Rat Posterior Parietal Cortex Research Articles

Acute Response and Neuroprotective Role of Myo/Nog Cells Assessed in a Rat Model of Focal Brain Injury.

Focal brain injury in the form of a needlestick (NS) results in cell death and induces a self-protective response flanking the lesion. Myo/Nog cells are identified by their expression of bone morphogenetic protein inhibitor Noggin, brain-specific angiogenesis inhibitor 1 (BAI1) and the skeletal muscle specific transcription factor MyoD. Myo/Nog cells limit cell death in two forms of retinopathy. In this study, we examined the acute response of Myo/Nog cells to a NS lesion that extended from the rat posterior parietal cortex to the hippocampus. Myo/Nog cells were identified with antibodies to Noggin and BAI1. These cells were the primary source of both molecules in the uninjured and injured brain. One day after the NS, the normally small population of Myo/Nog cells expanded approximately eightfold within a 1 mm area surrounding the lesion. Myo/Nog cells were reduced by approximately 50% along the lesion with an injection of the BAI1 monoclonal antibody and complement. The number of dying cells, identified by terminal deoxynucleotidyl transferase-mediated dUTP-biotin nick end labeling (TUNEL), was unchanged at this early time point in response to the decrease in Myo/Nog cells. However, increasing the number of Myo/Nog cells within the lesion by injecting BAI1-positive (+) cells isolated from the brains of other animals, significantly reduced cell death and increased the number of NeuN+ neurons compared to brains injected with phosphate buffered saline or exogenous BAI1-negative cells. These findings demonstrate that Myo/Nog cells rapidly react to injury within the brain and increasing their number within the lesion is neuroprotective.

Functional Differentiation of Dorsal and Ventral Posterior Parietal Cortex of the Rat: Implications for Controlled and Stimulus-Driven Attention.

The posterior parietal cortex (PPC) is important for visuospatial attention. The primate PPC shows functional differentiation such that dorsal areas are implicated in top-down, controlled attention, and ventral areas are implicated in bottom-up, stimulus-driven attention. Whether the rat PPC also shows such functional differentiation is unknown. Here, we address this open question using functional neuroanatomy and in vivo electrophysiology. Using conventional tract-tracing methods, we examined connectivity with other structures implicated in visuospatial attention including the lateral posterior nucleus of the thalamus (LPn) and the postrhinal cortex (POR). We showed that the LPn projects to the entire PPC, preferentially targeting more ventral areas. All parts of the PPC and POR are reciprocally connected with the strongest connections evident between ventral PPC and caudal POR. Next, we simultaneously recorded neuronal activity in dorsal and ventral PPC as rats performed a visuospatial attention (VSA ) task that engages in both bottom-up and top-down attention. Previously, we provided evidence that the dorsal PPC is engaged in multiple cognitive process including controlled attention (Yang etal. 2017). Here, we further showed that ventral PPC cells respond to stimulus onset more rapidly than dorsal PPC cells, providing evidence for a role in stimulus-driven, bottom-up attention.

Task phase-specific involvement of the rat posterior parietal cortex in performance of the TUNL task.

The posterior parietal cortex (PPC) participates in cognitive processes including working memory (WM), sensory evidence accumulation, and perceptually guided decision making. However, surprisingly little work has used temporally precise manipulations to dissect its role in different epochs of behavior taking place over short timespans, such as WM tasks. As a result, a consistent view of the temporally precise role of the PPC in these processes has not been described. In the present study, we investigated the temporally specific role of the PPC in the Trial-Unique, Nonmatching-to-Location (TUNL) task, a touchscreen-based, visuospatial WM task that relies on the PPC. To disrupt PPC activity in a temporally precise manner, we applied mild intracranial electrical stimulation (ICES). We found that intra-PPC ICES (100 μA) significantly impaired accuracy in TUNL without significantly altering response latency. Moreover, we found that the impairment was specific to ICES applied during the delay and test phases of TUNL. Consistent with previous reports showing delay- and choice-specific neuronal activity in the PPC, the results provide evidence that the rat PPC is required for maintaining memory representations of stimuli over a delay period as well as for making successful comparisons and choices between test stimuli. In contrast, the PPC appears not to be critical for initial encoding of sample stimuli. This pattern of results may indicate that early encoding of visual stimuli is independent of the PPC or that the PPC becomes engaged only when visual stimuli are spatially complex or involve memory or decision making.

Sense and Action across the Layers of the Rat Posterior Parietal Cortex.

The integration of sensory stimuli and ongoing motor outputs is called sensorimotor integration, and it occurs in motor as well as sensory cortices, where it helps to guide motor output ([Petersen, 2019][1]). The rodent whisker system is an excellent system in which to study the interaction of

Organization of Posterior Parietal-Frontal Connections in the Rat.

Recent investigations of the rat posterior parietal cortex (PPC) suggest that this region plays a central role in action control together with the frontal cortical areas. Posterior parietal-frontal cortical connections have been described in rats, but little is known about whether these connections are topographically organized as in the primate. Here, we injected retrograde and anterograde tracers into subdivisions of PPC as well as the frontal midline and orbital cortical areas to explore possible topographies within their connections. We found that PPC projects to several frontal cortical areas, largely reciprocating the densest input received from the same areas. All PPC subdivisions are strongly connected with the secondary motor cortex (M2) in a topographically organized manner. The medial subdivision (medial posterior parietal cortex, mPPC) has a dense reciprocal connection with the most caudal portion of M2 (cM2), whereas the lateral subdivision (lateral posterior parietal cortex, lPPC) and the caudolateral subdivision (PtP) are reciprocally connected with the intermediate rostrocaudal portion of M2 (iM2). Sparser reciprocal connections were seen with anterior cingulate area 24b. mPPC connects with rostral, and lPPC and PtP connect with caudal parts of 24b, respectively. There are virtually no connections with area 24a, nor with prelimbic or infralimbic cortex. PPC and orbitofrontal cortices are also connected, showing a gradient such that mPPC entertains reciprocal connections mainly with the ventral orbitofrontal cortex (OFC), whereas lPPC and PtP are preferentially connected with medial and central portions of ventrolateral OFC, respectively. Our results thus indicate that the connections of PPC with frontal cortices are organized in a topographical fashion, supporting functional heterogeneity within PPC and frontal cortices.

Functional Architecture and Encoding of Tactile Sensorimotor Behavior in Rat Posterior Parietal Cortex.

The posterior parietal cortex (PPC) in rodents is reciprocally connected to primary somatosensory and vibrissal motor cortices. The PPC neuronal circuitry could thus encode and potentially integrate incoming somatosensory information and whisker motor output. However, the information encoded across PPC layers during refined sensorimotor behavior remains largely unknown. To uncover the sensorimotor features represented in PPC during voluntary whisking and object touch, we performed loose-patch single-unit recordings and extracellular recordings of ensemble activity, covering all layers of PPC in anesthetized and awake, behaving male rats. First, using single-cell receptive field mapping, we revealed the presence of coarse somatotopy along the mediolateral axis in PPC. Second, we found that spiking activity was modulated during exploratory whisking in layers 2-4 and layer 6, but not in layer 5 of awake, behaving rats. Population spiking activity preceded actual movement, and whisker trajectory endpoints could be decoded by population spiking, suggesting that PPC is involved in movement planning. Finally, population spiking activity further increased in response to active whisker touch but only in PPC layers 2-4. Thus, we find layer-specific processing, which emphasizes the computational role of PPC during whisker sensorimotor behavior.SIGNIFICANCE STATEMENT The posterior parietal cortex (PPC) is thought to merge information on motor output and sensory input to orchestrate interaction with the environment, but the function of different PPC microcircuit components is poorly understood. We recorded neuronal activity in rat PPC during sensorimotor behavior involving motor and sensory pathways. We uncovered that PPC layers have dedicated function: motor and sensory information is merged in layers 2-4; layer 6 predominantly represents motor information. Collectively, PPC activity predicts future motor output, thus entailing a motor plan. Our results are important for understanding how PPC computationally processes motor output and sensory input. This understanding may facilitate decoding of brain activity when using brain-machine interfaces to overcome loss of function after, for instance, spinal cord injury.

Performance of the trial-unique, delayed non-matching-to-location (TUNL) task depends on AMPA/Kainate, but not NMDA, ionotropic glutamate receptors in the rat posterior parietal cortex.

Working memory (WM), the capacity for short-term storage and manipulation of small quantities of information, depends on fronto-parietal circuits. However, the function of the posterior parietal cortex (PPC) in WM has gone relatively understudied in rodents. Recent evidence calls into question whether the PPC is necessary for all forms of WM. Thus, the present experiment examined the role of the rat PPC in the Trial-Unique Non-matching-to-Location (TUNL) task, a touchscreen-based visuospatial WM task that relies on the rat medial prefrontal cortex (mPFC). Temporary inactivation of the PPC caused by bilateral infusions of muscimol and baclofen significantly impaired accuracy and increased the number of correction trials performed, indicating that the PPC is necessary for performance of TUNL. Additionally, we investigated the effects of blocking NMDA or non-NMDA parietal ionotropic glutamate receptors on TUNL and found that, in contrast to the prefrontal cortex, NMDA receptors in the PPC are not necessary for TUNL performance, whereas blockade of AMPA/Kainate receptors significantly impaired accuracy. These results indicate that performance of the TUNL task depends on the PPC but that NMDA receptor signaling within this brain area is not necessary for intact performance.

Posterior Parietal Cortex Guides Visual Decisions in Rats.

Neurons in putative decision-making structures can reflect both sensory and decision signals, making their causal role in decisions unclear. Here, we tested whether rat posterior parietal cortex (PPC) is causal for processing visual sensory signals or instead for accumulating evidence for decision alternatives. We disrupted PPC activity optogenetically during decision making and compared effects on decisions guided by auditory versus visual evidence. Deficits were largely restricted to visual decisions. To further test for visual dominance in PPC, we evaluated electrophysiological responses after individual sensory events and observed much larger response modulation after visual stimuli than auditory stimuli. Finally, we measured trial-to-trial spike count variability during stimulus presentation and decision formation. Variability decreased sharply, suggesting that the network is stabilized by inputs, unlike what would be expected if sensory signals were locally accumulated. Our findings suggest that PPC plays a causal role in processing visual signals that are accumulated elsewhere.SIGNIFICANCE STATEMENT Defining the neural circuits that support decision making bridges a gap between our understanding of simple sensorimotor reflexes and our understanding of truly complex behavior. However, identifying brain areas that play a causal role in decision making has proved challenging. We tested the causal role of a candidate component of decision circuits, the rat posterior parietal cortex (PPC). Our interpretation of the data benefited from our use of animals trained to make decisions guided by either visual or auditory evidence. Our results suggest that PPC plays a causal role specifically in visual decision making and may support sensory aspects of the decision, such as interpreting the visual signals so that evidence for a decision can be accumulated elsewhere.

Single neuron activity and theta modulation in the posterior parietal cortex in a visuospatial attention task.

The posterior parietal cortex (PPC) is implicated in directing and maintaining visual attention to locations in space. We hypothesized that the PPC also engages other cognitive processes in the transformation of behaviorally relevant visual inputs into appropriate actions, for example, monitoring of multiple locations, selection of responses to locations in space, and monitoring the outcome of response selections. We recorded single cells and local field potentials in the rat PPC during performance on a novel visuospatial attention (VSA) task that requires visually monitoring locations in space in order to make appropriate stimulus-guided locomotor responses. In each trial, rats attended to four locations on the floor of a maze. A randomly chosen location was briefly illuminated. Approach to the correct target location was followed by food reward. We observed that PPC activity correlated with multiple phases of the VSA task, including monitoring for stimulus onset, detection of a target, spatial location of the target, and target choice. A substantial proportion of cells with behavioral correlates were also modulated by outcome of the trial. Our analyses of local field potentials revealed strong oscillatory rhythms in the theta frequency band, and more than a third of PPC neurons were phase locked to theta oscillations. As in other brain regions, theta power correlated with running speed. Peak theta power was higher in superficial layers than deep layers providing evidence against volume conduction from the hippocampus. In addition, theta power was sensitive to the outcome of a choice. Theta power was significantly higher following incorrect choices compared with correct choices, possibly providing a prediction error signal. Our study provides evidence that the rat PPC has multiple roles in the translation of visual information into appropriate behavioral actions. © 2016 Wiley Periodicals, Inc.

Secondary visual cortex is critical to the expression of surprise-induced enhancements in cue associability in rats.

Considerable evidence indicates that reinforcement prediction error, the difference between the obtained and expected reinforcer values, modulates attention to potential cues for reinforcement. The surprising delivery or omission of a reinforcer enhances the associability of the stimuli that were present when the error was induced, so that they more readily enter into new associations in the future. Previous research from our laboratory identified brain circuit elements critical to the enhancement of stimulus associability by omission of an expected event and to the subsequent expression of that altered associability in more rapid learning. A key finding was that the rat posterior parietal cortex was essential during the encoding, consolidation and retrieval of associability memories that were altered by the surprising omission of an expected event in a serial prediction task. Here, we found that the function of adjacent secondary visual cortex was critical only to the expression of altered cue associability in that same task. This specialization of function is discussed in the context of broader cortical and subcortical networks for modulation of attention in associative learning, as well as recent anatomical investigations that suggest that the rodent posterior parietal cortex overlaps with and may subsume secondary visual cortex.

The role of rat posterior parietal cortex in coordinating spatial representations during place avoidance in dissociated reference frames on a continuously rotating arena (Carousel)

On the Carousel maze, rats are trained to avoid a sector of a circular rotating arena, punishable by a mild electric foot-shock. In the room frame (RF) variant, the punishable sector remains stable relative to the room, while in the arena frame (AF) version, the sector rotates with the arena. The rats therefore need to disregard local olfactory, tactile and self-motion cues in RF condition and distal extra-maze landmarks in the AF task. In both primates and rodents, the coordination of various spatial reference frames is thought to depend on the posterior parietal cortex (PPC). We have previously shown that PPC-lesioned rats can solve both variants of the Carousel avoidance task. Here we aimed to determine the effects of bilateral thermocoagulation lesion of the PPC in Long–Evans rats on the ability to transition between multiple spatial strategies. The rats were first trained in five sessions in one condition and then another five sessions in the other. The following training schemes were used: RF to AF, RF to RF reversal (sector on the opposite side), and AF to RF. We found a PPC lesion-associated impairment in the transition from the AF to RF task, but not vice versa. Furthermore, PPC lesion impaired performance in RF reversal. In accordance to the literature, we also found an impairment in navigation guided by intra-maze visuospatial cues, but not by extra-maze cues in the water maze. Therefore, the PPC lesion-induced impairment is neither specific to distant cues nor to allocentric processing. Our results thus indicate a role of the PPC in the flexibility in spatial behaviors guided by visual orientation cues.

A category-free neural population supports evolving demands during decision-making.

The posterior parietal cortex (PPC) receives diverse inputs and is involved in a dizzying array of behaviors. These multiple behaviors could rely on distinct categories of neurons specialized to represent particular variables or could rely on a single population of PPC neurons that is leveraged in different ways. To distinguish these possibilities, we evaluated rat PPC neurons recorded during multisensory decisions. Novel tests revealed that task parameters and temporal response features were distributed randomly across neurons, without evidence of categories. This suggests that PPC neurons constitute a dynamic network that is decoded according to the animal’s current needs. To test for an additional signature of a dynamic network, we compared moments when behavioral demands differ: decision and movement. Our novel state-space analysis revealed that the network explored different dimensions during decision and movement. These observations suggest that a single network of neurons can support the evolving behavioral demands of decision-making.

Interaction of egocentric and world-centered reference frames in the rat posterior parietal cortex.

Navigation requires coordination of egocentric and allocentric spatial reference frames and may involve vectorial computations relative to landmarks. Creation of a representation of target heading relative to landmarks could be accomplished from neurons that encode the conjunction of egocentric landmark bearings with allocentric head direction. Landmark vector representations could then be created by combining these cells with distance encoding cells. Landmark vector cells have been identified in rodent hippocampus. Given remembered vectors at goal locations, it would be possible to use such cells to compute trajectories to hidden goals. To look for the first stage in this process, we assessed parietal cortical neural activity as a function of egocentric cue light location and allocentric head direction in rats running a random sequence to light locations around a circular platform. We identified cells that exhibit the predicted egocentric-by-allocentric conjunctive characteristics and anticipate orienting toward the goal.

Deviance detection by a P3-like response in rat posterior parietal cortex

To better understand sensory processing in frontal and parietal cortex of the rat, and to further assess the rat as a model of human frontal-parietal processing, we recorded local field potentials (LFPs) from microelectrode arrays implanted in medio-dorsal frontal, and posterior parietal cortex of awake rats as they were presented with a succession of frequent “standard” tones and infrequent “oddball” tones. Extending previous results from surface recordings we found, after controlling for the frequencies of the standard and oddball tones, that rat frontal and parietal-evoked LFPs (eLFPs) exhibit significantly larger N1 (~40 ms latency), P2 (~100 ms), N2 (~160 ms), P3E (~200–240 ms), and P3L (~300–500 ms) amplitudes after an oddball tone. These neural oddball effects could contribute to the automatic allocation of attention to rare stimuli. To determine whether these enhanced responses to rare stimuli could be accounted for in terms of stimulus-specific neural adaptation (SSA), we also recorded during single-tone control sessions involving frequent standard, or infrequent oddball beeps alone. We compared the difference between rare-tone and frequent-tone response amplitudes in the two-tone context (oddball effect) or single-tone context which isolates the contribution of SSA (SSA effect). An analysis of variance (ANOVA) revealed a significant main effect of tone context on rare-tone response enhancements, showing that the rare-tone enhancements were stronger in the two-tone context than the single-tone context. This difference between tone contexts was greatest at the early P3E peak (200–240 ms post-beep) in parietal cortex, suggesting true deviance detection by this evoked response component, which cannot be accounted for in terms of simple models of SSA.

Functional Split between Parietal and Entorhinal Cortices in the Rat

Posterior parietal cortex (PPC) and medial entorhinal cortex (MEC) are important elements of the neural circuit for space, but whether representations in these areas are controlled by the same factors is unknown. We recorded single units simultaneously in PPC and MEC of freely foraging rats and found that a subset of PPC cells are tuned to specific modes of movement irrespective of the animals' location or heading, whereas grid cells in MEC expressed static spatial maps. The behavioral correlates of PPC cells switched completely when the same animals ran in a spatially structured maze or when they ran similar stereotypic sequences in an open arena. Representations in PPC were similar in identical mazes in different rooms where grid cells completely realigned their firing fields. The data suggest that representations in PPC are determined by the organization of actions while cells in MEC are driven by spatial inputs.

Interconnected cortical networks between primary somatosensory cortex septal columns and posterior parietal cortex in rat

Visual and somesthetic cues are used for spatial processing in the posterior parietal cortex (PPC) of the mammalian brain. In rats, somatic information collected by the mystacial whiskers is critically involved in constructing a neural representation of the external space. Here, we delineated the topography of the cortical pathway from the primary somatosensory cortex (SI) that may deliver vibrissal cues to PPC for spatial processing. For anterograde tracing, we made small injections of biotinylated dextran amine (BDA) into SI barrel cortex. The injections in the regions directly above the septal compartments produced dense terminals in PPC, whereas injections above the center of the barrels resulted in sparse terminals. For retrograde tracing, we made large injections of cholera toxin subunit B (CtB) in PPC. Retrogradely labeled neurons within SI barrel cortex formed multiple, parallel strips. In layer IV, these strips of labeled neurons were confined within the septal rows, extending from barrel arc position 0 to 5. In the extragranular layers, labeled neurons were clustered primarily within the vertical extensions of the septal rows and extended to the edges of neighboring barrel columns. Based on these findings, in which SI projections to PPC arise mainly from the septal columns, we hypothesize that septal columns may form interconnected cortical networks that engage in spatial information processing contingent on somestheic cues.

Interconnected cortical networks between primary somatosensory cortex septal columns and posterior parietal cortex in rat

AbstractVisual and somesthetic cues are used for spatial processing in the posterior parietal cortex (PPC) of the mammalian brain. In rats, somatic information collected by the mystacial whiskers is critically involved in constructing a neural representation of the external space. Here, we delineated the topography of the cortical pathway from primary somatosensory cortex (SI) that may deliver vibrissal cues to PPC for spatial processing. For anterograde tracing, we made small injections of biotinylated dextran amine (BDA) into SI barrel cortex. The injections in the regions directly above the septal compartments produced dense terminals in PPC, whereas injections above the center of the barrels resulted in sparse terminals. For retrograde tracing, we made large injections of cholera toxin subunit B (CtB) in PPC. Retrogradely labeled neurons within SI barrel cortex formed multiple, parallel strips. In layer IV, these strips of labeled neurons were confined within the septal rows, extending from barrel arc position 0 to 5. In the extragranular layers, labeled neurons were clustered primarily within the vertical extensions of the septal rows and extended to the edges of neighboring barrel columns. Based on these findings, in which SI projections to PPC arise mainly from the septal columns, we hypothesize that septal columns may form interconnected cortical networks that engage in spatial information processing contingent on somestheic cues. J. Comp. Neurol. 519:405–419, 2011. © 2011 Wiley‐Liss, Inc.

Low frequency oscillations in rat posterior parietal cortex are differentially activated by cues and distractors

The posterior parietal cortex (PPC) is hypothesized to detect visual cues among competing distractors. Anatomical and neurophysiologic evidence indicates that the rat PPC is part of a network of brain areas involved in directed attention, specifically when new task parameters or conditions are introduced. Here, we test the hypothesis that changes in the local field potential (LFP) of the PPC of rats performing a sustained attention task reflect aspects of detection. Two event-related potentials were observed during detection: the P300 response and the contingent negative variation (CNV). Spectrogram analysis also indicated a detection-specific increase in alpha power in the retention interval of this task. This is consistent with observations from human studies, which indicate that tasks requiring a subject to withhold a response produced a pronounced synchronization of alpha rhythms during the delay, and desynchronization during retrieval. We also found cycles of alpha synchrony and desynchrony in response to a periodic distractor. These cycles were most pronounced in the initial trial block of the distractor when the false alarm rate was highest, and as task performance improved these cycles significantly diminished. This result suggests that alpha cycling in the PPC represent neural activity critical for learning to inhibit distractors. The occurrence of alpha synchronization and desynchronization to attention-demanding stimuli, in addition to the P300 and CNV responses observed during detection, is evidence that rat PPC is involved in sustained attention, particularly in the presence of distractors.

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.

Lesion of posterior parietal cortex in rats does not disrupt place avoidance based on either distal or proximal orienting cues

A current topic in neurobiology is the study of the role of various brain structures in processing of spatial information. The present study was aimed at elucidating the role of the rat posterior parietal cortex in performing a place avoidance task. Two variants of the task were used: an arena frame task, in which animals were trained to avoid a sector defined by local cues bound to the surface of a rotating arena, and the room frame task, in which the shock sector was defined with respect to distal room landmarks. The results showed that both control and lesioned rats were able to efficiently solve both tasks, while locomotion was not altered. These results suggest that the posterior parietal cortex is not crucial for the processing of either proximal or distal cues in place avoidance.