Using hippocampal-striatal loops for spatial navigation and goal-directed decision-making

The hippocampus plays a central role in spatial representation, declarative and episodic memory. In this area, so-called place cells possess high spatial selectivity, firing preferentially when the individual is within a small area of the environment. Interestingly, it has been found in rats that these cells can be active also when the animal is outside the location or context of their corresponding place field producing so-called "forward sweeps". These typically occur at decision points during task execution and seem to be utilized, among other things, for the evaluation of potential alternative paths. Anticipatory firing is also found in the ventral striatum, a brain area that is strongly interconnected with the hippocampus and is known to encode value and reward. In this paper, we describe a biologically based computational model of the hippocampal-ventral striatum circuit that implements a goal-directed mechanism of choice, with the hippocampus primarily involved in the mental simulation of possible navigation paths and the ventral striatum involved in the evaluation of the associated reward expectancies. The model is validated in a navigation task in which a rat is placed in a complex maze with multiple rewarding sites. We show that the rat mentally activates place cells to simulate paths, estimate their value, and make decisions, implementing two essential processes of model-based reinforcement learning algorithms of choice: look-ahead prediction and the evaluation of predicted states.     

Active navigation and orientation-free spatial representations

In this study, we examined the orientation dependency of spatial representations following various learning conditions. We assessed the spatial representations of human participants after they had learned a complex spatial layout via map learning, via navigating within a real environment, or via navigating through a virtual simulation of that environment. Performances were compared between conditions involving (1) multiple- versus single-body orientation, (2) active versus passive learning, and (3) high versus low levels of proprioceptive information. Following learning, the participants were required to produce directional judgments to target landmarks. Results showed that the participants developed orientation-specific spatial representations following map learning and passive learning, as indicated by better performance when tested from the initial learning orientation. These results suggest that neither the number of vantage points nor the level of proprioceptive information experienced are determining factors; rather, it is the active aspect of direct navigation that leads to the development of orientation-free representations.     

PandaEPL: A library for programming spatial navigation experiments

Recent advances in neuroimaging and neural recording techniques have enabled researchers to make significant progress in understanding the neural mechanisms underlying human spatial navigation. Because these techniques generally require participants to remain stationary, computer-generated virtual environments are used. We introduce PandaEPL, a programming library for the Python language designed to simplify the creation of computer-controlled spatial-navigation experiments. PandaEPL is built on top of Panda3D, a modern open-source game engine. It allows users to construct three-dimensional environments that participants can navigate from a first-person perspective. Sound playback and recording and also joystick support are provided through the use of additional optional libraries. PandaEPL also handles many tasks common to all cognitive experiments, including managing configuration files, logging all internal and participant-generated events, and keeping track of the experiment state. We describe how PandaEPL compares with other software for building spatial-navigation experiments and walk the reader through the process of creating a fully functional experiment.     

Different spatial terms in navigation system for Chinese drivers

There has been some research on the comparison of spatial frame of reference system (FoRs) preferences between different areas in China. The goal of this research was to find out whether different habits could affect the performance in driving with the navigation system, with a dual-task experimental paradigm applied. Forty participants from South and North China (10 male and 10 female in each area group) were required to finish spatial term judgment task while playing simulated driving game. The results showed that Southern participants responded more quickly to relative FoRs terms than absolute terms during driving, but there was no significant difference between these two kinds of terms for Northerners--similarly to their habits. All participants' driving actions were disturbed by the spatial term judgment. Unexpectedly, the male participants drove better when responding to relative terms than absolute terms, while female performed conversely. These findings would be useful in navigation systems design for Chinese.     

Neural evidence supports a novel framework for spatial navigation

The spatial knowledge used for human navigation has traditionally been separated into three categories: landmark, route, and survey knowledge. While behavioral research has retained this framework, it has become increasingly clear from recent neuroimaging studies that such a classification system is not adequate for understanding the brain. This review proposes a new framework, with a taxonomy based on the cognitive processes and subprocesses involved in spatial navigation. The neural correlates of spatial memory can inform our understanding of the cognitive processes involved in human navigation, and conversely, the specific task demands of an experiment can inform the interpretation of neuroimaging results. This review examines the neural correlates of each cognitive process separately, to provide a closer inspection of each component of spatial navigation. While landmark, route, and survey knowledge are still important components of human navigation, the neural correlates are not neatly ascribed to these three categories. The present findings provide motivation for a more detailed examination of the cognitive processes engaged during wayfinding.     

Cue combination in human spatial navigation

This project investigated the ways in which visual cues and bodily cues from self-motion are combined in spatial navigation. Participants completed a homing task in an immersive virtual environment. In Experiments 1A and 1B, the reliability of visual cues and self-motion cues was manipulated independently and within-participants. Results showed that participants weighted visual cues and self-motion cues based on their relative reliability and integrated these two cue types optimally or near-optimally according to Bayesian principles under most conditions. In Experiment 2, the stability of visual cues was manipulated across trials. Results indicated that cue instability affected cue weights indirectly by influencing cue reliability. Experiment 3 was designed to mislead participants about cue reliability by providing distorted feedback on the accuracy of their performance. Participants received feedback that their performance with visual cues was better and that their performance with self-motion cues was worse than it actually was or received the inverse feedback. Positive feedback on the accuracy of performance with a given cue improved the relative precision of performance with that cue. Bayesian principles still held for the most part. Experiment 4 examined the relations among the variability of performance, rated confidence in performance, cue weights, and spatial abilities. Participants took part in the homing task over two days and rated confidence in their performance after every trial. Cue relative confidence and cue relative reliability had unique contributions to observed cue weights. The variability of performance was less stable than rated confidence over time. Participants with higher mental rotation scores performed relatively better with self-motion cues than visual cues. Across all four experiments, consistent correlations were found between observed weights assigned to cues and relative reliability of cues, demonstrating that the cue-weighting process followed Bayesian principles. Results also pointed to the important role of subjective evaluation of performance in the cue-weighting process and led to a new conceptualization of cue reliability in human spatial navigation.     

Parahippocampal and retrosplenial contributions to human spatial navigation

Spatial navigation is a core cognitive ability in humans and animals. Neuroimaging studies have identified two functionally defined brain regions that activate during navigational tasks and also during passive viewing of navigationally relevant stimuli such as environmental scenes: the parahippocampal place area (PPA) and the retrosplenial complex (RSC). Recent findings indicate that the PPA and RSC have distinct and complementary roles in spatial navigation, with the PPA more concerned with representation of the local visual scene and RSC more concerned with situating the scene within the broader spatial environment. These findings are a first step towards understanding the separate components of the cortical network that mediates spatial navigation in humans.     

Quantitative and qualitative sex differences in spatial navigation

We examined sex differences in spatial navigation performance using an ecologically relevant experimental paradigm in which virtual maze-like museums are projected in front of a treadmill. Thirty-two 20-30-year-old adults (16 women/16 men) performed a way-finding task in city-block (straight corridors) or variable (irregular corridors) topographies while walking on the treadmill. Sex differences in spatial navigation performance were reduced in variable topographies, suggesting less reliance on spatial relational learning among women. Also, spatial geometric knowledge of the mazes continued to be higher in men after all participants had attained perfect place-finding performance. Results indicate that sex differences in spatial navigation performance are modulated by interactions between environmental demands and sex differences in spatial processing.     

Synergy of stimulus-driven salience and goal-directed prioritization: Evidence from the spatial blink

In the spatial blink paradigm, participants search for a target of a designated color in a rapidly presented stream of letters at fixation. Target identification is typically impaired if a peripheral distractor appears shortly before the target, inducing a spatial blink, but impairment is observed only when the distractor also shares the sought-for color. Such results reveal an important top-down influence on the capture of attention. In the present experiments, we examined the influence of the bottom-up transients associated with the appearance and disappearance of distractors in the spatial blink paradigm. Onsets and offsets alone are incapable of inducing a spatial blink, but we found that the presence of such transients did enhance the effects of target-color-matched distractors. The results reveal important synergistic interactions between top-down and bottom-up factors involved in attentional capture.     

Differences in spatial knowledge acquired from maps and navigation

Models of the spatial knowledge people acquire from maps and navigation and the procedures required for spatial judgments using this knowledge are proposed. From a map, people acquire survey knowledge encoding global spatial relations. This knowledge resides in memory in images that can be scanned and measured like a physical map. From navigation, people acquire procedural knowledge of the routes connecting diverse locations. People combine mental simulation of travel through the environment and informal algebra to compute spatial judgments. An experiment in which subjects learned an environment from navigation or from a map evaluates predictions of these models. With moderate exposure, map learning is superior for judgments of relative location and straight-line distances among objects. Learning from navigation is superior for orienting oneself with respect to unseen objects and estimating route distances. With extensive exposure, the performance superiority of maps over navigation vanishes. These and other results are consonant with the proposed mechanisms.     

The effects of spatial representation on memory for verbal navigation instructions

Three experiments investigated effects of mental spatial representation on memory for verbal navigation instructions. The navigation instructions referred to a grid of stacked matrices displayed on a computer screen or on paper, with or without depth cues, and presented as two-dimensional diagrams or a three-dimensional physical model. Experimental instructions either did or did not promote a three-dimensional mental representation of the space. Subjects heard navigation instructions, immediately repeated them, and then followed them manually on the grid. In all display and experimental instruction conditions, memory for the navigation instructions was reduced when the task required mentally representing a three-dimensional space, with movements across multiple matrices, as compared with a two-dimensional space, with movements within a single matrix, even though the words in the navigation instructions were identical in all cases. The findings demonstrate that the mental representation of the space influences immediate verbatim memory for navigation instructions.     

Visuo‐spatial working memory in navigation

Two experiments employed dual task techniques to explore the role of working memory in route learning and subsequent route retrieval. Experiment 1 involved contrasting performance of two groups of volunteers respectively learning a route from a series of map segments or a series of visually presented nonsense words. Both groups performed learning and recognition under articulatory suppression or concurrent spatial tapping. Both concurrent tasks had an overall disruptive effect on each learning task. However, spatial tapping disrupted route recognition rather more than did articulatory suppression, while the nonsense word recognition was impaired more by articulatory suppression than by concurrent spatial tapping. Experiment 2 again used dual task methodology, but explored route learning by asking volunteers to follow the experimenter through the winding streets of a medieval European town centre. Retrieval involved following the same route while the experimenter followed and noted errors in navigation. Overall the results partially replicated those of Experiment 1 in that both concurrent tasks interfered with route learning. However, volunteers with high spatial ability appeared more affected by the concurrent spatial tapping task, whereas low spatial subjects appeared more affected by the concurrent articulatory suppression task. Results are interpreted to suggest that different aspects of working memory are involved in learning a route from a map with a greater emphasis on visuo‐spatial resources, but in tasks set in real environments where many cues of a varied nature are available, only high spatial ability subjects appear to rely heavily upon the visuospatial component of working memory. Copyright © 2002 John Wiley & Sons, Ltd.     

Newborns' preference for goal-directed actions

The central role of sensory-motor representations in cognitive functions is almost universally accepted. However, determining the link between motor execution and its sensory counterpart and when, during ontogenesis, this link originates are still under investigation. The aim of the present study is to investigate whether at birth this link is already present and 2-day-old newborns are able to discriminate between visual cues indicating goal-directed or non-goal-directed actions. Here, with a preferential looking technique, a hand grasping a ball was the observed movement and we orthogonally manipulated the three factors necessary to successfully reach the goal: (a) presence of the ball, (b) direction of the arm movement, and (c) hand shaping. Results indicated that newborns orient more frequently and look longer at a hand shape for whole hand prehension but only when the movement is directed away from the body and toward the external world. In addition, newborns prefer the away from the body movement only when the object is present. We argue that newborns prefer a movement directed toward the external world only when it may develop into a purposeful movement because of the presence of the to-be-grasped object. Overall, our results support the existence of primitive sensory-motor associations since the first days after birth.     

Egocentric organization of spatial activities in imagined navigation

Studies on spatial frameworks suggest that the way we locate objects in imagined environments is influenced by the physical and functional properties of the world and our body. The present study provides evidence that such an influence also characterizes imagined navigation. In Experiment 1, participants followed spatial directions to construct an imagined path, while either keeping constant or updating their orientation at each step. A pattern of step times diagnostic of spatial frameworks was obtained in the updated-orientation but not in the constant-orientation condition. In Experiment 2, participants performed the updated-orientation condition with two levels of external support for the reference frame being used. Step times conformed to the predictions of spatial frameworks in both conditions. Both experiments also provided support that the processes involved in imagined navigation exhibit the operator-operand dynamics of other mental skills previously documented in the mental arithmetic domain. These results reinforce Piaget's (1954) notion that spatial displacements and integer arithmetic share a set of structural characteristics     

Electrophysiological correlates of high-level perception during spatial navigation

We studied the electrophysiological basis of object recognition by recording scalp electroencephalograms while participants played a virtual-reality taxi driver game. Participants searched for passengers and stores during virtual navigation in simulated towns. We compared oscillatory brain activity in response to store views that were targets or nontargets (during store search) or neutral (during passenger search). Even though store category was solely defined by task context (rather than by sensory cues), frontal electrophysiological activity in the low frequency bands (primarily in the θ [4–8 Hz] band) reliably distinguished between the target, nontarget, and neutral store views. These results implicate low-frequency oscillatory brain activity in frontal regions as an important variable in the study of the cognitive processes involved in object recognition, categorization, and other forms of high-level perception.     

空间导航的测量及其在认知老化中的应用

空间导航是日常生活所必需的高级认知功能, 参与空间导航的海马及内嗅皮层等脑区易受到老化的影响并导致结构萎缩或功能紊乱。早期研究多利用动物实验、纸笔测验、现实环境等实验范式考察老年人的空间导航老化特点。由于具有与现实环境相似的场景、兼容磁共振成像扫描以及导航者可以与场景交互等优点, 虚拟现实技术被越来越多地被应用到空间导航的老化研究中, 并进一步揭示了海马等内侧颞叶脑区在空间导航老化中的重要作用。     

边界促进空间导航的认知神经机制

边界是指在人的视野中占据较大比例,且具有立体拓展平面的障碍物,对于人类和动物的空间导航行为具有极大的促进作用。认知发展研究发现儿童早期(1岁半～2岁)通过加工边界的空间几何结构实现物体定位,并且随着年龄的发展逐渐学会利用边界的高度信息(3.1岁～4.7岁)、长度信息(4～5岁)、视觉阻碍性信息(5岁)等完成空间导航。基于这些认知过程,神经影像学研究主要以成人为研究被试,发现大脑中的内侧颞叶和顶叶脑区在边界加工中有着不同功能作用。具体而言,边界的空间几何结构及构成要素(高度、长度和角度)体位置的学习和提取则由海马负责。但是,仍存在一些研究问题值得未来深入S探c讨i。e第n一c,e拓展深化边界促进与后顶叶之间的功能交互。第三,密切关注大脑对场地边界与场地中心编码的心理或神经表征的区别和联系。第四,重点考察阿尔兹海默症有关基因易感人群在基于边界导航的行为受损情况。最后,延伸探讨边界在长时记忆、时间知觉、视觉空间、社交网络等领域的影响机制。     

空间导航的脑网络基础和调控机制

空间导航在生活中时刻发生,空间能力衰退是阿尔兹海默症的重要早期表现。早期关于空间导航神经机制的研究主要关注单个脑区的特异性功能,但这些脑区如何交互以整合不同模态的信息支持复杂导航行为尚不清楚。脑成像技术、脑网络建模方法和神经调控手段的发展,为在脑网络水平理解人类空间导航的认知神经机制提供了重要研究手段。本研究试图融合空间导航认知神经机制研究的最新进展,借助脑网络建模、大数据分析、微电流刺激等前沿研究手段,研究空间导航脑网络的关键拓扑属性特征(如模块化、核心节点等),探寻该功能特异性神经网络的重要影响因素和调控机制,并构建空间导航的脑网络理论模型。研究成果将有利于理解人类复杂导航行为的脑网络基础,为阿尔兹海默症等相关认知障碍脑疾病的筛查和诊断提供重要参考。