB model. The key predictor of the Episodic Buffer model, memory load effects, was not supported and provides the only evidence against one of the two proposed models. Future work should examine the role of the posterior parietal cortex in spontaneous episodic retrieval to assess the validity of the AtoM model. Advanced imaging analysis techniques should be used to determine functional connectivity between the PPC and frontal and temporal memory regions.