Coding of intention in the posterior parietal cortex

To look at or reach for what we see, spatial information from the visual system must be transformed into a motor plan. The posterior parietal cortex (PPC) is well placed to perform this function, because it lies between visual areas, which encode spatial information, and motor cortical areas. The PPC contains several subdivisions, which are generally conceived as high-order sensory areas. Neurons in area 7a and the lateral intraparietal area fire before and during visually guided saccades. Other neurons in areas 7a and 5 are active before and during visually guided arm movements. These areas are also active during memory tasks in which the animal remembers the location of a target for hundreds of milliseconds before making an eye or arm movement. Such activity could reflect either visual attention or the intention to make movements. This question is difficult to resolve, because even if the animal maintains fixation while directing attention to a peripheral location, the observed neuronal activity could reflect movements that are planned but not executed. To address this, we recorded from the PPC while monkeys planned either reaches or saccades to a single remembered location. We now report that, for most neurons, activity before the movement depended on the type of movement being planned. We conclude that PPC contains signals related to what the animal intends to do.     

Multimodal integration for the representation of space in the posterior parietal cortex

The posterior parietal cortex has long been considered an 'association' area that combines information from different sensory modalities to form a cognitive representation of space. However, until recently little has been known about the neural mechanisms responsible for this important cognitive process. Recent experiments from the author's laboratory indicate that visual, somatosensory, auditory and vestibular signals are combined in areas LIP and 7a of the posterior parietal cortex. The integration of these signals can represent the locations of stimuli with respect to the observer and within the environment. Area MSTd combines visual motion signals, similar to those generated during an observer's movement through the environment, with eye-movement and vestibular signals. This integration appears to play a role in specifying the path on which the observer is moving. All three cortical areas combine different modalities into common spatial frames by using a gain-field mechanism. The spatial representations in areas LIP and 7a appear to be important for specifying the locations of targets for actions such as eye movements or reaching; the spatial representation within area MSTd appears to be important for navigation and the perceptual stability of motion signals.     

Electrical microstimulation distinguishes distinct saccade-related areas in the posterior parietal cortex

Electrical microstimulation (0.1-ms bipolar pulses at 500 Hz, current strength usually between 100 and 200 microA) was used to delineate saccade-related areas in the posterior parietal cortex of monkeys. Stimulation-induced saccades were found to be restricted to the lateral intraparietal area (area LIP) in the intraparietal sulcus (IPS) and a region on the medial aspect of the parietal lobe (area MP, medial parietal area), close to the caudal end of the cingulate sulcus, whereas stimulation of area 7a did not evoke eye movements. Two different types of evoked saccades were observed. Modified vector saccades, whose amplitude was modified by the position of the eyes at stimulation onset were the hallmark of sites in area LIP and area MP. The same sites were characterized by a propensity of single units active in the memory and presaccadic response segments of the memory saccade paradigm. Goal-directed saccades driving the eyes toward a circumscribed region relative to the head were largely restricted to a small strip of cortex on the lateral bank and the floor of the IPS (the intercalated zone), separating the representation of upward and downward directed saccades in LIP. Unlike stimulation in LIP or MP, stimulation in the intercalated zone gave rise to head, pinnae, facial, and shoulder movements accompanying the evoked saccades. We propose that the amplitude modification of vector saccades characterizing LIP and MP may reflect a spatially distributed head-centered coding scheme for saccades. On the other hand, the goal-directed saccades found in the intercalated zone could indicate the use of a spatially much more localized representation of desired location in head-centered space.     

Dissociation of attention in learning and action: Effects of lesions of the amygdala central nucleus, medial prefrontal cortex, and posterior parietal cortex.

Many associative learning theories assert that the predictive accuracy of events affects the allocation of attention to them. More reliable predictors of future events are usually more likely to control action based on past learning, but less reliable predictors are often more likely to capture attention when new information is acquired. Previous studies showed that a circuit including the amygdala central nucleus (CEA) and the cholinergic substantia innominata/nucleus basalis magnocellularis (SI/nBM) is important for both sustained attention guiding action in a five-choice serial reaction time (5CSRT) task and for enhanced new learning about less predictive cues in a serial conditioning task. In this study, the authors found that lesions of the cholinergic afferents of the medial prefrontal cortex interfered with 5CSRT performance but not with surprise-induced enhancement of learning, whereas lesions of cholinergic afferents of posterior parietal cortex impaired the latter effects but did not affect 5CSRT performance. CEA lesions impaired performance in both tasks. These results are consistent with the view that CEA affects these distinct aspects of attention by influencing the activity of separate, specialized cortical regions via modulation of SI/nBM. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Unilateral injury of posterior parietal cortex and spatial learning in hooded rats

The influences of bilateral or unilateral injuries within the posterior parietal cortex (PPC) upon spatial learning in a water maze were examined in three experiments. Place-learning and response-learning were investigated in a four-alley 'Greek-cross' shaped water maze with extra-maze visual cues available. No differences were detected on any of several measures sensitive to learning between the lesion groups on the place-learning task. Microanalysis of behavior within trials revealed that animals with either bilateral or right unilateral PPC injuries committed significantly more total errors, initial alley entrance ('reference memory') errors, and re-entry ('working memory') errors in the response-learning paradigm than did either the control or left PPC-injured rats. No differences were detected between the latter two groups on these measures. Unilateral lesions resulted in asymmetrical placing responses ipsilateral to the injury 10 days after surgery whereas bilateral injuries resulted in asymmetrical placing with mixed directionality. The acquisition of the response-learning problem in the absence of visual cues was studied on animals prepared with unilateral lesions and housed post-operatively either in isolation or in a 'complex environment.' In the absence of visual cues both right and left PPC-injured rats committed more errors than sham controls, and differential post-surgical housing did not attenuate these impairments. These same animals were trained on the landmark navigation task. Although no differences appeared between the lesion groups, a generalized but transient facilitation of learning was observed in animals housed in the 'complex' environment. Unilateral injuries placed in sham controls failed to disturb retention of the landmark navigation strategy. Because none of the PPC-injured animals were deficient in the landmark task, a result which is contrary to observations in other laboratories, the influence of post-surgical recovery interval upon acquisition of the landmark navigation strategy was explored. Animals were prepared with right PPC injuries and trained following either a 5 or 35 day recovery interval. Only those animals limited to the short recovery interval proved to have a spatial deficit in the landmark task. It is concluded that injuries in the PPC of either hemisphere disturb egocentric spatial functions. However, animals with left PPC injuries are able to compensate by using allocentric visual cues if they are available. It is due to the special role played by the right PPC in complex visuospatial functions that animals with this injury are unable to compensate.     

Role of the posterior parietal association cortex in the processing of spatial event information.

Humans and monkeys with damage to the parietal association cortex (PC) exhibit deficits on visual-spatial tasks that are often dependent upon the attentional requirements of the task at hand. In order to test whether there is a correspondence between humans and rats in terms of the function of PC in the processing of spatial information, separate groups of rats were trained in two list-learning tasks, a win–stay task and a win–shift task, on an eight-arm radial maze. It is assumed that these two tasks vary in degree of required attention effort, because the win–shift rule is considerably easier for the rat to learn than the win–stay rule. One group of animals in each condition was given preoperative training before receiving bilateral aspiration lesions of PC. Another group in each condition received PC lesions prior to training. The results indicated that animals with PC lesions are unable to acquire either the win–stay or the win–shift rule. However, of the animals with preoperative training, only the win–stay trained animals were deleteriously affected by the brain lesion. The win–shift animals showed no postoperative decline in performance. Because rats with PC lesions are only impaired on the more demanding spatial task, these results are consistent with the human clinical observations. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Specific changes in conditioned responding following neurotoxic damage to the posterior parietal cortex.

The central nucleus (CN) of the amygdala and basal forebrain cholinergic projections to the posterior parietal cortex (PPC) are involved in regulating changes in attentional processing of conditioned stimuli. In a previous study, lesions of the CN produced a deficit in conditioned orienting behavior (rearing on the hind legs) when a visual stimulus was paired with food. Unconditioned orienting (rearing to nonreinforced presentations of the stimulus) and conditioned food cup behavior were unaffected. The present study examined the contribution of the PPC to attentional orienting behavior. Damage to the PPC did not affect orienting behavior but produced deficits in food cup behavior. These findings help define the specific contributions of the PPC to attentional processing and associative learning. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Differential activation of right superior parietal cortex and intraparietal sulcus by spatial and nonspatial attention

Malignant brain tumors (primary and metastatic) are apparently resistant to most therapeutic efforts. Several randomized trials have provided evidence supporting the efficacy of radiation therapy. Attempts at improving the results of external beam radiotherapy include altered fractionation, radiation sensitizers and concomitant chemotherapy. In low-grade gliomas, all clinical studies with radiotherapy have employed conventional dose fractionation regimens. In high-grade gliomas, hypofractionation schedules represent effective palliative regimens in poor prognosis subsets of patients; short-term survival in these patients has not allowed to evaluate late toxicity. In tumors arising within the central nervous system, hyperfractionated irradiation exploits the differences in repair capacity between tumour and late responding normal tissues. It may allow for higher total dose and may result in increased tumor cell kill. Accelerated radiotherapy may reduce the repopulation of tumor cells between fractions. It may potentially improve tumor control for a given dose level, provided that there is no increase in late normal tissue injury. In supratentorial malignant gliomas, superiority of accelerated hyperfractionated over conventionally fractionated schedules was observed in a randomized trial; however, the gain in survival was less than 6 months. At present no other randomized trial supports the preferential choice for altered fractionation irradiation. Also in pediatric brainstem tumors there are no data to confirm the routine use of hyperfractionated irradiation, and significant late sequelae have been reported in the few long-term survivors. Shorter treatment courses with accelerated hyperfractionated radiotherapy may represent a useful alternative to conventional irradiation for the palliation of brain metastases. Different considerations have been proposed to explain this gap between theory and clinical data. Patients included in dose/effect studies are not stratified by prognostic factors and other treatment-related parameters. This observation precludes any definite conclusion about the relative role of conventional and of altered fractionation. New approaches are currently in progress. More prolonged radiation treatments, up to higher total doses, could delay time to tumor progression and improve survival in good prognosis subsets of patients; altered fractionation may be an effective therapeutic tool to achieve this goal.     

Matching patterns of activity in primate prefrontal area 8a and parietal area 7ip neurons during a spatial working memory task

Single-unit recording studies of posterior parietal neurons have indicated a similarity of neuronal activation to that observed in the dorsolateral prefrontal cortex in relation to performance of delayed saccade tasks. A key issue addressed in the present study is whether the different classes of neuronal activity observed in these tasks are encountered more frequently in one or the other area or otherwise exhibit region-specific properties. The present study is the first to directly compare these patterns of neuronal activity by alternately recording from parietal area 7ip and prefrontal area 8a, under the identical behavioral conditions, within the same hemisphere of two monkeys performing an oculomotor delayed response task. The firing rate of 222 posterior parietal and 235 prefrontal neurons significantly changed during the cue, delay, and/or saccade periods of the task. Neuronal responses in the two areas could be distinguished only by subtle differences in their incidence and timing. Thus neurons responding to the cue appeared earliest and were more frequent among the task-related neurons within parietal cortex, whereas neurons exhibiting delay-period activity accounted for a larger proportion of task-related neurons in prefrontal cortex. Otherwise, the task-related neuronal activities were remarkably similar. Cue period activity in prefrontal and parietal cortex exhibited comparable spatial tuning and temporal duration characteristics, taking the form of phasic, tonic, or combined phasic/tonic excitation in both cortical populations. Neurons in both cortical areas exhibited sustained activity during the delay period with nearly identical spatial tuning. The various patterns of delay-period activity-tonic, increasing or decreasing, alone or in combination with greater activation during cue and/or saccade periods-likewise were distributed to both cortical areas. Finally, similarities in the two populations extended to the proportion and spatial tuning of presaccadic and postsaccadic neuronal activity occurring in relation to the memory-guided saccade. The present findings support and extend evidence for a faithful duplication of receptive field properties and virtually every other dimension of task-related activity observed when parietal and prefrontal cortex are recruited to a common task. This striking similarity attests to the principal that information shared by a prefrontal region and a sensory association area with which it is connected is domain specific and not subject to hierarchical elaboration, as is evident at earlier stages of visuospatial processing.     

Activity in posterior parietal cortex following somatosensory stimulation in man: magnetoencephalographic study using spatio-temporal source analysis

The cyclin-dependent kinase (Cdk) inhibitor p21Waf1/Cip1/Sdi1, important for p53 tumor suppressor-dependent cell growth control in humans and other organisms, mediates G1/S-phase arrest through inhibition of cyclin-dependent kinases (Cdks). The enzymatic activity of these kinases is essential for progress through the cell division cycle and one level of cell cycle regulation is exerted through inhibition of Cdks by a family of small proteins, including p21. Cdk inhibition requires a sequence of approximately 60 amino acids within the p21 NH2-terminus. Using proteolytic mapping, matrix-assisted laser desorption/ionization (MALDI) mass spectrometry, HPLC and size-exclusion chromatography, we show that p21, active as a Cdk inhibitor, exists in an extended, non-globular conformation in the absence of its biological target and that p21 lacks the hallmarks of stable secondary and tertiary structure. We have developed an efficient approach to obtain detailed proteolytic maps that takes advantage of the high accuracy and sensitivity of MALDI mass spectrometry. Our method allows a proteolytic map to be obtained from a single mass spectrum for fragments produced from a single proteolytic reaction.     

Covert orienting of visuospatial attention in children with developmental coordination disorder

It is still unclear whether impairments in visuospatial processing in children with developmental coordination disorder (DCD) are a consequence of their motor deficits or are independent of them. In two experiments, 20 children with DCD and 20 matched controls were tested on the covert orienting of a visuospatial attention task (COVAT). Experiment 1 used a COVAT with peripheral cues and an 80% probability that targets would appear at the cued location. While the results suggested a deficit in the disengage operation of orienting covert attention for the DCD group, they were difficult to reconcile with models of covert orienting and the results of past research. Experiment 2 tested subjects on two new versions of the COVAT: the first used peripheral cues and no probability information (exogenous mode), and the second used central cues and an 80% probability that targets would appear at the cued location (endogenous mode). The DCD group displayed attentional orienting deficits only for the endogenous mode. These results suggest that impairments in the endogenous control of visuospatial attention are independent of motor deficits in DCD.     

Gating of neuronal responses in macaque primary visual cortex by an attentional spotlight

Neuronal responses were recorded from the striate cortex of monkeys trained to perform visual discrimination at locations in the visual field to which their attention was drawn. A subset of neurons showed vigorous responses to visual stimuli for trials in which the monkey was directing its attention to the respective receptive field location. In trials where attention is directed elsewhere, responses to the same stimuli were significantly reduced. In some cells the early response component was not modulated by attention, but later components were affected by the locus of attention. The results suggest the operation of a feedback in the paradigm that spotlights a topographically restricted area of V1 for further processing at higher levels.     

Olfactory neuronal responses in the primate orbitofrontal cortex: analysis in an olfactory discrimination task

1. The primate orbitofrontal cortex receives inputs from the primary olfactory (pyriform) cortex and also from the primary taste cortex. To investigate how olfactory information is encoded in the orbitofrontal cortex, the responses of single neurons in the orbitofrontal cortex and surrounding areas were recorded during the performance of an olfactory discrimination task. In the task, the delivery of one of eight different odors indicated that the monkey could lick to obtain a taste of sucrose. If one of two other odors was delivered from the olfactometer, the monkey had to refrain from licking, otherwise he received a taste of saline. 2. Of the 1,580 neurons recorded in the orbitofrontal cortex, 3.1% (48) had olfactory responses and 34 (2.2%) responded differently to the different odors in the task. The neurons responded with a typical latency of 180 ms from the onset of odorant delivery. 3. Of the olfactory neurons with differential responses in the task, 35% responded solely on the basis of the taste reward association of the odorants. Such neurons responded either to all the rewarded stimuli, and none of the saline-associated stimuli, or vice versa. 4. The remaining 65% of these neurons showed differential selectivity for the stimuli based on the odor quality and not on the taste reward association of the odor. 5. The findings show that the olfactory representation within the orbitofrontal cortex reflects for some neurons (65%) which odor is present independently of its association with taste reward, and that for other neurons (35%), the olfactory response reflects (and encodes) the taste association of the odor. The additional finding that some of the odor-responsive neurons were also responsive to taste stimuli supports the hypothesis that odor-taste association learning at the level of single neurons in the orbitofrontal cortex enables such cells to show olfactory responses that reflect the taste association of the odor.     

Electrical responses of the auditory area of the cerebellar cortex to acoustic stimulation

Single unit activity from the VI and VII lobuli of the cerebellar vermis cortex was studied following acoustical stimulation with sound signals of different parameters. Cerebellar neurons, as compared to those from the auditory system, showed low selectivity to sound frequency, intensity and duration. However, about 2/3 of the neurons were selectively sensitive to interaural time and intensity differences; about 1/3 of neurons showed a specific response to signals simulating sound motion in a definite direction. Thus, cerebellar neurons seem to be mainly responsive to those sound parameters which are essential for sound localization.     

Functional specialization in auditory cortex: responses to frequency-modulated stimuli in the cat's posterior auditory field

The mammalian auditory cortex contains multiple fields but their functional role is poorly understood. Here we examine the responses of single neurons in the posterior auditory field (P) of barbiturate- and ketamine-anesthetized cats to frequency-modulated (FM) sweeps. FM sweeps traversed the excitatory response area of the neuron under study, and FM direction and the linear rate of change of frequency (RCF) were varied systematically. In some neurons, sweeps of different sound pressure levels (SPLs) also were tested. The response magnitude (number of spikes corrected for spontaneous activity) of nearly all field P neurons varied with RCF. RCF response functions displayed a variety of shapes, but most functions were of low-pass characteristic or peaked at rather low RCFs (<100 kHz/s). Neurons with strong responses to high RCFs (high-pass or nonselective RCF response function characteristics) all displayed spike count-SPL functions to tone burst onsets that were monotonic or weakly nonmonotonic. RCF response functions and best RCFs often changed with SPL. For most neurons, FM directional sensitivity, quantified by a directional sensitivity (DS) index, also varied with RCF and SPL, but the mean and width of the distribution of DS indices across all neurons was independent of RCF. Analysis of response timing revealed that the phasic response of a neuron is triggered when the instantaneous frequency of the sweep reaches a particular value, the effective Fi. For a given neuron, values of effective Fi were independent of RCF, but depended on FM direction and SPL and were associated closely with the boundaries of the neuron's frequency versus amplitude response area. The standard deviation (SD) of the latency of the first spike of the response decreased with RCF. When SD was expressed relative to the rate of change of stimulus frequency, the resulting index of frequency jitter increased with RCF and did so rather uniformly in all neurons and largely independent of SPL. These properties suggest that many FM parameters are represented by, and may be encoded in, orderly temporal patterns across different neurons in addition to the strength of responses. When compared with neurons in primary and anterior auditory fields, field P neurons respond better to relatively slow FMs. Together with previous studies of responses to modulations of amplitude, such as tone onsets, our findings suggest more generally that field P may be best suited for processing signals that vary relatively slowly over time.     

Projection pattern of single corticocortical fibers from the parietal cortex to the motor cortex

Arborization of single corticocortical (CC) axons projecting from the parietal cortex to the motor cortex (Mx) was analysed using an intraaxonal staining technique in the cat. Stem axons arising from cell bodies in area 5 ramified repeatedly into numerous terminal branches in the Mx, forming 2-6 patches (0.2-0.8 mm in diameter) separated by a terminal-free gap. Axon terminals were distributed mainly in layers II and III and sparsely in layers V, VI and I. This feature is quite similar to that of thalamocortical axons and other corticocortical fibres. Thus the patchy organization may be a basic input structure for afferents of the Mx and play a role in generation of adequate motor output patterns in the Mx.     

Lateralization of emotionality in right parietal cortex of the rat.

Lesions of right parietal cortex in the rat increase activity in the open field compared with left parietal lesions, especially after section of the corpus callosum. Left or right motor or medial frontal cortex lesions do not have a lateralized effect. This evidence of a localized asymmetry between the cerebral hemispheres strongly implies that right parietal cortex has a role in emotionality in this species. Our findings suggest a functional similarity to right parietal cortex in man. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Landmark control and updating of self-movement cues are largely maintained in head direction cells after lesions of the posterior parietal cortex.

Head direction (HD) cells discharge as a function of the rat's directional orientation with respect to its environment. Because animals with posterior parietal cortex (PPC) lesions exhibit spatial and navigational deficits, and the PPC is indirectly connected to areas containing HD cells, we determined the effects of bilateral PPC lesions on HD cells recorded in the anterodorsal thalamus. HD cells from lesioned animals had similar firing properties compared to controls and their preferred firing directions shifted a corresponding amount following rotation of the major visual landmark. Because animals were not exposed to the visual landmark until after surgical recovery, these results provide evidence that the PPC is not necessary for visual landmark control or the establishment of landmark stability. Further, cells from lesioned animals maintained a stable preferred firing direction when they foraged in the dark and were only slightly less stable than controls when they self-locomoted into a novel enclosure. These findings suggest that PPC does not play a major role in the use of landmark and self-movement cues in updating the HD cell signal, or in its generation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Covert visual attention modulates face-specific activity in the human fusiform gyrus: fMRI study

Several lines of evidence demonstrate that faces undergo specialized processing within the primate visual system. It has been claimed that dedicated modules for such biologically significant stimuli operate in a mandatory fashion whenever their triggering input is presented. However, the possible role of covert attention to the activating stimulus has never been examined for such cases. We used functional magnetic resonance imaging to test whether face-specific activity in the human fusiform face area (FFA) is modulated by covert attention. The FFA was first identified individually in each subject as the ventral occipitotemporal region that responded more strongly to visually presented faces than to other visual objects under passive central viewing. This then served as the region of interest within which attentional modulation was tested independently, using active tasks and a very different stimulus set. Subjects viewed brief displays each comprising two peripheral faces and two peripheral houses (all presented simultaneously). They performed a matching task on either the two faces or the two houses, while maintaining central fixation to equate retinal stimulation across tasks. Signal intensity was reliably stronger during face-matching than house matching in both right- and left-hemisphere predefined FFAs. These results show that face-specific fusiform activity is reduced when stimuli appear outside (vs. inside) the focus of attention. Despite the modular nature of the FFA (i.e., its functional specificity and anatomic localization), face processing in this region nonetheless depends on voluntary attention.     

Activation of the visual cortex in motivated attention.

Functional activation (measured with fMRI) in occipital cortex was more extensive when participants view pictures strongly related to primary motive states (i.e., victims of violent death, viewer-directed threat, and erotica). This functional activity was greater than that observed for less intense emotional (i.e., happy families or angry faces) or neutral images (i.e., household objects, neutral faces). Both the extent and strength of functional activity were related to the judged affective arousal of the different picture contents, and the same pattern of functional activation was present whether pictures were presented in color or in grayscale. It is suggested that more extensive visual system activation reflects "motivated attention," in which appetitive or defensive motivational engagement directs attention and facilitates perceptual processing of survival-relevant stimuli. (PsycINFO Database Record (c) 2010 APA, all rights reserved)