Journal ot VeStibUlar Kesean.:u, I, ki..), PI'" ./,.
Copyright © 1997 Elsevier Science Inc.
Printed in the USA. All rights reserved
0957-4271/97 $17.00 + .00
PH S09S7-4271(96)00183-8
Original Contribution
HOW TO CONSTRUCT AND MOVE A CAT'S NECK
W. Graf,* E. Keshner,t F. J. R. Richmond,+ Y. Shinoda,§ K. Statler,11
and Y.
*CNRS-College de France, 75231 Paris Cedex OS, France, tRehabilitation Institute of Chicago, Chicago, Illinois
60611, USA :j::Department of Physiology, Queen's University, Kingston, Ontario K7L 3N6, Canada, §Department
of Physiology, School of Medicine, Tokyo Medical and Dental University, Yushima, Tokyo 113, Japan,
IIDepartment of Physiology and Biophysics, Northwestern University, Chicago, Illinois 60611, USA,
of Physiology, Tokyo Medical College, Shinjuku, Tokyo 160, Japan
Reprint address: Werner M. Graf, CNRS-LPPA, College de France, 11 place Marcelin Berthelot, F-75231, Paris
Cedex 05, France. Tel:+33-1-44-27-16-30; Fax: +33-1-44-27-13-82; E-mail: wg@ccr.jussieu.fr
Abstract - Extensive information has been ac-
ulated over the past several years about the
sensory-motor system, in particular
relating to cats. Using still x-ray and cinera-
analysis, the skeletal geometry of
md.-neCK posture in three dimensions--when an
is resting, actively orienting, or locomot-
described. From these descriptions, cervi:..
vertebral, and craniocervical joint biome-
for all three rotational dimensions are
lantified. These behavioral data on muscle and
movements have been incorporated in a
biomechanical, functional anatomical model of the
Ilead-neck movement system. Individual as well
groups of neck muscles have been measured in
and their kinematics determined. The role
several reflex and voluntary behavioral contexts,
including muscle co-contractions. Having
lished how each movement is accomplished, the
neuronal sensory-motor reflex basis of head-neck
system stabilization in space is addressed. The ves-
tibular system is largely responsible for acquisi-
tion and maintenance of upright posture. The bi-
lateral semicircular canais (horizontai, anterior,
posterior) and otoliths utriculus) feed
information differentially to specific neck mus-
cles: these connections are reviewed with regard
to the origin of the reflex arc from each receptor
to its destination of specific muscles. Behavioral
data from normal animals, and from animals
whose vestibular receptor systems are selectively
lesioned, will be reviewed to complement the func-
tional interpretation of the sensory-motor trans-
formations. Finally, the requirements for space-
time coordinated cat head-neck movements will
be synthesized, based on biomechanics, muscle ki-
nematics, canal/otolith connectivity, and selective
lesion experiments. © 1997 Elsevier Science Inc.
D Keywords - head-neck; stabilization; cat;
sensory-motor; posture.
Introduction
The head-neck movement system provides an
excellent model for studying sensory-motor
transformations, because most of the contribut-
ing sensory and motor systems are well-
researched and described. This review provides
a complete description of three areas necessary
to describe postural and reflex control of the
head-neck movement system: head-neck joint
biomechanics, cervical muscle geometry/kine-
matics, and the vestibular sensory input to neck
muscles subserving vestibulocolIi::.' reJ1exes. SkeJ-
etal geometry during movement 11as been mea-
sured by x-ray and cineradiography (1-4), and
neck muscle kinematics have been determined
for severa] reflex and voluntary behavioral con-
texts (1,2,5-9). These data have been incorpo-
rated in the most thorough head-neck move-
ment model to date.
A detailed description of the vestibular sen-
sory input to neck muscles and behavioral data
219