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John H. Lawrence III and T. Richard Nichols

Muscle actions are often defined within a single anatomical reference plane. Yet animals must control posture and movement within a three-dimensional (3-D) environment, responding to a 3-D array of perturbing forces. Based on information gained regarding the 3-D muscle mechanics at the cat ankle joint complex (companion paper), we decided to study how alterations in the 3-D AJC orientation might affect ankle joint postural control. We used a 6 degree-of-freedom force-moment sensor to assess the affect of ankle joint orientation on the 3-D isometric joint torques evoked by electrical stimulation of muscles crossing the ankle joint complex (AJC) in the deeply anesthetized cat. An orthogonal axis system was established at the designated ankle rotation center, such that pitch (flexion-extension), yaw (abduction-adduction), and roll (inversion-eversion) axis torques were calculated. Experimental results suggest that both the magnitude and sign of extra-sagittal torques from the gastrocnemius muscles are joint angle dependent. Also, the hind limb levering system stabilizes the AJC against large yaw and roll rotations away from the control position.

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Thomas J. Burkholder and T. Richard Nichols

Postural regulation is an important part of a variety of motor tasks, including quiet standing and locomotion. Muscle length feedback, both the autogenic length feedback arising from a muscle's own spindles, and heterogenic length feedback, arising from its agonists and antagonists, is a strong modulator of muscle force and well suited to postural maintenance. The effects of this reflex feedback on 3-D force generation and limb mechanics are not known. In this paper, we present a mechanical model for relating 3-D changes in cat hindlimb posture to changes in muscle lengths. These changes in muscle length are used to estimate changes in both intrinsic muscle force generation and muscle activation by length feedback pathways. Few muscles are found to have directly agonist mechanical actions, and most differ by more than 20°. Endpoint force fields are largely uniform across the space investigated. Both autogenic and heterogenic feedback contribute to whole limb resistance to perturbation, autogenic pathways being most dramatic. Length feedback strongly reinforced a restoring force in response to end-point displacement.

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John H. Lawrence III and T. Richard Nichols

Muscle actions are often defined with respect to a single anatomical reference plane based on a “predominant” functional activity. Yet animals must control posture and movement within a three-dimensional (3-D) environment, exerting control over more than one reference plane when responding to a 3-D array of perturbing forces. Consequently, enhanced knowledge concerning the 3-D torque capabilities of certain appendicular muscles might provide for greater understanding of the biomechanical basis for motor control. We propose that the cat postural control mechanism utilizes the inherent 3-D mechanical actions of ankle flexors and extensors to maintain extra-saggital joint stiffness. We used a 6 degree-of-freedom force-moment sensor to assess the effect of ankle joint orientation on the 3-D nature of isometric joint torques evoked by electrical stimulation of muscles crossing the AJC in the deeply anesthetized cat. An orthogonal axis system was established at the designated ankle rotation center, such that pitch (defining flexion-extension), yaw (abduction-adduction), and roll (inversion-eversion) axis torques were calculated. Experimental results show that the classical cat ankle flexor and extensors evoke large extra-sagittal torques as well. Also, the hind limb levering system stabilizes the AJC against large yaw and roll rotations away from the control position.

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Sondra G. Siegel, T. Richard Nichols and Timothy C. Cope

Cutaneous reflexes have been described primarily according to their actions in the flexion/extension plane. It is shown here, by measuring electromyography and isometric force in decerebrate cats, that ankle muscles are activated in relation to their actions in the abduction/adduction plane during sural nerve (SNR) and crossed-extension (XER) reflexes. Ankle adductors (tibialis posterior, extensor digitorum longus, and flexors digitorum and hallucis longus) were active in XER, but not in SNR. Muscles producing ankle abduction (medial and lateral gastrocnemii and peroneus longus and brevis) were often activated in both reflexes, and medial gastrocnemius and peroneus longus were consistently more strongly activated in SNR than in XER. This differential pattern of muscle activation results in greater abduction torque at the ankle in SNR than in XER. These data demonstrate reflex organization in relation to the multidirectional torque generated by individual muscles.

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Alan J. Sokoloff, Timothy C. Cope, T. Richard Nichols and Arthur W. English

Differences in the directions of torque produced by motor units might be used by the nervous system to coordinate posture and movement. Here we report plantar flexion and abduction isometric torques exerted at the ankle by 158 motor units in the cat medial gastrocnemius (MG) muscle. In five cats, motor unit torque direction differed by an average of 10°. Torque direction was weakly correlated with unit contraction time in 3 of 5 experiments, with tetanic force in 3 of 5 experiments, and with conduction velocity in 1 of 5 experiments. The direction of whole muscle torque, however, was constant at all levels of MG activation produced in sural and crossed extension reflexes. Thus, although there is a range in the direction of torque produced by motor units in the cat MG, we find no evidence for the ordering of motor units according to torque direction during MG activation.