Most models assume that top-down attention enhances the gain of sensory neurons tuned to behaviorally relevant stimuli (on-target gain). However, theoretical work suggests that when targets and distracters are highly similar, attention should enhance the gain of neurons that are tuned away from the target, because these neurons better discriminate neighboring features (off-target gain). While it is established that off-target neurons support difficult fine discriminations, it is unclear if top-down attentional gain can be optimally applied to informative off-target sensory neurons or if gain is always applied to on-target neurons, regardless of task demands. To test the optimality of attentional gain in human visual cortex, we used functional magnetic resonance imaging and an encoding model to estimate the response profile across a set of hypothetical orientation-selective channels during a difficult discrimination task. The results suggest that top-down attention can adaptively modulate off-target neural populations, but only when the discriminanda are precisely specified in advance. Furthermore, logistic regression revealed that activation levels in off-target orientation channels predicted behavioral accuracy on a trial-by-trial basis. Overall, these data suggest that attention does not only increase the gain of sensory-evoked responses, but may bias population response profiles in an optimal manner that respects both the tuning properties of sensory neurons and the physical characteristics of the stimulus array.

1. The purpose of this study is to define the cortical regions that subserve voluntary saccadic eye movements and spatial working memory in humans. 2. Regional cerebral blood flow (rCBF) during performance of oculomotor tasks was measured with [15O]-H2O positron emission tomography (PET). Eleven well-trained, healthy young adults performed the following tasks: visual fixation, visually guided saccades, antisaccades (a task in which subjects made saccades away from rather than toward peripheral targets), and either an oculomotor delayed response (ODR, a task requiring memory-guided saccades after a delay period) or a conditional antisaccade task (a task in which the color of the peripheral target determined whether a saccade toward or away from the target was required). An additional six subjects performed a sequential hand movement task to compare localization of hand-related motor cortex and the frontal eye fields (FEFs) and of the hand- and eye-movement-related regions of the supplementary motor area (SMA). 3. Friston's statistical parametric mapping (SPM) method was used to identify significant changes in rCBF associated with task performance. Because SPM does not take advantage of the anatomic information available in magnetic resonance (MR) scans, each subject's PET scan was registered to that individual's MR scan, after which all PET and MR studies were transformed to conform to a standard reference MR image set. Subtraction images were visually inspected while overlayed on the reference MR scan to which PET images had been aligned, in order to confirm anatomic localization of significant rCBF changes. 4. Compared with visual fixation, performing visually guided saccades led to a significant bilateral activation in FEF, cerebellum, striate cortex, and posterior temporal cortex. Right posterior thalamus activation was also observed. 5. The visually guided saccade task served as the comparison task for the ODR, antisaccade, and conditional antisaccade tasks for identification of task-related changes in rCBF beyond those associated with saccade execution. Performance on the ODR task was associated with a bilateral increase of rCBF in FEFs, SMA, dorsolateral prefrontal cortex (DLPFC), and posterior parietal cortex. The cortical regions of increased regional blood flow during the ODR task also showed increased rCBF during the antisaccade task; however, FEF and SMA activations were significant only in the right hemisphere. These findings closely parallel those of single-cell recording studies with behaving monkeys in indicating that FEF, DLPFC, SMA, and posterior parietal cortex perform computational activity for voluntary purposive saccades. 6. Comparison of PET scans obtained during performance of eye movement and hand movement tasks indicated that peak activations in FEF were located approximately 2 cm lateral and 1 cm anterior to those of hand-related motor cortex. The oculomotor area of SMA, the supplementary eye field (SEF), was located approximately 7-8 mm anterior and superior to the hand-related area of SMA. 7. During performance of antisaccade and ODR tasks, rCBF was significantly lower in ventromedial prefrontal cortex (PFC), along the rectus gyrus, and in ventral anterior cingulate cortex than during the visually guided saccade and fixation tasks. During the antisaccade task, the ventral region of lower rCBF involved medial structures including left ventral striatum and bilateral medial temporal-limbic cortex. During the ODR task, the ventral aspect of the region of lower rCBF extended laterally, rather than medially, to include the temporal poles. The lower blood flow observed in ventromedial PFC during both the antisaccade and ODR tasks, relative to the visually guided saccade and fixation tasks, suggests that modulation of output from ventromedial PFC to limbic cortex and the striatum may play a role in the voluntary control of saccadic eye movements, possibly in the suppression of responses that would interrupt

Functional Magnetic Resonance Imaging (fMRI) was used to identify a small area in the human posterior fusiform gyrus that responds selectively to faces (PF). In the same subjects, phase-encoded rotating and expanding checkerboards were used with fMRI to identify the retinotopic visual areas V1, V2, V3, V3A, VP and V4v. PF was found to lie anterior to area V4v, with a small gap present between them. Further recordings in some of the same subjects used moving low-contrast rings to identify the visual motion area MT. PF was found to lie ventral to MT. In addition, preliminary evidence was found using fMRI for a small area that responded to inanimate objects but not to faces in the collateral sulcus medial to PF. The retinotopic visual areas and MT responded equally to faces, control randomized stimuli, and objects. Weakly face-selective responses were also found in ventrolateral occipitotemporal cortex anterior to V4v, as well as in the middle temporal gyrus anterior to MT. We conclude that the fusiform face area in humans lies in non-retinotopic visual association cortex of the ventral form-processing stream, in an area that may be roughly homologous in location to area TF or CITv in monkeys.

The microvascular relationships important to surgery of aneurysms in the anterior communicating region were defined in 50 cadaver brains. The recurrent artery of Heubner was frequently exposed before the A-1 segment in defining the neck on anterior cerebral aneurysms because it commonly courses anterior to A-1. It arose from the A-2 segment of the anterior cerebral artery (ACA) in 78% and most commonly terminated in the area of the anterior perforated substance, and lateral to it in the Sylvian fissure. The anterior communicating artery (ACoA) frequently gave rise to perforating arteries which terminated in the superior surface of the optic chiasm and above the chiasm in the anterior hypothalamus. This finding contrasts with previous reports that no perforating branches arise from the communicating artery. The proximal half of the A=1 segment was a richer source of perforating arteries than the distal half. The A-1 branches most commonly terminated in the anterior perforated substance, the optic chiasm, and the region of the optic tract. The ACoA increased in size as the difference in the diameter between the right and left A-1 segments increased. Frequent variants such as double or triple ACoA's, triple A-2 segments, and duplication of the A-1 segments were encountered. The clinical consequences of occlusion of the recurrent artery and of the perforators from the ACoA and medial and lateral segment of A-1 are reviewed.

Perception of biological motions plays a major adaptive role in identifying, interpreting, and predicting the actions of others. It may therefore be hypothesized that the perception of biological motions is subserved by a specific neural network. Here we used fMRI to verify this hypothesis. In a group of 10 healthy volunteers, we explored the hemodynamic responses to seven types of visual motion displays: drifting random dots, random dot cube, random dot cube with masking elements, upright point-light walker, inverted point-light walker, upright point-light walker display with masking elements, and inverted point-light walker display with masking elements. A gradient in activation was observed in the occipitotemporal junction. The responses to rigid motion were localized posteriorly to those responses elicited by nonrigid motions. Our results demonstrate that in addition to the posterior portion of superior temporal sulcus, the left intraparietal cortex is involved in the perception of nonrigid biological motions.

Posterior parietal cortex (PPC) is thought to play a critical role in decision making, sensory attention, motor intention, and/or working memory. Research on the PPC in non-human primates has focused on the lateral intraparietal area (LIP) in the intraparietal sulcus (IPS). Neurons in LIP respond after the onset of visual targets, just before saccades to those targets, and during the delay period in between. To study the function of posterior parietal cortex in humans, it will be crucial to have a routine and reliable method for localizing specific parietal areas in individual subjects. Here, we show that human PPC contains at least two topographically organized regions, which are candidates for the human homologue of LIP. We mapped the topographic organization of human PPC for delayed (memory guided) saccades using fMRI. Subjects were instructed to fixate centrally while a peripheral target was briefly presented. After a further 3-s delay, subjects made a saccade to the remembered target location followed by a saccade back to fixation and a 1-s inter-trial interval. Targets appeared at successive locations "around the clock"(same eccentricity, approximately 30 degrees angular steps), to produce a traveling wave of activity in areas that are topographically organized. PPC exhibited topographic organization for delayed saccades. We defined two areas in each hemisphere that contained topographic maps of the contra-lateral visual field. These two areas were immediately rostral to V7 as defined by standard retinotopic mapping. The two areas were separated from each other and from V7 by reversals in visual field orientation. However, we leave open the possibility that these two areas will be further subdivided in future studies. Our results demonstrate that topographic maps tile the cortex continuously from V1 well into PPC.

In noninvasive neuroimaging, neural activity is inferred from local fluctuations in deoxyhemoglobin. A fundamental question of functional magnetic resonance imaging (fMRI) is whether the inferred neural activity is driven primarily by synaptic or spiking activity. The answer is critical for the interpretation of the blood oxygen level-dependent (BOLD) signal in fMRI. Here, we have used well-established visual-system circuitry to create a stimulus that elicits synaptic activity without associated spike discharge. In colocalized recordings of neural and metabolic activity in cat primary visual cortex, we observed strong coupling between local field potentials (LFPs) and changes in tissue oxygen concentration in the absence of spikes. These results imply that the BOLD signal is more closely coupled to synaptic activity.

Positron emission tomography (PET) was used to identify cortical and subcortical regions involved in the control of reaching to visual targets. Regional cerebral blood flow (rCBF) was measured in eight healthy subjects using H2(15)O PET during the performance of three different tasks. All tasks required central fixation while a 400-ms target was flashed every 5 s at a random location around a virtual circle centered on the fixation target. Additional instructions differed according to the task:(i) visual detection of the target without overt responses;(ii) immediate pointing to the most recent target in the sequence, and (iii) pointing to the previous target in the sequence. By design, the two motor tasks differed in the cognitive processing required. In each trial of immediate pointing, the spatial location of only the most recent target needed to be processed. In each trial of pointing to the previous, instead, while the most recent target was stored in memory for the movement of the next trial, the previous target had to be retrieved from memory to direct the current movement. Limb trajectories were comparable between the two motor tasks in terms of most spatiotemporal parameters examined. Significant rCBF increases were identified using analysis of covariance and t statistics. Compared with visual detection there was activation of primary sensorimotor cortex, ventrolateral precentral gyrus, inferior frontal gyrus in the opercular region, supramarginal gyrus, and middle occipital gyrus, all these sites in the hemisphere (left) contralateral to the moving limb, and cerebellar vermis, during both immediate pointing and pointing to the previous. During immediate pointing there was additional activation of left inferior parietal lobule close to the intraparietal sulcus, and when compared with pointing to the previous, dorsolateral prefrontal cortex bilaterally. During pointing to the previous, instead, there was additional activation of supplementary motor cortex, anterior and midcingulate, and inferior occipital gyrus in the left hemisphere; superior parietal lobule, supramarginal gyrus, and posterior hippocampus in the right hemisphere; lingual gyri and cerebellar hemispheres bilaterally; anterior thalamus; and pulvinar. The activation of two partially distinct cerebral networks in these two motor tasks reflects the different nature of signal processing involved. In particular, the specific activation of intraparietal sulcus and prefrontal cortex in immediate pointing appears characteristic of a network for visuospatial working memory. By contrast, the corticolimbic network engaged in pointing to the previous could mediate spatial attention and the sequence of encoding, recording, and decoding of spatial memories required by a dual task with two competing targets.

The primate visual system is believed to comprise two main pathways: a ventral pathway for conscious perception and a dorsal pathway that can process visual information and guide action without accompanying conscious knowledge. Evidence for this theory has come primarily from studies of neurological patients and animals. Using fMRI, we show here that even though observers are completely unaware of test object images owing to interocular suppression, their dorsal cortical areas demonstrate substantial activity for different types of visual objects, with stronger responses to images of tools than of human faces. This result also suggests that in binocular rivalry, substantial information in the suppressed eye can escape the interocular suppression and reach dorsal cortex.

We have addressed the question of whether the brain's capacity to resolve an ambiguous retinal image depends upon the activity of early visual areas or whether it involves the investment of the received image with higher order cognitive hypotheses. To resolve the issue, we have used the technique of positron emission tomography to detect increases in regional cerebral blood flow (rCBF) in the brains of humans while they perceive the simple figures described by Schumann (1900) and by Kanizsa (1979). These figures produce striking percepts of surfaces or contours variously described as illusory, subjective, cognitive, or anomalous because they depend upon the brain's ability to complete the figures. If such completion is due to higher order cognitive processes or a combination of higher order and early areas, then, one might expect areas of increased rCBF outside the occipital lobe when subjects perceive these figures. However, if completion is mediated entirely by early visual areas, then the increases in rCBF will be restricted to these regions. Our results show that the perception of subjective contours is associated with significant activity in early visual areas only, particularly in area V2, leading us to conclude that the occipital cortex can contribute to the perception of these stimuli without higher order cognitive influence specific to the completion task.

OBJECTIVE This study was undertaken to identify brain structures associated with emotion in normal elderly subjects. METHOD Eight normal subjects aged 55-78 years were shown film clips intended to provoke the emotions of happiness, fear, or disgust as well as a neutral state. During emotional activation, regional cerebral blood flow was measured with the use of [15O]H2O positron emission tomography imaging, and subjective emotional responses were recorded. Data were analyzed by subtracting the values during the neutral condition from the values in the various emotional activations. RESULTS The stimuli produced a general activation in visual pathways that included the primary and secondary visual cortex, involving regions associated with object and spatial recognition. In addition, the specific emotions produced different regional limbic activations, which suggests that different pathways may be used for different types of emotional stimuli. CONCLUSIONS Emotional activation in normal elderly subjects was associated with increases in blood flow in limbic and paralimbic brain structures. Brain activation may be specific to the emotion being elicited but probably involves complex sensory, association, and memory circuitry. Further studies are needed to identify activations that are specific for emotion.