The hippocampus appears to be crucial for long-term episodic memory, yet its precise role remains elusive. Electrophysiological studies in rodents offer a useful starting point for developing models of hippocampal processing in the spatial domain. Here we review one such model that points to an essential role for the hippocampus in the construction of mental images. We explain how this neural-level mechanistic account addresses some of the current controversies in the field, such as the role of the hippocampus in imagery and short-term memory, and discuss its broader implications for the neural bases of episodic memory.

ABSTRACT:The hippocampus plays a crucial role within the neural systems for long-term memory, but little if any role in the short-term retention of some types of stimuli. Nonetheless, the hippocampus may be specialized for allocentric topographical processing, which impacts on short-term memory or even perception. To investigate this we developed performance-matched tests of perception (match-to-sample) and shortterm memory (2 s delayed-match-to-sample) for the topography and for the nonspatial aspects of visual scenes. Four patients with focal hippocampal damage and one with more extensive damage, including right parahippocampal gyrus, were tested. All five patients showed impaired topographical memory and spared nonspatial processing in both memory and perception. Topographical perception was profoundly impaired in the patient with parahippocampal damage, mildly impaired in two of the hippocampal cases, and clearly preserved in the other two hippocampal cases (including one with dense amnesia). Our results suggest that the hippocampus supports allocentric topographical processing that is indispensable when appropriately tested after even very short delays, while the presence of the sample scene can allow successful topographical perception without it, possibly via a less flexible parahippocampal representation. V V C 2006 Wiley-Liss, Inc.

Mixed evidence exists for executive dysfunction in autism spectrum disorders (ASD). This may be because of the nature of the tasks used, the heterogeneity of participants, and difficulties with recruiting appropriate control groups. A comprehensive battery of 'executive' tests was administered to 22 individuals with Asperger syndrome and 22 well-matched controls. Performance was analysed both between groups and on an individual basis to identify outliers in both the ASD and control groups. There were no differences between the groups on all 'classical' tests of executive function. However, differences were found on newer tests of executive function. Specifically, deficits in planning, abstract problem solving and especially multitasking. On the tests that discriminated the groups, all of the ASD individuals except one were identified as significantly impaired (i.e. below the 5th percentile of the control mean) on at least one executive measure. This study provides evidence for significant executive dysfunction in Asperger syndrome. Greatest dysfunction appeared in response initiation and intentionality at the highest level-the ability to engage and disengage actions in the service of overarching goals. These deficits are best observed through using more recent, ecologically valid tests of executive dysfunction. Moreover, performance on these measures correlated with autistic symptomatology. Keywords: Executive functions; Autism spectrum disorder; Problem solving; Multitasking; Prefrontal cortex 2 Executive processes in Asperger syndromeGoldsmiths Research Online 'Executive function' is traditionally used as an umbrella term for abilities such as planning,working memory, impulse control, inhibition and shifting set, as well as the initiation and monitoring of action (Roberts, Robbins, & Weiskrantz, 1998;Stuss & Knight, 2002). Animal, behavioural and neuropsychological studies have linked these functions to frontal structures of the brain, and to prefrontal cortex in particular. Additionally, a number of neurodevelopmental disorders have been linked to executive dysfunction, including autism spectrum disorder (ASD; see Russell, 1997). The core features of autism are abnormalities of social interaction, impairments in verbal and non-verbal communication and a restricted repertoire of interests and activities, all present from early childhood (American Psychiatric Association, 1994). If executive dysfunction is a central component of ASD then this would have important implications for diagnosis, intervention and our theoretical understanding of the syndrome. However, the presence, or causal nature of executive dysfunction to autism is much debated. Systematic reviews of the literature reveal mixed evidence for executive dysfunction in ASD, although difficulties in planning, mental flexibility and generativity have been documented (see Pennington, & Ozonoff, 1996, for review). Mixed findings may arise from a number of issues, including the nature of the tasks administered as well as the nature of the autism a...

The ability of humans to predict and explain other people's behaviour by attributing to them independent mental states, such as desires and beliefs, is considered to be due to our ability to construct a 'Theory of Mind'. Recently, several neuroimaging studies have implicated the medial frontal lobes as playing a critical role in a dedicated 'mentalizing' or 'Theory of Mind' network in human brains. Here, we report a patient, G.T., who suffered an exceptionally rare form of stroke-bilateral anterior cerebral artery infarction, without rupture or the complications associated with anterior communicating artery aneurysms. Detailed high-resolution neuroanatomical investigations revealed extensive damage to the medial frontal lobes bilaterally, including regions identified to be critical for 'Theory of Mind' by functional neuroimaging of healthy human subjects. For the first time in such a patient, we carried out a thorough assessment of her cognitive profile including, critically, an experimental investigation of her performance on a range of tests of 'Theory of Mind'. G.T. had a dysexecutive syndrome characterized by impairments in planning and memory, as well as a tendency to confabulate. Importantly, however, she did not have any significant impairment on tasks probing her ability to construct a 'Theory of Mind', demonstrating that the extensive medial frontal regions destroyed by her stroke are not necessary for this function. These findings have important implications for the functional anatomy of 'Theory of Mind', as well as our understanding of medial frontal function. Possible reasons for the discrepancies between our results and neuroimaging studies are discussed. We conclude that our findings urge caution against using functional imaging as the sole method of establishing cognitive neuroanatomy.

It is well-established that active rehearsal increases the efficacy of memory consolidation. It is also known that complex events are interpreted with reference to prior knowledge. However, comparatively little attention has been given to the neural underpinnings of these effects. In healthy adults humans, we investigated the impact of effortful, active rehearsal on memory for events by showing people several short video clips and then asking them to recall these clips, either aloud (Experiment 1) or silently while in an MRI scanner (Experiment 2). In both experiments, actively rehearsed clips were remembered in far greater detail than unrehearsed clips when tested a week later. In Experiment 1, highly similar descriptions of events were produced across retrieval trials, suggesting a degree of semanticization of the memories had taken place. In Experiment 2, spatial patterns of BOLD signal in medial temporal and posterior midline regions were correlated when encoding and rehearsing the same video. Moreover, the strength of this correlation in the posterior cingulate predicted the amount of information subsequently recalled. This is likely to reflect a strengthening of the representation of the video's content. We argue that these representations combine both new episodic information and stored semantic knowledge (or "schemas"). We therefore suggest that posterior midline structures aid consolidation by reinstating and strengthening the associations between episodic details and more generic schematic information. This leads to the creation of coherent memory representations of lifelike, complex events that are resistant to forgetting, but somewhat inflexible and semantic-like in nature.

Planning spatial paths through our environment is an important part of everyday life and is supported by a neural system including the hippocampus and prefrontal cortex. Here we investigated the precise functional roles of the components of this system in humans by using fMRI as participants performed a simple goal-directed route-planning task. Participants had to choose the shorter of two routes to a goal in a visual scene that might contain a barrier blocking the most direct route, requiring a detour, or might be obscured by a curtain, requiring memory for the scene. The participant's start position was varied to parametrically manipulate their proximity to the goal and the difference in length of the two routes. Activity in medial prefrontal cortex, precuneus, and left posterior parietal cortex was associated with detour planning, regardless of difficulty, whereas activity in parahippocampal gyrus was associated with remembering the spatial layout of the visual scene. Activity in bilateral anterior hippocampal formation showed a strong increase the closer the start position was to the goal, together with medial prefrontal, medial and posterior parietal cortices. Our results are consistent with computational models in which goal proximity is used to guide subsequent navigation and with the association of anterior hippocampal areas with nonspatial functions such as arousal and reward expectancy. They illustrate how spatial and nonspatial functions combine within the anterior hippocampus, and how these functions interact with parahippocampal, parietal, and prefrontal areas in decision making and mnemonic function.

Theta frequency oscillations in the 6-to 10-Hz range dominate the rodent hippocampal local field potential during translational movement, suggesting that theta encodes self-motion. Increases in theta power have also been identified in the human hippocampus during both real and virtual movement but appear as transient bursts in distinct high-and low-frequency bands, and it is not yet clear how these bursts relate to the sustained oscillation observed in rodents. Here, we examine depth electrode recordings from the temporal lobe of 13 presurgical epilepsy patients performing a selfpaced spatial memory task in a virtual environment. In contrast to previous studies, we focus on movement-onset periods that incorporate both initial acceleration and an immediately preceding stationary interval associated with prominent theta oscillations in the rodent hippocampal formation. We demonstrate that movementonset periods are associated with a significant increase in both low (2-5 Hz)-and high (6-9 Hz)-frequency theta power in the human hippocampus. Similar increases in low-and high-frequency theta power are seen across lateral temporal lobe recording sites and persist throughout the remainder of movement in both regions. In addition, we show that movement-related theta power is greater both before and during longer paths, directly implicating human hippocampal theta in the encoding of translational movement. These findings strengthen the connection between studies of theta-band activity in rodents and humans and offer additional insight into the neural mechanisms of spatial navigation.theta | hippocampus | navigation | spatial memory | intracranial EEG T he rodent hippocampal local field potential (LFP) is dominated by 6-to 10-Hz theta oscillations during translational movement (1, 2). Both the power (1, 3) and frequency (3-6) of theta are positively correlated with running speed. Theta oscillations might therefore encode self-motion information and contribute to the generation of spatially modulated firing patterns (7-9). A critical concern for contemporary neuroscience is to establish whether this hypothesis can be translated across species. During navigation, intracranial recordings from depth electrodes in the human hippocampus have shown that theta is more prevalent during movement than during stationary periods (10-12) and that theta power increases with movement speed (13). In addition, increases in movement-related theta power are seen across the neocortex (10,11,14). These findings support the hypothesis that human theta oscillations might encode self-motion information. However, it has also been demonstrated that human theta-band activity typically occurs in transient bursts distributed throughout movement, in contrast to the continuous high-amplitude oscillation observed in the rodent (15,16). Moreover, these studies identified movement-related oscillations within both higher and lower frequency theta bands (17)(18)(19)(20). Hence, it is not yet clear how theta oscillations in the human hippocampus relate to thos...

When we visualize scenes, either from our own past or invented, we impose a viewpoint for our "mind's eye" and we experience the resulting image as spatially coherent from that viewpoint. The hippocampus has been implicated in this process, but its precise contribution is unknown. We tested a specific hypothesis based on the spatial firing properties of neurons in the hippocampal formation of rats, that this region supports the construction of spatially coherent mental images by representing the locations of the environmental boundaries surrounding our viewpoint. Using functional magnetic resonance imaging, we show that hippocampal activation increases parametrically with the number of enclosing boundaries in the imagined scene. In contrast, hippocampal activity is not modulated by a nonspatial manipulation of scene complexity nor to increasing difficulty of imagining the scenes in general. Our findings identify a specific computational role for the hippocampus in mental imagery and episodic recollection.