VESTIBULAR SIGNALS Jan Van Gisbergen
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Transcript of VESTIBULAR SIGNALS Jan Van Gisbergen
VESTIBULAR SIGNALS
Jan Van Gisbergen
PhD COURSE SENSORY SYSTEMSUtrecht, September 29, 2008
detection of self motion sensing body orientation in space visual perception in earth-centric coordinates
SCOPE
Functions and limitations of vestibular sensors
Ambiguity problem of the otoliths
Solution to the ambiguity problem
Visual-vestibular fusion
Transformations from head to body reference frame
Visual space perception in static tilt
Bayesian model
MULTISENSORY INTEGRATION
VESTIBULAR SENSORS
functions &
limitations
VESTIBULAR SENSORS
canals
otoliths
SEMICIRCULAR CANALS
measure head rotation
CANAL GEOMETRY
Three perpendicular canals measure head rotation in 3 dimensions
BEST ROTATION AXES
a = anterior canal
p = posterior canal
h = horizontal canal
rotation in direction of arrow excites afferents from a given canal
BEST ROTATION AXIS OF CANAL AFFERENT
Yakushin (2006) JNP
OPTIMAL ROTATION AXES OF CANAL AFFERENTS
DYNAMICS OF CANAL SIGNALS
Insensitive to low-frequency rotations (high pass filter)
Canal afferent fiber in 8th nerve• High resting discharge• Codes cupula deviation
CONSTANT ROTATION IN DARKNESS
• rotation percept decays
• after stop, percept of rotation in opposite direction
• reflects cupular mechanics
OTOLITHS
sensitive to linear acceleration during translation and to tilt, due to the pull of gravity
HAIRCELL ACTIVATION
• each haircell is connected to separate nerve fiber
• deflecting cilia toward kinocilium depolarizes haircell
POLARISATION VECTOR
Eron et al. (2008) J. Neurophysiol.
each otolith cell has cosine tuning:
indicates head orientation where pull of gravity has maximum effect
tilt stimuli:
nose down left ear down nose up
OTOLITHS
sensitive to tilt and translation
POLARISATION VECTORS
Fernandez and Goldberg (1976) JNP
AMBIGUITY PROBLEMOF THE OTOLITHS
OTOLITH SIGNAL IS AMBIGUOUS
hair cells cannot distinguish tilt and translation
SOMATOGRAVIC ILLUSION
pilot is upright, but feels tilted
(only in darkness)
AMBIGUITY PROBLEM
otolith signal may have various causes:
• translation (a)• force of gravity due to tilt (g)• combination of a and g
How can the brain resolve this ambiguity ?
inverse problem
CANAL- OTOLITH INTERACTION MODEL
• canals detect rotation during tilt changes
• their signal helps to decompose otolith signal
Angelaki et al. (1999)
CANAL–OTOLITH INTERACTION MODEL
basic principle:- tilt stimulates otoliths AND canals- translation stimulates only otoliths
Merfeld and Zupan (2002) J. Neurophysiology
tilt angle
linear acceleration
angular velocity
percepts during rotation about a tilted axis
(OVAR)
Vingerhoets et al. (2006) J. Neurophysiol.
Vingerhoets et al. (2007) J. Neurophysiol.
TESTING THE MODEL
THE ACTUAL MOTION
- rotation about tilted axis
- in darkness
- constant velocity
MODEL PREDICTIONS
rotation signal decays gradually
wrong interpretation otolith signal: illusory translation percept
SCHEMATIC SUMMARY OF RESULTS
confirms prediction
rotation percept
translation percept
Actual motion:
Percept:
TRANSLATION AND ROTATION PERCEPT DATA
rotation percept
translation percept
VISUAL – VESTIBULAR FUSION
FUSION OF VISUAL AND VESTIBULAR SIGNALS FOR DETECTION OF
EGOMOTION
• brain interprets whole field motion as due to egomotion
• can detect constant velocity motion (low pass)
• complements vestibular motion detection (high pass)
CANALS ARE HIGH PASS
ROTATION IN LIGHT: VISUAL CONTRIBUTION
• Rotation percept in the light is veridical
• Visual system detects low frequencies
• Canals detect high frequencies
circular vection
CONVERGENCE IN VESTIBULAR NUCLEUS
visual system detects low frequencies
canals detect high frequencies
OPTIC FLOW PROVIDES LOW FREQUENCY INPUT FROM LINEAR MOTION
• otoliths detect linear acceleration
• optic flow can induce linear egomotion (train illusion!!)
TRANSFORMATIONS FROM HEAD TO BODY
REFERENCE FRAME
BALANCE
1. Otoliths measure head position in space
2. To maintain balance, we must know body position in space
3. Which mechanisms are involved?
BALANCE
1. Measure head position in space (HS)
2. Measure position head on trunk (HB)
3. Compute position of body in space (BS):
BS = HS - HB
EFFECT OF ELECTRICAL OTOLITH STIMULATION
•experiment in darkness
•results in body tilt
Why?
HS changes
HB is not changed
BS changes, subject corrects
BS = HS - HB
VESTIBULAR - NECK PROPRIOCEPTIVE INTERACTIONS
neuron b codes linear motion in body-centered reference frame (accounts for neck signals)
neuron c codes motion in head-reference frame
Angelaki (2008) Ann Rev Neuroscience
monkey moves on sled in various directions
cell recording in cerebellum (FN)
VISUAL SPACE PERCEPTION IN
STATIC TILT
VISUAL VERTICAL
1. How can we determine the orientation of visual objects relative to the direction of gravity, even when we are tilted in darkness?
2. What is the role of the vestibular system?
SENSING THE DIRECTION OF GRAVITY
Two different tasks:
1. Set line to vertical (SVV)
2. Estimate your body tilt (SBT)
Van Beuzekom & Van Gisbergen (2000) J. Neurophysiol.
Van Beuzekom et al. (2001) Vision Res.
Kaptein & Van Gisbergen (2004, 2005) J. Neurophysiol.
De Vrijer et al. (2008) J. Neurophysiol.
experiments in darkness
ACCURACY vs PRECISION
Accuracy:
How close is the response to the true value?
Precision:
How reproducible is the response?
darts analogy:
ACCURACY AND PRECISION IN LINE TASK (SVV)
accuracy
precision
De Vrijer et al. (2008) J. Neurophysiol.
De Vrijer et al. (2008) in progress
ACCURACY IN LINE TASK
due to underestimation of body tilt?
NO UNDERESTIMATION OF BODY TILT
SVV SBT
• Subjects know quite well how they are tilted (SBT)
• Yet, their line settings undercompensate for tilt (SVV)
Van Beuzekom et al. (2001) Vision Res.
Kaptein and Van Gisbergen (2004) J. Neurophysiol.
PRECISION IN LINE TASK
is scatter in SVV simply reflection of noise in body tilt signal?
De Vrijer et al. (2008) J. Neurophysiol.
De Vrijer et al. (2008) in progress
SVV LESS NOISY THAN SBT
De Vrijer et al. in progress
psychometric experiments at 0o and 90o tilt:
SVV LESS NOISY THAN SBT
SUMMARY SBT AND SVV DATA
Two paradoxical findings:
1. subject knows tilt angle, yet makes biased line settings
2. more certain about line setting than about body tilt
estimate body tilt (SBT) adjust line to vertical (SVV)
SBT DATA SHOW:
• An unbiased head tilt signal is available
• Noise increases with tilt angle
SIGNALS REQUIRED FOR SPATIAL VISION
retinal signal
to compute line in space (Ls), brain must combine info about line orientation on retina (LR) and head tilt (HS)
head-tilt signal
SIMPLY USING RAW TILT SIGNAL …
would not explain SVV bias !!spatial vision would be accurate, but noisy
raw tilt signal
A BAYESIAN PERSPECTIVE
IDEAL OBSERVER MODEL
IDEAL OBSERVER STRATEGY
1) Use sensory data: noisy tilt signal suggests range of possible tilt angles (likelihood)
2) Use prior knowledge: we know that large tilt angles are very uncommon (prior)
3) Most likely tilt angle (posterior) is product of likelihood and prior
Eggert (1998) PhD Thesis, Munich
MacNeilage et al. (2007) Exp. Brain Res.
De Vrijer et al. (2008) J. Neurophysiol.
IDEAL OBSERVER STRATEGY
Tilt prior has 2 effects on SVV:
• Less noise
• Bias at large tilt
WHY WOULD THIS MAKE SENSE?
1) Less noise in spatial vision
2) Downside: bias at large tilts
3) Average performance improves (large tilts are rare)
DEMO
BIAS EFFECT INCREASES WITH TILT
no bias
De Vrijer et al. (2008) J. Neurophysiology
no bias
small bias
small bias
large bias
large bias
MODEL PARAMETERS
1) head tilt noise level in upright
2) increase of head tilt noise with tilt
3) prior width
4) eye torsion amplitude
MODEL FITS: SVV ACCURACY
MODEL FITS: SVV ACCURACY
< 0
QUESTIONS1) The canal-related neuron illustrated in the paper by Yakushin (see slide 10 in lecture) can be
identified by its excitatory and inhibitory responses to sinusoidal rotation about various axis orientations relative to the head.
a) What would happen if the same head-fixed rotations would be applied, but now with the animal in supine position, rather than in the prone position shown in the figure? Explain your answer.
b) Sketch expected response patterns for a neuron getting its input from the right anterior canal, when using the same rotation axes as shown in the lecture sheet?
c) Estimate the frequency of rotation and explain why its choice is important
2) In the scheme showing how combining a signal Hs and a signal coding Hb can yield a signal body orientation in space (Bs), the two input signals have opposite signs (see slide 38).
a) Explain why this makes sense, given the fact that we can tilt the head and still remain upright
b) Explain why electrical otolith stimulation does cause perturbation of balance
MODEL EXPLANATION OF NOISE LEVELS:SVV vs SBT PRECISION
• SVV is less noisy than the SBT (remarkable, but explained by model)
• SBT becomes more noisy at larger tilt (supports model assumption)
• SBT noise levels compatible with head-tilt fit results
CONCLUSION
Accuracy-precision trade-off in spatial vision:
• Bayesian strategy reduces noise at small tilts
• causes systematic errors at large tilts
CENTRIFUGE EXPERIMENT
proefpersoon wordt in donker langdurig rondgedraaid op centrifuge
• draait nog steeds rond maar voelt geen rotatie meer
• zit in feite rechtop maar voelt zich gekanteld
OVAR
TWO TRANSFORMATIONS OF PRIMARY VESTIBULAR SIGNALS
1. Signals from the canals (ω) are used to decompose signals from otolith afferents (α) into gravitational (g, orientation) and translational (f) components.
2. Gravitational estimates are also used to transform head-fixed angular velocity signals from the
semicircular canals (ω) into inertial velocity, i.e., space-referenced angular velocity (ωs)