To reduce the information gap between human neuroimaging and macaque physiology and anatomy, we mapped fMRI signals produced by moving and stationary stimuli (random dots or lines) in fixating monkeys. Functional sensitivity was increased by a factor of approximately 5 relative to the BOLD technique by injecting a contrast agent (monocrystalline iron oxide nanoparticle [MION]). Areas identified as motion sensitive included V2, V3, MT/V5, vMST, FST, VIP, and FEF (with moving dots), as well as V4, TE, LIP, and PIP (with random lines). These regions sensitive for moving dots are largely in agreement with monkey single unit data and (except for V3A) with human fMRI results. Moving lines activate some regions that have not been previously implicated in motion processing. Overall, the results clarify the relationship between the motion pathway and the dorsal stream in primates.

We used high-field (3T) functional magnetic resonance imaging (fMRI) to label cortical activity due to visual spatial attention, relative to flattened cortical maps of the retinotopy and visual areas from the same human subjects. In the main task, the visual stimulus remained constant, but covert visual spatial attention was varied in both location and load. In each of the extrastriate retinotopic areas, we found MR increases at the representations of the attended target. Similar but smaller increases were found in V1. Decreased MR levels were found in the same cortical locations when attention was directed at retinotopically different locations. In and surrounding area MT+, MR increases were lateralized but not otherwise retinotopic. At the representation of eccentricities central to that of the attended targets, prominent MR decreases occurred during spatial attention.

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Stereopsis, the perception of depth from small differences between the images in the two eyes, provides a rich model for investigating the cortical construction of surfaces and space. Although disparity-tuned cells have been found in a large number of areas in macaque visual cortex, stereoscopic processing in these areas has never been systematically compared using the same stimuli and analysis methods. In order to examine the global architecture of stereoscopic processing in primate visual cortex, we studied fMRI activity in alert, fixating human and macaque subjects. In macaques, we found strongest activation to near/far compared to zero disparity in areas V3, V3A, and CIPS. In humans, we found strongest activation to the same stimuli in areas V3A, V7, the V4d topolog (V4d-topo), and a caudal parietal disparity region (CPDR). Thus, in both primate species a small cluster of areas at the parieto-occipital junction appears to be specialized for stereopsis.

The frontal eye field (FEF) is one of several cortical regions thought to modulate sensory inputs. Moreover, several hypotheses suggest that the FEF can only modulate early visual areas in the presence of a visual stimulus. To test for bottom-up gating of frontal signals, we microstimulated subregions in the FEF of two monkeys and measured the effects throughout the brain with functional magnetic resonance imaging. The activity of higher-order visual areas was strongly modulated by FEF stimulation, independent of visual stimulation. In contrast, FEF stimulation induced a topographically specific pattern of enhancement and suppression in early visual areas, but only in the presence of a visual stimulus. Modulation strength depended on stimulus contrast and on the presence of distractors. We conclude that bottom-up activation is needed to enable topdown modulation of early visual cortex and that stimulus saliency determines the strength of this modulation.Contemporary hypotheses propose that feedback signals from areas in frontal and parietal cortex exert control over the processing of incoming visual information (1-5). Several models suggest that these signals are gated by bottom-up stimulation (6-9). In these models, feedback signals only influence neurons activated by visual input, just as has been observed for attentional effects, which are known to be strongest for neurons well driven by a visual stimulus (10)(11)(12). No causal evidence exists, however, to support these hypotheses, with the * To whom correspondence should be addressed. wim@nmr.mgh.harvard.edu. In our first experiment, the goal was to detect the functional consequences of EM of the FEF in the absence of a visual stimulus, using stimulation levels below those needed to evoke saccades. We first obtained anatomical ( fig. S1B) and behavioral evidence (Fig. 1A and fig. S2) in two monkeys that several chronically implanted microelectrodes were positioned in the FEF. Before each fMRI experiment, we stimulated these electrodes inside the MR scanner to determine the threshold needed to evoke saccades and to identify the saccade end point, or movement field (MF), of each FEF stimulation site (Fig. 1A and fig. S2). During the actual fMRI experiment, the monkeys carried out a passive fixation task while we alternated between epochs of no-EM and epochs of EM, at a stimulation level of 50% of the saccade-inducing amplitude. The use of this method in awake animals allowed us to titrate the stimulation to functionally relevant levels (14), an advantage compared to a previous study in anesthetized animals (15).The left column of Fig Our first experiment indicated that FEF-EM increased fMRI activity in higher-order visual areas known to be directly connected to the FEF. If feedback effects are gated by visual stimulation, however, one also predicts FEF-EM effects in visual areas separated from the FEF by multiple synapses, in the presence of a visual stimulus. In a second experiment, we therefore placed high-contrast, colored, moving gratings in the ...

We compared three-dimensional structure-from-motion (3D-SFM) processing in awake monkeys and humans using functional magnetic resonance imaging. Occipital and midlevel extrastriate visual areas showed similar activation by 3D-SFM stimuli in both species. In contrast, intraparietal areas showed significant 3D-SFM activation in humans but not in monkeys. This suggests that human intraparietal cortex contains visuospatial processing areas that are not present in monkeys.To reconstruct the third dimension from a two-dimensional (2D) retinal image, our brain uses binocular as well as monocular cues such as shading, texture, and occlusion. Both humans and monkeys are also able to extract the 3D structure of an object from motion parallax cues that activate occipitoparietal areas in both species (1-3). Because neurons in the middle temporal area (MT/V5) are sensitive for speed gradients that reflect planes tilted in depth (4, see also 5), this area might play a crucial role in the extraction of depth from motion. Supporting evidence has been gleaned from a functional magnetic resonance imaging (fMRI) study showing 3D-SFM sensitivity in the human MT/V5 complex (hMT/V5ϩ) (6 ). These human fMRI results raise a first question: To what extent can they be generalized to the primate visual system? Furthermore, anatomical evidence suggests that there might be functional differences between human and monkey intraparietal cortex: The intraparietal sulcus separates area 5 from area 7 in the monkey, whereas in humans these two areas belong to the superior parietal lobe. In addition, in humans, the intraparietal lobe separates area 7 from areas 39 and 40, which have no clear counterpart in monkeys (7 ). Therefore, our second goal was to determine whether monkey intraparietal cortex is as important for motion-dependent depth processing as implied by human imaging (6 ).To address these issues, we turned to recently developed fMRI techniques (8) in awake (9-12) rather than anesthetized (13-14 ) monkeys. By performing human fMRI with virtually identical experimental procedures as in the awake monkey fMRI experiments, reliable interspecies comparisons could be made.The stimuli were displays of nine randomly connected lines, rotating in depth, that created a clear 3D percept (movie S1). Control stimuli consisted of the same displays that were either static or moving in one plane. We controlled for potential attentional differences between the 3D and 2D conditions by using a 1-back task in humans, as well as a demanding high-acuity fixation task (8, 9) in both species.In line with earlier reports (4-6, 13), area MT/V5 was activated more by 3D than by 2D moving random-line displays. In addition, the area in the fundus of the superior temporal sulcus (FST) also exhibited significant 3D-SFM sensitivity (Fig. 1A). Figure 2A shows the (3D -2D) pattern of activation for a single human subject, and the average activation for a group of eight subjects is shown in Fig. 2B. These results are similar to those obtained in an earlier study in which so...

Human area V1 offers an excellent opportunity to study, using functional MRI, a range of properties in a specific cortical visual area, whose borders are defined objectively and convergently by retinotopic criteria. The retinotopy in V1 (also known as primary visual cortex, striate cortex, or Brodmann's area 17) was defined in each subject by using both stationary and phase-encoded polar coordinate stimuli. Data from V1 and neighboring retinotopic areas were displayed on f lattened cortical maps. In additional tests we revealed the paired cortical representations of the monocular ''blind spot.'' We also activated area V1 preferentially (relative to other extrastriate areas) by presenting radial gratings alternating between 6% and 100% contrast. Finally, we showed evidence for orientation selectivity in V1 by measuring transient functional MRI increases produced at the change in response to gratings of differing orientations. By systematically varying the orientations presented, we were able to measure the bandwidth of the orientation ''transients'' (45°).

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