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Volume 16, Number 22,
Issue of November 15, 1996
pp. 7297-7307
Copyright ©1996 Society for Neuroscience
Organization of Octopus Arm Movements: A Model System for
Studying the Control of Flexible Arms
Received May 30, 1996; revised Aug. 13, 1996; accepted Aug. 26, 1996.
Yoram Gutfreund1,
Tamar Flash2,
Yosef Yarom1,
Graziano Fiorito3,
Idan Segev1, and
Binyamin Hochner1
1 Department of Neurobiology and Center for Neuronal
Computation, Institute of Life Sciences, Hebrew University, Jerusalem
91904, Israel, 2 Department of Applied Mathematics, The
Weizmann Institute of Science, Rehovot 76100, Israel, and
3 Laboratorio di Neurobiologia, Stazione Zoologica ``A.
Dohrn'' di Napoli, Napoli 80121, Italy
Octopus arm movements provide an extreme example of controlled
movements of a flexible arm with virtually unlimited degrees of
freedom. This study aims to identify general principles in the
organization of these movements. Video records of the movements of
Octopus vulgaris performing the task of reaching toward
a target were studied. The octopus extends its arm toward the target by
a wave-like propagation of a bend that travels from the base of the arm
toward the tip. Similar bend propagation is seen in other octopus arm
movements, such as locomotion and searching. The kinematics (position
and velocity) of the midpoint of the bend in three-dimensional space
were extracted using the direct linear transformation algorithm. This
showed that the bend tends to move within a single linear plane in a
simple, slightly curved path connecting the center of the animal's
body with the target location. Approximately 70% of the reaching
movements demonstrated a stereotyped tangential velocity profile. An
invariant profile was observed when movements were normalized for
velocity and distance. Two arms, extended together in the same
behavioral context, demonstrated identical velocity profiles. The
stereotyped features of the movements were also observed in spontaneous
arm extensions (not toward an external target). The simple and
stereotypic appearance of the bend trajectory suggests that the
position of the bend in space and time is the controlled variable. We
propose that this strategy reduces the immense redundancy of the
octopus arm movements and hence simplifies motor control.
Key words:
movement control;
muscular-hydrostat;
kinematics;
octopus;
cephalopods;
motor program;
flexible arm;
arm extension;
reaching movements
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