How Listing's Law May Emerge from Neural Control of Reactive Saccades

We hypothesize that Listing's Law emerges as a result of two key properties of the saccadic sensory-motor system: 1) The visual sensory apparatus has a 2-D topology and 2) motor synergists are synchronized. The theory is tested by showing that eye attitudes that obey Listing's Law are achieved in a 3-D saccadic control system that translates visual eccentricity into synchronized motor commands via a 2-D spatial gradient. Simulations of this system demonstrate that attitudes assumed by the eye upon accurate foveation tend to obey Listing's Law.Office of Naval Research (N00014-92-J-1309, N00014-95-1-1409); Air Force Office of Scientific Research (90-0083

A Biomechanical Model of Human Oculomotor Plant Kinematics Based Upon Geometric Algebra

A biomechanical model of the human oculomotor plant kinematics in 3-D as a function of muscle length changes is presented. It can represent a range of alternative interpretations of the data as a function of one parameter. The model is free from such deficits as singularities and the nesting of axes found in alternative formulations such as the spherical wrist (Paul, l98l). The equations of motion are defined on a quaternion based representation of eye rotations and are compact atnd computationally efficient.Air Force Office of Scientific Research (90-0128, F49620-92-J-0225); Defense Advanced Research Projects Agency (AFOSR 90-0083); Office of Naval Research (N00014-92-J-l309

Neural Control of Rhythmic Coordinated Movements

How do humans and other animals accomplish coordinated movements? How are novel combinations of limb joints rapidly assembled into new behavioral units that move together in in-phase or anti-phase movement patterns during complex movement tasks? A neural model simulates data from human bimanual coordination tasks. As in the data, anti-phase oscillations at low frequencies switch to in-phase oscillations at high frequencies, in-phase oscillations occur both at low and high frequencies, phase fluctuations occur at the anti-phase in-phase transition, a "seagull effect" of larger errors occurs at intermediate phases, and oscillations slip toward in-phase and anti-phase when driven at intermediate phases.Air Force Office of Scientific Research (90-0128, F49620-92-J-0225, F49620-92-J-0499, 90-0083); Office of Naval Research (N00014-92-J-1309, N00014-92-J-1309); National Science Foundation (IIU-90-24877); Army Research Office (DAAL03-88-K-0088

A Neural Pattern Generator that Exhibits Frequency-Dependent In-Phase and Anti-Phase Oscillations

This article describes a. neural pattern generator based on a cooperative-competitive feedback neural network. The two-channel version of the generator supports both in-phase and anti-phase oscillations. A scalar arousal level controls both the oscillation phase and frequency. As arousal increases, oscillation frequency increases and bifurcations from in-phase to anti-phase, or anti-phase to in-phase oscillations can occur. Coupled versions of the model exhibit oscillatory patterns which correspond to the gaits used in locomotion and other oscillatory movements by various animals.Air Force Office of Scientific Research (90-0128, 90-0175); National Science Foundation (IRI-90-24877); Army Research Office (DAAL03-88-k-0088

Neural Control of Interlimb Oscillations I: Human Bimanual Coordination

How do humans and other animals accomplish coordinated movements? How are novel combinations of limb joints rapidly assembled into new behavioral units that rnove together in in-phase or anti-phase movement patterns during complex movement tasks? A neural central pattern generator (CPG) model simulates data from human bimanual coordination tasks. As in the data, anti-phase oscillations at low frequencies switch to in-phase oscillations at high frequencies, in-phase oscillation occur both at low and high frequencies, phase fluctuations occur at the anti-phase in-phase transition, a "seagull effect" of larger errors occurs at intermediate phases, and oscillations slip toward in-phase and anti-phase when driven at intermediate phases. These oscillations and bifurcations are emergent properties of the CPG model in response to volitional inputs. The CPC model is a version of the Ellias-Grossberg oscillator. Its neurons obey Hodgkin-Huxley type equations whose excitatory signals operate on a faster time scale than their inhibitory signals in a recurrent on-center off-surround anatomy. When an equal cornmand or GO signal activates both model channels the model CPC: can generate both in-phase and anti-phase oscillations at different GO amplitudes. Phase transitions frorn either in-phase to anti-phase oscillations, or from anti-phase to in- phase oscillations, can occur in different pararncter ranges, as the GO signal increases.Air Force Office of Scientific Research (F49620-92-J-0499, 90-0083, F49620-92-J-0225, 90-0128); Office of Naval Research (N00014-92-J-1309); Army Research Office (DAAL03-0088); National Science Foundation (IRI-90-24877

Neural Control of Interlimb Coordination and Gait Timing in Bipeds and Quadrupeds

1) A large body of behavioral data conceming animal and human gaits and gait transitions is simulated as emergent properties of a central pattern generator (CPG) model. The CPG model incorporates neurons obeying Hodgkin-Huxley type dynamics that interact via an on-center off-surround anatomy whose excitatory signals operate on a faster time scale than their inhibitory signals. A descending cornmand or arousal signal called a GO signal activates the gaits and controL their transitions. The GO signal and the CPG model are compared with neural data from globus pallidus and spinal cord, among other brain structures. 2) Data from human bimanual finger coordination tasks are simulated in which anti-phase oscillations at low frequencies spontaneously switch to in-phase oscillations at high frequencies, in-phase oscillations can be performed both at low and high frequencies, phase fluctuations occur at the anti-phase in-phase transition, and a "seagull effect" of larger errors occurs at intermediate phases. When driven by environmental patterns with intermediate phase relationships, the model's output exhibits a tendency to slip toward purely in-phase and anti-phase relationships as observed in humans subjects. 3) Quadruped vertebrate gaits, including the amble, the walk, all three pairwise gaits (trot, pace, and gallop) and the pronk are simulated. Rapid gait transitions are simulated in the order--walk, trot, pace, and gallop--that occurs in the cat, along with the observed increase in oscillation frequency. 4) Precise control of quadruped gait switching is achieved in the model by using GO-dependent modulation of the model's inhibitory interactions. This generates a different functional connectivity in a single CPG at different arousal levels. Such task-specific modulation of functional connectivity in neural pattern generators has been experimentally reported in invertebrates. Phase-dependent modulation of reflex gain has been observed in cats. A role for state-dependent modulation is herein predicted to occur in vertebrates for precise control of phase transitions from one gait to another. 5) The primary human gaits (the walk and the run) and elephant gaits (the amble and the walk) are sirnulated. Although these two gaits are qualitatively different, they both have the same limb order and may exhibit oscillation frequencies that overlap. The CPG model simulates the walk and the run by generating oscillations which exhibit the same phase relationships. but qualitatively different waveform shapes, at different GO signal levels. The fraction of each cycle that activity is above threshold quantitatively distinguishes the two gaits, much as the duty cycles of the feet are longer in the walk than in the run. 6) A key model properly concerns the ability of a single model CPG, that obeys a fixed set of opponent processing equations to generate both in-phase and anti-phase oscillations at different arousal levels. Phase transitions from either in-phase to anti-phase oscillations, or from anti-phase to in-phase oscillations, can occur in different parameter ranges, as the GO signal increases.Air Force Office of Scientific Research (90-0128, F49620-92-J-0225, 90-0175); National Science Foundation (IRI-90-24877); Office of Naval Research (N00014-92-J-1309); Army Research Office (DAAL03-88-K-0088); Advanced Research Projects Agency (90-0083

Neural Control of Interlimb Oscillations I: Human Bimanual Coordination

How do humans and other animals accomplish coordinated movements? How are novel combinations of limb joints rapidly assembled into new behavioral units that rnove together in in-phase or anti-phase movement patterns during complex movement tasks? A neural central pattern generator (CPG) model simulates data from human bimanual coordination tasks. As in the data, anti-phase oscillations at low frequencies switch to in-phase oscillations at high frequencies, in-phase oscillation occur both at low and high frequencies, phase fluctuations occur at the anti-phase in-phase transition, a "seagull effect" of larger errors occurs at intermediate phases, and oscillations slip toward in-phase and anti-phase when driven at intermediate phases. These oscillations and bifurcations are emergent properties of the CPG model in response to volitional inputs. The CPC model is a version of the Ellias-Grossberg oscillator. Its neurons obey Hodgkin-Huxley type equations whose excitatory signals operate on a faster time scale than their inhibitory signals in a recurrent on-center off-surround anatomy. When an equal cornmand or GO signal activates both model channels the model CPC: can generate both in-phase and anti-phase oscillations at different GO amplitudes. Phase transitions frorn either in-phase to anti-phase oscillations, or from anti-phase to in- phase oscillations, can occur in different pararncter ranges, as the GO signal increases.Air Force Office of Scientific Research (F49620-92-J-0499, 90-0083, F49620-92-J-0225, 90-0128); Office of Naval Research (N00014-92-J-1309); Army Research Office (DAAL03-0088); National Science Foundation (IRI-90-24877

A Neural Pattern Generator that Exhibits Arousal-Dependant Human Gait Transitions

A neural pattern generator based upon a non-linear cooperative-competitive feedback neural network is presented. It can generate the two standard human gaits: the walk and the run. A scalar arousal or GO signal causes a bifurcation from one gait to the next. Although these two gaits are qualitatively different, they both have the same limb order and may exhibit oscillation frequencies that overlap. The model simulates the walk and the run via qualitatively different waveform shapes. The fraction of cycle that activity is above threshold distinguishes the two gaits, much as the duty cycles of the feet are longer in the walk than in the run.Air Force Office of Scientific Research (90-0128, F49620-92-J-0225, 90-0175); National Science Foundation (IRI-90-24877); Office of Naval Research (N00014-92-J-1309); Defense Advanced Research Projects Agency (AFOSR 90-0083

Quadruped Gait Transitions from a Neural Pattern Generator with Arousal Modulated Interactions

A four-channel neural pattern generator is described in which both the frequency and the relative phase of oscillations are controlled by a scalar arousal or GO signal. The generator is used to simulate quadruped gaits; in particular, rapid transitions are simulated in the order - walk, trot, pace, and gallop - that occurs in the cat. Precise switching control is achieved by using an arousal dependent modulation of the model's inhibitory interactions. This modulation generates a different functional connectivity in a single network at different arousal levels.Air Force Office of Scientific Research (90-0128, F49620-92-J-0225, 90-0175); National Science Foundation (IRI-90-24877); Office of Naval Research (N00014-92-J-1309); Defense Advanced Research Projects Agency (AFOSR 90-0083

Neural Control of Interlimb Oscillations II. Biped and Quadruped Gaits and Bifurications

Behavioral data concerning animal and human gaits and gait transitions are simulated as emergent properties of a central pattern generator (CPG) model. The CPG model is a version of the Ellias-Grossberg oscillator. Its neurons obey Hodgkin-Huxley type equations whose excitatory signals operate on a faster time scale than their inhibitory signals in a recurrent on-center off-surround anatomy. A descending command or GO signal activates the gaits and triggers gait transitions as its amplitude increases. A single model CPG can generate both in-phase and anti-phase oscillations at different GO amplitudes. Phase transition from either in-phase to anti-phase oscillations, or from anti-phase to in-phase oscillations, can occur in different parameter ranges, as the GO signal increases. Quadruped vertebrate gaits, including the amble, the walk, all three pairwise gaits (trot, pace, and gallop), and the pronk are simulated using this property. Rapid gait transitions are simulated in the order walk, trot, pace, and gallop that occurs in the cat, along with the observed increase in oscillation frequency. Precise control of quadruped gait switching uses GO-dependent. modulation of inhibitory interactions, which generates a different functional anatomy at different arousal levels. The primary human gaits (the walk and the run) and elephant gaits (the amble and the walk) are simulated, without modulation, by oscillations with the same phase relationships but different waveform shapes at different GO signal levels, much as the duty cycles of the feet are longer in the walk than in the run. Relevant neural data from spinal cord, globus palliclus, and motor cortex, among other structures, are discussedArmy Research Office (DAAL03-88-K-0088); Advanced Research Projects Agency (90-0083); National Science Foundation (IRI-90-24877); Office of Naval Research (N00014-92-J-1309); Air Force Office of Scientific Research (F49620-92-J-0499, F49620-92-J-0225, 90-0128