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P E Sharp

Publications and source records attributed to P E Sharp.

At least 19 recordsLinked to original sources
PubMed2001

The anatomical and computational basis of the rat head-direction cell signal.

As a rat navigates through space, neurons called head-direction (HD) cells provide a signal of the rat's momentary directional heading. Although partly guided by landmarks, the cells also show a remarkable ability to track directional heading based on angular head movement. Theoretical models suggest that the HD cells are linked together to form an attractor network, and that cells which signal angular velocity update the directional setting of the attractor. Recently, cell types similar to those required theoretically have been discovered in the lateral mammillary and dorsal tegmental nuclei. Lesion and anatomical data suggest these nuclei might constitute the postulated attractor-path integration mechanism, and that they provide the HD cell signal to cortical areas where it has been observed.

Animals↗
PubMed2001

Head direction, place, and movement correlates for cells in the rat retrosplenial cortex.

The retrosplenial cortex is strongly connected with brain regions involved in spatial signaling. To test whether it also codes space, single cells were recorded while rats navigated in an open field. As in earlier work (L. L. Chen, L. H. Lin, C. A. Barnes, & B. L. McNaughton, 1994; L. L. Chen, L. H. Lin, E. J. Green, C. A. Barnes, & B. L. McNaughton, 1994), the authors found head direction cells with properties similar to those in other areas. These cells were slightly anticipatory. Another cell type fired to particular combinations of location, direction, and movement, which suggested that they may fire whenever the rat approaches a particular location, using a particular locomotor behavior. The remaining cells could not be clearly categorized but also showed a significant correlation with one or more of the spatial-movement variables examined. The fact that the retrosplenial cortex contains spatial and movement-related signals and is connected with the motor cortex suggests that it may play a role in path integration or navigational motor planning.

Animals↗
PubMed2001

Angular velocity and head direction signals recorded from the dorsal tegmental nucleus of gudden in the rat: implications for path integration in the head direction cell circuit.

When a rat navigates through space, head direction (HD) cells provide an ongoing signal of the rat's directional heading. It is thought that these cells rely, in part, on angular path integration of the rat's head movements. This integration requires that the HD cell system receive information about angular head movements and that this information be combined with the current directional signal, to generate the next "predicted" direction. Recent data suggest that the dorsal tegmental nucleus (DTN) may play a critical role in helping to generate the HD cell signal. To test this, recordings were made from cells in the DTN in freely moving rats. The following cell types were found: (a) "classic" HD cells, (b) angular velocity cells, and (c) cells that fired as a function of both head direction and angular velocity. Thus, DTN cells exhibit firing characteristics that are critical to the neural circuit hypothesized for generation of the HD cell signal.

Acceleration↗
PubMed1999

The anterior thalamic head-direction signal is abolished by bilateral but not unilateral lesions of the lateral mammillary nucleus.

Head-direction (HD) cells are neurons that signal a rat's directional heading in the horizontal plane. Evidence suggests that the lateral mammillary nucleus (LMN) may play an important role in generating the HD signal. Here, we examined the role of LMN in the HD circuit by comparing the effects of unilateral and bilateral LMN lesions on the activity of HD cells in the anterodorsal thalamus (AD). HD cells were recorded from AD in freely behaving rats. In the middle of the recording session, the rat received either bilateral or unilateral lesions of LMN. Immediately after the lesion, we continued recording the same HD cell in AD that had been recorded before the lesion. Additional cells were recorded from lesioned animals for up to 3 weeks after the lesion. We found that bilateral lesions of LMN permanently abolish HD cells in AD. After bilateral lesions, AD exhibits unusual rhythmic oscillations and velocity-correlated spike activity. Unilateral lesions of LMN did not abolish HD cells in AD. After unilateral lesions, the firing properties of HD cells in AD become more similar to those of HD cells in the intact hemisphere of LMN. We discuss the implications of these findings for understanding the role of LMN in the HD circuit.

Action Potentials↗
PubMed1999

Comparison of the timing of hippocampal and subicular spatial signals: implications for path integration.

Cells in several portions of the hippocampal formation show location-related firing, so that the momentary rate of each cell signals the spatial location of a freely moving rat. Insight into how these signals are generated, and how they travel around the hippocampal circuitry, can be gained by examination of the exact timing of the locational signal. Here, this was investigated for both hippocampal and subicular cells. For this, several aspects of the spatial firing pattern of each cell were examined over a series of time shifts, in which spikes were paired with locations occupied by the animal in either the immediate past, present, or future. Results showed that subicular cells appear to anticipate future locations by approximately 50 to 70 msec, on average. In contrast, hippocampal cells were best correlated with positions about 30 to 40 msec in the future. However, this timing, for hippocampal cells only, was related to the average session running speed, so that the cells were correlated with future locations at slow speeds, but lagged behind (were correlated with past locations) at high speeds. These data support the idea that both subicular and hippocampal cells use a path integration mechanism to generate their spatial signal (since both can anticipate future location). For the hippocampal cells the mechanism does not, apparently, take into account speed information, however. Also, the data suggest that the subicular signal cannot be the result of simple transmission of spatial information from the hippocampus to the subiculum, since this would predict that the subicular signal should correlate with later positions than the hippocampal signal.

Action Potentials↗
PubMed1999

Complimentary roles for hippocampal versus subicular/entorhinal place cells in coding place, context, and events.

At least two important questions are posed by the existence of hippocampal place cells. The first of these has to do with how the complex, abstract properties exhibited by these cells can be explained mechanistically. The second has to do with the implications of place cells for our conception of the broader role of the hippocampus in spatial and other behaviors. Here, evidence is reviewed that: (1) Hippocampal cells show different "maps" (place cell representations) for each environment the animal visits and, in fact, can show multiple maps even for any one environment. The choice of the current map for any one environment depends on environmental, contextual, and event-related variables. (2) Cells in the subiculum and entorhinal cortex also show location-specific firing patterns (like hippocampal place cells), but show the same pattern for each environment the animal visits. A model is presented that is a variant of hippocampus-based path integration models developed by McNaughton and colleagues. In this version, the subiculum and entorhinal cortex work together to form a single, universal map that is used for each environment, and that can exhibit path integration abilities. The universal subicular/entorhinal representation is postulated to assist the hippocampal layer to rapidly form new environment and context specific "maps" for each new environment/temporal context ("episode") the animal experiences. In this view, hippocampal layer activity is always obligatorily spatial, due to the input from the entorhinal universal "map." However, the fact that the hippocampus generates a new map in response to global, non-spatial, contextual attributes of each situation, means that the hippocampus is always coding non-spatial aspects of a situation using its obligatorily spatial code. This brings the hippocampal place cell activity in to line with the broader role that has been postulated for the hippocampus in learning and memory functions.

Animals↗
PubMed1999

Subicular place cells expand or contract their spatial firing pattern to fit the size of the environment in an open field but not in the presence of barriers: comparison with hippocampal place cells.

Cells in the hippocampus and subiculum signal spatial location in fundamentally different ways. Specifically, hippocampal cells show environment-specific spatial patterns, whereas subicular cells show the same pattern in each environment. In this study, cell firing patterns were recorded in both a large square and in a smaller square located within the large square. For some groups, portions of the small square were left in place during exposure to the large square, thus forming partial barriers. Subicular cell patterns during exposure to the large square were expanded versions of those in the small square. Hippocampal cells were likely to change their pattern completely ("remap") during exposure. However, when the barriers were left in place, cells in both areas retained the same pattern while rats were in the small square, regardless of whether they also had access to the entire large square area. Thus, subicular cells can change the size of their spatial pattern to fit the environment but will not do so across barriers.

Action Potentials↗
PubMed1998

Role of the lateral mammillary nucleus in the rat head direction circuit: a combined single unit recording and lesion study.

We recorded head direction (HD) cells from the lateral mammillary nucleus (LMN) and anterior thalamus (ATN) of freely behaving rats and also made bilateral lesions of LMN while recording HD cells from ATN. We discovered that the tuning functions of LMN HD cells become narrower during contraversive head turns, but not ipsiversive head turns, compared to when the head is not turning. This narrowing effect does not occur for ATN HD cells. We also found that the HD signal in LMN leads that in ATN by about 15-20 ms. When LMN was lesioned bilaterally, HD cells in ATN immediately lost their directional firing properties and never recovered them. Based on these findings, we argue that LMN may be an essential component of an attractor-integrator network that participates in generating the HD signal.

Animals↗
PubMed1997

Subicular cells generate similar spatial firing patterns in two geometrically and visually distinctive environments: comparison with hippocampal place cells.

Cells in both the hippocampus and the subiculum show location related firing patterns, so that the momentary firing rate of a cell is related to the spatial location of a freely moving rat as it navigates in an environment. Since the subiculum receives a strong anatomical projection from the hippocampus, it seems possible that the subicular cell spatial patterns are simply driven by the spatial signals from hippocampal place cells. Data presented here, however, suggest that the two areas code space in fundamentally different ways. Here, spatial firing patterns of individual hippocampal and subicular cells were studied as rats navigated in two different environments. The two chambers were a cylinder and a square, of equal area. For some rats the two chambers were painted to have similar visual stimulus characteristics, while for others, the two were very different. The subicular cells showed very similar firing patterns in the two chambers, regardless of whether they were visually similar or different. In contrast, as predicted based on the findings of earlier studies, hippocampal place cells showed different patterns in the two (again, regardless of their visual similarity). These results suggest that the subicular cells have the ability to transfer a single, abstract spatial representation from one environment to another. This pattern is stretched to fit within the boundaries of the current environment. Thus, the subicular cells seem to provide a generic representation of the geometric relationships between different locations in an environment. It seems possible that this representation may contribute to some navigational abilities exhibited by animals, such as dead reckoning, and novel route generation in unfamiliar environments. In contrast, it appears that hippocampal place cells provide a spatial representation-which is unique for each environment and which is strongly influenced by the exact details and overall context of the situation.

Animals↗
PubMed1997

Anticipatory time intervals of head-direction cells in the anterior thalamus of the rat: implications for path integration in the head-direction circuit.

Head-direction cells are neurons that signal a rat's directional heading in the horizontal plane. Head-direction cells in the anterior thalamus are anticipatory, so that their firing rate is better correlated with the rat's future head direction than with the present or past head direction. We recorded single-unit activity from head-direction cells in the anterior thalamus of freely moving rats. We measured the time interval by which each individual cell anticipated the rat's future head direction, which we refer to as the cell's anticipatory time interval (ATI). Head-direction cells in the anterior thalamus anticipated the rat's future head direction by an average ATI of approximately 17 ms. However, different anterior thalamic cells consistently anticipated the future head direction by different ATIs ranging between 0 and 50 ms. We found that the ATI of an anterior thalamic head-direction cell was correlated with several parameters of the cell's directional tuning function. First, cells with long ATIs sometimes appeared to have two peaks in their directional tuning function, whereas cells with short ATIs always had only one peak. Second, the ATI of a cell was negatively correlated with the cell's peak firing rate, so that cells with longer ATIs fired at a slower rate than cells with shorter ATIs. Third, a cell's ATI was correlated with the width of its directional tuning function, so that cells with longer ATIs had broader tuning widths than cells with shorter ATIs. These relationships between a cell's ATI and its directional tuning parameters could not be accounted for by artifactual broadening of the tuning function, which occurs for cells that fire in correlation with the future (rather than present) head direction. We found that when the rat's head is turning, the shape of an anterior thalamic head-direction cell's tuning function changes in a systematic way, becoming taller, narrower, and skewed. This systematic change in the shape of the tuning function may be what causes anterior thalamic cells to effectively anticipate the rat's future head direction. We propose a neural circuit mechanism to account for the firing behavior we have observed in our experiments, and we discuss how this circuit might serve as a functional component of a neural system for path integration of the rat's directional heading.

Analysis of Variance↗
PubMed1996

Neural network modeling of the hippocampal formation spatial signals and their possible role in navigation: a modular approach.

Cells throughout the hippocampal formation show striking spatial firing correlates as a rat navigates through space. These cells are thought to play a critical role in orchestrating the navigational abilities of the animals, since damage to the hippocampal formation causes spatial learning deficits. Here, we present a theoretical framework aimed at explaining how the different spatial signals are generated, as well as how they may help guide navigational behavior. Earlier work from our laboratory has presented a simple model for how the location-related signals exhibited by hippocampal place cells could be generated, based on convergent sensory information. Here, the results of this work are combined with two more recent models, to provide a more comprehensive theoretical framework. Specifically, we present 1) A neural network model of head direction cells, based on the idea that the directional signals are generated using a path integration mechanism. Cells which combine directional and angular head velocity information project onto the head direction cells, to "update" the current directional signal. This model reproduces the basic phenomenon of direction-specific firing, as well as the anticipatory nature of this firing, reported for some head direction cells. 2) A network simulation of how the hippocampal spatial signals could be used to orchestrate instrumental learning. Here, place and directional signals converge onto motor cells, each of which are thus driven to fire to specific combinations of location and directional heading. Each active motor cell generates a small leftward or rightward "step" of the simulated animal. When the simulated goal is encountered, recently active synapses are strengthened, so that goal-directed trajectories are "stamped in". We have found these models useful in helping to clarify our thinking about the proposed theoretical principles, as well as in generating testable predictions.

Animals↗
PubMed1996

Visual and vestibular influences on head-direction cells in the anterior thalamus of the rat.

As a rat navigates through space, head-direction cells provide an ongoing signal of its momentary directional heading. This directional signal is thought to be generated, in part, by a dead-reckoning mechanism that uses angular motion information to constantly update the directional representation. This study investigated what kinds of angular motion information might be used for dead reckoning. Anterior thalamic head-direction cells were recorded from rats in a rotatable, cylindrical chamber, which could independently deliver visual motion cues and vestibular cues. Results suggest that both of these angular motion cues have an influence on head-direction cells and may thus be used for dead reckoning. The authors conclude that vestibular and visual movement cues work interactively, along with visual landmarks and motor signals, to determine the directional frame of reference.

Animals↗
PubMed1995

Simulation of spatial learning in the Morris water maze by a neural network model of the hippocampal formation and nucleus accumbens.

Cells in the hippocampal formation show spatial firing correlates thought to be critical to the role played by this structure in spatial learning. Place cells in the hippocampus proper show location-specific activity, whereas cells in the postsubiculum fire as a function of momentary directional heading. One question which has received little attention is how these spatial signals are used by motor structures to actually guide spatial behavior. Here we present a model of how one kind of spatial behavior, instrumental learning in the Morris water maze, could be guided by the spatial information in the hippocampal formation. For this, we concentrate on the hippocampal projection to the nucleus accumbens, which is strongly implicated in instrumental learning. In the model, simulated firing patterns of place cells and head direction cells activate "motor" cells in the "accumbens." Each motor cell causes a particular locomotor movement in a simulated rat. In this way, the "rat" locomotes through the simulated environment. Each step places the animal in a slightly different location and directional orientation, which, in turn, activates a different set of place and head direction cells, thus causing the next locomotor response, and so on. Connection strengths between cells are initially set randomly. When the animal encounters the reward location, however, connections are altered, so that recently active synapses are strengthened. Thus, successful moves in a particular locational and directional context are "stamped in." Simulated rats show rapid learning, similar in many ways to that of actual rats. In particular, they generate efficient routes to the goal after minimal experience, and can do so from somewhat novel starting positions. Consideration of the model architecture shows that 1) combined use of directional and place information is an example of a linearly inseparable problem and that 2) some types of novel route generation, often thought to require a "cognitive mapping" strategy, can be generated from the S-R type model used here.

Animals↗
PubMed1995

Anticipatory head direction signals in anterior thalamus: evidence for a thalamocortical circuit that integrates angular head motion to compute head direction.

Several regions in the rat brain contain neurons known as head-direction cells, which fire only when the rat's head is facing in a specific direction. Head-direction cells are influenced only by the direction of the head with respect to the static environmental surroundings, and not by the position of the head relative to the body. Each head-direction cell has its own preferred direction of firing, so that together, the population of cells provides a continuous signal of momentary directional heading. Here, head-direction cells were recorded from the post-subicular cortex (PSC) and anterodorsal nucleus (ADN) of the thalamus of freely moving rats. Cell activity was analyzed in relation to both momentary head direction, and the angular velocity of head turns. Head-direction cells in PSC maintained the same directional firing preference, regardless of the angular head velocity. By contrast, head-direction cells in ADN systematically shifted their directional firing preference, as a function of angular head velocity. The ADN cells always shifted their directional tuning peak to the left during clockwise head turns, and to the right during counterclockwise head turns. These results suggest that ADN neurons anticipate the future direction of the head, whereas PSC neurons encode the present direction of the head. Based on these findings, we hypothesize that neurons in PSC and ADN are reciprocally connected to form a thalamocortical circuit, which computes the directional position of the rat's head by integrating the angular motion of the head over time.

Animals↗
PubMed1995

Influences of vestibular and visual motion information on the spatial firing patterns of hippocampal place cells.

Hippocampal place cells show location-specific firing as animals locomote in an environment. A possible explanation for these place fields is that each cell is simply driven by environmental sensory inputs available in its field. This cannot provide the full explanation, however, since cells can maintain stable place fields even in the absence of reliable environmental orienting cues. This suggests the cells are also influenced by movement-related information, since this is the only available, ongoing indicator of current location when external orienting cues are not present. Two candidates for the movement-related information are vestibular activation, and visual motion. To test for these influences, place cells were recorded as animals locomoted in a cylindrical apparatus that was made so that its wall (painted with vertical black and white stripes) and floor could be independently rotated, to provide visual motion and vestibular inputs, respectively. The results showed that both these inputs could influence place fields. Sometimes they caused a predictable locational shift, so that the field rotated its location on the apparatus floor in a way that was compatible with the movement indicated by the vestibular and/or visual motion input. This updating was most reliably obtained when the two inputs were presented in combination. In other cases, the apparatus rotations caused unpredictable changes in firing characteristics, so that cells either stopped firing, or developed place fields that were altered in overall size, shape, and eccentricity. Interestingly, the probability of these changes increased with experience with the rotational manipulations, suggesting a learned component.

Animals↗
PubMed1994

Spatial correlates of firing patterns of single cells in the subiculum of the freely moving rat.

Hippocampal lesions cause spatial learning deficits, and single hippocampal cells show location-specific firing patterns, known as place fields. This suggests the hippocampus plays a critical role in navigation by providing an ongoing indication of the animal's momentary spatial location. One question that has received little attention is how this locational signal is used by downstream brain regions to orchestrate actual navigational behavior. As a first step, we have examined the spatial firing correlates of cells in the dorsal subiculum as rats navigate in an open-field, pellet-searching task. The subiculum is one of the few major output zones for the hippocampus, and it, in turn, projects to numerous other brain areas, each thought to be involved in various learning and memory functions. Most subicular cells showed a robust locational signal. The patterns observed were different from those in the hippocampus, however, in that cells tended to fire throughout much of the environment, but showed graded, location-related rate modulation, such that there were some localized regions of high firing and other regions with relatively low firing. There were slight quantitative differences between the proximal (adjacent to the hippocampus) and distal (farther from the hippocampus) subicular regions, with distal cells showing slightly higher average firing rates, spatial signaling, and firing field size. This was of interest since these two regions have different efferent connections. Examination of spike trains allowed classification of cells into bursting, nonbursting, and theta (putative interneuron) categories, and this is similar to subicular cell types identified in vitro. Interestingly, the bursting and nonbursting types did not differ detectably in spatial firing properties, suggesting that differences in intrinsic membrane properties do not necessitate differences in coding of environmental inputs. The results suggest that the subiculum transmits a robust, highly distributed spatial signal to each of its projection areas, and that this signal is transmitted in both a bursting and nonbursting mode.

Animals↗