The relatively low degree of complexity of behaviour in
newborn animals makes them a very attractive model for
experimental analysis (Campbell and Marbey, 1972;
Groothuis, 1993; Eilam, 1994 b). Altricial rodents are typically
born furless, have closed eyes and ears, are relatively immobile
and are poorly supported by their limbs. Within 2–3 weeks, fur
covers the body, the eyes and ears open and well-developed
legs support the body. At this time, the pups exhibit marked
hyperactivity to a degree that is rarely matched later in life
(Spear and Barke, 1983). In the present study, we used this
period of extensive changes to determine how gerbils develop
their specialized mode of locomotion, in what order the
movement types emerge and how the synchronous stepping
typical of gerbils evolves in ontogeny.
The morphology of an animal limits the range of movements
that it may execute, thus providing a constraint to certain
locomotor activities (Taylor, 1989). Since the morphological,
physiological and behavioural demands imposed by walking
and running increase markedly as speed, endurance and body
size increase (Garland, 1983; Hildebrand, 1980, 1989), it is
assumed that each animal will utilize its specific morphology
to optimize locomotion (Alexander, 1989, 1992; Dagg, 1973;
Eilam, 1994a). Rodents are an excellent group for assessing
functional morphology, as they display a variety of body
morphologies (Mendelssohn and Yom-Tov, 1987) and modes
of locomotion (Djawdan and Garland, 1988). Since body mass
and body morphology undergo extensive changes during the
postnatal development of rodents, it is intriguing to determine
whether motor performance is also transformed in the course
of ontogeny. Consequently, the present work was aimed at
studying the postnatal development of quadrupedal activity in
Wagner’s gerbil (Gerbillus dasyurus).
Gerbils, like other small cursorial mammals, run using a
specialized mode of saltatory locomotion termed the ‘bound’
(Dagg, 1973; Gambaryan, 1974). In bounding, gerbils leap
with their relatively long hindlegs and land on their relatively
short forelegs, with each pair of legs acting synchronously.
Bounders typically store elastic energy in their trunk, by
arching it dorsoventrally when landing on the forelegs and
unarching it when thrusting with the hindlegs in order to leap
(Biewener, 1989). In accordance with this mode of locomotion,
bounders share a typical body morphology: their hindlegs are
longer than their forelegs and their trunk is arched in order to
position the centre of body mass above the hindlegs, thus
reducing yaw, pitch or roll torque (Dagg, 1973; Gabriel, 1984).
While previous studies of postnatal behaviour in gerbils
concentrated on the emergence of reflexes (Fox, 1964; Kaplan
and Hyland, 1972) or on general biological perspectives
(Valentine and Baudoin, 1980; Cabana et al. 1990), the present
work focuses on describing the development of the bound and
363
The Journal of Experimental Biology 198, 363–372 (1995)
Printed in Great Britain © The Company of Biologists Limited 1995
During the first two postnatal weeks, Wagner’s gerbil
( Gerbillus dasyurus) pups employed stepping sequences
considered to be more basic than in the later stages of
development and displayed alternate stepping of the legs in
each girdle. In the third postnatal week, the adult mode of
locomotion, the bound, became prominent and
synchronous stepping dominated locomotion, gradually
replacing alternate stepping. Motor performance of the
gerbil pups corroborates previous studies in following a
developmental motor gestalt termed ‘warm-up’, which
involves an ordered incorporation of movements along
three distinct spatial dimensions. In the present results, the
validity of warm-up has been extended to a quantitative
perspective: the order in which movement types reached
their peak performance was identical to the order of their
emergence in ontogeny. Transient modes of locomotion
were also employed by gerbil pups during postnatal
development, in accordance with changes in body
morphology, indicating that there exists a causal link
between body design and specific modes of locomotion.
Key words: locomotion, walking, running, bound, stride, warm-up,
rodent, gerbil, Gerbillus dasyurus .
Introduction
Summary
POSTNATAL DEVELOPMENT OF SYNCHRONOUS STEPPING IN THE GERBIL
(GERBILLUS DASYURUS)
HAGIT BLUMBERG-FELDMAN AND DAVID EILAM*
Department of Zoology, George S. Wise Faculty of Life-Sciences, Tel-Aviv University, Ramat-Aviv 69 978, Israel
Accepted 12 August 1994
*Author for correspondence.