Implanting electrical devices in the nervous system to treat neural diseases is becoming very common. The success of these brain-machine interfaces depends on the electrodes that come into contact with the neural tissue. Here we show that conventional tungsten and stainless steel wire electrodes can be coated with carbon nanotubes using electrochemical techniques under ambient conditions. The carbon nanotube coating enhanced both recording and electrical stimulation of neurons in culture, rats and monkeys by decreasing the electrode impedance and increasing charge transfer. Carbon nanotube-coated electrodes are expected to improve current electrophysiological techniques and to facilitate the development of long-lasting brain-machine interface devices.
To find a target object in a crowded scene, a face in a crowd for example, the visual system might turn the neural representation of each object on and off in a serial fashion, testing each representation against a template of the target item. Alternatively, it might allow the processing of all objects in parallel but bias activity in favor of those neurons that represent critical features of the target, until the target emerges from the background. To test these possibilities, we recorded neurons in area V4 of monkeys freely scanning a complex array to find a target defined by color, shape, or both. Throughout the period of searching, neurons gave enhanced responses and synchronized their activity in the gamma range whenever a preferred stimulus in their receptive field matched a feature of the target, as predicted by parallel models. Neurons also gave enhanced responses to candidate targets that were selected for saccades, or foveation, reflecting a serial component of visual search. Thus, serial and parallel mechanisms of response enhancement and neural synchrony work together to identify objects in a scene. To find a target object in a crowded scene, a face in a crowd for example, the visual system might turn the neural representation of each object on and off in a serial fashion, testing each representation against a template of the target item. Alternatively, it might allow the processing of all objects in parallel but bias activity in favor of those neurons that represent critical features of the target, until the target emerges from the background. To test these possibilities, we recorded neurons in area V4 of monkeys freely scanning a complex array to find a target defined by color, shape, or both. Throughout the period of searching, neurons gave enhanced responses and synchronized their activity in the gamma range whenever a preferred stimulus in their receptive field matched a feature of the target, as predicted by parallel models. Neurons also gave enhanced responses to candidate targets that were selected for saccades, or foveation, reflecting a serial component of visual search. Thus, serial and parallel mechanisms of response enhancement and neural synchrony work together to identify objects in a scene.
We review two studies aimed at understanding the role of prefrontal cortex (PFC) in the control of attention. The first study examined which attentional functions are critically dependent on PFC by removing PFC unilaterally and transecting the forebrain commissures in two macaques. The monkeys fixated a central cue and discriminated the orientation of a colored target grating presented among colored distracter gratings in either the hemifield affected by the PFC lesion or the normal control hemifield. When the cue was held constant for many trials, task performance in the affected hemifield was nearly normal. However, performance was severely impaired when the cue was switched frequently across trials. The monkeys were unimpaired in a pop-out task with changing targets that did not require top-down attentional control. Thus, the PFC lesion resulted in selective impairment in the monkeys' ability to switch top-down control. In the second study, we used fMRI to investigate the neural correlates of top-down control in humans performing tasks identical to those used in the monkey experiments. Several fronto-parietal and posterior visual areas showed enhanced activation when attention was switched, which was greater on color cueing (top-down) trials relative to popout trials. Taken together, our findings indicate that both frontal and parietal cortices are involved in generating top-down control signals for attentive switching, which may then be fed back to visual processing areas. The PFC in particular plays a critical role in the ability to switch attentional control on the basis of changing task demands.
Brightness changes can be induced in a static gray field by modulating the luminance of surrounding areas. We used this induction phenomenon to investigate the neural representation of perceived brightness. Extracellular recordings were made in striate cortex, the lateral geniculate nucleus (LGN), and the optic tract of anesthetized cats using stimuli that produced brightness induction. While a cell's receptive field (RF) was covered by uniform gray illumination, the luminance of rectangular flanking regions was modulated sinusoidally in time, inducing brightness changes in the RF. We looked for a correspondence between the modulation of a cell's response and stimulus conditions that did or did not produce perceptual changes in brightness. We found that the responses of retinal ganglion cell axons in the optic tract were never correlated with brightness. On the other hand, many neurons in striate cortex and a small fraction in the LGN responded in a phase-locked manner at the temporal frequency of the flank modulation, even though the flanks were 3-7 degrees beyond the edges of the RF. Only in striate cortex were cells found that had responses correlated with brightness under all stimulus conditions. These findings suggest that brightness information is explicitly represented in the responses of neurons in striate cortex as part of a neural representation of object surfaces.
Although neurons in primary visual cortex are sensitive to the spatial distribution and intensity of light, their responses have not been thought to correlate with the perception of brightness. Indeed, primary visual cortex is often described as an initial processing stage that sends information to higher cortical areas where perception of brightness, color, and form occurs. However, a significant percentage of neurons in primary visual cortex were shown to respond in a manner correlated with perceived brightness, rather than responding strictly to the light level in the receptive fields of the cells. This finding suggests that even at the first stage of visual cortical processing, spatial integration of information yields perceptual qualities that are only indirectly related to the pattern of illumination of the retina.
It is widely held that the frontal eye field (FEF) in prefrontal cortex (PFC) modulates processing in visual cortex with attention, although the evidence for a necessary role is equivocal. To help identify critical sources of attentional feedback to area V4, we surgically removed the entire lateral PFC, including the FEF, in one hemisphere and transected the corpus callosum and anterior commisure in two macaques. This deprived V4 of PFC input in one hemisphere while keeping the other hemisphere intact. In the absence of PFC, attentional effects on neuronal responses and synchrony in V4 were significantly reduced and the remaining effects of attention were delayed in time indicating a critical role of PFC. Conversely, distracters captured attention and influenced V4 responses. However, because the effects of attention in V4 were not eliminated by PFC lesions, other sources of top-down attentional control signals to visual cortex must exist outside of PFC.
Brain imaging, electrical stimulation, and neurophysiological studies have all implicated the prefrontal cortex (PFC) in the top-down control of attention. Specifically, feedback from PFC has been proposed to bias activity in visual cortex in favor of attended stimuli over irrelevant distracters. To identify which attentional functions are critically dependent on PFC, we removed PFC unilaterally in combination with transection of the corpus callosum and anterior commissure in two macaques. In such a preparation, the ipsilesional hemisphere is deprived of top-down feedback from PFC to visual cortex, and the contralesional hemisphere can serve as an intact normal control. Monkeys were trained to fixate a central cue and discriminate the orientation of a colored target grating presented among colored distracter gratings in either the hemifield affected by the PFC lesion or the normal control hemifield. Locations of the targets and distracters were varied, and the color of the central cue specified the color of the target on each trial. The behavioral response was a bar release, and thus attentional impairments could be distinguished from impaired oculomotor control. When the cue was held constant for many trials, task performance in the affected hemifield was nearly normal. However, the monkeys were severely impaired when the cue was switched frequently across trials. The monkeys were unimpaired in a pop-out task with changing targets that did not require top-down attentional control. The PFC thus appears to play a critical role in the ability to flexibly reallocate attention on the basis of changing task demands.
Recent studies have suggested that V1 neurons extract figures from their backgrounds, in that they respond better to interior features of figures than to equivalent features of background stimuli. This is reportedly true even when the figure boundaries are distant from the borders of the classical receptive field (RF). To test the role of V1 neurons in figure-ground segregation, we recorded their responses to texture figures on texture backgrounds, centered on the RF. The texture elements of the figures remained identical across trials, and figure boundaries were defined by orientation differences between the elements in the background texture relative to elements in the figure. For nearly all neurons (98/102), responses to a large texture figure did not differ from the responses to a uniform-texture background. Although many neurons gave enhanced responses to texture boundaries, this occurred only when the boundaries were within or close to the RF borders. Similar effects were found in V2. For neurons in V1, the limited spatial extent of the contextual modulation was not increased either at low stimulus contrast or when the animal was rewarded for detecting an orientation-defined figure. Thus, V1 neurons appear to signal texture boundaries rather than figures per se. Unexpectedly, many V1 neurons gave significant long-latency responses to texture stimuli located entirely outside the classical RF, up to 5°f rom the RF border in some cases. However, these responses did not depend on the stimulus forming a figure that contained the RF. Although V1 neurons are influenced by stimuli outside the classical RF, they do not appear to segregate figures from ground.
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