Monkey superior colliculus represents rapid eye movements in a two-dimensional motor map

1993 ◽  
Vol 69 (3) ◽  
pp. 965-979 ◽  
Author(s):  
K. Hepp ◽  
A. J. Van Opstal ◽  
D. Straumann ◽  
B. J. Hess ◽  
V. Henn

1. Although the eye has three rotational degrees of freedom, eye positions, during fixations, saccades, and smooth pursuit, with the head stationary and upright, are constrained to a plane by ListingR's law. We investigated whether Listing's law for rapid eye movements is implemented at the level of the deeper layers of the superior colliculus (SC). 2. In three alert rhesus monkeys we tested whether the saccadic motor map of the SC is two dimensional, representing oculocentric target vectors (the vector or V-model), or three dimensional, representing the coordinates of the rotation of the eye from initial to final position (the quaternion or Q-model). 3. Monkeys made spontaneous saccadic eye movements both in the light and in the dark. They were also rotated about various axes to evoke quick phases of vestibular nystagmus, which have three degrees of freedom. Eye positions were measured in three dimensions with the magnetic search coil technique. 4. While the monkey made spontaneous eye movements, we electrically stimulated the deeper layers of the SC and elicited saccades from a wide range of initial positions. According to the Q-model, the torsional component of eye position after stimulation should be uniquely related to saccade onset position. However, stimulation at 110 sites induced no eye torsion, in line with the prediction of the V-model. 5. Activity of saccade-related burst neurons in the deeper layers of the SC was analyzed during rapid eye movements in three dimensions. No systematic eye-position dependence of the movement fields, as predicted by the Q-model, could be detected for these cells. Instead, the data fitted closely the predictions made by the V-model. 6. In two monkeys, both SC were reversibly inactivated by symmetrical bilateral injections of muscimol. The frequency of spontaneous saccades in the light decreased dramatically. Although the remaining spontaneous saccades were slow, Listing's law was still obeyed, both during fixations and saccadic gaze shifts. In the dark, vestibularly elicited fast phases of nystagmus could still be generated in three dimensions. Although the fastest quick phases of horizontal and vertical nystagmus were slower by about a factor of 1.5, those of torsional quick phases were unaffected. 7. On the basis of the electrical stimulation data and the properties revealed by the movement field analysis, we conclude that the collicular motor map is two dimensional. The reversible inactivation results suggest that the SC is not the site where three-dimensional fast phases of vestibular nystagmus are generated.(ABSTRACT TRUNCATED AT 400 WORDS)

1994 ◽  
Vol 72 (3) ◽  
pp. 1171-1180 ◽  
Author(s):  
J. Hore ◽  
S. Watts ◽  
D. Tweed

1. Overarm throwing is a skilled multijoint movement with potentially many degrees of freedom. Considering only the arm > or = 7 degrees of freedom are involved (shoulder 3, elbow 2, wrist 2). For each arm segment 3 degrees of freedom are potentially required to specify its angular position (orientation) at any moment during a throw. Simplification of the control problem for the CNS would occur if there were constraints on these degrees of freedom. The objective was to determine whether such constraints exist at ball release when throwing at targets in different directions using only the arm. 2. The angular positions in three dimensions of the distal phalanx of the middle finger, the hand, the forearm, and the upper arm were simultaneously recorded with search coils as subjects sat with a fixed trunk and threw balls at nine targets in an approximate +/- 40 degree work space. Ball release was signaled by microswitches on the proximal and distal phalanges of the middle finger (proximal and distal triggers). 3. On throwing at any one target the hand at ball release adopted a similar orientation for each throw, i.e., for a particular vertical and horizontal angular position the hand adopted a similar torsional position. On throwing at targets throughout the work space, angular position (rotation) vectors describing hand positions in space at ball release were confined to a two-dimensional surface rather than a three-dimensional volume. This constraint in hand torsion occurred near and at ball release but not throughout the entire throw. It was not due to mechanical factors because such a surface was not obtained when subjects deliberately twisted their arms when throwing. Thus at ball release during a "natural" throw the hand was constrained to 2 of its possible 3 angular degrees of freedom. 4. The same constraint was also found for finger, forearm, and upper arm angular positions in space at ball release as determined at both the proximal and distal triggers. A consequence is that at ball release the entire arm was constrained to 2 of its possible 7 degrees of freedom. 5. The two-dimensional position vector surface for each arm segment was similar to that obtained when pointing with a straight arm at the same targets. In both cases they showed torsion and were twisted like the surface obtained by rotations around the horizontal and vertical axes of a Fick gimbal. However, in some subjects the throwing surfaces were tilted from the vertical.(ABSTRACT TRUNCATED AT 400 WORDS)


1998 ◽  
Vol 79 (4) ◽  
pp. 2082-2096 ◽  
Author(s):  
Hiroshi Aizawa ◽  
Robert H. Wurtz

Aizawa, Hiroshi and Robert H. Wurtz. Reversible inactivation of monkey superior colliculus. I. Curvature of saccadic trajectory. J. Neurophysiol. 79: 2082–2096, 1998. The neurons in the intermediate layers of the monkey superior colliculus (SC) that discharge before saccadic eye movements can be divided into at least two types, burst and buildup neurons, and the differences in their characteristics are compatible with different functional contributions of the two cell types. It has been suggested that a spread of activity across the population of the buildup neurons during saccade generation may contribute to the control of saccadic eye movements. The influence of any such spread should be on both the horizontal and vertical components of the saccade because the map of the movement fields on the SC is a two-dimensional one; it should affect the trajectory of saccade. The present experiments used muscimol injections to inactivate areas within the SC to determine the functional contribution of such a spread of activity on the trajectory of the saccades. The analysis concentrated on saccades made to areas of the visual field that should be affected primarily by alteration of buildup neuron activity. Muscimol injections produced saccades with altered trajectories; they became consistently curved after the injection, and successive saccades to the same targets had similar curvatures. The curved saccades showed changes in their direction and speed at the very beginning of the saccade, and for those saccades that reached the target, the direction of the saccade was altered near the end to compensate for the initially incorrect direction. Postinjection saccades had lower peak speeds, longer durations, and longer latencies for initiation. The changes in saccadic trajectories resulting from muscimol injections, along with the previous observations on changes in speed of saccades with such injections, indicate that the SC is involved in influencing the eye position during the saccade as well as at the end of the saccade. The changes in trajectory when injections were made more rostral in the SC than the most active burst neurons also are consistent with a contribution of the buildup neurons to the control of the eye trajectory. The results do not, however, support the hypothesis that the buildup neurons in the SC act as a spatial integrator.


2008 ◽  
Vol 62 (1) ◽  
Author(s):  
Peter C. Chu

The Navy’s mine impact burial prediction model creates a time history of a cylindrical or a noncylindrical mine as it falls through air, water, and sediment. The output of the model is the predicted mine trajectory in air and water columns, burial depth/orientation in sediment, as well as height, area, and volume protruding. Model inputs consist of parameters of environment, mine characteristics, and initial release. This paper reviews near three decades’ effort on model development from one to three dimensions: (1) one-dimensional models predict the vertical position of the mine’s center of mass (COM) with the assumption of constant falling angle, (2) two-dimensional models predict the COM position in the (x,z) plane and the rotation around the y-axis, and (3) three-dimensional models predict the COM position in the (x,y,z) space and the rotation around the x-, y-, and z-axes. These models are verified using the data collected from mine impact burial experiments. The one-dimensional model only solves one momentum equation (in the z-direction). It cannot predict the mine trajectory and burial depth well. The two-dimensional model restricts the mine motion in the (x,z) plane (which requires motionless for the environmental fluids) and uses incorrect drag coefficients and inaccurate sediment dynamics. The prediction errors are large in the mine trajectory and burial depth prediction (six to ten times larger than the observed depth in sand bottom of the Monterey Bay). The three-dimensional model predicts the trajectory and burial depth relatively well for cylindrical, near-cylindrical mines, and operational mines such as Manta and Rockan mines.


1999 ◽  
Vol 82 (5) ◽  
pp. 2612-2632 ◽  
Author(s):  
Pierre A. Sylvestre ◽  
Kathleen E. Cullen

The mechanics of the eyeball and its surrounding tissues, which together form the oculomotor plant, have been shown to be the same for smooth pursuit and saccadic eye movements. Hence it was postulated that similar signals would be carried by motoneurons during slow and rapid eye movements. In the present study, we directly addressed this proposal by determining which eye movement–based models best describe the discharge dynamics of primate abducens neurons during a variety of eye movement behaviors. We first characterized abducens neuron spike trains, as has been classically done, during fixation and sinusoidal smooth pursuit. We then systematically analyzed the discharge dynamics of abducens neurons during and following saccades, during step-ramp pursuit and during high velocity slow-phase vestibular nystagmus. We found that the commonly utilized first-order description of abducens neuron firing rates (FR = b + kE + rE˙, where FR is firing rate, E and E˙ are eye position and velocity, respectively, and b, k, and r are constants) provided an adequate model of neuronal activity during saccades, smooth pursuit, and slow phase vestibular nystagmus. However, the use of a second-order model, which included an exponentially decaying term or “slide” (FR = b + kE + rE˙ + uË − c[Formula: see text]), notably improved our ability to describe neuronal activity when the eye was moving and also enabled us to model abducens neuron discharges during the postsaccadic interval. We also found that, for a given model, a single set of parameters could not be used to describe neuronal firing rates during both slow and rapid eye movements. Specifically, the eye velocity and position coefficients ( r and k in the above models, respectively) consistently decreased as a function of the mean (and peak) eye velocity that was generated. In contrast, the bias ( b, firing rate when looking straight ahead) invariably increased with eye velocity. Although these trends are likely to reflect, in part, nonlinearities that are intrinsic to the extraocular muscles, we propose that these results can also be explained by considering the time-varying resistance to movement that is generated by the antagonist muscle. We conclude that to create realistic and meaningful models of the neural control of horizontal eye movements, it is essential to consider the activation of the antagonist, as well as agonist motoneuron pools.


1987 ◽  
Vol 58 (4) ◽  
pp. 832-849 ◽  
Author(s):  
D. Tweed ◽  
T. Vilis

1. This paper develops three-dimensional models for the vestibuloocular reflex (VOR) and the internal feedback loop of the saccadic system. The models differ qualitatively from previous, one-dimensional versions, because the commutative algebra used in previous models does not apply to the three-dimensional rotations of the eye. 2. The hypothesis that eye position signals are generated by an eye velocity integrator in the indirect path of the VOR must be rejected because in three dimensions the integral of angular velocity does not specify angular position. Computer simulations using eye velocity integrators show large, cumulative gaze errors and post-VOR drift. We describe a simple velocity to position transformation that works in three dimensions. 3. In the feedback control of saccades, eye position error is not the vector difference between actual and desired eye positions. Subtractive feedback models must continuously adjust the axis of rotation throughout a saccade, and they generate meandering, dysmetric gaze saccades. We describe a multiplicative feedback system that solves these problems and generates fixed-axis saccades that accord with Listing's law. 4. We show that Listing's law requires that most saccades have their axes out of Listing's plane. A corollary is that if three pools of short-lead burst neurons code the eye velocity command during saccades, the three pools are not yoked, but function independently during visually triggered saccades. 5. In our three-dimensional models, we represent eye position using four-component rotational operators called quaternions. This is not the only algebraic system for describing rotations, but it is the one that best fits the needs of the oculomotor system, and it yields much simpler models than do rotation matrix or other representations. 6. Quaternion models predict that eye position is represented on four channels in the oculomotor system: three for the vector components of eye position and one inversely related to gaze eccentricity and torsion. 7. Many testable predictions made by quaternion models also turn up in models based on other mathematics. These predictions are therefore more fundamental than the specific models that generate them. Among these predictions are 1) to compute eye position in the indirect path of the VOR, eye or head velocity signals are multiplied by eye position feedback and then integrated; consequently 2) eye position signals and eye or head velocity signals converge on vestibular neurons, and their interaction is multiplicative.(ABSTRACT TRUNCATED AT 400 WORDS)


2008 ◽  
Vol 99 (5) ◽  
pp. 2602-2616 ◽  
Author(s):  
Marion R. Van Horn ◽  
Pierre A. Sylvestre ◽  
Kathleen E. Cullen

When we look between objects located at different depths the horizontal movement of each eye is different from that of the other, yet temporally synchronized. Traditionally, a vergence-specific neuronal subsystem, independent from other oculomotor subsystems, has been thought to generate all eye movements in depth. However, recent studies have challenged this view by unmasking interactions between vergence and saccadic eye movements during disconjugate saccades. Here, we combined experimental and modeling approaches to address whether the premotor command to generate disconjugate saccades originates exclusively in “vergence centers.” We found that the brain stem burst generator, which is commonly assumed to drive only the conjugate component of eye movements, carries substantial vergence-related information during disconjugate saccades. Notably, facilitated vergence velocities during disconjugate saccades were synchronized with the burst onset of excitatory and inhibitory brain stem saccadic burst neurons (SBNs). Furthermore, the time-varying discharge properties of the majority of SBNs (>70%) preferentially encoded the dynamics of an individual eye during disconjugate saccades. When these experimental results were implemented into a computer-based simulation, to further evaluate the contribution of the saccadic burst generator in generating disconjugate saccades, we found that it carries all the vergence drive that is necessary to shape the activity of the abducens motoneurons to which it projects. Taken together, our results provide evidence that the premotor commands from the brain stem saccadic circuitry, to the target motoneurons, are sufficient to ensure the accurate control shifts of gaze in three dimensions.


1991 ◽  
Vol 35 (01) ◽  
pp. 40-57
Author(s):  
Nickolas Vlahopoulos ◽  
Michael M. Bernitsas

The dynamic behavior of a nonintegral riser bundle is studied parametrically. The dynamics of each component-riser is analyzed by a three-dimensional, nonlinear, large deflection, small strain model with coupled bending and torsion. Component-risers are slender, thin-walled, extensible or inextensible tubular beam-columns, subject to response and deformation dependent hydrodynamic loads. The con-nector equations of equilibrium are used to derive the connector forces and moments. Substructuring can thus be achieved even though in three dimensions connectors do not impose linearly dependent deflections at substructure interfaces. The developed time incremental and iterative finite-element computer code is used to analyze the effects of water depth, distribution of connectors, distance between component risers and number of finite elements in the numerical model. The problem of total CPU (central processor unit) time and the advantages of substructuring are discussed by running cases of up to 1094 degrees of freedom.


Author(s):  
Nicholas H. Wasserman

Contemporary technologies have impacted the teaching and learning of mathematics in significant ways, particularly through the incorporation of dynamic software and applets. Interactive geometry software such as Geometers Sketchpad (GSP) and GeoGebra has transformed students' ability to interact with the geometry of plane figures, helping visualize and verify conjectures. Similar to what GSP and GeoGebra have done for two-dimensional geometry in mathematics education, SketchUp™ has the potential to do for aspects of three-dimensional geometry. This chapter provides example cases, aligned with the Common Core State Standards in mathematics, for how the dynamic and unique features of SketchUp™ can be integrated into the K-12 mathematics classroom to support and aid students' spatial reasoning and knowledge of three-dimensional figures.


Author(s):  
Jeffrey S. Oishi ◽  
Geoffrey M. Vasil ◽  
Morgan Baxter ◽  
Andrew Swan ◽  
Keaton J. Burns ◽  
...  

The magnetorotational instability (MRI) occurs when a weak magnetic field destabilizes a rotating, electrically conducting fluid with inwardly increasing angular velocity. The MRI is essential to astrophysical disc theory where the shear is typically Keplerian. Internal shear layers in stars may also be MRI-unstable, and they take a wide range of profiles, including near-critical. We show that the fastest growing modes of an ideal magnetofluid are three-dimensional provided the shear rate, S , is near the two-dimensional onset value, S c . For a Keplerian shear, three-dimensional modes are unstable above S  ≈ 0.10 S c , and dominate the two-dimensional modes until S  ≈ 2.05 S c . These three-dimensional modes dominate for shear profiles relevant to stars and at magnetic Prandtl numbers relevant to liquid-metal laboratory experiments. Significant numbers of rapidly growing three-dimensional modes remainy well past 2.05 S c . These finding are significant in three ways. First, weakly nonlinear theory suggests that the MRI saturates by pushing the shear rate to its critical value. This can happen for systems, such as stars and laboratory experiments, that can rearrange their angular velocity profiles. Second, the non-normal character and large transient growth of MRI modes should be important whenever three-dimensionality exists. Finally, three-dimensional growth suggests direct dynamo action driven from the linear instability.


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