Role of the parietal cortex in long-term representation of spatial information in the rat

The processing of spatial information in the brain requires a network of structures within which the hippocampus plays a prominent role by elaborating an allocentric representation of space. The parietal cortex has long been suggested to have a complementary function. An overview of lesion and unit recording data in the rat indicates that the parietal cortex is involved in different aspects of spatial information processing including allocentric and egocentric processing. More specifically, the data suggest that the parietal cortex plays a fundamental role in combining visual and motion information, a process that would be important for an egocentric-to-allocentric transformation process. Furthermore, the parietal cortex may also have a role in the long-term storage of representation although this possibility needs further evidence. The data overall show that the parietal cortex occupies a unique position in the brain at the interface of perception and representation.     

The roles of the medial prefrontal cortex and hippocampus in a spatial paired-association task

Although the roles of both the hippocampus and the medial prefrontal cortex (mPFC) have been suggested in a spatial paired-associate memory task, both areas were investigated separately in prior studies. The current study investigated the relative contributions of the hippocampus and mPFC to spatial paired-associate learning within a single behavioral paradigm. In a novel behavioral task, a pair of different objects appeared repeatedly across trials, but in different arms in a radial maze, and different rules were associated with those arms for reward. Specifically, in an "object-in-place" arm, the rat was required to choose a particular object associated with the arm. In a "location-in-place" arm, the animal was required to choose a certain within-arm location (ignoring the object occupying the location). Compared to normal animals, rats with ibotenic acid-based lesions in the hippocampus showed an irrecoverable impairment in performance in both object-in-place and location-in-place arms. When the mPFC was inactivated by muscimol (GABA(A) receptor agonist) in the normal animals with intact hippocampi, they showed the same severe impairment as seen in the hippocampal lesioned rats only in object-in-place arms. The results confirm that the hippocampus is necessary for a biconditional paired-associate task when space is a critical component. The mPFC, however, is more selectively involved in the object-place paired-associate task than in the location-place paired-associate task. The current task powerfully demonstrates an experimental situation in which both the hippocampus and mPFC are required and may serve as a useful paradigm for investigating the neural mechanisms of object-place association.     

Reduced spiking in entorhinal cortex during the delay period of a cued spatial response task

Intrinsic persistent spiking mechanisms in medial entorhinal cortex (mEC) neurons may play a role in active maintenance of working memory. However, electrophysiological studies of rat mEC units have primarily focused on spatial modulation. We sought evidence of differential spike rates in the mEC in rats trained on a T-maze, cued spatial delayed response task. Animals begin at the base of the T-maze where a 1-sec white noise and visual light cue are presented on the left or right side of the maze. Rats are rewarded for responding toward the cued direction. In correct trials, we observed decreased spike rates during the delay period, the time interval between cue presentation and reward delivery. Firing-rate histograms show significant decreases during the delay period compared to 5-sec windows from both pre-cue and post-reward periods. We analyzed how running speed and trajectory specificity correlated to spike rate. Twice as many cells were responsive to cue alone compared to running speed. Trajectory specificity did not relate significantly to firing rate. Decreased spike rate may reflect active maintenance in other structures inhibiting mEC. Alternately, the reduction may reflect decreases in background activity during enhanced attention and cholinergic modulation. Lastly, animals often ran through the T-maze choice-point with varying speed. We calculated the spatial posterior probability density from spike rates during these choice-point passes. Slow passes through the choice point were characterized by greater probability of decoding to the reward locations on correct trials compared to quick passes on the maze consistent with similar "look-ahead" properties previously reported in the hippocampus and ventral striatum.     

Circadian phase-shifted rats show normal acquisition but impaired long-term retention of place information in the water task

It is thought that circadian rhythms may influence learning and memory processes. However, research supporting this view does not dissociate a mnemonic impairment from other performance deficits. Furthermore, published reports do not specify the type of memory system influenced by the circadian system. The present study assessed the effects of phase shifting on acquisition and expression of place navigation in the water maze, a task sensitive to hippocampal dysfunction. The results showed that phase-shifting circadian rhythms in rats impaired the expression of place information on a retention test but not initial acquisition or encoding of place information. These results suggest that disruption of circadian rhythms may impair consolidation of previously encoded hippocampal place information.     

Human topological task adapted for rats: Spatial information processes of the parietal cortex

Human research has shown that lesions of the parietal cortex disrupt spatial information processing, specifically topological information. Similar findings have been found in non-humans. It has been difficult to determine homologies between human and non-human mnemonic mechanisms for spatial information processing because methodologies and neuropathology differ. The first objective of the present study was to adapt a previously established human task for rats. The second objective was to better characterize the role of parietal cortex (PC) and dorsal hippocampus (dHPC) for topological spatial information processing. Rats had to distinguish whether a ball inside a ring or a ball outside a ring was the correct, rewarded object. After rats reached criterion on the task (>95%) they were randomly assigned to a lesion group (control, PC, and dHPC). Animals were then re-tested. Post-surgery data show that controls were 94% correct on average, dHPC rats were 89% correct on average, and PC rats were 56% correct on average. The results from the present study suggest that the parietal cortex, but not the dHPC processes topological spatial information. The present data are the first to support comparable topological spatial information processes of the parietal cortex in humans and rats.     

Social effects on rat spatial choice in an open field task

Pairs of rats foraged in trials either together or separately in an open field apparatus for pellets hidden in discreet locations in a 5 × 5 matrix. Trial duration was either 1 or 4 min. The tendency to choose locations that had earlier been visited by another rat was examined by comparing the choices made in the presence and absence of the other rat. Rats avoided visits to locations that had earlier been visited by the other rat, but only if they had also visited the same location earlier in a short duration trial. This pattern of results is consistent with earlier findings from experiments using the radial arm maze. Furthermore, when rats did visit locations that had earlier been visited by the other rat in a long duration trial, they tended to be locations that had been visited longer ago by the other rat than would be expected. This suggests a forgetting function for social memories. These data provide evidence that the social memory reported in earlier studies using the radial-arm maze can be found in other experimental paradigms and that at least some of its properties are common in the two paradigms.     

The posterior parietal cortex and long-term memory representation of spatial information

The hypothesis to be explored in this chapter is based on the assumption that the posterior parietal cortex (PPC) is directly involved in representing a subset of the spatial features associated with spatial information processing and plays an important role in perceptual memory as well as long-term memory encoding, consolidation, and retrieval of spatial information. After presentation of the anatomical location of the PPC in rats, the nature of PPC representation based on single spatial features, binding of visual features associated with visual spatial attention, binding of object-place associations associated with acquisition and storage of associations where one of the elements is a spatial component, and binding of ideothetic and allothetic information in long-term memory is discussed. Additional evidence for a PPC role in mediation of spatial information in long-term storage is offered. Finally, the relationship between the PPC and the hippocampus from a systems and dynamic point view is presented.     

Temporary inactivation of the dorsal entorhinal cortex impairs acquisition and retrieval of spatial information

We tested the effects of temporary inactivation of the dorsal entorhinal cortex on spatial discrimination using a conditioned cue preference (CCP) paradigm. The three phases of the procedure were: pre-exposure: unreinforced exploration of the center platform and two adjacent arms of an eight-arm radial maze; training: rats were confined to the ends of the two arms on alternate days – one arm always contained food and the other never contained food; testing: unreinforced exploration of the center platform and the two arms. Rats that received bilateral infusions of saline into the dorsal entorhinal cortex before the training trials or before the test trial spent significantly more time in the arm that previously contained food than in the arm that never contained food, demonstrating that they had acquired and were able to express information that discriminated between the two adjacent maze arms. In contrast, rats that received bilateral, intra-entorhinal infusions of muscimol, a gamma-aminobutyric acid_a (GABA_a) agonist, before either training or testing spent equal amounts of time in the two arms, indicating that they failed to acquire and were unable to express this information. Interactions between the entorhinal cortex and hippocampus in the acquisition and expression of the information required for this discrimination are discussed.     

Microinjections of leupeptin in the frontal cortex or dorsal hippocampus block spatial learning in the rat

Adult male Long-Evans hooded rats were given bilateral microinjections of 18 mM leupeptin or saline through cannulae implanted with tips aimed at the frontal cortex or CA1 or CA3 hippocampal cell fields. Five minutes following injections animals were allowed to complete an eight-arm radial maze. The acquisition criterion required that the animal make 7 correct choices of the first 8, and 8 correct choices of the first 10, for five consecutive sessions. Leupeptin slowed acquisition of the eight-arm radial maze task when injected into the CA1 and CA3 hippocampal fields or the frontal cortex, but did not influence spontaneous activity. These results suggest that earlier reports of the amnestic effect of leupeptin when administered into the lateral cerebral ventricle may have been due to effects within the cortex and hippocampus. The present experiment represents the first attempt to identify behaviorally those brain areas in which leupeptin acts to alter learning.     

Parietal cortex, navigation, and the construction of arbitrary reference frames for spatial information

The registration of spatial information by neurons of the parietal cortex takes on many forms. In most experiments, spatially modulated parietal activity patterns are found to take as their frame of reference some part of the body such as the retina. However, recent findings obtained in single neuron recordings from both rat and monkey parietal cortex suggest that the frame of reference utilized by parietal cortex may also be abstract or arbitrary in nature. Evidence in rats comes from work indicating that parietal activity in freely behaving rodents is organized according to the space defined by routes taken through an environment. In monkeys, evidence for an object-centered frame of reference has recently been presented. The present work reviews single neuron recording experiments in parietal cortex of freely behaving rats and considers the potential contribution of parietal cortex in solving navigational tasks. It is proposed that parietal cortex, in interaction with the hippocampus, plays a critical role in the selection of the most appropriate route between two points and, in addition, produces a route-based positional signal capable of guiding sensorimotor transitions.     

Neural coding of reward magnitude in the orbitofrontal cortex of the rat during a five-odor olfactory discrimination task

The orbitofrontal cortex (OBFc) has been suggested to code the motivational value of environmental stimuli and to use this information for the flexible guidance of goal-directed behavior. To examine whether information regarding reward prediction is quantitatively represented in the rat OBFc, neural activity was recorded during an olfactory discrimination “go”/“no-go” task in which five different odor stimuli were predictive for various amounts of reward or an aversive reinforcer. Neural correlates related to both actual and expected reward magnitude were observed. Responses related to reward expectation occurred during the execution of the behavioral response toward the reward site and within a waiting period prior to reinforcement delivery. About one-half of these neurons demonstrated differential firing toward the different reward sizes. These data provide new and strong evidence that reward expectancy, regardless of reward magnitude, is coded by neurons of the rat OBFc, and are indicative for representation of quantitative information concerning expected reward. Moreover, neural correlates of reward expectancy appear to be distributed across both motor and nonmotor phases of the task.     

Strategic grouping in the spatial span memory task

Spatial short-term memory performance was examined in relation to participants' strategies. A total of 20 adult participants viewed and reproduced sequences of locations that varied in length (five, six, seven, or eight locations) and spatial separability (a manipulation of the configurations). In trial-by-trial self-reports, participants described five types of strategies. Chunking the spatial sequences into groups of three or four locations was the sole strategy associated with increased accuracy. Participants demonstrated considerable variability in the strategies that they selected, suggesting that cognitive resources are allocated to strategy selection, execution, and monitoring in the spatial span task. Spatially separable sequences were more accurately recalled than nonseparable sequences, independent of strategic grouping, suggesting two levels of grouping in the spatial span task.     

Strategic grouping in the spatial span memory task

Spatial short-term memory performance was examined in relation to participants’ strategies. A total of 20 adult participants viewed and reproduced sequences of locations that varied in length (five, six, seven, or eight locations) and spatial separability (a manipulation of the configurations). In trial-by-trial self-reports, participants described five types of strategies. Chunking the spatial sequences into groups of three or four locations was the sole strategy associated with increased accuracy. Participants demonstrated considerable variability in the strategies that they selected, suggesting that cognitive resources are allocated to strategy selection, execution, and monitoring in the spatial span task. Spatially separable sequences were more accurately recalled than nonseparable sequences, independent of strategic grouping, suggesting two levels of grouping in the spatial span task.     

On the representation of task information in task switching: Evidence from task and dimension switching

Task switching research has revealed that task changes lead to a performance switch cost. The present study focuses on the organization of task components in the task set. Three different views of task set organization have been distinguished and evidence in favor of each of these has been reported in the literature. In four experiments, we orthogonally varied the categorization task (magnitude and parity) and the stimulus dimension on which the categorization was to be made. Experiments 1, 2, and 4 used Stroop-like number stimuli, whereas Experiment 3 used global-local stimuli to define the stimulus dimension. In Experiments 2-4, the cue-stimulus interval was also varied. The findings showed that a change of any component resulted in a cost, without any reliable difference in the size of these costs. These results are consistent with the flat view on task-set organization, which assumes that the task set binds all elements in an unstructured representation, which is completely reconfigured each time a change to the task set is required. The implications of these findings are discussed in relation to other findings and the different views on task-set organization.     

Modes of spatial coding in the Simon task*

Many models of the Simon effect assume that categorical spatial representations underlie the phenomenon. The present study tested this assumption explicitly in two experiments, both of which involved eight possible spatial positions of imperative stimuli arranged horizontally on the screen. In Experiment 1, the eight stimulus locations were marked with eight square boxes that appeared at the same time during a trial. Results showed gradually increasing Simon effects from the central locations to the outer locations. In Experiment 2, the eight stimulus locations consisted of a combination of three frames of spatial reference (hemispace, hemifield, and position relative to the fixation), with each frame appearing in different timings. In contrast to Experiment 1, results showed an oscillating pattern of the Simon effect across the horizontal positions. These findings are discussed in terms of grouping factors involved in the Simon task. The locations seem to be coded as a single continuous dimension when all are visible at once as in Experiment 1, but they are represented as a combination of the lateral categories (“left” vs. “right”) with multiple frames of reference when the reference frames are presented successively as in Experiment 2.     

Processing of place information in stop consonants

One of the basic questior, s that models of speech perception must answer concerns the conditions under which various cues will be extracted from a stimulus and the nature of the mechanisms which mediate this process. Two selective adaptation experiments were carried out to explore this question for the phonetic feature of place of articulation in both syllableinitial and syllable-final positions. In the first experiment, CV and VC stimuli were constructed with complete overlap in their second- and third-formant transitions. Despite this essentially complete overlap, no adaptation effects were found for a VC adaptor and a CV test series (or vice versa). In the second experiment, various vowel, vowel-like, and VC-like adaptors were used. The VC-like adaptors did have a significant effect on the CV category boundary, while the vowel and vowel-like stimuli did not. These results are interpreted within both one- and twolevel models of selective adaptation. These models are distinguished by whether selective adaptation is assumed to affect a single auditory level of processing or to affect both an auditory level and a later phonetic level. However, both models incorporate detectors at the auditory level which respond whenever particular formant transitions are present. These auditory detectors are not sensitive to the position of the consonant transition information within the syllable.     

Rate of information segregation in developmentally dyslexic children

Slowed processing of sequential perceptual information is related to developmental dyslexia. We investigated this unimodally and crossmodally in developmentally dyslexic children and controls ages 8-12 years. The participants judged whether two spatially separate trains of brief stimuli, presented at various stimulus onset asynchronies (SOA) in one or two senses, were synchronous or not. The stimulus trains consisted of light flashes in vision, clicks in audition, and indentations of the skin in the tactile sense. The dyslexic readers required longer SOAs than controls for successful performance in all six comparisons. The crossmodal spatiotemporal resolution of the groups differed more than unimodal performance. The dyslexic readers' segregation performance was also less differentiated than that of the controls. Our results show that not only sensory but also polysensory nonverbal information processing is temporally impaired in dyslexic children.