Spatial Navigation Based on Novelty Mediated Autobiographical Memory

Abstract. This paper presents a method for spatial navigation performed mainly on past experiences. The past experiences are remembered in their temporal context, i.e. as episodes of events. The learned episodes form an ac-tive autobiography that determines the future navigation behaviour. The epi-sodic and autobiographical memories are modelled to resemble the memory formation process that takes place in the rat hippocampus. The method im-plies naturally inferential reasoning in the robotic framework that may make it more flexible for navigation in unseen environments. The relation between novelty and life-long exploratory (latent) learning is shown to be important and therefore is incorporated into the learning process. As a result, active au-tobiography formation depends on latent learning while individual trials might be reward driven. The experimental results show that learning mediat-ed by novelty provides a flexible and efficient way to encode spatial informa-tion in its contextual relatedness and directionality. Therefore, performing a novel task is fast but solution is not optimal. In addition, learning becomes naturally a continuous process- encoding and retrieval phase have the same underlying mechanism, and thus do not need to be separated. Therefore, building a “life long ” autobiography is feasible.

The spectro-contextual encoding and retrieval theory of episodic memory.

The spectral fingerprint hypothesis, which posits that different frequencies of oscillations underlie different cognitive operations, provides one account for how interactions between brain regions support perceptual and attentive processes (Siegel etal., 2012). Here, we explore and extend this idea to the domain of human episodic memory encoding and retrieval. Incorporating findings from the synaptic to cognitive levels of organization, we argue that spectrally precise cross-frequency coupling and phase-synchronization promote the formation of hippocampal-neocortical cell assemblies that form the basis for episodic memory. We suggest that both cell assembly firing patterns as well as the global pattern of brain oscillatory activity within hippocampal-neocortical networks represents the contents of a particular memory. Drawing upon the ideas of context reinstatement and multiple trace theory, we argue that memory retrieval is driven by internal and/or external factors which recreate these frequency-specific oscillatory patterns which occur during episodic encoding. These ideas are synthesized into a novel model of episodic memory (the spectro-contextual encoding and retrieval theory, or "SCERT") that provides several testable predictions for future research

Segmentation of experience and episodic memory across species

How continuous ongoing perceptual experience is processed by the brain and mind to form unique episodes in memory is a key scientific question. Recent work in Psychology and Neuroscience has proposed that humans perceptually segment continuous ongoing experience into meaningful units, which allows the successful formation of episodic memories. Despite accumulating work demonstrating that non- human animals also display a capability of episodic-‘like’ memory, whether non-human animals segment continuous ongoing experience into ‘meaningful’ episodic units is a question that has not been fully explored. Hence, the main goal of the research in this thesis aims to address whether a comparable segmentation process (or processes) of continuous ongoing experience occurs for non-human animals in their formation of episodic-like memory, as it does for humans in their formation of episodic memory. Chapter 2 argues that, similarly to humans, rats can use top-down like prediction-error processing in segmenting for subsequent memory to guide behaviour in an episodic-like spontaneous object recognition task. Chapter 3 suggests that mice readily incorporate conspecific-contextual information using episodic-like memory processing, indicating that conspecifics can act as a segmentation cue for non-human animals. Chapter 4 highlights that humans and rodents may similarly segment continuous ongoing experience during turns made around spatial boundaries. Chapter 5 argues that individual place cells can represent content of episodic nature, with the theoretical implication of this being discussed in relation to episodic memory. Thus, the results presented in this thesis, as well as re-interpretation of previous literature, would argue in favour of non-humans segmenting their experience for episodic-like memory. Finally, the evidence is evaluated in the context of whether episodic-like memory in non-human animals is simply just episodic memory as experienced in humans

Scene construction impairments in Alzheimer's disease – A unique role for the posterior cingulate cortex

Episodic memory dysfunction represents one of the most prominent and characteristic clinical features of patients with Alzheimer's disease (AD), attributable to the degeneration of medial temporal and posterior parietal regions of the brain. Recent studies have demonstrated marked impairments in the ability to envisage personally relevant events in the future in AD. It remains unclear, however, whether AD patients can imagine fictitious scenes free from temporal constraints, a process that is proposed to rely fundamentally upon the integrity of the hippocampus. The objective of the present study was to investigate the capacity for atemporal scene construction, and its associated neural substrates, in AD. Fourteen AD patients were tested on the scene construction task and their performance was contrasted with 14 age- and education-matched healthy older Control participants. Scene construction performance was strikingly compromised in the AD group, with significant impairments evident for provision of contextual details, spatial coherence, and the overall richness of the imagined experience. Voxel-based morphometry analyses based on structural MRI revealed significant associations between scene construction capacity and atrophy in posterior parietal and lateral temporal brain structures in AD. In contrast, scene construction performance in Controls was related to integrity of frontal, parietal, and medial temporal structures, including the parahippocampal gyrus and posterior hippocampus. The posterior cingulate cortex (PCC) emerged as the common region implicated for scene construction performance across participant groups. Our study highlights the importance of regions specialised for spatial and contextual processing for the construction of atemporal scenes. Damage to these regions in AD compromises the ability to construct novel scenes, leading to the recapitulation of content from previously experienced events

The distinct and overlapping brain networks supporting semantic and spatial constructive scene processing

Scene imagery features prominently when we recall autobiographical memories, imagine the future and navigate around in the world. Consequently, in this study we sought to better understand how scene representations are supported by the brain. Processing scenes involves a variety of cognitive processes that in the real world are highly interactive. Here, however, our goal was to separate semantic and spatial constructive scene processes in order to identify the brain areas that were distinct to each process, those they had in common, and the connectivity between regions. To this end, participants searched for either semantic or spatial constructive impossibilities in scenes during functional MRI. We focussed our analyses on only those scenes that were possible, thus removing any error detection that would evoke reactions such as surprise or novelty. Importantly, we also counterbalanced possible scenes across participants, enabling us to examine brain activity and connectivity for the same possible scene images under two different conditions. We found that participants adopted different cognitive strategies, which were reflected in distinct oculomotor behaviour, for each condition. These were in turn associated with increased engagement of lateral temporal and parietal cortices for semantic scene processing, the hippocampus for spatial constructive scene processing, and increased activation of the ventromedial prefrontal cortex (vmPFC) that was common to both. Connectivity analyses showed that the vmPFC switched between semantic and spatial constructive brain networks depending on the task at hand. These findings further highlight the well-known semantic functions of lateral temporal areas, while providing additional support for the previously-asserted contribution of the hippocampus to scene construction, and recent suggestions that the vmPFC may play a key role in orchestrating scene processing

Representational dynamics across multiple timescales in human cortical networks

Human cognition occurs at multiple timescales, including immediate processing of the ongoing experiences and slowly drifting higher-level thoughts. To understand how the brain selects and represents these various types of information to guide behavior, this thesis examined representational content within sensory regions, multiple demand (MD) network, and default mode network (DMN). Chapter 1 provides a background review of the current literature. It begins by reviewing experimental investigations of component visual processes that unfold over time. Next, the MD network is introduced as a collection of frontal and parietal regions involved in implementing cognitive control by assembling the required operations for task-relevant behavior. Finally, the DMN is introduced in the context of temporal processing hierarchies, with focus on its representation of situation models summarizing interactions among entities and the environment. The first experiment, presented in Chapter 2, used EEG/MEG to track multiple component processes of selective attention. Five distinct processing operations with different time-courses were quantified, including representation of visual display properties, target location, target identity, behavioral significance, and finally, possible reactivation of the attentional template. Chapter 3 used fMRI to examine neural representations of task episodes, which are temporally organized sequences of steps that occur within a given context. It was found that MD and visual regions showed sensitivity to the fine structure of the contents within a task. DMN regions showed gradual change throughout the entire task, with increased activation at the offset of the entire episode. Chapter 4 analyzed activation profiles of DMN regions using six diverse tasks to examine their functional convergence during social, episodic, and self-referential thought. Results supported proposals of separate subsystems, yet also suggest integration within the DMN. The final chapter, Chapter 5, provides an extended discussion of theoretical concepts related to the three experiments and proposes possible avenues for further research

Neural processes underpinning episodic memory

Episodic memory is the memory for our personal past experiences. Although numerous functional magnetic resonance imaging (fMRI) studies investigating its neural basis have revealed a consistent and distributed network of associated brain regions, surprisingly little is known about the contributions individual brain areas make to the recollective experience. In this thesis I address this fundamental issue by employing a range of different experimental techniques including neuropsychological testing, virtual reality environments, whole brain and high spatial resolution fMRI, and multivariate pattern analysis. Episodic memory recall is widely agreed to be a reconstructive process, one that is known to be critically reliant on the hippocampus. I therefore hypothesised that the same neural machinery responsible for reconstruction might also support ‘constructive’ cognitive functions such as imagination. To test this proposal, patients with focal damage to the hippocampus bilaterally were asked to imagine new experiences and were found to be impaired relative to matched control participants. Moreover, driving this deficit was a lack of spatial coherence in their imagined experiences, pointing to a role for the hippocampus in binding together the disparate elements of a scene. A subsequent fMRI study involving healthy participants compared the recall of real memories with the construction of imaginary memories. This revealed a fronto-temporo-parietal network in common to both tasks that included the hippocampus, ventromedial prefrontal, retrosplenial and parietal cortices. Based on these results I advanced the notion that this network might support the process of ‘scene construction’, defined as the generation and maintenance of a complex and coherent spatial context. Furthermore, I argued that this scene construction network might underpin other important cognitive functions besides episodic memory and imagination, such as navigation and thinking about the future. It is has been proposed that spatial context may act as the scaffold around which episodic memories are built. Given the hippocampus appears to play a critical role in imagination by supporting the creation of a rich coherent spatial scene, I sought to explore the nature of this hippocampal spatial code in a novel way. By combining high spatial resolution fMRI with multivariate pattern analysis techniques it proved possible to accurately determine where a subject was located in a virtual reality environment based solely on the pattern of activity across hippocampal voxels. For this to have been possible, the hippocampal population code must be large and non-uniform. I then extended these techniques to the domain of episodic memory by showing that individual memories could be accurately decoded from the pattern of activity across hippocampal voxels, thus identifying individual memory traces. I consider these findings together with other recent advances in the episodic memory field, and present a new perspective on the role of the hippocampus in episodic recollection. I discuss how this new (and preliminary) framework compares with current prevailing theories of hippocampal function, and suggest how it might account for some previously contradictory data

From cognitive maps to spatial schemas

A schema refers to a structured body of prior knowledge that captures common patterns across related experiences. Schemas have been studied separately in the realms of episodic memory and spatial navigation across different species and have been grounded in theories of memory consolidation, but there has been little attempt to integrate our understanding across domains, particularly in humans. We propose that experiences during navigation with many similarly structured environments give rise to the formation of spatial schemas (for example, the expected layout of modern cities) that share properties with but are distinct from cognitive maps (for example, the memory of a modern city) and event schemas (such as expected events in a modern city) at both cognitive and neural levels. We describe earlier theoretical frameworks and empirical findings relevant to spatial schemas, along with more targeted investigations of spatial schemas in human and non-human animals. Consideration of architecture and urban analytics, including the influence of scale and regionalization, on different properties of spatial schemas may provide a powerful approach to advance our understanding of spatial schemas

On the role of the hippocampus in the acquisition, long-term retention and semanticisation of memory

Institute for Adaptive and Neural ComputationA consensus on how to characterise the anterograde and retrograde memory processes that are lost or spared after hippocampal damage has not been reached. In this thesis, I critically re-examine the empirical literature and the assumptions behind current theories. I formulate a coherent view of what makes a task hippocampally dependent at acquisition and how this relates to its long-term fate. Findings from a neural net simulation indicate the plausibility of my proposals. My proposals both extend and constrain current views on the role of the hippocampus in the rapid acquisition of information and in learning complex associations. In general, tasks are most likely to require the hippocampus for acquisition if they involve rapid, associative learning about unfamiliar, complex, low salience stimuli. However, none of these factors alone is sufficient to obligatorily implicate the hippocampus in acquisition. With the exception of associations with supra-modal information that are always dependent on the hippocampus, it is the combination of factors that is important. Detailed, complex information that is obligatorily hippocampally-dependent at acquisition remains so for its lifetime. However, all memories are semanticised as they age through the loss of detailed context-specific information and because generic cortically-represented information starts to dominate recall. Initially hippocampally dependent memories may appear to become independent of the hippocampus over time, but recall changes qualitatively. Multi-stage, lifelong post-acquisition memory processes produce semanticised re-representations of memories of differing specificity and complexity, that can serve different purposes. The model simulates hippocampal and cortical interactions in the acquisition and maintenance of episodic and semantic events, and behaves in accordance with my proposals. In particular, conceptualising episodic and semantic memory as representing points on a continuum of memory types appears viable. Support is also found for proposals on the relative importance of the hippocampus and cortex in the rapid acquisition of information and the acquisition of complex multi-model information; and the effect of existing knowledge on new learning. Furthermore, episodic and semantic events become differentially dependent on cortical and hippocampal components. Finally, as a memory ages, it is automatically semanticised and becomes cortically dependent

Constructing, Perceiving, and Maintaining Scenes: Hippocampal Activity and Connectivity.

In recent years, evidence has accumulated to suggest the hippocampus plays a role beyond memory. A strong hippocampal response to scenes has been noted, and patients with bilateral hippocampal damage cannot vividly recall scenes from their past or construct scenes in their imagination. There is debate about whether the hippocampus is involved in the online processing of scenes independent of memory. Here, we investigated the hippocampal response to visually perceiving scenes, constructing scenes in the imagination, and maintaining scenes in working memory. We found extensive hippocampal activation for perceiving scenes, and a circumscribed area of anterior medial hippocampus common to perception and construction. There was significantly less hippocampal activity for maintaining scenes in working memory. We also explored the functional connectivity of the anterior medial hippocampus and found significantly stronger connectivity with a distributed set of brain areas during scene construction compared with scene perception. These results increase our knowledge of the hippocampus by identifying a subregion commonly engaged by scenes, whether perceived or constructed, by separating scene construction from working memory, and by revealing the functional network underlying scene construction, offering new insights into why patients with hippocampal lesions cannot construct scenes