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L H Snyder

Publications and source records attributed to L H Snyder.

At least 19 recordsLinked to original sources

Nonspatial saccade-specific activation in area LIP of monkey parietal cortex.

We present evidence that neurons in the lateral intraparietal area (LIP) of monkey posterior parietal cortex (PPC) are activated by the instruction to make an eye movement, even in the complete absence of a spatial target. This study employed a visually guided motor task that dissociated the type of movement to make (saccade or reach) from the location where the movement was to be made. Using this task, animals were instructed to prepare a specific type of movement prior to knowing the spatial location of the movement target. We found that 25% of the LIP neurons recorded in two animals were activated significantly more by the instruction to prepare a saccade than by the instruction to prepare a reach. This finding indicates that LIP is involved in more than merely spatial attention and provides further evidence for nonspatial effector-specific signal processing in the dorsal stream.

Animals↗

Intention-related activity in the posterior parietal cortex: a review.

Over the last few years it is becoming increasingly apparent that an important role of the posterior parietal cortex is to process sensory information for the purpose of planning actions. We review studies showing that a large component of neural activity in area LIP is related to planning saccades and activity in a nearby parietal reach region (PRR) to reaches. This intention related activity dominates the delay period in delayed movement tasks, and also comprises a substantial component of the transient response. These findings, along with additional anatomical and physiological evidence, lends support to the idea that different cortical areas within the PPC represent plans for different actions. We also found strong modulation of activity when movement plans were changed without changes in the locus of attention. This result suggests that PPC, which has been postulated to play a role in shifting attention, may also play a role in changing movement intentions. Sensory related activity was also present in these tasks and may be related to the stimulus or to attention. These experiments show that there are intention and sensory related activities in the PPC consistent with its proposed role in sensory-motor transformations. These studies also show that care must be taken to measure intention-related signals and not assume that all task dependent modulation in the PPC reflects attention.

Movement↗

Coordinate transformations for eye and arm movements in the brain.

Recent work on the coding of spatial information in the brain has significantly advanced our knowledge of sensory to motor transformations on several fronts. The encoding of information referenced to the retina (eye-centered) but modulated by eye position, called a gain field representation, has proved to be very common throughout parietal and occipital cortex. The use of an eye-centered representation as a working memory of spatial location is problematic if the eyes move during the memory period. Details regarding the manner in which the brain solves this problem are beginning to emerge. Finally, the discovery of eye-centered representations of ongoing or intended arm movements has changed the way we think about the order of operations in the sensory to motor coordinate transformation.

Animals↗

Computational approaches to sensorimotor transformations.

Behaviors such as sensing an object and then moving your eyes or your hand toward it require that sensory information be used to help generate a motor command, a process known as a sensorimotor transformation. Here we review models of sensorimotor transformations that use a flexible intermediate representation that relies on basis functions. The use of basis functions as an intermediate is borrowed from the theory of nonlinear function approximation. We show that this approach provides a unifying insight into the neural basis of three crucial aspects of sensorimotor transformations, namely, computation, learning and short-term memory. This mathematical formalism is consistent with the responses of cortical neurons and provides a fresh perspective on the issue of frames of reference in spatial representations.

Animals↗

Saccade-related activity in the parietal reach region.

In previous experiments, we showed that cells in the parietal reach region (PRR) in monkey posterior parietal cortex code intended reaching movements in an eye-centered frame of reference. These cells are more active when an arm compared with an eye movement is being planned. Despite this clear preference for arm movements, we now report that PRR neurons also fire around the time of a saccade. Of 206 cells tested, 29% had perisaccadic activity in a delayed-saccade task. Two findings indicate that saccade-related activity does not reflect saccade planning or execution. First, activity is often peri- or postsaccadic but seldom presaccadic. Second, cells with saccade-related activity were no more likely to show strong saccadic delay period activity than cells without saccade-related activity. These findings indicate that PRR cells do not take part in saccade planning. Instead, the saccade-related activity in PRR may reflect cross-coupling between reach and saccade pathways that may be used to facilitate eye-hand coordination. Alternatively, saccade-related activity may reflect eye position information that could be used to maintain an eye-centered representation of intended reach targets across eye movements.

Animals↗

Reach plans in eye-centered coordinates.

The neural events associated with visually guided reaching begin with an image on the retina and end with impulses to the muscles. In between, a reaching plan is formed. This plan could be in the coordinates of the arm, specifying the direction and amplitude of the movement, or it could be in the coordinates of the eye because visual information is initially gathered in this reference frame. In a reach-planning area of the posterior parietal cortex, neural activity was found to be more consistent with an eye-centered than an arm-centered coding of reach targets. Coding of arm movements in an eye-centered reference frame is advantageous because obstacles that affect planning as well as errors in reaching are registered in this reference frame. Also, eye movements are planned in eye coordinates, and the use of similar coordinates for reaching may facilitate hand-eye coordination.

Animals↗

The contributions of vestibular signals to the representations of space in the posterior parietal cortex.

Vestibular signals play an important role in spatial orientation, perception of object location, and control of self-motion. Prior physiological research on vestibular information processing has focused on brainstem mechanisms; relatively little is known about the processing of vestibular information at the level of the cerebral cortex. Recent electrophysiological experiments examining the use of vestibular canal signals in two different perceptual tasks are described: computation of self motion and localization of visual stimuli in a world-centered reference frame. These two perceptual functions are mediated by different parts of the posterior parietal cortex, the former in the dorsal aspect of the medial superior temporal area (MSTd) and the latter in area 7a.

Animals↗

Separate body- and world-referenced representations of visual space in parietal cortex.

In order to direct a movement towards a visual stimulus, visual spatial information must be combined with postural information. For example, directing gaze (eye plus head) towards a visible target requires the combination of retinal image location with eye and head position to determine the location of the target relative to the body. Similarly, world-referenced postural information is required to determine where something lies in the world. Posterior parietal neurons recorded in monkeys combine visual information with eye and head position. A population of such cells could make up a distributed representation of target location in an extraretinal frame of reference. However, previous studies have not distinguished between world-referenced and body-referenced signals. Here we report that modulations of visual signals (gain fields) in two adjacent cortical fields, LIP and 7a, are referenced to the body and to the world, respectively. This segregation of spatial information is consistent with a streaming of information, with one path carrying body-referenced information for the control of gaze, and the other carrying world-referenced information for navigation and other tasks that require an absolute frame of reference.

Animals↗

Posterior parietal areas specialized for eye movements (LIP) and reach (PRR) using a common coordinate frame.

The posterior parietal cortex (PPC) has long been considered a sensory area specialized for spatial awareness and the directing of attention. However, a new, far reaching concept is now emerging that this area is involved in integrating sensory information for the purpose of planning action. Moreover, experiments by our group and others over the last two decades indicate that PPC is in fact anatomically organized with respect to action. PPC also is an 'association' cortex which must combine different sensory modalities which are coded in different coordinate frames. We have found, at least for two different cortical areas within PPC, that different sensory signals are brought into a common coordinate frame. This coordinate frame codes locations with respect to the eye, but also gain modulates the activity by eye and body position signals. An interesting feature of this coordinate representation at the population level is that it codes concurrently target locations in multiple coordinate frames (eye, head, body and world). Depending on how this population of neurons is sampled, different coordinate transformations can be accomplished by the same population of neurons.

Animals↗

Change in motor plan, without a change in the spatial locus of attention, modulates activity in posterior parietal cortex.

The lateral intraparietal area (LIP) of macaque monkey, and a parietal reach region (PRR) medial and posterior to LIP, code the intention to make visually guided eye and arm movements, respectively. We studied the effect of changing the motor plan, without changing the locus of attention, on single neurons in these two areas. A central target was fixated while one or two sequential flashes occurred in the periphery. The first appeared either within the response field of the neuron being recorded or else on the opposite side of the fixation point. Animals planned a saccade (red flash) or reach (green flash) to the flash location. In some trials, a second flash 750 ms later could change the motor plan but never shifted attention: second flashes always occurred at the same location as the preceding first flash. Responses in LIP were larger when a saccade was instructed (n = 20 cells), whereas responses in PRR were larger when a reach was instructed (n = 17). This motor preference was observed for both first flashes and second flashes. In addition, the response to a second flash depended on whether it affirmed or countermanded the first flash; second flash responses were diminished only in the former case. Control experiments indicated that this differential effect was not due to stimulus novelty. These findings support a role for posterior parietal cortex in coding specific motor intention and are consistent with a possible role in the nonspatial shifting of motor intention.

Animals↗

Coding of intention in the posterior parietal cortex.

To look at or reach for what we see, spatial information from the visual system must be transformed into a motor plan. The posterior parietal cortex (PPC) is well placed to perform this function, because it lies between visual areas, which encode spatial information, and motor cortical areas. The PPC contains several subdivisions, which are generally conceived as high-order sensory areas. Neurons in area 7a and the lateral intraparietal area fire before and during visually guided saccades. Other neurons in areas 7a and 5 are active before and during visually guided arm movements. These areas are also active during memory tasks in which the animal remembers the location of a target for hundreds of milliseconds before making an eye or arm movement. Such activity could reflect either visual attention or the intention to make movements. This question is difficult to resolve, because even if the animal maintains fixation while directing attention to a peripheral location, the observed neuronal activity could reflect movements that are planned but not executed. To address this, we recorded from the PPC while monkeys planned either reaches or saccades to a single remembered location. We now report that, for most neurons, activity before the movement depended on the type of movement being planned. We conclude that PPC contains signals related to what the animal intends to do.

Animals↗

Multimodal representation of space in the posterior parietal cortex and its use in planning movements.

Recent experiments are reviewed that indicate that sensory signals from many modalities, as well as efference copy signals from motor structures, converge in the posterior parietal cortex in order to code the spatial locations of goals for movement. These signals are combined using a specific gain mechanism that enables the different coordinate frames of the various input signals to be combined into common, distributed spatial representations. These distributed representations can be used to convert the sensory locations of stimuli into the appropriate motor coordinates required for making directed movements. Within these spatial representations of the posterior parietal cortex are neural activities related to higher cognitive functions, including attention. We review recent studies showing that the encoding of intentions to make movements is also among the cognitive functions of this area.

Animals↗

Behavior and physiology of the macaque vestibulo-ocular reflex response to sudden off-axis rotation: computing eye translation.

The vestibulo-ocular reflex (VOR) has historically been considered a computationally simple reflex: to stabilize images on the retina against imposed head rotation, the eyes must be counterrotated by an equal amount in the opposite direction. During almost any head rotation, however, the eyes are also translated. We show that the VOR compensates for 90% of this translation, and suggest a computational scheme by which this is done, based on a temporal dissection of the VOR response to sudden head rotation. An initial response that corrects only for imposed rotation is refined by a series of three temporally delayed corrections of increasing complexity. The first correction takes only head rotation and viewing distance into account; the second, head rotation, viewing distance, and otolith translation; and the third, head rotation, viewing distance, otolith translation, and translation of the eyes relative to the otoliths. Responses of type I gaze velocity Purkinje (GVP) cells in the cerebellar flocculus and ventral paraflocculus of rhesus monkeys were recorded during sudden head rotation. We show that cell discharge was modulated both by axis location and by viewing distance, suggesting that GVP cells play a role in the VOR response to rotation-induced eye translation.

Animals↗

Head position signals used by parietal neurons to encode locations of visual stimuli.

The mechanism for object location in the environment, and the perception of the external world as stable when eyes, head and body are moved, have long been thought to be centred on the posterior parietal cortex. However, head position signals, and their integration with visual and eye position signals to form a representation of space referenced to the body, have never been examined in any area of the cortex. Here we show that the visual and saccadic activities of parietal neurons are strongly affected by head position. The eye and head position effects are equivalent for individual neurons, indicating that the modulation is a function of gaze direction, regardless of whether the eyes or head are used to direct gaze. These data are consistent with the idea that the posterior parietal cortex contains a distributed representation of space in body-centred coordinates.

Animals↗

Coordinate transformations in the representation of spatial information.

Coordinate transformations are an essential aspect of behavior. They are required because sensory information is coded in the coordinates of the sensory epithelia (e.g. retina, skin) and must be transformed to the coordinates of muscles for movement. In this review we will concentrate on recent studies of visual-motor transformations. The studies show that representations of space are distributed, being specified in the activity of many cells rather than in the activity of individual cells. Furthermore, these distributed representations appear to be derived by a specific operation, which systematically combines visual signals with eye and head position signals.

Animals↗

Changes in vestibulo-ocular reflex (VOR) anticipate changes in vergence angle in monkey.

The axis of head rotation is usually different from the axis of eye rotation. Geometrical considerations show that the eyes translate with respect to visual targets located near the head. In order to accurately stabilize retinal images against this translation, the vestibulo-ocular reflex (VOR) must be modulated inversely with the target's distance. The neural correlate of viewing distance used to modulate the VOR is not known. Since vergence angle is related inversely to viewing distance, an efference copy of instantaneous vergence angle or proprioceptive afferent information from extraocular muscles could be used to adjust the VOR. To examine this hypothesis, we compared the time-course of changes in the VOR with the time-course of changes in vergence angle. The VOR was induced by briefly rotating monkeys about a vertical axis at different times during the execution of vergence eye movements. We found that the amplitude of the VOR changed systematically during the course of a vergence eye movement. On average, the changes in the amplitude of vestibular induced eye movements anticipated changes in vergence angle by 50 msec, but in some instances, up to 200 msec of anticipation was observed. These data suggest that a central command signal rather than an afferent or efferent copy of vergence eye position was used to modulate the VOR.

Animals↗

Effect of viewing distance and location of the axis of head rotation on the monkey's vestibuloocular reflex. I. Eye movement responses.

1. The vestibuloocular reflex (VOR) stabilizes images on the retina against movements of the head in space. Viewing distance, target eccentricity, and location of the axis of rotation may influence VOR responses because rotation of the head about most axes in space rotates and translates the eyes relative to visual targets. To study the VOR response to combined rotation and translation, monkeys were placed on a rate table and rotated briefly in the dark about a vertical axis that was located in front of or behind the eyes. The monkeys fixated a near or far visual target that was extinguished before the rotation. Eye movements were recorded from both eyes by the use of the search coil technique. 2. Peak eye velocity evoked by the VOR was linearly related to vergence angle for any axis of rotation. The percent change in the VOR with near target viewing relative to far target viewing at a vergence angle of 20 degrees was linearly related to the location of the axis of rotation. Axes located behind the eyes produced positive changes in VOR amplitude, and axes located in front of the eyes produced negative changes in VOR amplitude. An axis of rotation located in the coronal plane containing the centers of rotation of the eyes produced no modification of VOR amplitude. For any axis, the VOR compensated for approximately 90% of the translation of the eye relative to near targets. 3. The initial VOR response was not correct in magnitude but was refined by a series of three temporally delayed corrections of increasing complexity. The earliest VOR-evoked eye movement (10-20 ms after rotation onset) was independent of viewing distance and rotational axis location. In the next 100 ms, eye speed appeared to be sequentially modified three times: within 20 ms by viewing distance; within 30 ms by otolith translation; and within 100 ms by eye translation relative to the visual target. 4. These data suggest a formal model of the VOR consisting of four channels. Channel 1 conveys an unmodified head rotation signal with a pure delay of 10 ms. Channel 2 conveys an angular head velocity signal, modified by viewing distance with a pure delay of 20 ms, but invariant with respect to the location of the axis of rotation. Channel 3 conveys a linear head velocity signal, dependent on the location of the axis of rotation, that is modified by viewing distance with a pure delay of 30 ms.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Vertical vestibuloocular reflex in cat: asymmetry and adaptation.

1. We studied eye velocity during the first 2 s of the vertical vestibuloocular reflex (VOR) elicited from cats placed on their sides (90 degrees roll position) and rotated about an earth vertical axis. Vestibular stimuli were presented in the dark and consisted of brief trapezoidal velocity profiles. Eye movements were recorded with a magnetic search coil, and eye velocity was analyzed with high temporal resolution. 2. The first 2 s of upward or downward eye velocity after the onset of head rotation was characterized and compared. Adaptive changes in VOR gain (eye/head velocity) were then induced, and upward and downward eye velocity responses were again compared. 3. The early time course of the vertical VOR was complex. After a latency of approximately 15 ms, eye velocity increased rapidly until it was equal in magnitude and opposite in direction to head velocity. The peak eye velocity decayed within less than 1 s to a plateau of slow-phase eye velocity (SPEV) equal to approximately -0.6 times the head velocity. Peak upward and downward eye velocity was symmetric. The transition from peak to plateau was more rapid for the downward VOR (slow phases downward) than for the upward VOR (slow phases upward). The plateau attained by upward SPEV was approximately 15% higher than the plateau attained by downward SPEV. 4. VOR gain adaptation was symmetric. The percentage change in adapted upward eye velocity equalled the percentage change in adapted downward eye velocity. Both peak and plateau SPEV adapted, but peak eye velocity adapted less than plateau eye velocity. VOR latency was unchanged by adaptation. 5. The trajectory of the VOR response to steps of head velocity could be divided into an invariant and a variant interval. The invariant interval consisted of the initial approximately 15 ms of the eye movement. Neither direction of head movement (upward vs. downward) nor adaptation of the VOR gain effected the eye movement trajectory during the invariant interval. The variant interval began approximately 30 ms after the onset of head movement and approximately 15 ms after the onset of eye movement. In unadapted animals, downward eye speed exceeded upward eye speed during the variant interval. In adapted animals, eye speed during the variant interval, but not during the invariant interval, diverged from eye speed in the unadapted state. We suggest that the initial invariant interval (approximately 15 ms) of the eye movement response trajectory may represent the direct response of the classically described three-neuron arc.(ABSTRACT TRUNCATED AT 400 WORDS)

Adaptation, Physiological↗