Escolar Documentos
Profissional Documentos
Cultura Documentos
REVIEW
Recent advances in integrative studies of locomotion have revealed sev- through Newtons laws, accelerates in the
eral general principles. Energy storage and exchange mechanisms discov- opposite direction. Yet, studies of walking,
ered in walking and running bipeds apply to multilegged locomotion and running, swimming, and flying indicate that
even to flying and swimming. Nonpropulsive lateral forces can be sizable, the spatial and temporal dynamics of force
but they may benefit stability, maneuverability, or other criteria that application are not as simple as they might
become apparent in natural environments. Locomotor control systems first appear. From whale sperm to sperm
combine rapid mechanical preflexes with multimodal sensory feedback whales, locomotion is almost always pro-
and feedforward commands. Muscles have a surprising variety of functions duced by appendages that oscillate or by
in locomotion, serving as motors, brakes, springs, and struts. Integrative bodies that undulate, pulse, or undergo peri-
approaches reveal not only how each component within a locomotor staltic waves. As a consequence, although an
system operates but how they function as a collective whole. animal may appear to move forward at a
steady speed, the forces it exerts on the en-
Locomotion, movement through the environ- Frequently, model organisms are chosen be- vironment are anything but constant.
ment, is the behavior that most dictates the cause they perform some function exception- As a general illustration of the complexi-
morphology and physiology of animals. Evo- ally well. When performance is exaggerated, ties of the temporal and spatial variation of
lutionary pressures for efficient, rapid, adjust- structure-function relations become more ob- locomotor forces, we show examples from a
able, or just plain reliable movement often vious. From studies on specialized animals, diverse assortment of animals in Fig. 1. First,
push the envelope of organism design. Biol- we can extrapolate to other systems in which consider legged locomotion on land, where
ogists have long been attracted to locomotor the properties of interest are not present in the each leg pushes against the ground, produc-
extremes because they provide especially extreme, but in which the principles of func- ing an equal and opposite ground reaction
clear examples from which to determine tion are fundamentally the same. Effective force (Fig. 1A). Two basic mechanisms have
structure-function relations. It is not a coin- comparison demands data on many model sys- been proposed to explain the different patterns
cidence, for example, that David Keilin first tems. Fortunately, the experimental and com- of time-variant forces measured during walk-
discovered cytochromes within insect flight putational tool kit available to study locomotion ing and running (Fig. 1B) (4 6). When ani-
muscle, a tissue that exhibits the highest has expanded explosively in recent years (1), mals walk, the body vaults up and over each
known metabolic rate, or that J. Z. Young making comparisons possible among a large stiff leg in an arc, analogous to an inverted
discovered a giant axon in a squid, an animal variety of animals, both in the lab and in nature. pendulum (Fig. 1B). Kinetic energy in the
capable of rapid escape responses through jet As we review recent advances in the in- first half of the stance phase is transformed
propulsion. Other fundamental discoveries re- tegrative and comparative study of animal into gravitational potential energy, which is
garding central pattern generators, visual pro- locomotion, four themes emerge. First, the partially recovered as the animals body falls
cessing, skeletal remodeling, and many other spatiotemporal dynamics of locomotion are forward and downward in the second half of
important physiological phenomena originated complicated but understandable on the basis the stance phase (4, 7, 8). To travel faster,
from studies of locomotion. Locomotion is not, of a few common principles, including com- legged animals change to running gaits that
however, the simple net outcome of isolated mon mechanisms of energy exchange and the are analogous to bouncing on a pogo stick
specializations in individual cells and tissues. use of force for propulsion, stability, and (Fig. 1B) (4, 7, 9 12). As a leg strikes the
Although it is possible to deconstruct the me- maneuverability. Second, the locomotory per- ground in a running gait, kinetic and gravita-
chanics of locomotion into a simple cascade formance of animals in natural habitats reflects tional potential energy is temporarily stored
brain activates muscles, muscles move skele- trade-offs between different ecologically im- as elastic strain energy in muscles, tendons,
ton, skeleton performs work on external portant aspects of behavior and is affected by and ligaments and then is nearly all recovered
worldsuch a unidirectional framework fails the physical properties of the environment. during the propulsive second half of the
to incorporate essential dynamic properties that Third, the control of locomotion is not a linear stance phase. Remarkably, these basic mech-
emerge from feedback operating between and cascade, but a distributed organization requiring anisms of energy conservation have been
within levels. One key challenge in the study of both feedforward motor patterns and neural and demonstrated in a wide variety of animals
locomotion is to determine how each individual mechanical feedback. Fourth, muscles perform that differ in leg number, posture, body
component within a locomotor system operates, many different functions in locomotion, a view shape, body mass, or skeleton type (1315),
while at the same time discovering how they expanded by the integration of muscle phys- including humans, kangaroos, dogs, lizards,
function collectively as an integrated whole. iology with whole-animal mechanics. These crabs, and cockroaches (4, 16 18).
An integrative approach to locomotion fo- emergent themes are by no means an exhaus- Surprisingly, time-variant forces and en-
cuses on the interactions between the muscu- tive list (2, 3), and they leave room for future ergy exchange in the lateral direction could
lar, skeletal, nervous, respiratory, and circu- reviews. In particular, we note that important be equally important. The legs of sprawled-
latory systems. These systems possess func- advances in energetics, exercise physiology, posture animals such as insects (Fig. 1C)
tional properties that emerge only when they molecular biomechanics, biophysical ecolo- (18 20), crabs (14), crayfish (2123), spi-
interact with each other and the environment. gy, and many aspects of neurobiology are ders (24), and geckos (17) generate substan-
beyond the scope of this review. tial lateral forces while in contact with the
Department of Integrative Biology, University of Cal-
ground. The time course of these lateral forc-
ifornia, Berkeley, CA 94720, USA.
Forcing the Issue es is consistent with the hypothesis that elas-
*To whom correspondence should be addressed. E-
At its most fundamental level, locomotion is tic energy storage and recovery may occur
mail: flymanmd@socrates.berkeley.edu deceptively simple. An organism exerts a within the horizontal plane (25, 26), orthog-
Authors listed in alphabetical order. force on the external environment and, onal to the direction of motion. In addition,