The proprioceptive coding of multidirectional ankle joint movements was investigated, focusing in particular on the question as to how accurately the direction of a movement is encoded when all the proprioceptive information from all the muscles involved in the actual movement is taken into account. During ankle movements imposed on human subjects, the activity of 30 muscle spindle afferents originating in the extensor digitorum longus, tibialis anterior, extensor hallucis longus and peroneus lateralis muscles was recorded from the lateral peroneal nerve using the microneurographic technique. In the first part of the study, it was proposed to investigate whether muscle spindle afferents have a preferred direction, as previously found to occur in the case of cortical cells, and to analyze the neural coding of the movement trajectories using a "population vector model." This model is based on the idea that neuronal coding can be analyzed in terms of a series of vectors, each based on specific movement parameters. In the present case, each vector gives the mean contribution of a population of muscle spindle afferents within one directionally tuned muscle. A given population vector points in the "preferred sensory direction" of the muscle to which it corresponds, and its length is the mean frequency of all the afferents within that muscle. Our working hypothesis was that the sum of these weighted vectors points in the same direction as the ongoing movement. The results show that each muscle spindle afferent, and likewise each muscle, has a specific preferred sensory direction, as well as a preferred sensory sector within which it is capable of sending sensory information to the central nervous system. Interestingly, the results also demonstrate that the preferred directions are the same as the directions of vibration-induced illusions. In addition, the results show that the neuronal population vector model describes the multipopulation proprioceptive coding of spatially oriented 2D limb movements, even at the peripheral sensory level, based on the sum vectors calculated from all the muscles involved in the movement. In an accompanying paper, the coding of more complex 2D movements such as those involved in drawing rectilinear and curvilinear geometrical shapes was investigated.

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The aim of the present study was to establish if there exists reflex connections from ligamentous structures in cervical facet joints and the fusimotor system of dorsal neck muscles. In seven cats, anaesthetized with alpha-chloralose, bradykinin (BK) of concentrations between 12 and 50 microg was injected into the facet joint between C1 and C2. Recordings were made from single muscle spindle afferents (MSA) originating in contralateral trapezius and splenius muscles (TrSp). Fusimotor induced changes in the sensitivity of the muscle spindle afferents were assessed by recording the responses to sinusoidal stretches of the TrSp muscles. The mean rate of discharge and the depth of modulation of a fitted sine were taken as quantitative estimates of the response. A total of 25 MSAs were recorded, and 21 of these showed clear-cut alterations in their responses to the sinusoidal stretches following Bk. injections into contralateral facet joint. The majority of the responding afferents (13/21) showed changes in their responses indicating an increased activity of static fusimotoneurones, although responses of dynamic and mixed static and dynamic nature were also seen. Local anaesthetics applied to the intraarticular receptors abolished the effects. Injection (i.v.) of a general anaesthetic (pentobarbital) abolished the effects. The results show that there exist reflex connections between receptors in cervical facet joints and fusimotoneurones of dorsal neck muscles, and this might be of importance in the pathophysiology behind whiplash associated disorders (WAD).

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The aim of the present study was to further investigate the contribution of primary muscle spindle feedback to proprioception and higher brain functions, such as movement trajectory recognition. For this purpose, complex illusory movements were evoked in subjects by applying patterns of muscle tendon vibration mimicking the natural Ia afferent pattern. Ia afferent messages were previously recorded using microneurographic method from the six main muscle groups acting on the ankle joint during imposed "writing like" movements. The mean Ia afferent pattern was calculated for each muscle group and used as a template to pilot each vibrator. Eleven different vibratory patterns were applied to ten volunteers. Subjects were asked both to copy the perceived illusory movements by hand on a digitizing tablet and to recognize and name the corresponding graphic symbol. The results show that the Ia afferent feedback of a given movement evokes the illusion of the same movement when it is applied to the subject via the appropriate pattern of muscle tendon vibration. The geometry and the kinematic parameters of the imposed and illusory movements are very similar and the so-called "two-thirds power law" is present in the reproduction of the vibration-induced illusory movements. Vibrations within the "natural" frequency range of Ia fibres firing (around 30 Hz) produce clear illusions of movements in all the tested subjects. In addition, increasing the mean frequency of the vibration patterns resulted in a linear increase in the size of the illusory movements. Lastly, the subjects were able to recognize and name the symbols evoked by the vibration-induced primary muscle spindle afferent patterns in 83% of the trials. These findings suggest that the "proprioceptive signature" of a given movement is associated with the corresponding "perceptual signature". The neural mechanisms possibly underlying the sensory to perceptual transformation are discussed in the general framework of "the neuronal population vector model".