Does proprioception contribute to the sense of verticality?

The representation of the vertical can be assessed by measuring the visual, tactile or perception of the postural vertical (PV). It is well accepted that visceral graviceptors and Golgi tendon receptors contribute to perception of the PV, whereas the role of muscular proprioception (Ia muscle spindles) remains to be clarified. The objective of this study was to analyze whether or not muscular proprioception contributes to the representation of verticality. We hypothesized that the modulation of proprioception by appropriate tendinomuscular vibration may tilt the PV. We present two experiments that explore this hypothesis. PV was firstly measured in the pitch plane in twelve healthy subjects in two conditions: baseline and vibration of both Achilles tendons during six minutes, according to a counterbalanced presentation. PV orientation (mean) and dispersion (standard deviation) were calculated on the six measurements per condition. Vibration of the Achilles tendons induced a systematic backward tilt of PV (2.7 degrees +/- 0.8 degrees on average; P < 10(-3)), and no significant changes regarding PV dispersion. A clear post-effect was found for PV orientation in subjects who started the experiment with the condition vibration on. The possibility that PV could also be modulated by a much shorter duration of vibration (less than 8 s of vibration) was analyzed in a second experiment using a similar general procedure but a simplified experimental plane in other subjects. Again a backward PV tilt was induced by the vibration, but with a magnitude lower than that found in the first experiment, and with no post-effect. Since body geometry was unchanged, these two studies mean that proprioception contributes to the representation of the vertical. In conclusion, this study brings new insights to our understanding of the sense of verticality and contributes to the interpretation of backward falls induced by vibration. The induced backward fall is partly due to alignment of the erect posture with a backward-tilted referential of verticality.