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Life Science Open Day | 2006 | Weizmann Institute of Science

972 8 934 4574


Department of
Neurobiology
972 8 934 6099
ehud.ahissar@weizmann.ac.il
Sensation is an active process. For example, while viewing and touching, the eyes and
hands are constantly moving. These movements prevent receptor adaptation, but also
optimize sensory processing. Studying the computational basis of such active sensing
is difficult, because of closed-loop interactions between motor control and sensory
processing. Thus, most research on sensory processing and motor control is conducted
in an open loop condition, in which either motor control or sensory processing is
eliminated.
Sensory information is encoded by both spatial and temporal cues, and accurate
representations of the world depend on both types of information. Rats move their
whiskers repetitively to explore the environment, and to detect, localize and identify
objects. Sensory information, in turn, affects future motor movements. How this motor-
sensory-motor functional loop is implemented across neuronal loops of the whisker
system is not yet known. Our present focus is on the principles of motor control of sensor
movements; peripheral encoding induced by sensor movements, and central decoding
of the sensory information.
Whisking and object localization
During exploration, localization and identification of objects rats actively move
their whiskers back-and-forth, in a stereotypical pattern termed whisking. Recently,
we developed methods of characterizing this behavior in detail in freely-moving and
behaving animals (Knutsen et al, 2005). We found that rats control whisking variables,
such as amplitude, velocity and duration, separately and in a context dependent manner.
We have designed perceptual paradigms in order to assess the acuity with which rats
can localize objects using their whiskers. We have found that whisking behavior is
important for localization, and that it is not merely sufficient for rats to contact objects
to determine their locations: the whiskers must be able to move, independently of
head movements, in order for rats to accurately localize. Future studies will be aimed
at understanding how whisking facilitates the encoding of object location, and how it
relates to neural activity.
Active sensing mediated by multiple motor-
sensory-motor loops
Ehud Ahissar
Sebastian Haidarliu
Knarik Bagdasarian
Naama Rubin
Per Knutsen
Moshe Fried
Chunxiu Yu
Avraham Saig
Erez Simony
www.weizmann.ac.il/ neurobiology/labs/
ahissar/ahissare.htm
Fig 1. Whisking behavior during object localization A. Movie frames from a localization
trial. The rat was trained to determine which of the two vertical poles was more posterior. B. The
behavioral setup. C. Whisker movements were tracked from high-speed video frames and the angle
of the whiskers relative to the head plotted. The angles of the right and left C2 whiskers are plotted;
thick line indicates moments a whisker contacted the poles.
Encoding of active vibrissal touch in trigeminal
ganglion
Using artificial whisking in anesthetized rats, we found that
vibrissal information is carried by three types of neurons in
the trigeminal ganglion: Whisking cells, Touch cells and
Whisking/Touch cells, which convey specific signals related to
active whisking and touch: whisking motion, contact timing,
and combined whisking-touch information. We found that the
horizontal coordinate of object location is encoded primarily
by spike timing (Szwed et al., 2003), the vertical by neuron
identity, and the radial primarily by firing rate (Szwed et al.,
2006).
Parallel processing in thalamus
To understand how these signals are processed by the brain to
localize and identify objects, we recorded from individual neurons
from the three major thalamic nuclei of the whisker system, each
belonging to a different afferent pathway - paralemniscal, a
recently discovered pathway (extralemniscal), and the lemniscal
pathway. We found that different sensory signals related to
active touch are conveyed separately via the thalamus by these
three parallel afferent pathways. The paralemniscal conveys
sensor motion (whisking) signals via the POm, the extralemniscal
conveys contact (touch) signals via VPMvl, and the lemniscal
pathway conveys combined whisking-touch signals via VPMdm.
Since each of these pathways project to a different target in
the brain, and eventually closes the sensory-motor loop along
a separate path, the functional segregation we observed
raises a novel possibility for hierarchical processing: Instead of
periphery-to-cortex hierarchical sensory processing which leads
to cortex-to-periphery hierarchical motor processing, as current
dogmas suggest, the brain might implement the processing of
active touch along nested motor-sensory-motor loops, organized
hierarchically. In the case of vibrissal processing, our results
suggest that sensor-motion control is implemented along a
low-order loop that includes the paralemniscal pathway, object
localization along a higher-order loop via the extralemniscal
pathway, and object identification along a further higher-order
loop via the lemniscal pathway (Fig.2, Yu et al., in press).
Layer-specific responses in cortex
While studying cortical representations of active touch
and object location, we found that the dynamics of neuronal
responses were affected by the presence of an object in a
layer specific manner. In both Whisking and Touch conditions,
deep neurons (layers 4 and 5a) exhibited facilitation during the
whisking train, while superficial neurons (layer 2/3) exhibited
depression. In layers 2/3 and 4, responses were stronger during
touch trials than during whisking in air. In layer 5a response
strengths were similar for both conditions. Overall, our results
indicate that, in the cortex, adaptation does not depend only on
the level of activity or the frequency of its repetition, but rather
on the nature of the sensory information that is conveyed by
that activity.
Selected publications
Shulz, D.E., Sosnik, R., Ego, V., Haidarliu,S. and Ahissar, E. (2000)
A neuronal analogue of state-dependent learning. Nature, 403,
549-553.
Ahissar, E., Sosnik, R. and Haidarliu, S. (2000) Transformation
from temporal to rate coding in somatosensory thalamocortical
system. Nature, 406, 302-306.
Gamzu, E. and Ahissar, E. (2001) Importance of temporal cues for
tactile spatial frequency discrimination. J. Neurosci., 21, 7416-
7427.
Ahissar, E. and Arieli, A. (2001) Figuring space by time. Neuron,
32, 185-201.
Ahissar, E. and Kleinfeld, D. (2003) Closed-loop neuronal
computations: focus on vibrissa somatosensation in rat.
Cerebral Cortex, 13, 53-62.
Szwed, M., Bagdasarian, K., and Ahissar, E. (2003). Encoding of
vibrissal active touch. Neuron, 40, 621-630.
Knusten, PM., Derdikman, D., and Ahissar, E. (2005). Tracking
whisker and head movements in unrestrained behaving
rodents. J Neurophysiol., 93, 294-301.
Szwed, M., Bagdasarian, K., Blumenfeld, B., Barak, O.,
Derdikman, D., and Ahissar, E. (2006). Responses of trigeminal
ganglion neurons to the radial distance of contact during active
vibrissal touch. J Neurophysiol., 95, 791-802.
Golomb, D., Ahissar, E., and Kleinfeld, D. (2006) Coding of
stimulus frequency by latency in thalamic networks through the
interplay of GABAb-mediated feedback and stimulus shape. J
Neurophysiol., 1735 -1750.
Zacksenhouse, M., Ahissar, E. (2006) Temporal decoding
by phase-locked loops: Unique features of circuit-level
implementations and their significance for vibrissal information
processing. Neural Comput., (In press)
Yu, C., Derdikman, D., Haidarliu, S., and Ahissar, E. (2006).
Parallel thalamic pathways for whisking and touch signals in the
rat. PLoS Biol., (In press)
Acknowledgements
Ehud Ahissar holds the Helen Diller Family professorial Chair in
Neurobiology. Our work is supported by ISF, BSF, Minerva,
BMBF and HFSP Grants; Edith C. Blum Foundation and Ms
Esther Smidof Foundation
Fig 2. Proposed multiple motor-sensory-motor loops Whisking signals
(W) are conveyed by the paralemniscal pathway via the POm and
involve whisking control (when). Touch signals (T) are conveyed
by the extralemniscal pathway via VPMvl, and involve processing of
object location (where). Combined whisking-touch signals (WT) are
conveyed by the lemniscal pathway via VPMdm, and involve processing
of object identity (what).

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