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C L Colby

Publications and source records attributed to C L Colby.

At least 19 recordsLinked to original sources

Stimulus-response incompatibility activates cortex proximate to three eye fields.

We used functional magnetic resonance imaging (fMRI) to investigate cortical activation during the performance of three oculomotor tasks that impose increasing levels of cognitive demand. (1) In a visually guided saccade (VGS) task, subjects made saccades to flashed targets. (2) In a compatible task, subjects made leftward and rightward saccades in response to foveal presentation of the uppercase words "LEFT" or "RIGHT." (3) In a mixed task, subjects made rightward saccades in response to the lowercase word "left" and leftward saccades in response to the lowercase word "right" on incompatible trials (60%). The remaining 40% of trials required compatible responses to uppercase words. The VGS and compatible tasks, when compared to fixation, activated the three cortical eye fields: the supplementary eye field (SEF), the frontal eye field (FEF), and the parietal eye field (PEF). The mixed task, when compared to the compatible task, activated three additional cortical regions proximate to the three eye fields: (1) rostral to the SEF in medial frontal cortex; (2) rostral to the FEF in dorsolateral prefrontal cortex (DLPFC); (3) rostral and lateral to the PEF in posterior parietal cortex. These areas may contribute to the suppression of prepotent responses and in holding novel visuomotor associations in working memory.

Adult↗

Visual, saccade-related, and cognitive activation of single neurons in monkey extrastriate area V3A.

Area V3A is an extrastriate visual area that provides a major input to parietal cortex. To identify the sensory, saccade-related, and cognitive signals carried by V3A neurons, we recorded from single units in alert monkeys during performance of fixation and memory guided saccade tasks. We found that visual responses to stationary stimuli in area V3A were affected by the behavioral relevance of the stimulus. The amplitude of the visual response differed between the memory-guided saccade task, in which the monkey had to use the information provided by the stimulus to guide its behavior, and the fixation task. For 18% (29/163) of V3A neurons, the response was significantly enhanced in the memory-guided saccade task as compared with that in the fixation task. For 8% (13/163) of V3A neurons, the amplitude of response in the memory-guided saccade task was significantly suppressed. We also observed task-related modulation of activity prior to stimulus onset. Among the V3A neurons (37/163) that showed significant differences between tasks in prestimulus activity, the majority (89%; 33/37) showed greater prestimulus activity in the memory-guided saccade task. Task-related increases in prestimulus activity in the memory-guided saccade task were not always matched by increases in the sensory response, indicating that visual responses and prestimulus activity can be modulated independently. Activity in the memory period was suppressed compared with prestimulus activity for 83% (49/59) of the V3A neurons that showed a significant difference in activity (59/197) between these two epochs. For some neurons, memory-period activity dropped even below the baseline level in the fixation task, indicating that there may be an active suppression mechanism. Many V3A neurons (75%, 148/197) also had activity in the saccade epoch. This activity was most prominent immediately after the saccade. Postsaccadic activity was observed even when testing was carried out in total darkness, indicating that this activity reflects, at least in part, extraretinal signals and is not simply a response to visual reafference. These results indicate that several kinds of signals are carried by single neurons in extrastriate area V3A. Moreover, activity in V3A is subject to modulation by extraretinal factors, including attention, anticipation, memory, and saccadic eye movements.

Animals↗

Cortical networks subserving pursuit and saccadic eye movements in humans: an FMRI study.

High-field (3 Tesla) functional magnetic resonance imaging (MRI) was used to investigate the cortical circuitry subserving pursuit tracking in humans and compare it to that for saccadic eye movements. Pursuit performance, relative to visual fixation, elicited activation in three areas known to contribute to eye movements in humans and in nonhuman primates: the frontal eye field, supplementary eye field, and intraparietal sulcus. It also activated three medial regions not previously identified in human neuroimaging studies of pursuit: the precuneus and the anterior and posterior cingulate cortices. All six areas were also activated during saccades. The spatial extent of activation was similar for saccades and pursuit in all but two regions: spatial extent was greater for saccades in the superior branch of the frontal eye field and greater for pursuit in posterior cingulate cortex. This set of activations for smooth pursuit parallels the network of oculomotor areas characterized in nonhuman primates and complements recent studies showing that common cortical networks subserve oculomotor functions and spatial attention in humans.

Adolescent↗

Space and attention in parietal cortex.

The space around us is represented not once but many times in parietal cortex. These multiple representations encode locations and objects of interest in several egocentric reference frames. Stimulus representations are transformed from the coordinates of receptor surfaces, such as the retina or the cochlea, into the coordinates of effectors, such as the eye, head, or hand. The transformation is accomplished by dynamic updating of spatial representations in conjunction with voluntary movements. This direct sensory-to-motor coordinate transformation obviates the need for a single representation of space in environmental coordinates. In addition to representing object locations in motoric coordinates, parietal neurons exhibit strong modulation by attention. Both top-down and bottom-up mechanisms of attention contribute to the enhancement of visual responses. The saliance of a stimulus is the primary factor in determining the neural response to it. Although parietal neurons represent objects in motor coordinates, visual responses are independent of the intention to perform specific motor acts.

Animals↗

Ventral intraparietal area of the macaque: congruent visual and somatic response properties.

In a previous report, we described the visual response properties in the ventral intraparietal area (area VIP) of the awake macaque. Here we describe the somatosensory response properties in area VIP and the patterns of correspondence between the responses of single neurons to independently administered tactile and visual stimulation. VIP neurons responded to visual stimulation only or to visual and tactile stimulation. Of 218 neurons tested, 153 (70%) were bimodal in the sense that they responded to stimuli that were independently applied in either sensory modality. Unimodal visual and bimodal neurons were intermingled within the recording area and could not be distinguished on the basis of their visual response properties alone. Most of the cells with a tactile receptive field (RF) responded well to light touch or air puffs. The distribution of RF locations principally emphasized the head (85%), with approximately equivalent representations of the upper and lower face areas. The tactile and visual RFs were aligned in a congruent manner, with the intersection of the visual vertical and horizontal meridian having its tactile counterpart in the nose/mouth area. Small foveal visual RFs were paired with small tactile RFs on the muzzle, and peripheral visual RFs were associated with tactile RFs on the side of the head or body. Most cells showed a strong sensitivity to moving stimuli, and the preferred directions of visual and tactile motion coincided in 85% of bimodal cells. In some cases, bimodal responses patterns were complementary: cells responding to motion in depth toward the monkey had responses, whereas cells responding to motion in depth away form the monkey had responses. Other forms of bimodal response congruence included orientation selectivity, and , , and / response types. The large proportion of bimodal tactile and visual neurons with congruent response properties in area VIP indicates that there are important functional differences between area VIP and other dorsal stream areas involved in the analysis of motion. We suggest that VIP is involved in the construction of a multisensory, head-centered representation of near extrapersonal space.

Animals↗

Spatial representations for action in parietal cortex.

Parietal cortex contains multiple representations of visual space. Single neurons in area LIP encode attended locations relative to the fovea, while some VIP neurons encode stimulus location relative to the head and some MIP neurons may encode location relative to the arm. These multiple representations are tailored to guide specific kinds of actions: eye movements, head movements and arm movements, respectively. The function of parietal cortex is to signal the location of attended objects relative to the observer. It does so in order to allow the organism to act on its environment. The many different kinds of actions that can be performed are likely to be supported by these very different kinds of spatial representations.

Animals↗

Visual, presaccadic, and cognitive activation of single neurons in monkey lateral intraparietal area.

1. Posterior parietal cortex contains neurons that are visually responsive and active in relation to saccadic eye movements. We recorded from single neurons in a subregion of parietal cortex, the lateral intraparietal area (LIP), in alert rhesus monkeys. To characterize more completely the circumstances under which LIP neurons are responsive, we used five tasks designed to test the impact of sensory, motor, and cognitive factors. We obtained quantitative data in multiple tasks in 91 neurons. We measured neural activity during central fixation and in relation to stimulus onset and saccade onset. 2. LIP neurons have visual responses to the onset of a stationary stimulus in the receptive field. These visual responses occurred both in tasks that require a subsequent eye movement toward the stimulus and in tasks in which eye movements are not permitted, indicating that this activity is sensory rather than presaccadic. 3. Visual responses were enhanced when the monkey had to use information provided by the stimulus to guide its behavior. The amplitude of the sensory response to a given stimulus was increased in a task in which the monkey would subsequently make a saccade to the location signaled by the stimulus, as compared with the amplitude of the visual response in a simple fixation task. 4. The visual response was also enhanced when the monkey attended to the stimulus without looking at it. This result shows that enhancement does not reflect saccade preparation because the response is enhanced even when the monkey is not permitted to make a saccade. Instead, enhancement reflects the allocation of attention to the spatial locus of the receptive field. 5. Many LIP neurons had saccade-related activity in addition to their visual responses. The visual response for most neurons was stronger than the saccade-related activation. 6. Saccade-related activity was independent of visual activity. Similar presaccadic activity was observed in trials that included a recent visual stimulus (memory-guided saccade task) and in trials with no visual stimulus (learned saccade task). 7. We observed increases in activity during fixation in tasks in which the monkey could anticipate the onset of a behaviorally significant stimulus. LIP neurons usually showed low levels of background firing in the fixation task during the period before stimulus onset. This background activity was increased in the peripheral attention and memory-guided saccade tasks during the period when the monkey was waiting for a behaviorally relevant stimulus to appear. 8. The results from these several tasks indicate that LIP neurons are activated in a variety of circumstances and are not involved exclusively in sensory processing or motor planning. The modulation of sensory responses by attention and anticipation suggests that cognitive factors play a major role in parietal function.

Animals↗

Ventral intraparietal area of the macaque: anatomic location and visual response properties.

1. The middle temporal area (MT) projects to the intraparietal sulcus in the macaque monkey. We describe here a discrete area in the depths of the intraparietal sulcus containing neurons with response properties similar to those reported for area MT. We call this area the physiologically defined ventral intraparietal area, or VIP. In the present study we recorded from single neurons in VIP of alert monkeys and studied their visual and oculomotor response properties. 2. Area VIP has a high degree of selectivity for the direction of a moving stimulus. In our sample 72/88 (80%) neurons responded at least twice as well to a stimulus moving in the preferred direction compared with a stimulus moving in the null direction. The average response to stimuli moving in the preferred direction was 9.5 times as strong as the response to stimuli moving in the opposite direction, as compared with 10.9 times as strong for neurons in area MT. 3. Many neurons were also selective for speed of stimulus motion. Quantitative data from 25 neurons indicated that the distribution of preferred speeds ranged from 10 to 320 degrees/s. The degree of speed tuning was on average twice as broad as that reported for area MT. 4. Some neurons (22/41) were selective for the distance at which a stimulus was presented, preferring a stimulus of equivalent visual angle and luminance presented near (within 20 cm) or very near (within 5 cm) the face. These neurons maintained their preference for near stimuli when tested monocularly, suggesting that visual cues other than disparity can support this response. These neurons typically could not be driven by small spots presented on the tangent screen (at 57 cm). 5. Some VIP neurons responded best to a stimulus moving toward the animal. The absolute direction of visual motion was not as important for these cells as the trajectory of the stimulus: the best stimulus was one moving toward a particular point on the face from any direction. 6. VIP neurons were not active in relation to saccadic eye movements. Some neurons (10/17) were active during smooth pursuit of a small target. 7. The predominance of direction and speed selectivity in area VIP suggests that it, like other visual areas in the dorsal stream, may be involved in the analysis of visual motion.

Animals↗

The updating of the representation of visual space in parietal cortex by intended eye movements.

Every eye movement produces a shift in the visual image on the retina. The receptive field, or retinal response area, of an individual visual neuron moves with the eyes so that after an eye movement it covers a new portion of visual space. For some parietal neurons, the location of the receptive field is shown to shift transiently before an eye movement. In addition, nearly all parietal neurons respond when an eye movement brings the site of a previously flashed stimulus into the receptive field. Parietal cortex both anticipates the retinal consequences of eye movements and updates the retinal coordinates of remembered stimuli to generate a continuously accurate representation of visual space.

Animals↗

Oculomotor control and spatial processing.

In the past year research in the oculomotor system has concentrated on some hitherto neglected areas, and also caused a re-evaluation of several long-standing concepts. Careful studies of the translational (otolith) vestibulo-ocular reflex and the torsional system have demonstrated their importance. A re-evaluation of the role of the superior colliculus in the generation of saccades has provided evidence for its participation in the feedback process. New studies of the interaction of eye movements and visual processing have shown that the brain can compensate for the visual effects of eye movements and maintain a retinotopic representation of visual space for the saccadic system.

Animals↗

Heterogeneity of extrastriate visual areas and multiple parietal areas in the macaque monkey.

The definition of visual areas remains a key problem in the effort to elucidate cortical functions. Visual areas vary along a number of dimensions and are increasingly difficult to define according to traditional criteria at higher levels of the hierarchy. Three recently discovered areas in monkey parietal association cortex illustrate a new approach to this problem. Their definition depends on assessment of neuronal response properties in the alert, behaving animal combined with precise reconstruction of recording sites. This approach permits recognition of functionally distinct areas in the absence of retinotopic maps.

Animals↗

The neuroanatomy and neurophysiology of attention.

Attention is a distributed process. The activity of neurons in many brain structures can be modulated by the attentional state of the animal. Attention directed toward a particular external stimulus is often reflected in an enhancement of the sensory response to that stimulus. Enhancement is spatially selective for neurons in many areas and explicitly eye-movement related in most. Attention directed toward the internal representation of a stimulus may be associated with a prolongation of neural activity. These modulations of neuronal responsiveness underscore the dynamic nature of neural processing. Competition between left- and right-brain structures in the control of attention is common. While attention is perceived as a unitary process, it is subserved by many brain structures. Given the wide distribution of attentional processes, it is not surprising that children diagnosed as having attentional deficits show considerable diversity in symptoms and etiology.

Animals↗

Topographical organization of cortical afferents to extrastriate visual area PO in the macaque: a dual tracer study.

We have examined the origin and topography of cortical projections to area PO, an extrastriate visual area located in the parieto-occipital sulcus of the macaque. Distinguishable retrograde fluorescent tracers were injected into area PO at separate retinotopic loci identified by single-neuron recording. The results indicate that area PO receives retinotopically organized inputs from visual areas V1, V2, V3, V4, and MT. In each of these areas the projection to PO arises from the representation of the periphery of the visual field. This finding is consistent with neurophysiological data indicating that the representation of the periphery is emphasized in PO. Additional projections arise from area MST, the frontal eye fields, and several divisions of parietal cortex, including four zones within the intraparietal sulcus and a region on the medial dorsal surface of the hemisphere (MDP). On the basis of the laminar distribution of labeled cells we conclude that area PO receives an ascending input from V1, V2, and V3 and receives descending or lateral inputs from all other areas. Thus, area PO is at approximately the same level in the hierarchy of visual areas as areas V4 and MT. Area PO is connected both directly and indirectly, via MT and MST, to parietal cortex. Within parietal cortex, area PO is linked to particular regions of the intraparietal sulcus including VIP and LIP and two newly recognized zones termed here MIP and PIP. The wealth of connections with parietal cortex suggests that area PO provides a relatively direct route over which information concerning the visual field periphery can be transmitted from striate and prestriate cortex to parietal cortex. In contrast, area PO has few links with areas projecting to inferior temporal cortex. The pattern of connections revealed in this study is consistent with the view that area PO is primarily involved in visuospatial functioning.

Animals↗

Corticotectal circuit in the cat: a functional analysis of the lateral geniculate nucleus layers of origin.

1. The dorsal lateral geniculate nucleus (LGN) of the cat is a major thalamic relay between the retina and several visual cortical areas. These cortical areas in turn project to the superior colliculus (SC). The aim of the present experiment was to determine which LGN layers provide a necessary input to the corticotectal circuit. 2. Individual layers of the LGN were reversibly inactivated by microinjection of cobalt chloride during recording of visual responses in the retinotopically corresponding part of the superior colliculus. 3. For cells driven through the contralateral eye, inactivation of layer A or the medial interlaminar nucleus (MIN) had little effect on visual responsiveness in the superior colliculus. In contrast, inactivation of layer C abolished visual responses at one-quarter of the SC recording sites, reduced responses at another quarter, and left half of the recording sites unaffected. 4. For cells driven through the ipsilateral eye, inactivation of layer C1 or the MIN had no effect. Inactivation of layer A1 uniformly reduced visual responses in the superior colliculus and usually abolished them entirely. 5. These results are compatible with previous work showing that cortical input to the SC originates from Y-cells. They indicate that two of the five Y-cell containing layers (A1 and C) provide major inputs to the corticotectal circuit. The results suggest that layer A1 is functionally allied to layer C as well as to layer A.

Animals↗

The responses of single cells in the lateral geniculate nucleus of the rhesus monkey to color and luminance contrast.

The responses of parvocellular and magnocellular cells in the monkey lateral geniculate nucleus (LGN) were assessed to red-green color substitution at various luminance ratios. Parvocellular LGN cells lacking well defined color selectivity could be rendered unresponsive to red-green light exchange at specific luminance ratios. By contrast, magnocellular LGN cells could not be silenced at any luminance ratio. The difference between these two types of cells is attributable to the greater sensitivity of magnocellular cells. The results suggest that magnocellular cells are well suited for the detection of any spatially localized change, be it the result of luminance or wavelength change.

Action Potentials↗