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Perception of the Body in Space: Mechanisms

Laurence R. Young

10.1002/cphy.cp010322

Source: Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes

Originally published: 1984

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Perception of Orientation Based on Multiple Sensory Modalities
1.1 Semicircular Canals
1.2 Otolith Organs
1.3 Somatosensory Cues
1.4 Limb Position
2 Psychophysical Measures of Perception of Orientation and Motion
3 Angular Acceleration
3.1 Rotation in Dark
3.2 Active Versus Passive Movement
3.3 Torsion‐Pendulum Model
3.4 Thresholds
3.5 Oculogyral Illusion
3.6 Adaptation
3.7 Caloric and Alcohol Effects
4 Linear Motion and Gravity
4.1 Nature of Linear Accelerometers
4.2 Static Orientation to Vertical
4.3 Contributions of Nonlabyrinthine Sensors to Perception of Tilt
4.4 Reduced Somatosensory Cues
4.5 Labyrinthine Defective Subjects
4.6 Amplified Somatosensory or Postural Cues
5 Dynamic Response of Otolith System
5.1 Ambiguity of Subjective Response to Acceleration
5.2 Linear Acceleration Steps
5.3 Sinusoidal Linear Acceleration
6 Combined Rotational and Linear Accelerations
6.1 Consistent and Inconsistent Vestibular Signals
6.2 Centripetal Acceleration
6.3 Cross‐Coupled Angular Accelerations; Coriolis Illusion
6.4 Rotation About Off‐Vertical Axes
7 Visual Effects on Perceived Orientation—Models
7.1 Static Visual Orientation
7.2 Moving Visual Fields
7.3 Static Visual‐ Vestibular Interaction
7.4 Vection and Dynamic Visual‐Vestibular Interaction
8 Spatial Orientation in Altered Environments
8.1 Motion Sickness
9 Summary
Figure 1. Figure 1.

Definition of axes for linear and angular motion.

From Hixon et al. 89
Figure 2. Figure 2.

Schematic representation of spatial orientation process. “True state” vector, X, consisting of linear and angular positions and velocities, is produced by changes resulting from three sources: unforced behavior of body, Xprocessed by A, commanded body changes, commands U processed by B, and unmeasured disturbances, R. Various sensors are each responsive, especially to one or more components of measured state. Symbol “∧” indicates an estimate of the vector; ω, angular velocity; f, specific force, gravity minus linear accleration; θ, angle. Measured state signals are combined with “expected state,” , derived from a presumed internal model of the body, in optimum estimator to produce estimate of orientation, .

From Young 174
Figure 3. Figure 3.

Subjective estimates of angular displacement (produced by triangular velocity wave form) are greater for retrospective than for concurrent estimates.

Adapted from Guedry 76
Figure 4. Figure 4.

Subjective angular velocity decays and may reverse during prolonged constant‐velocity rotation. Sudden stop after prolonged rotation elicits transient, oppositely directed, postrotatory response.
Figure 5. Figure 5.

Subjective velocity follows actual velocity only for 1st 10 s of constant angular acceleration, then plateaus and actually decays.

Adapted from Guedry and Lauver 183
Figure 6. Figure 6.

Normalized torsion‐pendulum response, X, for system with long time constant, TL = 15 s. A: rising response to acceleration step is detected when it reaches Xmin. B: decaying response following velocity step has duration tμ.
Figure 7. Figure 7.

Time to detect, tdet, step of yaw angular acceleration of magnitude α increases sharply for acceleration levels less than 2–3 deg/s2.

Adapted from Guedry 76
Figure 8. Figure 8.

Theoretical response of the torsion‐pendulum cupula model to threshold velocity step, Ut(t) = e−t/18, to threshold acceleration step, at(t) = 1 − e−t/18, and to a combined stimulus c(t) = K[Ut(t) + at(t)]. In theory, calculated cupula response for c(t) never exceeds threshold response to velocity or acceleration threshold steps alone and would be undetectable for K as large as unity. Measured thresholds for c(t) tended to values of K below 0.75, lending support to signal‐detection model rather than hard limit model for threshold.

Adapted from Ormsby 131
Figure 9. Figure 9.

Frequency response of adaptation model for subjective sensation during yaw angular motion: subjective angular velocity/input angular velocity equals 24.8e−0.3ss2/(s + 25)(s + 0.0625)(s + .033).

From Young and Oman 178
Figure 10. Figure 10.

Exaggerated cupula displacement during A: angular acceleration; B: caloric stimulation; C: first phase of alcohol nystagmus (PAN I).
Figure 11. Figure 11.

Measurement of relative inclination of luminous line that subjects set to apparent vertical when they are tilted laterally. Range and median shown for 13 subjects. Overestimation of tilt at small angles is the E or Müller effect shown by some subjects. The underestimation A or Aubert effect at large angles is more pronounced.

Adapted from Bischoff 9, using data of Udo de Haes 157
Figure 12. Figure 12.

Measurements of angle of a line set to apparent vertical, plotted against lateral component of specific force for various body tilt angles (ϕ) and g levels. Apparent tilt varies according to shear force; it also increases with compressive force component.

Adapted from Correia et al. 36,37
Figure 13. Figure 13.

Schematic representation of perceived pitch categories in various gravitational fields. The z‐axis component of specific force determines whether actual pitch is underestimated (category A) or overestimated (category E).

From Ormsby and Young 132
Figure 14. Figure 14.

Alteration of z‐component of specific force to yield observed errors in perceived pitch. See text for explanation.

From Ormsby and Young 132
Figure 15. Figure 15.

Model predictions for perceived lateral tilt angle as function of actual tilt (as in Fig. 11) compared with data taken at 1 g and 2 g.

From Ormsby and Young 132
Figure 16. Figure 16.

Comparison between settings of line to perceived horizontal by normals and labyrinthine defectives (LDs) in air and when tactile cues were reduced by submersion in water.

Adapted from Graybiel 71
Figure 17. Figure 17.

Latency to detection of step of horizontal linear acceleration for subjects upright. See text for equation of model.

From Meiry 112
Figure 18. Figure 18.

Latency to detection of steps of vertical linear acceleration for subjects upright. Model is regression hyperbola for 8 subjects, yielding minimum response time of 0.37 s and velocity constant of 0.022 g‐s.

From Melvill‐Jones and Young 117
Figure 19. Figure 19.

Phase angle frequency response for perceived velocity vs, input velocity. Vertical lines showing ±1 SD.

Dynamic ocular counterrolling vs. lateral specific force, measured by Kellogg 184. From Young and Meiry 187
Figure 20. Figure 20.

Illustration of perception of pitch during constant acceleration.

From Benson 181
Figure 21. Figure 21.

Shift of direction of specific force vector during constant velocity centrifuge rotation produces slow shift in direction of estimated vertical or horizontal.

Adapted from Young 186 and Graybiel and Brown 67
Figure 22. Figure 22.

Coordinated turn leading to error of spatial orientation.
Figure 23. Figure 23.

Illustration of cross‐coupled stimulation. Rolling head movement, ωx, during sustained yaw rotation, ωz, leads to erroneous transient perception of yaw and pitch velocity.

From Benson 181
Figure 24. Figure 24.

Schematic representation of compensation principle for effect of head tilt on retinal image or proximal stimulus. ϕ, Interference function; ϕ−1, compensatory function; S, mapping function of sensory channel.

Adapted from Bischoff 9
Figure 25. Figure 25.

Flow‐chart representation of visual‐vestibular interaction.

From Young 174
Figure 26. Figure 26.

Model of optic‐vestibular orientation to the vertical.

Adapted from Bischoff 9
Figure 27. Figure 27.

Sensory conflict model for resolution of visual‐vestibular interaction. A: dual input conflict model; B: conflict measure and weighting function.

From Zacharias and Young 180


Figure 1.

Definition of axes for linear and angular motion.

From Hixon et al. 89


Figure 2.

Schematic representation of spatial orientation process. “True state” vector, X, consisting of linear and angular positions and velocities, is produced by changes resulting from three sources: unforced behavior of body, Xprocessed by A, commanded body changes, commands U processed by B, and unmeasured disturbances, R. Various sensors are each responsive, especially to one or more components of measured state. Symbol “∧” indicates an estimate of the vector; ω, angular velocity; f, specific force, gravity minus linear accleration; θ, angle. Measured state signals are combined with “expected state,” , derived from a presumed internal model of the body, in optimum estimator to produce estimate of orientation, .

From Young 174


Figure 3.

Subjective estimates of angular displacement (produced by triangular velocity wave form) are greater for retrospective than for concurrent estimates.

Adapted from Guedry 76


Figure 4.

Subjective angular velocity decays and may reverse during prolonged constant‐velocity rotation. Sudden stop after prolonged rotation elicits transient, oppositely directed, postrotatory response.



Figure 5.

Subjective velocity follows actual velocity only for 1st 10 s of constant angular acceleration, then plateaus and actually decays.

Adapted from Guedry and Lauver 183


Figure 6.

Normalized torsion‐pendulum response, X, for system with long time constant, TL = 15 s. A: rising response to acceleration step is detected when it reaches Xmin. B: decaying response following velocity step has duration tμ.



Figure 7.

Time to detect, tdet, step of yaw angular acceleration of magnitude α increases sharply for acceleration levels less than 2–3 deg/s2.

Adapted from Guedry 76


Figure 8.

Theoretical response of the torsion‐pendulum cupula model to threshold velocity step, Ut(t) = e−t/18, to threshold acceleration step, at(t) = 1 − e−t/18, and to a combined stimulus c(t) = K[Ut(t) + at(t)]. In theory, calculated cupula response for c(t) never exceeds threshold response to velocity or acceleration threshold steps alone and would be undetectable for K as large as unity. Measured thresholds for c(t) tended to values of K below 0.75, lending support to signal‐detection model rather than hard limit model for threshold.

Adapted from Ormsby 131


Figure 9.

Frequency response of adaptation model for subjective sensation during yaw angular motion: subjective angular velocity/input angular velocity equals 24.8e−0.3ss2/(s + 25)(s + 0.0625)(s + .033).

From Young and Oman 178


Figure 10.

Exaggerated cupula displacement during A: angular acceleration; B: caloric stimulation; C: first phase of alcohol nystagmus (PAN I).



Figure 11.

Measurement of relative inclination of luminous line that subjects set to apparent vertical when they are tilted laterally. Range and median shown for 13 subjects. Overestimation of tilt at small angles is the E or Müller effect shown by some subjects. The underestimation A or Aubert effect at large angles is more pronounced.

Adapted from Bischoff 9, using data of Udo de Haes 157


Figure 12.

Measurements of angle of a line set to apparent vertical, plotted against lateral component of specific force for various body tilt angles (ϕ) and g levels. Apparent tilt varies according to shear force; it also increases with compressive force component.

Adapted from Correia et al. 36,37


Figure 13.

Schematic representation of perceived pitch categories in various gravitational fields. The z‐axis component of specific force determines whether actual pitch is underestimated (category A) or overestimated (category E).

From Ormsby and Young 132


Figure 14.

Alteration of z‐component of specific force to yield observed errors in perceived pitch. See text for explanation.

From Ormsby and Young 132


Figure 15.

Model predictions for perceived lateral tilt angle as function of actual tilt (as in Fig. 11) compared with data taken at 1 g and 2 g.

From Ormsby and Young 132


Figure 16.

Comparison between settings of line to perceived horizontal by normals and labyrinthine defectives (LDs) in air and when tactile cues were reduced by submersion in water.

Adapted from Graybiel 71


Figure 17.

Latency to detection of step of horizontal linear acceleration for subjects upright. See text for equation of model.

From Meiry 112


Figure 18.

Latency to detection of steps of vertical linear acceleration for subjects upright. Model is regression hyperbola for 8 subjects, yielding minimum response time of 0.37 s and velocity constant of 0.022 g‐s.

From Melvill‐Jones and Young 117


Figure 19.

Phase angle frequency response for perceived velocity vs, input velocity. Vertical lines showing ±1 SD.

Dynamic ocular counterrolling vs. lateral specific force, measured by Kellogg 184. From Young and Meiry 187


Figure 20.

Illustration of perception of pitch during constant acceleration.

From Benson 181


Figure 21.

Shift of direction of specific force vector during constant velocity centrifuge rotation produces slow shift in direction of estimated vertical or horizontal.

Adapted from Young 186 and Graybiel and Brown 67


Figure 22.

Coordinated turn leading to error of spatial orientation.



Figure 23.

Illustration of cross‐coupled stimulation. Rolling head movement, ωx, during sustained yaw rotation, ωz, leads to erroneous transient perception of yaw and pitch velocity.

From Benson 181


Figure 24.

Schematic representation of compensation principle for effect of head tilt on retinal image or proximal stimulus. ϕ, Interference function; ϕ−1, compensatory function; S, mapping function of sensory channel.

Adapted from Bischoff 9


Figure 25.

Flow‐chart representation of visual‐vestibular interaction.

From Young 174


Figure 26.

Model of optic‐vestibular orientation to the vertical.

Adapted from Bischoff 9


Figure 27.

Sensory conflict model for resolution of visual‐vestibular interaction. A: dual input conflict model; B: conflict measure and weighting function.

From Zacharias and Young 180
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Article Title: Perception of the Body in Space: Mechanisms
 

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Laurence R. Young. Perception of the Body in Space: Mechanisms. Compr Physiol 2011, Supplement 3: Handbook of Physiology, The Nervous System, Sensory Processes: 1023-1066. First published in print 1984. doi: 10.1002/cphy.cp010322