Tom Hartley

The Well-Worn Route and the Path Less Traveled: Distinct Neural Bases of Route Following and Wayfinding in Humans

Finding ones way in a large-scale environment may engage different cognitive processes than following a familiar route. The neural bases of these processes were investigated using functional MRI (fMRI). Subjects found their way in one virtual-reality town and followed a well-learned route in another. In a control condition, subjects followed a visible trail. Within subjects, accurate wayfinding activated the right posterior hippocampus. Between-subjects correlations with performance showed that good navigators (i.e., accurate wayfinders) activated the anterior hippocampus during wayfinding and head of caudate during route following. These results coincide with neurophysiological evidence for distinct response (caudate) and place (hippocampal) representations supporting navigation. We argue that the type of representation used influences both performance and concomitant fMRI activation patterns.

Brain areas sensitive to coherent visual motion

Detection of coherent motion versus noise is widely used as a measure of global visual-motion processing. To localise the human brain mechanisms involved in this performance, functional magnetic resonance imaging (fMRI) was used to compare brain activation during viewing of coherently moving random dots with that during viewing spatially and temporally comparable dynamic noise. Rates of reversal of coherent motion and coherent-motion velocities (5 versus 20 deg s−1) were also compared. Differences in local activation between conditions were analysed by statistical parametric mapping. Greater activation by coherent motion compared to noise was found in V5 and putative V3A, but not in V1. In addition there were foci of activation on the occipital ventral surface, the intraparietal sulcus, and superior temporal sulcus. Thus, coherent-motion information has distinctive effects in a number of extrastriate visual brain areas. The rate of motion reversal showed only weak effects in motion-sensitive areas. V1 was better activated by noise than by coherent motion, possibly reflecting activation of neurons with a wider range of motion selectivities. This activation was at a more anterior location in the comparison of noise with the faster velocity, suggesting that 20 deg s−1 is beyond the velocity range of the V1 representation of central visual field. These results support the use of motion-coherence tests for extrastriate as opposed to V1 function. However, sensitivity to motion coherence is not confined to V5, and may extend beyond the classically defined dorsal stream.

Space in the Brain: how the hippocampal formation supports spatial cognition

Over the past four decades, research has revealed that cells in the hippocampal formation provide an exquisitely detailed representation of an animals current location and heading. These findings have provided the foundations for a growing understanding of the mechanisms of spatial cognition in mammals, including humans. We describe the key properties of the major categories of spatial cells: place cells, head direction cells, grid cells and boundary cells, each of which has a characteristic firing pattern that encodes spatial parameters relating to the animals current position and orientation. These properties also include the theta oscillation, which appears to play a functional role in the representation and processing of spatial information. Reviewing recent work, we identify some themes of current research and introduce approaches to computational modelling that have helped to bridge the different levels of description at which these mechanisms have been investigated. These range from the level of molecular biology and genetics to the behaviour and brain activity of entire organisms. We argue that the neuroscience of spatial cognition is emerging as an exceptionally integrative field which provides an ideal test-bed for theories linking neural coding, learning, memory and cognition.

Predictions derived from modelling the hippocampal role in navigation.

Abstract. A computational model of the lesion and single unit data from navigation in rats is reviewed. The model uses external (visual) and internal (odometric) information from the environment to drive the firing of simulated hippocampal place cells. Constraints on the functional form of these inputs are drawn from experiments using an environment of modifiable shape. The place cell representation is used to guide navigation via the creation of a representation of goal location via Hebbian modification of synaptic strengths. The model includes consideration of the phase of firing of place cells with respect to the theta rhythm of hippocampal EEG. A series of predictions for behavioural and single-unit data in rats are derived from the input and output representations of the model.

Infant emmetropization: longitudinal changes in refraction components from nine to twenty months of age.

Rapid emmetropization is described in pediatrically normal infants from 9 months of age during the following year. The infants, obtained from various categories of the Cambridge population screening program, provided a broad range of refractive errors. The large group of 254 nonanisometropic infants studied allowed the mean rate of change and dependence on the initial refraction value to be determined. Refraction was measured by cycloplegic retinoscopy. Rapid emmetropization changes occurred in the following refractive components: mean spherical equivalent (MSE), astigmatism magnitude, the horizontal astigmatism component, the infants most positive meridian, and the infants most negative meridian. The MSE and astigmatism rates of change (diopters/year), were highly dependent on their respective initial powers (r=—0.61 and r=—0.76). The percentage weighted mean proportional rate of change for MSE was - 30% (SE 4%) and for astigmatism magnitude it was - 59% (SE 14%). There was much individual variation, with some exhibiting fast emmetropization and others not. The MSE and astigmatism changes, however, were almost independent of each other. The refractive errors of the most positive and most negative meridians emmetropize because they are both derived from the MSE and half the astigmatism. With-the-rule astigmatism was more prevalent than against-the-rule astigmatism at 9 months of age, and with-the-rule astigmatism exhibited a significantly greater proportional rate of change. The relationship of emmetropization and refractive screening is considered. A new component “MOMS” is introduced, the maximum ocular meridional separation when both eyes are considered. Thus incorporating astigmatism and anisometropia may be a good single indicator of conditions associated with later amblyopia. The almost independent emmetropization of the MSE and astigmatism components is an important result to consider in theories of emmetropization, refractive screening, clinical prescribing, and the evaluation of infants in treatment trials.

Selective interference with verbal short-term memory for serial order information: A new paradigm and tests of a timing-signal hypothesis

Many recent computational models of verbal short-term memory postulate a separation between processes supporting memory for the identity of items and processes supporting memory for their serial order. Furthermore, some of these models assume that memory for serial order is supported by a timing signal. We report an attempt to find evidence for such a timing signal by comparing an “item probe” task, requiring memory for items, with a “list probe” task, requiring memory for serial order. Four experiments investigated effects of irrelevant speech, articulatory suppression, temporal grouping, and paced finger tapping on these two tasks. In Experiments 1 and 2, irrelevant speech and articulatory suppression had a greater detrimental effect on the list probe task than on the item probe task. Reaction time data indicated that the list probe task, but not the item probe task, induced serial rehearsal of items. Phonological similarity effects confirmed that both probe tasks induced phonological recoding of visual inputs. Experiment 3 showed that temporal grouping of items during list presentation improved performance on the list probe task more than on the item probe task. In Experiment 4, paced tapping had a greater detrimental effect on the list probe task than on the item probe task. However, there was no differential effect of whether tapping was to a simple or a complex rhythm. Overall, the data illustrate the utility of the item probe/list probe paradigm and provide support for models that assume memory for serial order and memory for items involve separate processes. Results are generally consistent with the timing-signal hypothesis but suggest further factors that need to be explored to distinguish it from other accounts.

Anterior Hippocampus and Goal-Directed Spatial Decision Making

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 participants 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.

The Hippocampal Role in Spatial Memory and the Familiarity-Recollection Distinction: A Case Study

Memory for object locations and for events (comprising the receipt of an object) was tested in a case of developmental amnesia with focal hippocampal damage. Tests used virtual reality environments and forced-choice recognition with foils chosen to equalize the performance of control participants across conditions. Memory for the objects received was unimpaired, but the context of their receipt was forgotten. Memory for short lists of object locations was unimpaired when tested from the same viewpoint as presentation but impaired when tested from a shifted viewpoint. Same-view performance was disrupted by changing the background scene. These results are consistent with Jon having preserved matching to fixed sensory-bound representations but impaired reconstructed or manipulable representations underlying shifted-viewpoint recognition and episodic recollection.

Topographical Short-Term Memory Differentiates Alzheimer's Disease From Frontotemporal Lobar Degeneration

We used a recently developed test of spatial memory—the Four Mountains Test—to investigate the core cognitive processes underpinning topographical disorientation in patients with amnestic mild cognitive impairment (a‐MCI) and mild Alzheimers disease (AD). Performance of these clinical groups was compared with age‐matched controls, patients with frontotemporal lobar degeneration (FTLD), and patients with subjective memory impairments. We investigated the perception (concurrent match‐to‐sample) and short‐term retention (2‐s delayed match‐to‐sample) of the configuration of topographical features in computer‐generated landscapes shown from different viewpoints. Thirty‐one patients were tested (7 AD, 6 a‐MCI, 7 temporal variant FTLD, 5 frontal variant FTLD, 6 subjective memory impairment) and 25 age‐ and gender‐matched controls. Brain MRI was available for 27 patients; medial temporal lobe atrophy was assessed using a visual rating scale. Patients with a‐MCI or mild AD were impaired on topographical short‐term memory, but not perception. No other group differences were found on the topographical subtests. Notably, patients with temporal variants of FTLD performed normally, regardless of the laterality of damage. Subtests for the perception and retention of nonspatial aspects of the landscapes (weather conditions, seasonal and daily variations in lighting and color) were poor at differentiating the patient groups. These results indicate a core deficit in representing topographical layout, even for very short durations, within the context of more general long‐term memory impairments found in AD, and suggest that this function is particularly sensitive to the earliest stages of the disease.

Complementary memory systems: competition, cooperation and compensation

Spatial navigation depends on dissociable memory systems that have distinct neural bases and employ different forms of representation. One system gradually acquires reliable sequences of responses to given situations (e.g. repeatedly following a fixed route), and depends on the striatum. The other develops flexible representations permitting novel responses (e.g. finding new shortcuts), and depends on the hippocampus. Voermans and colleagues explore the interaction between these two systems using functional neuroimaging and behavioural measures in a clinical population.