Integrating what and when across the primate medial temporal lobe

Episodic memory or memory for the detailed events in our lives is critically dependent on structures of the medial temporal lobe (MTL). A fundamental component of episodic memory is memory for the temporal order of items within an episode. To understand the contribution of individual MTL structures to temporal-order memory, we recorded single-unit activity and local field potential from three MTL areas (hippocampus and entorhinal and perirhinal cortex) and visual area TE as monkeys performed a temporal-order memory task. Hippocampus provided incremental timing signals from one item presentation to the next, whereas perirhinal cortex signaled the conjunction of items and their relative temporal order. Thus, perirhinal cortex appeared to integrate timing information from hippocampus with item information from visual sensory area TE.     

Predictive codes for forthcoming perception in the frontal cortex

Incoming sensory information is often ambiguous, and the brain has to make decisions during perception. "Predictive coding" proposes that the brain resolves perceptual ambiguity by anticipating the forthcoming sensory environment, generating a template against which to match observed sensory evidence. We observed a neural representation of predicted perception in the medial frontal cortex, while human subjects decided whether visual objects were faces or not. Moreover, perceptual decisions about faces were associated with an increase in top-down connectivity from the frontal cortex to face-sensitive visual areas, consistent with the matching of predicted and observed evidence for the presence of faces.     

Encoding of spatial location by posterior parietal neurons

The cortex of the inferior parietal lobule in primates is important for spatial perception and spatially oriented behavior. Recordings of single neurons in this area in behaving monkeys showed that the visual sensitivity of the retinotopic receptive fields changes systematically with the angle of gaze. The activity of many of the neurons can be largely described by the product of a gain factor that is a function of the eye position and the response profile of the visual receptive field. This operation produces an eye position-dependent tuning for locations in head-centered coordinate space.     

Top-down versus bottom-up control of attention in the prefrontal and posterior parietal cortices

Attention can be focused volitionally by "top-down" signals derived from task demands and automatically by "bottom-up" signals from salient stimuli. The frontal and parietal cortices are involved, but their neural activity has not been directly compared. Therefore, we recorded from them simultaneously in monkeys. Prefrontal neurons reflected the target location first during top-down attention, whereas parietal neurons signaled it earlier during bottom-up attention. Synchrony between frontal and parietal areas was stronger in lower frequencies during top-down attention and in higher frequencies during bottom-up attention. This result indicates that top-down and bottom-up signals arise from the frontal and sensory cortex, respectively, and different modes of attention may emphasize synchrony at different frequencies.     

Spatial representation in the entorhinal cortex

As the interface between hippocampus and neocortex, the entorhinal cortex is likely to play a pivotal role in memory. To determine how information is represented in this area, we measured spatial modulation of neural activity in layers of medial entorhinal cortex projecting to the hippocampus. Close to the postrhinal-entorhinal border, entorhinal neurons had stable and discrete multipeaked place fields, predicting the rat's location as accurately as place cells in the hippocampus. Precise positional modulation was not observed more ventromedially in the entorhinal cortex or upstream in the postrhinal cortex, suggesting that sensory input is transformed into durable allocentric spatial representations internally in the dorsocaudal medial entorhinal cortex.     

Neuroscience. Where the brain monitors the body

Researchers haven't known exactly where in the brain all the signals that allow an animal to keep track of its limbs are located, but now a team may have located some of the neurons that first make these multisensory connections. On page 1782, researchers report evidence that a small region of the parietal cortex of the monkey brain known as area 5 may enable the monkey to integrate many sources of information about its body and thereby update its mental model of what the body is doing. The researchers based this conclusion on their finding that some area 5 neurons fire at their fastest rates when the visual feedback from a monkey's arm matches the sensory feedback, an indication that the neurons are sensitive to both streams of information.     

Neuroscience. More to seeing than meets the eye

A single event may elicit several different sensory stimuli such as vision, sound, and touch. But how does the brain know which of the many different stimuli arriving in the sensory cortex of the brain are connected? In her Perspective, de Gelder discusses new findings showing that when a touch is applied on the same side of the body as a visual cue, vision is enhanced (Macaluso et al.). She explains that this effect is due to neurons projecting from the somatosensory (touch) area of the sensory cortex back to the visual cortex. These neurons keep the visual cortex informed about tactile stimuli elicited at the same time as the visual stimuli.     

Neuronal activity in the lateral intraparietal area and spatial attention

Although the parietal cortex has been implicated in the neural processes underlying visual attention, the nature of its contribution is not well understood. We tracked attention in the monkey and correlated the activity of neurons in the lateral intraparietal area (LIP) with the monkey's attentional performance. The ensemble activity in LIP across the entire visual field describes the spatial and temporal dynamics of a monkey's attention. Activity subtending a single location in the visual field describes the attentional priority at that area but does not predict that the monkey will actually attend to or make an eye movement to that location.     

Backward spreading of memory-retrieval signal in the primate temporal cortex

Bidirectional signaling between neocortex and limbic cortex has been hypothesized to contribute to the retrieval of long-term memory. We tested this hypothesis by comparing the time courses of perceptual and memory-retrieval signals in two neighboring areas in temporal cortex, area TE (TE) and perirhinal cortex (PRh), while monkeys were performing a visual pair-association task. Perceptual signal reached TE before PRh, confirming its forward propagation. In contrast, memory-retrieval signal appeared earlier in PRh, and TE neurons were then gradually recruited to represent the sought target. A reasonable interpretation of this finding is that the rich backward fiber projections from PRh to TE may underlie the activation of TE neurons that represent a visual object retrieved from long-term memory.     

Comment on "Top-down versus bottom-up control of attention in the prefrontal and posterior parietal cortices"

Buschman and Miller (Reports, 30 March 2007, p. 1860) described the activity of ensembles of neurons in parietal and frontal cortex of monkeys performing visual search for targets that were easy or hard to distinguish from distractors. However, their conclusions are called into question by discrepancies between their results and publications from other laboratories measuring the same neural process.     

Modulation of human visual cortex by crossmodal spatial attention

A sudden touch on one hand can improve vision near that hand, revealing crossmodal links in spatial attention. It is often assumed that such links involve only multimodal neural structures, but unimodal brain areas may also be affected. We tested the effect of simultaneous visuo-tactile stimulation on the activity of the human visual cortex. Tactile stimulation enhanced activity in the visual cortex, but only when it was on the same side as a visual target. Analysis of effective connectivity between brain areas suggests that touch influences unimodal visual cortex via back-projections from multimodal parietal areas. This provides a neural explanation for crossmodal links in spatial attention.     

Visual stimuli and evoked responses in the rat

Rats were presented with a series of visual stimuli, each described by shape, size, and position. Cortical recordings were made in medial and lateral positions of rat visual cortex. Average responses were calculated and correlated. This analysis revealed that information regarding size was contained in electrical activity from medial areas, and informtion regarding shape in electrical activity from lateral areas.     

A map of visual space induced in primary auditory cortex

Maps of sensory surfaces are a fundamental feature of sensory cortical areas of the brain. The relative roles of afferents and targets in forming neocortical maps in higher mammals can be examined in ferrets in which retinal inputs are directed into the auditory pathway. In these animals, the primary auditory cortex contains a systematic representation of the retina (and of visual space) rather than a representation of the cochlea (and of sound frequency). A representation of a two-dimensional sensory epithelium, the retina, in cortex that normally represents a one-dimensional epithelium, the cochlea, suggests that the same cortical area can support different types of maps. Topography in the visual map arises both from thalamocortical projections that are characteristic of the auditory pathway and from patterns of retinal activity that provide the input to the map.     

Experimentally induced visual projections into auditory thalamus and cortex

Retinal cells have been induced to project into the medial geniculate nucleus, the principal auditory thalamic nucleus, in newborn ferrets by reduction of targets of retinal axons in one hemisphere and creation of alternative terminal space for these fibers in the auditory thalamus. Many cells in the medial geniculate nucleus are then visually driven, have large receptive fields, and receive input from retinal ganglion cells with small somata and slow conduction velocities. Visual cells with long conduction latencies and large contralateral receptive fields can also be recorded in primary auditory cortex. Some visual cells in auditory cortex are direction selective or have oriented receptive fields that resemble those of complex cells in primary visual cortex. Thus, functional visual projections can be routed into nonvisual structures in higher mammals, suggesting that the modality of a sensory thalamic nucleus or cortical area may be specified by its inputs during development.     

Physiological evidence for serial processing in somatosensory cortex

Removal of the representation of a specific body part in the postcentral cortex of the macaque resulted in the somatic deactivation of the corresponding body part in the second somatosensory area. In contrast, removal of the entire second somatosensory area had no grossly detectable effect on the somatic responsivity of neurons in the postcentral cortex. This direct electrophysiological evidence for serial cortical processing in somesthesia is similar to that found earlier for vision and, taken together with recent anatomical evidence, suggests that there is a common cortical plan for the processing of sensory information in the various sensory modalities.     

How visual stimuli activate dopaminergic neurons at short latency

Unexpected, biologically salient stimuli elicit a short-latency, phasic response in midbrain dopaminergic (DA) neurons. Although this signal is important for reinforcement learning, the information it conveys to forebrain target structures remains uncertain. One way to decode the phasic DA signal would be to determine the perceptual properties of sensory inputs to DA neurons. After local disinhibition of the superior colliculus in anesthetized rats, DA neurons became visually responsive, whereas disinhibition of the visual cortex was ineffective. As the primary source of visual afferents, the limited processing capacities of the colliculus may constrain the visual information content of phasic DA responses.     

Reorganization of retinotopic cortical maps in adult mammals after lesions of the retina

The organization of the visual cortex has been considered to be highly stable in adult mammals. However, 5 degrees to 10 degrees lesions of the retina in the contralateral eye markedly altered the systematic representations of the retina in primary and secondary visual cortex when matched inputs from the ipsilateral eye were also removed. Cortical neurons that normally have receptive fields in the lesioned region of the retina acquired new receptive fields in portions of the retina surrounding the lesions. The capacity for such changes may be important for normal adjustments of sensory systems to environmental contingencies and for recoveries from brain damage.     

Fast-forward playback of recent memory sequences in prefrontal cortex during sleep

As previously shown in the hippocampus and other brain areas, patterns of firing-rate correlations between neurons in the rat medial prefrontal cortex during a repetitive sequence task were preserved during subsequent sleep, suggesting that waking patterns are reactivated. We found that, during sleep, reactivation of spatiotemporal patterns was coherent across the network and compressed in time by a factor of 6 to 7. Thus, when behavioral constraints are removed, the brain's intrinsic processing speed may be much faster than it is in real time. Given recent evidence implicating the medial prefrontal cortex in retrieval of long-term memories, the observed replay may play a role in the process of memory consolidation.     

Selective attention from voluntary control of neurons in prefrontal cortex

Animals can learn to voluntarily control neuronal activity within various brain areas through operant conditioning, but the relevance of that control to cognitive functions is unknown. We found that rhesus monkeys can control the activity of neurons within the frontal eye field (FEF), an oculomotor area of the prefrontal cortex. However, operantly driven FEF activity was primarily associated with selective visual attention, and not oculomotor preparation. Attentional effects were untrained and were observed both behaviorally and neurophysiologically. Furthermore, selective attention correlated with voluntary, but not spontaneous, fluctuations in FEF activity. Our results reveal a specific association of voluntarily driven neuronal activity with "top-down" attention and suggest a basis for the use of neurofeedback training to treat disorders of attention.     

Two functional channels from primary visual cortex to dorsal visual cortical areas

Relationships between the M and P retino-geniculo-cortical visual pathways and "dorsal" visual areas were investigated by measuring the sources of local excitatory input to individual neurons in layer 4B of primary visual cortex. We found that contributions of the M and P pathways to layer 4B neurons are dependent on cell type. Spiny stellate neurons receive strong M input through layer 4Calpha and no significant P input through layer 4Cbeta. In contrast, pyramidal neurons in layer 4B receive strong input from both layers 4Calpha and 4Cbeta. These observations, along with evidence that direct input from layer 4B to area MT arises predominantly from spiny stellates, suggest that these different cell types constitute two functionally specialized subsystems.